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TAXONOMICAL REVISIONS

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REPUBLIK INDONESIA
REPUBLIC OF INDONESIA
LEMBAGA ILMU PENGETAHUAN INDONESIA (L.LP.L)
INDONESIAN INSTITUTE OF SCIENCES

FLORA MALESIANA

BEING
AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA /
INCLUDING KEYS FOR DETERMINATION / DIAGNOSTIC DESCRIPTIONS |
REFERENCES TO THE LITERATURE|/|SYNONYMY]/AND DISTRIBUTION /
AND NOTES ON THE ECOLOGY OF
ITS WILD AND COMMONLY CULTIVATED PLANTS

PUBLISHED
UNDER THE AUSPICES OF LEMBAGA BIOLOGI NASIONAL
BOTANIC GARDENS OF INDONESIA / BOGOR / JAVA AND
OF THE RIJKSHERBARIUM / LEYDEN / NETHERLANDS

PREPARED
ON ANINTERNATIONAL CO-OPERATIVE BASIS UNDER THE SUPERVISION OF
SEVERAL DIRECTORS OF BOTANIC GARDENS/ KEEPERS OF HERBARIA
AND VARIOUS PROMINENT BOTANISTS

FOR THE PROMOTION OF
BOTANICAL SCIENCE AND THE CULTURAL ADVANCEMENT OF
THE PEOPLES OF SOUTH-EASTERN ASIA TO
THE SOUTHWEST PACIFIC REGION

SERIES I VOLUME 9

SPERMATOPHYTA

GENERAL EDITOR:

Dr C; G. G. J. VAN SITEENIS
DIRECTOR OF THE FOUNDATION ‘FLORA MALESIANA’

PUBLISHED BY:

MARTINUS NIJHOFF / DR W. JUNK PUBLISHERS
THE HAGUE / BOSTON / LONDON
WONT — iN Oks)

Distributors:
for the United States and Canada

Kluwer Boston, Inc.
190 Old Derby Street
Hingham, MA 02043
USA

for all other countries

Kluwer Academic Publishers Group
Distribution Center

P.O. Box 322

3300 AH Dordrecht

The Netherlands

Library of Congress Cataloging in Publication Data

(Revised for volume 9, part 3)
Main entry under title:

Flora Malesiana.

At head of title: Republic of Indonesia, Ministry of
Agriculture.

Issued in pts.

Includes bibliographical references.

1. Botany--Malay Archipelago--Collected works.
2. Phytogeography--Collected works. 3. Plant
collectors--Malay Archipelago--Collected works.
I. Steenis, Cornelis Gijsbert Gerrit Jan van, ed.

II. Indonesia. Departemen Pertanian. III. Series.

QK366.F56 582 .09598 (2p le

ISBN 90-247-2780-4 (this volume)

Publication dates

Part 1 27 Dec. 1979
Part 2 23 July 1982
Part 3 28 Jan. 1983

Copyright © 1983 by Martinus Nijhoff Publishers, The Hague.

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system,
or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise

without the prior written permission of the publisher,

Martinus Nijhoff Publishers, P.O. Box 566, 2501 CN The Hague, The Netherlands.

PRINTED IN THE NETHERLANDS

CONTENTS
ERROR ene Se bea CNG ia «6 aa «1a oleae ale ante d sr BSE bas ar aie (3)
UICTIINI Eo as ee a a wid ea C28OCE GE eek ss ee (5)
Dedication by R.E.G. Pichi Sermolli & C.G.G.J. van Steenis ...................- (7)
aia RISENTIS ORINIT SIDED ete aa. cee os 2 2's? Jas 3S P a eke ae ere Pee bee ee ee (45)

TAXONOMICAL REVISIONS

in alphabetical sequence

aeaemnimeene —T ENY WY, Be; ENDENENSOOEN eet sin ca ae as gs eta a i cre 1

eyperaceac—IP by: 3-H. Kern .&,H:-P..Nooteboom. 22825 2e> ene eee ss ee ee 107

Pcrocm pacene by PS. AShtonis... 33.9 2n;s ke ee ee oe eee 237

mehecne—F By: 1:P. Jessop). cia. 2 nsec ton oan. Ree eee eee 189
ADDENDA

to volumes 4—9

Addenda, corrigenda et emendanda by C.G.G.J. van Steenis ef al. ............... 553

INDEX

Index to scientific plant names by M.J. van Steenis-Kruseman & E.E. van Nieuwkoop 575

1906

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2 a escron purde ee a ore
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avyeuuts gleth ptgalrve A bile
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Alti~-w :
DV hettar,
fe

Dedicated to the memory of
ODOARDO BECCARI

DEDICATION

A dedication to ODOARDO BECCARY, the greatest botanist ever to study in Malesia, is long overdue.
Although best known as a plant taxonomist, his versatile genius extended far beyond the basic
field of this branch of Botany, his wide interest leading him to investigate the laws of evolution,
the interrelations between plants and animals, the connection between vegetation and environ-
ment, plant distribution, the cultivated and useful plants of Malesia and many other problems of
plant life. But, even if he devoted his studies to plants, in the depth of his mind he was primarily
a naturalist, and in his long, lonely and dangerous explorations in Malesia he was attracted to all
aspects of nature and human life, assembling, besides plants, an incredibly large number of collec-
tions and an invaluable wealth of drawings and observations in zoology, anthropology and ethnol-
ogy. He was indeed a naturalist, and one of the greatest of his time; but never in his mind were
the knowledge and beauty of Nature disjoined, and, as he was a true and complete naturalist, he
was at the same time a poet and an artist.

His Nelle foreste di Borneo, Viaggi e ricerche di un naturalista (1902), excellently translated into
English (in a somewhat abbreviated form) by Prof. E. GicLiotiand revised and edited by F.H.H.
GUILLEMARD as Wanderings in the great forests of Borneo (1904), is a treasure in tropical botany;
it is in fact an unrivalled introduction to tropical plant life and animals, man included. It is a most
readable book touching on all sorts of topics and we advise it to be studied by all young people
whose ambition it is to devote their life to tropical research.

In the last years of his life, BECCARI was rearranging his diaries, notes and observations of the
expeditions to eastern Malesia with the intention of publishing a second book on his explorations,
but very unfortunately death did not allow him to carry out his wish. He left only a revised copy
of his diaries and field notes which formed the bulk of the book Nuova Guinea, Selebes e Moluc-
che, published posthumously by his son NELLO BEccaRiin 1924. It is neither well known nor duly
appreciated outside Italy, since no translation has been published. Undoubtedly it lacks the glam-
our and freshness of the previous book, being devoid of the original and acute observations deriv-
ed from his field research, which are largely profuse in his Nelle foreste di Borneo, but it offers
a good and fascinating description of his adventurous travels with a wealth of interesting remarks,
and it is an invaluable documentation of the natural features of those almost unexplored coun-
tries, particularly of the characteristics and customs of their inhabitants.

This synthesis, however, is only part of his oeuvre and before entering on his achievements, let
us first look at his life and the development of his ideas and ideals.

BECCARI’s early youth was ill fated. He was born in Florence, in his father’s home in the Via
dei Benci at the corner of Borgo dei Greci, on November 16, 1843. His mother, ANTONIETTA MI-
Nuccl, from Radda in Chianti in Tuscany, died soon after his birth, and his father GruSEPPE BEC-
CARI, from an ancient family native of Rimini (Romagna), died in 1849 when ODOARDO was Six
years old; he was brought up by his maternal uncle Minuccio Minuccl. In April 1853 he entered
the Collegio ‘Ferdinando’ in Lucca, where his love for botany was nurtured by the Vice Rector
and Prefect of Studies, the Abbé IGNAzio MezzetTtT1! and by his Professor of Botany in the
Lyceum of Lucca and Director of the Botanic Garden, CESARE Biccui. The latter, aware of the
talent of his pupil and perhaps foreseeing his glorious future, in 1860 dedicated to him a new spe-
cies, Tulipa beccariana’, the first of the numerous plants and animals to be named in his honour.

Beccarr’s first collections date back to 1856, when he was a student of the College of Lucca
and still only 13 years old. During his stay there he assembled a herbarium, which was still in exis-

(1) In his honour, in 1871, BEccARI named a new genus of Annonaceae Mezzettia.
(2) Tulipa beccariana Biccut, Agg. Fl. Lucch. (1860) 21, nom. nud.; 1 Giardini 8 (1861) 50, t. 2.

(7)

FLORA MALESIANA

tence at the beginning of the present century. A search for it in Lucca as a separate herbarium
was unsuccessful, but several specimens with labels headed ‘Erbario Beccari’ are present in the
herbarium of IGNAZIO MEZZETTI, now kept in the Lyceum Machiavelli of Lucca, where BECCARI
attended secondary school until July 1861. Whether these specimens are part of the separate BEc-
CARI herbarium included in MEzzeTTI’s herbarium or duplicates of it, is uncertain. Other plants
collected in the period in which BEccarI was a student in Lucca are kept in WeBp’s herbarium
in Florence.

In August 1861 he published his first paper and in the autumn of the same year, when he was
18 years old, BECCARIcommenced his studies in the Faculty of Natural Science at the University
of Pisa. At first, perhaps under the influence of Biccut, he devoted his attention to the Cryp-
togams and already in 1862 BECCARIs name appears together with those of several eminent botan-
ists of the time, among the collectors of the ‘Erbario Crittogamico Italiano’, the classical exsiccata
with printed labels, founded in 1858 by GIUSEPPE DE Notaris of Genoa.

In the University of Pisa, BECCARI distinguished himself so much in botany, that the celebrated
botanist PlETRO SAvi made him an assistant to the chair of botany in January 1863, while he was
still an undergraduate. Dissatisfied with the conservatism of SAvi, however, he gave up his assis-
tantship and moved to the University of Bologna from where he took his degree in Natural Science
on July 1, 1864, by disputing his thesis on the structure of the lichen Arnoldia cyathodes Mass.
[ = Plectopsora cyathodes (Mass.) KORBER] with the famous professor in botany ANTONIO BERTO-
LONI.

Before his graduation, BEccaARthad already planned a long journey to far away regions, when
in June 1864, in the laboratory of Prof. GIOVANNI CAPELLINI, geologist at the University of Bo-
logna, he met Marquis Giacomo Doria, a young, impassioned naturalist, later patron and Maece-
nas of science, and founder of the Museo Civico di Storia Naturale at Genoa which bears his
name. The two young men, united by the same enthusiasm for scientific exploration in unknown
countries, soon fraternized. Shortly after his graduation, BECCARI visited Doria at Genoa, where
they decided to undertake a long exploration together and, counselled also by the celebrated
British naturalist JOHN BALL, they chose the Kingdom of Sarawak, in Borneo, as the destination
of their enterprise. As part of his preparations, BECCARI spent the period from February to April
1865 in the great British centres of botany, the British Museum in London and Kew Gardens, ob-
viously to acquaint himself roughly with the plants of Borneo. He met the HooKERs, CHARLES
Darwin and Sir JAMES BROOKE, the Rajah of Sarawak, who assured him of the assistance of his
nephew, the Tuan-muda, Sir CHARLES BROOKE, then governing the territory in his absence.

Thus prepared, young OpoarRpDo, at the age of 22, commenced his studies on the flora of the
Malesian tropics, which was to become his main life occupation, and in which he would rise to
the greatest heights as a scientific explorer, naturalist, and botanist. He sailed from Southampton
on April 4, 1865, and met Doria and his own brother, GIOVANNI BATTISTA BECCARI (who was on
his way to Japan), at Alexandria. From there they travelled by train to Suez and by boat to Aden
and then to Ceylon, where they spent a fortnight. There BECcARI visited the famous Botanic Gar-
dens at Peradeniya and climbed Mt Petrotallagalla, where he made his first personal acquaintance
with the tropical flora and started collecting. Via Penang island and Singapore, the voyagers arriv-
ed on June 19, 1865 at Kuching, capital of Sarawak, which they had selected as their base of opera-
tions. At the beginning they were guests of the Tuan-muda, Sir CHARLES BROOKE; later they settled
in a house of their own with servants, and also bought a small boat (‘sampan’) for their excursions
in the forests along the river. Soon BEccARIand Doria took up their botanical and zoological col-
lecting in the dense and primitive forests which at that time surrounded Kuching. BECCARI was
anxious to know the mountains and to collect intensively; thus he undertook the construction of a

(8)

Dedication

big hut in the forest of Gunong Mattang at an altitude of about 300 m with the intention of making
it the base for the explorations and collections in the primitive forests of Borneo. However, after
some months the health of Doria deteriorated to such a point that at the beginning of March 1866
he was forced to return to Italy. Thus, BEccARI, having accompanied his friend to Singapore, re-
mained alone to carry out the programme which he had planned with him and had hoped to realize
together. At the beginning of April he moved to his house in the forest of Gunong Mattang which
he called ‘Vallombrosa’, after the great monastery hidden in the dense mountain forest of Prato-
magno, east of Florence. Together with a Chinese cook and four Malesian boys he spent nearly
all the remaining months of 1866 there, except for four excursions and a period in October-No-
vember at Kuching to pack his collections. On one of these excursions to Gunong Poe he discov-
ered a new species of Rafflesia, the famous parasitic plant with vegetative parts extremely reduced
and a gigantic flower, the largest in the plant kingdom, apparently arising directly from the stem
of the host liana. The largest flower of the new species, Rafflesia tuan-mudae, so named in honour
of the Tuan-muda of Sarawak, Sir CHARLES BROOKE, attained about 56 cm in diameter.

In Nelle foreste di Borneo BECCARI gives many details of his hut at Mattang. He had cleverly
designed it for drying plants and preparing zoological specimens rather than for lodging; soon it
became an active and efficient laboratory, full of all sorts of products of nature. In his book he
described his primitive life there as very happy and fully suited to his temperament. With only a
cotton coat, trousers and a Chinese straw hat, mostly bare-footed, he carefully explored the sur-
rounding primary forest, assembling marvellous collections of plants and animals. Back at his
hut, he devoted many hours to arranging his collection, making drawings and descriptions and
recording those notes and observations which later became the basis of his fascinating book Nelle
foreste di Borneo.

At the beginning of 1867 BEccaRt abandoned the hut at Mattang and spent the first two months
at Kuching collecting in the surroundings, but chiefly arranging and packing his large collections.
In March 1867 he again undertook his adventurous wanderings with the intention of visiting the
interior of Sarawak. One of his trips from mid-March to the last days of May was devoted to the
exploration of Batang-Lupar and the lakes of Kapuas with the main purpose of hunting orang-
utan. He assembled there one of the best collections of these animals (skin, skeletons, heads and
skulls, and even a foetus) and a wealth of observations which allowed him to express the opinion
that the hominids did not originate in dense forest, like that of Borneo, and that the orang-utan,
particularly well adapted to an arboreous environment, would be, not an ancestor, but a collateral
of man. In his opinion, the hominids were derived from forerunners, allied to the great anthro-
poids of tropical Africa, with an anatomical conformation, particularly of the limbs, more suited
to evolve towards a biped gait and an erect habit and they had their origin in more open vegetation,
like that of some regions of tropical Africa, where we find the greatest number of large mammals
with rapid locomotion. Recent research in south-western Ethiopia seems to support this hypo-
thesis.

From August 12 to September 14, 1867, BEccarr collected in the district of Bintulu and in the
country of the Kayan. From there, he was looking forward to organizing an expedition to the inte-
rior regions of Sarawak, which at that time were still nearly unexplored and hardly visited by
Europeans; but his project found every possible difficulty and obstacle. Despite them, without
guide or interpreter, but with only four men and a small boat, he set out on September 15 from
Bintulu on his journey through the interior of Sarawak along the basins of the main rivers Bintulu,
Redjang and Batang Lupar and their tributaries, across the ridges of hills and mountains which
represent the watershed between them. The journey was made mostly sailing up, or down, the
rivers in various native paddle boats obtained from time to time from the natives, but also on foot

(9)

FLORA MALESIANA

to overcome some impassable rapids or to cross the ridge between two adjacent basins; often he
was forced to walk with difficulty in the stream beds, or to proceed slowly with a compass through
the dense forests; more than once he was in real danger, even near the end of his travels when,
having lost his compass, without food, in an unhabited region, he got lost for two days in a dense
forest. On November 20, 1867, BEccaARI arrived at Kuching where he concluded this long, hard
and risky enterprise.

In the first two years of his stay in Borneo BEccarr’s health remained excellent, but in the last
months it had been deteriorating. Already in June 1867 he had suffered the first attack of malaria
and later many others followed. Furthermore, in July of the same year he had observed the first
symptoms of elephantiasis on his right ankle. After the expedition to the interior of Sarawak he
spent two months in Kuching arranging and packing the collections he had made. He had planned
another long journey crossing the inland of Borneo from Kuching to Pontianak; but in January
his health worsened, and being unable to subdue the high fever which had troubled him for some
days, Beccariwas forced to undertake his homeward-bound voyage. He left Kuching on January
29, 1868 and arrived in Italy on March 2, after explorations in Borneo which had lasted almost
three years.

In Florence BEccARIwas the guest of his old friend from the College of Lucca, EmiLio MARCUC-
ci, who had taken up the profession of architect but had not given up his love of botany, and who
greatly assisted BECCaRI in that period in recovering his health. The house was located in Borgo
Tegolaio 48, very close to the Museum of Physics and Natural History, where BEccarti had prob-
ably assembled his collections. The house soon became a meeting place for young lovers of natural
history including LEvreR and Sommrer. Soon after his return from Borneo G. Doria and R.
GEsTRO, from the Civic Museum of Natural History of Genoa, were also his guests for several
days, evidently to be informed, in detail, about the large zoological collections he had assembled
in Borneo.

At that time BECCARI was very busy sorting out and working on his collections; he also made
agreements with collaborating specialists to study particular groups such as seagrasses (ASCHER-
SON 1871), pteridophytes (CESATI 1876), mosses (HAMPE 1872), lichens (VON KREMPELHUBER 1875)
and hepatics (DE NoTaris 1876), efc. (see Appendix 4); he probably also started distributing dupli-
cate specimens of his Bornean plants. However, together with his technical work, he carried on
with the study of his collection.

In March 1869, Beccari started, at his own expense, the publication of a new periodical, the
Nuovo Giornale Botanico Italiano, which was intended as a replacement of the Giornale Botanico
Italiano founded in 1844 by F. PARLATORE, but interrupted in 1852. BEccaARI edited three volumes
of the new periodical (1869, 1870, 1871) with the help of his friend Marcuccl, to whom he dedi-
cated the new genus Marcuccia (Annonaceae) as a sign of gratitude for the help received in editing
these volumes, particularly during his travels in Ethiopia in 1870. His first accounts of Bornean
plants appeared in early volumes of his journal and many other papers dealing with his collections
written by himself and other botanists were published in subsequent volumes. However, in spring
1871, when preparing for his expedition to New Guinea, BECCARI became aware of the difficulties
of editing a journal when abroad making long expeditions in distant regions and handed the man-
agement of the Nuovo Giornale Botanico Italiano over to T. CARUEL, who edited it until the end
of 1893, when the journal became the official publication of the Italian Botanical Society, which
it continues to be.

Fascinated by his primitive life in Borneo, BECCARI was not satisfied with city life. Probably
he had already developed the idea of undertaking a second journey to Malesia, when he received
an offer to join an Italian expedition to Ethiopia. He sailed on February 14, 1870, from Genoa and

(10)

Dedication

together with the zoologist Marquis ORAZIO ANTINORI and Prof. Arturo IssEL, geologist at the
University of Genoa, visited the Bay of Assab and later, on behalf of the Italian Geographical
Society, the country of Bogos. There he assembled a rich collection of plants (315 species of sper-
matophytes and pteridophytes and 289 species of mosses, algae, fungi and lichens) enumerated
and partly described in MARTELLI’ Florula Bogosensis (1886). He came back to Italy on October
20, 1870.

Soon after his return to Florence, BECCARI materialized his project to visit Malesia again and
after careful preparations, training himself in geodetics, astronomy and meteorology, he set out
on November 24, 1871 from Genoa for the island of New Guinea, accompanied by Count Luici
MarRIA D’ALBERTIS, an Italian nobleman who was passionately fond of hunting and natural
history. Their first visit was to West Java where they stayed for some time in the Botanic Gardens
at Bogor. The young director, Dr. RUDOLPH SCHEFFER, must have facilitated his exploring for a
few days on Mt Gedeh, with its Tjibodas mountain garden, and adjacent primary forest on Mts
Pangerango and Megamendong. Further stops were made at Flores and Timor in the Lesser Sunda
Islands, and the islands of Banda and Ambon, where they arrived on March 7, 1872, and enjoyed
the kind and helpful hospitality of Captain P.F. Kraatand his wife, the Italian lady AMALIA Ma-
LAN. After a short journey to Buru and Ceram for information they came back to Ambon, where
they organized the expedition to western New Guinea renting a small schooner, the ‘Bu-
rung-Laut’, of 25 tons with a crew of eight men.

On March 21, 1872, BEccARI and D’ALBERTIS sailed from Ambon to New Guinea, and having
touched the islands of Geser and Goram reached Kapaor on the west coast of the mainland. Later,
on April 30, they arrived at the small island of Sorong where they dismissed the Burung-Laut and
rented a hut in which they established their first scientific station, collecting chiefly along the
Ramoi river. They remained on Sorong island till July 15 when, having left their collections there
with one of their men as keeper, they sailed in a ramshackle indigenous sailing boat with a crew
of eleven Papuas to Dorei and later to Andai, where they arrived on August 7 after a long voyage,
full of adventures.

BeccaRI and D’ALBERTIS established their home and headquarters for the exploration of the
region in a large Papuan hut built on high palafittes near the Andai river surrounded by the forest,
with a splendid view of both the sea and the Arfak Mts. While D’ALBERTIS made a trip in the
mountains, Beccarti collected intensively in the vicinity of Andai, but on September 28 he moved
to Putat on the lower slopes of the Arfak Mts with the intention of exploring the higher regions.
Unfortunately on October 9, he was informed that D’ALBERTISs had fallen seriously ill and he was
compelled to return to Andai. The poor health of D’ ALBERTIS necessitated that the travellers return
to Ambon, but only on November 2 was BEccariable to find an indigenous boat to reach Sorong
and later Ambon. But in Sorong, unexpectedly, they found a schooner sent from Ambon in search
of them. After recovering, safe and sound, the collections left there four months before, they sail-
ed to Ambon, where they arrived on December 5, 1872. There BECCARI and D’ALBERTIS were
greatly surprised to find the Italian Royal Corvette ‘Vettor Pisani’ on which D’ ALBERTIS obtained
a passage, leaving his friend alone. Thus, BEccari concluded his first expedition to New Guinea,
during which, despite all sorts of difficulties and serious health troubles, he had assembled a col-
lection of about 700 species of plants and a rich amount of zoologica!, ethnographical and mine-
ralogical specimens.

BECCARI remained in Ambon for about two months to arrange and pack the collections. There,
as a guest of Captain KraaLand his wife, he soon regained his health and prepared a trip to the
Aru and Kei islands.

Beccari departed from Ambon on February 8, 1873, having obtained a passage on a Dutch

(11)

FLORA MALESIANA

Government steamer. Stricken with smallpox en route, BECcARI nonetheless reached the Aru is-
lands on February 22, and with his base on Wokam, he collected plants and animals and made
a topographical survey of the islands (see Appendix 1, C: Maps). On July 6, he moved to the Kei
islands in a big local sailing boat, a Bughis prahu, on which Beccari was the guest of its Chinese
master. But the boat suffered shipwreck on the east coast of Grand Kei. Fortunately he could save
all his collections and collecting equipment. BEccARtI found the flora of these islands unexpectedly
poor and after visiting Small Kei as well, he sailed on October 4 to Ambon with four men in a
small indigenous sailing boat of only 4 tons, bought at Dulan. Despite the premonitions of the
natives, the risky voyage of about 350 miles was successful and on October 23 he reached Ambon,
where he stayed for two weeks, partly to arrange his collections, but chiefly to recover his strength,
being again a guest of his friends, the KRAALs.

On November 5, 1873, BEccaRIsailed by the steamer ‘Koning Willem III’ towards the West Mo-
luccas (Buru and Ternate), proceeding via North Celebes towards Southwest Celebes, where he
disembarked at Makassar on November 18. He stayed in the region for nearly three months until
February 6, 1874. From there, as a paying passenger on an old Chinese boat of about 40 tons,
similar to a prahu, he went to the larger islands south and southeast of Celebes (Kabaena and
Muna) and to Kendari on the southeast coast of Celebes where he arrived on February 23. He re-
mained in this district for six months to collect and make topographical surveys, but the collec-
tions did not increase very much because the flora was not particularly interesting and because
the region was plagued with pirates on the sea, and head-hunters on land. He chiefly collected in-
land at Lepo-Lepo. Here he was informed that a Dutch vessel was looking for him at Kendari.
It was the Escort vessel ‘Sumatra’ of the Royal Dutch Navy, which had been sent from Makassar
in search of him, since it was rumoured that he was in danger from the pirates which infested the
sea of Kendari. BEccarthad already decided to leave Kendari and accepted with pleasure the kind
offer of the Captain of the vessel to take him and his men aboard to Makassar. Having packed
his collections he sailed from Kendari on August 10 and arrived at Makassar after a voyage of
five days.

In these last months Beccarr’s funds had been running out, but early in 1874 he had already
written from Makassar to his friend G. Dortain Genoa for financial help to carry out his project
of a second expedition to New Guinea. When he was back at Makassar on August 29 he received
the joyfull news that his friend had convinced the authorities of Genoa to contribute 15,000 lira
towards a new, second expedition to the great island.

Aware that the season was not suitable for sailing to New Guinea, BEccartsoon left Makassar
by the same steamer ‘Koning Willem III’ on which he had travelled from Ambon to Makassar
some months before. He proceeded to Bali, Surabaja, Semarang, and through the interior of Java
to Bogor, to recuperate and to sort out his collections. There, he also spent some days at Tjibodas
and on Mt Pangerango collecting. Unwearied, BEccarileft Jakarta on October 15, 1874, and via
Surabaja, Makassar, the island of Bima and Timor in the Lesser Sunda Islands he arrived at Ter-
nate island in the Moluccas on November 11. He remained there about 20 days and assembled
rich botanical and zoological collections in the primitive forest near the hut (named by him ‘Para-
disino’) which the Dutch Resident had built for him on the slopes of the volcano.

BEcCARI intended to organize his travel to New Guinea from Ternate, but soon he realized that
this was impossible, and on December 4 he left the island by the mail-steamer arriving at Ambon
three days later. There he prepared for his new expedition financed by the Province and the
Municipality of Genoa to West New Guinea, his old hunting grounds of 1872. He hired for his
voyage the brig-schooner ‘Deli’ with a crew of 10, and accompanied by 8 men and a young boy
for collecting plants and animals.

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Dedication

He sailed on January 22, 1875, and arrived at Sorong Island on February 1, establishing his base
in the schooner and making trips to Ramoi, Dorei Hum, Mt Morait and venturing inland from
Has as far as a river, the War Samson, not then recorded on the maps. Together with plants he
amassed a rich collection of birds. On March 5 he left Sorong and after a visit of some days to
Waigeu Island chiefly hunting for birds, he proceeded to Dorei and soon to Warbusi and Momi
on the west coast of Geelvink Bay mainly with the aim of obtaining some specimens of cassowa-
ries. Later, in April, always in the ‘Deli’, he went to the islands of the Bay which he had not pre-
viously visited, spending nearly twenty days on Japen, a week on the uninhabited Mios Num,
twenty days on Schouten Island and a week on a small island, Pulo Manim, near Mafor. On June
2 he arrived at Dorei where he found the Italian Corvette ‘Vettor Pisani’, and where he received
the warmest welcome. Having arranged his collections he prepared the expedition to the Arfak
Mts; on June 16 he started from Andai towards the mountains establishing his exploration base
at Hatam (1500 m) in the centre of the mountain group. On June 23 he climbed one of the summits
of the Arfak Mts (about 2000 m). He had planned to remain two months in the mountains, but
on July 12 he was forced to cut short his exploration and to come down to the coast because of
beri-beri among the crew of the schooner: two men had already died and the others were seriously
ill. Thus BEccaRI realized that there was no choice; he had to go back to Ternate: his second ex-
pedition to New Guinea was nearing its end; his dream of climbing and exploring the highest re-
gions of the Arfak Mts had vanished for a second time. On July 18 he left Mansinam, near Andai,
where the ‘Deli’ was riding at anchor, and on August 4, 1875, he arrived at Ternate, but in the
meantime the beri-beri had killed most of the crew. He remained at Ternate three months ar-
ranging his collections, notes and observations.

The scientific results of the second expedition to New Guinea were very important. Even if the
botanical specimens were not particularly numerous, the zoological collections were very plenti-
ful, especially the skins of birds which surpassed 2000 in number, and included a set of birds of
paradise which still remains one of the best of its kind. No less abundant were the ethnological
collections consisting of every sort of object in use by natives. Also an important set of approxi-
mately 200 Papuan skulls enriched the anthropological collections. But the expedition was also
very fruitful because of his untiring activity as a naturalist and explorer in making notes of every-
thing that attracted his attention and in studying all the aspects of the regions he visited. During
the expedition he had also made various topographic surveys which later allowed the geographer
Gurpo Cora (see Appendix 1, C: Maps) to draw maps of some regions of New Guinea, and he
had assembled a great wealth of botanical, zoological, ethnological and anthropological observa-
tions which are profusely reported in his letters published by E.H. GiGLioti, and in his book
Nuova Guinea, Selebes e Molucche. Full of interest are the observations on the characteristics and
origin of the Papuans, and on the life of birds, particularly those on the bower-birds of paradise,
Amblyornis inornata, and its ‘capanne e giardini’, which are carefully and at the same time poeti-
cally described by him in a paper full of interesting scientific and philosophical considerations,
pervaded with a deep-rooted love and admiration of Nature.

Learning that a Dutch expedition to New Guinea was being prepared, with the vessel ‘Soeraba-
ja’, with the aim of performing a bathymetric survey, BECCARIreceived permission to accompany
this. It lasted from November 11, 1875 till January 29, 1876, visiting Dorei, the Bay of Wanda-
men, the islands of Roon and Krudu, the Bay of Humboldt, the island Arimosa, Awek (Japen
I.), Dorei, Waigeu, Misool, the Bay of MacCluer, the Bay of Gouns, the island of Geser (off
Southeast Ceram), and Ambon. From there he returned on the mail steamer to Ternate. He stayed
there about a month to arrange and pack his latest collections and to ship them to Italy. This third
expedition to New Guinea had not yielded results as far as BEccaARr’s botanical and zoological in-

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FLORA MALESIANA

terests were concerned, but it had allowed him to assemble many ethnological and anthropological
notes, and to improve his topographic surveys.

On March 12, 1876, Beccart sailed from Ternate to Java, on the first stage of his homeward
voyage. He arrived, unexpectedly, at Florence on June 18, 1876, after about four years of bold
and glorious exploration.

On his return BECcARI was received with great honour. On July 14, 1876, the Municipality of
Florence bestowed the freedom of the city on him; some scientific societies, such as the Zoological
Society in London, and the Italian Anthropological Society, elected him an honorary member.
Other scientific associations, such as the Italian Geographical Society and the Tuscan Society of
Horticulture as well as the Faculty of Science of the Royal Institute of Advanced Studies of Flor-
ence awarded him a gold medal. But he was not affected by these honours and devoted his time
to his collections and to his friends in Florence and Genoa. However, the glamour of exploration
and the call of the wild were too strongly in his nature and after a year BECCARI made one further
long voyage to the Malesian islands.

He and Captain Count Enrico A. D’ALBERTIS, a cousin of his former companion, set out from
Genoa on October 14, 1877, on a trip, properly intended more for pleasure than for science, to
Australia, en route travelling through India from Bombay, Lahore, Delhi, Benares, Lucknow, to
Calcutta, touching Singapore and Kuching (December 1877), meeting in Australia FERDINAND
VON MUELLER, and proceeding to Tasmania and New Zealand.

On the return voyage he parted from D’ALBERTIS in Singapore and proceeded to Jakarta and
then to Bogor where he spent two weeks, preparing a collecting trip in Central West Sumatra.

Sailing from Jakarta on 28 May 1878, he arrived in early June via Padang and Padang Pandjang
at Mt Singalang, a primary-forest-clad, long-extinct volcano of nearly 2900 m height. Here he had
a hut built, as before in Sarawak and Ternate, which he made his headquarters. The hut, named
by him ‘Bellavista’, was placed above the limit of cultivation and on the lower fringe of the primi-
tive forest, at an altitude of about 1700 m. He remained there from June 12 to early August, mak-
ing rich collections on the flanks and on the top of the volcano. Later he set his base in a house
in the village of Ajer Mantcior at the base of Mt Singalang till September 20. After a short stay
in Padang to arrange his collections, he undertook a journey on October 4 in the provinces and
on October 22 he sailed from Padang to Bangkok where he arrived on November 10. During his
travels in West Sumatra (see map of his itineraries in BECCARI 1930) he assembled large botanical
and zoological collections; the largest were made on Mt Singalang, the harvest of plants running
to a thousand numbers in all. Amongst them were the famous Rafflesia arnoldii and the then un-
known, largest, erect aroid in the world, Amorphophallus titanum, a really colossal herbaceous
plant, the tuber being up to 53 cm in diameter, the inflorescence more than 1.5 m high, the lamina
of leaf covering a surface of about 15 m in circumference and the petiole attaining about 29 cm
diameter at the base. From Bangkok he began his homeward journey to Italy, arriving in Florence
on December 28, 1878, thus concluding the last of his fascinating explorations in Malesia.

Reviewing the results of his botanical activities in the six years exploration in the field through
almost the whole of Malesia, it is evident that BEccarr’s exploration in Sarawak was the most
fruitful and thorough, with the huge number of over 4000 collections in two and a half years. The
great virtue and value of his collections can only be properly estimated if one takes into considera-
tion that Beccari collected species rather than specimens, and that he seldom collected a species
twice. Each species was studied, dissected and annotated on the spot and mostly carried flowers
and fruit. The Sumatran collection again was rather large, about 1000 numbers in five months
travel, especially when one considers that BECcCaARr’s interests were wide; in Sumatra he also dedi-
cated time to the study of agriculture, forest products and fruit trees, as he had done in Borneo.

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Dedication

The amount of these collections clearly contrasts with less than 1000 numbers in the three years
spent in the Moluccas, Celebes and New Guinea (see Appendix 4), although really there are more,
as most collections from the Kei and Aru islands and Kendari in Southeast Celebes are unnum-
bered and unlisted. As the flora of at least New Guinea is not less rich in proportion to that of
Borneo or Sumatra, the reasons for this contrast can only be explained by Beccarr’s activity in
the field. Firstly, we must consider that he was a thorough collector and disliked gathering occa-
sional or incomplete specimens; he preferred always to stop some days to collect systematically
in a place which he considered botanically interesting, rather than to gather here and there along
his path, en route, when moving from one place to another in his long expeditions. In a word,
he preferred to collect intensively rather than extensively. In Borneo and in Sumatra where he had
huts for drying, labelling and drawing his specimens (‘Vallombrosa’ and ‘Bellavista’ respectively)
his collections were more numerous than in New Guinea, Celebes and Moluccas when he frequent-
ly moved his collecting base, sometimes being forced to do so because threatened by native
head-hunters, or by pirates. Finally, we must also bear in mind that the second expedition to New
Guinea was made thanks to the financial support of the Province and Municipality of Genoa, se-
cured on the warmest recommendation of Giacomo Doria, his friend and zoologist of the expedi-
tion to Borneo. BEccaARI knew that he longed to enlarge the zoological collections of the Civic
Museum of Natural History which he had founded in 1867, and, in his profound honesty, he felt
bound to assemble large zoological collections for the Genoa Museum and to put zoology before
botany. Indeed, the zoological collections were very rich in quality and in quantity, while the bo-
tanical collections were not particularly numerous. Another reason for this contrast can also be
found in Beccarr’s health which was worse in eastern Malesia than in Borneo and Sumatra.

In 1878 Beccari was still only 35 years old, but had accumulated an unrivalled, immense
amount of material, great scientific-botanical experience and knowledge of the Malesian flora,
in fact had proved himself the greatest explorer of his time. He would prove himself also to be
the greatest botanist in the elaboration of his results, surpassing BLUME in the width of the field
he covered, including plant-geography, ecology and biology.

Having concluded his explorations in Malesia, BEccart devoted the rest of his life to the study
of his collections and of palms, except for an unhappy experience as Director of the Botanical Col-
lections and Garden of the Royal Museum of Physics and Natural History of Florence in 1878—
1879, and a short journey to Ethiopia soon after.

The vicissitudes of BEccaRr’s life as Director of the Botanical Collections and Garden cannot
be understood without knowledge of some of the events in the history of the Florence Museum
and the sale of BEccarr’s collections. The Royal Imperial Museum of Physics and Natural History
in Florence was founded in 1775 by PreTRo LEOPOLDO Di LorENA, Grand Duke of Tuscany, and
was installed in a building in Via Romana, not far from the Palazzo Pitti, the palace of the Grand
Duke, at the base of the great and famous Giardino di Boboli. Part of this was soon designated
as the Botanical Garden of the Museum. Thanks to the great interest of the Grand Duke in Natural
Science, the scientific collections were greatly increased in the years thereafter and the Museum
was subdivided into various sections (Cabinets) and in 1789 an astronomical observatory was also
installed there. From then on all the Museum complex was usually named ‘La Specola’ by the Flo-
rentines. The botanical section consisted of the so-called Botanical Collections (herbaria, carpo-
logical collections, vegetable products, wax models and fossil plants) and the Botanical Garden.
After various events dominated by the historical course of Tuscany in the first half of the 19th
century, FiLippo PARLATORE was appointed in 1842 director of the Collections and Garden. He
greatly contributed to the growth of the herbaria and to the organization of the department. In
1854 Pump BARKER WEBB died in Paris and bequeathed his invaluable herbarium and library to

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FLORA MALESIANA

the Florence Museum, together with an annual income of 6945.58 lira (derived from the sale of
a palace in Paris) and known as Wesp’s Legacy, for their maintenance and increase. In 1859 the
Government of Tuscany established the Istituto di Studi Superiori Pratici e di Perfezionamento
in Firenze, which had its centre near the Monastery of San Marco; and the Museum of Physics
and Natural History with its collections and the botanical garden, although situated on the op-
posite side of the river Arno, became part of that Institute as the seat of the Faculty of Science.
However, the Museum continued to have a director of its own, and in 1868 PARLATORE was ap-
pointed to that office. Very unfortunately, the Institute of Advanced Studies, which only became
the University of Florence in 1923, had no Rector responsible for the scientific and didactic activi-
ty, but only an Administrative Board which determined the course of events in the Museum in
the following years without an adequate knowledge of the problems and needs of scientific re-
search. In 1860, after various vicissitudes, the Giardino dei Semplici, founded in 1545 and one
of the most ancient in the world, became state property, and, in 1869, together with the adjoining
buildings (originally the stables of the Grand Duke) was assigned to the Institute of the Advanced
Studies being situated near the centre of the Institute at San Marco. In 1872 the Italian Govern-
ment, the Province and the Commune of Florence signed a convention for the enlargement of the
Institute of Advanced Studies, and the Board of the Institute decided to move some of the Cabi-
nets of the Museum to the centre of the Institute. The latter also foresaw the removal of the Botan-
ical Collections and Garden of the Museum from La Specola to the Giardino dei Semplici and
pertinent buildings at San Marco, in order to have the botanical department nearer to the Institute,
and to maintain only one garden, the Giardino dei Semplici, by far more famous than that of the
Museum. Thus, during 1877 and 1880 the Cabinets of Chemistry, Physics, Geology and Mineralo-
gy were shifted from the Museum to San Marco. Only in 1879 did the Commune of Florence ac-
tually hand over the Giardino dei Semplici to the Institute of Advanced Studies. The latter decided
that the Botanical Collections and Garden ought soon to be moved to San Marco. However, they
were conveyed there only several years later owing to the opposition of several botanists, among
whom BEccarI, and other personalities, who were against the removal of the botanical collec-
tions, as they considered it, for several reasons, to be a great error.

Fitippo PARLATORE died on September 9, 1877. He was the last Director of the Museum of Phys-
ics and Natural History in Florence as the Institute had decided that the Dean of the Faculty was
to hold the directorship of the Museum. However, the office of Director of the Botanical Collec-
tions and Garden was vacant, and according to general opinion BECcaRI was the best qualified
and most worthy successor to the work of PARLATORE, who had so greatly enlarged the herbarium
and library and raised them to the level of the greatest museums in the world. But BEccaRr’s ap-
pointment was strongly opposed by the Dean of the Science Faculty and the Board of the Institute
of Advanced Studies, particularly because BECCARI was firmly convinced that the Director of the
Botanical Collections and Garden should have no hand in teaching. On October 14, 1877, hardly
more than a month after PARLATORE’s death, BECCARI undertook his travels with E. D’ALBERTIS,
sketched above, and at that time no resolution had been taken. Only on March 26, 1878, while
BECCARI was journeying in Australia, was he, in spite of the opposition, appointed Director of
the Botanical Collections and Garden of the Royal Museum of Physics and Natural History of
Florence, with the duty of supervising the practical phytographic research of the students. BECcA-
Ri returned to Florence from the exploration of Sumatra on December 28, 1878, and soon took
up the office of Director.

In the preceding years BEccaARI had organized, worked and studied on his own in the field of
botany, and his brilliant achievements were naturally a one-man show. In the field he had to make
his own decisions, and learned to do so immediately. He had no rivals and had always very subor-

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Dedication

dinate personnel whom he could command. That was very different from the situation in which
he was now placed as a Director, with a graded staff accustomed to some privileges acquired dur-
ing the long directorship of PARLATORE. On the other hand Beccari was indefatigable and tidy
in his work and he required everybody to be active and precise in carrying out his duty. He wanted
to infuse new life into the operations of the botanical collections and garden, introducing methods
that differed from the traditional ones. Very soon his reforms became unpopular among several
of the staff, who felt that he lacked respect for the memory of his predecessor; this state of affairs
acerbated the hostility towards him of the Institute of Advanced Studies. But such hostility was
not something to scare him.

It is clear, of course, that BECCARI, with his enormous drive and ambition, proved by his unique
exploration and study of the tropical floras, thinking big, botanically and otherwise, wanted to
raise the Florentine centre into an institute which could compete with the leading world herbaria.
For this there was excellent opportunity as the Florence botanical collections had already acquired
the enormous and (still) most important herbarium of PHiLIp BARKER WEBB, bristling with types
of old collections, with funds attached for its maintenance, and further the important herbaria
of CESALPINO, MICHELI, TARGIONI, and PARLATORE. To these could now be added his own numer-
ous collections from the East, with its enormous mass of duplicates for the further enrichment
of the Florence herbarium by exchange. There was, therefore, every reason and opportunity to
fulfill his ambition.

When Beccaritook on the directorship, he was aware that the botanical collections were under
threat of removal from the Museum to the buildings adjoining the Giardino dei Semplici near the
centre of the Institute of Advanced Studies. At first he was rather in favour of the project since
he thought that the Institute had large funds for the construction of new buildings and that they
would be better and more suitable for the collections than those of the Museum. But when he dis-
covered that the funds of the Institute were scarce, and the buildings were the old stables of the
Grand Duke of Tuscany, very humid, unsuitable for both the herbarium and library, and far
worse than those of the Museum, he became a most obstinate and relentless opponent to the
removal of the collections. His hostility was the primary cause of a wide gulf between him and
the Institute Board. On the other hand Beccari began to understand that his ambition of raising
the Florence Botanical Collections to the level of the other great European herbaria would be dif-
ficult to realize. But this controversy was not the reason for his resignation as Director, at least
not the main one. Indeed Beccarr’s resignation was for a different reason.

On one of his visits to Java during his travels he had been requested by the Dutch East Indies
Government to sell his collections to the Bogor (Buitenzorg) Herbarium for the cash payment of
a considerable amount of money and his appointment as botanical explorer in the Garden, or a
life annuity of 5000 lira. The offer was alluring, but BEccaRI wished his collections to remain in
Italy and to spend the rest of his life in Florence, attending to their study. However, his own estate
was seriously compromised owing to the expenses for his long expeditions, and before his depar-
ture for the last journey to Australia, New Zealand and Sumatra (1877—1878) BEccARI undertook
negotiations with the Florence Institute of Advanced Studies for the sale of his Malesian collec-
tions. The Institute asked Marquis G. DortA, the Director of the Civic Museum of Natural History
for an appraisal of BEccaRr’s botanical collections, which were estimated at 75,065 lira. On the
basis of this valuation and considering the offer of the Dutch East Indies Government, the Insti-
tute of Advanced Studies offered to buy all his botanical collections from Malesia against an annu-
ity of 5000 lira for the rest of his life. BEccaRt accepted this offer but on the condition that the
collections were entrusted to the Museum of Physics and Natural History of Florence and that
he was entitled to have them at his disposal during his lifetime; in exchange he would assume

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FLORA MALESIANA

responsibility for their study and conservation. During his journey, in May 1878, when he was
in Batavia prior to his expedition to West Sumatra, BECCARI was informed that the Board of the
Institute of Advanced Studies had accepted his conditions and had officially approved the pur-
chase of his collections. Consequently he refused the offer of the Dutch East Indies Govern-
ment.

When Beccaricame back to Florence and took up the directorship of the Botanical Collections
and Garden of the Museum, he ought to have accepted the contract but learnt that the Board of
the Institute of Advanced Studies intended to use the money from the Wess legacy for the pay-
ment of his life annuity. BECCARI was greatly disappointed on hearing this decision, since he had
thought that his life annuity would be paid by different Institute funds: he disliked the idea that
the WEBB collections were to be deprived of nearly all their endowment until his own death. Fur-
thermore, he knew that the WEBB legacy was the main source of income of the botanical depart-
ment of the Museum and without it, his ambition to make the Florence herbarium one of the
greatest in the world and a leading centre of tropical botany could not be accomplished. He under-
stood too that under these conditions, the sale of his collections was incompatible with the duty
of his office as Director. Indeed, he got a personal benefit from the sale, while as Director of the
Collections and Garden it was his duty to avoid that these were deprived of a large amount of
money necessary for their maintenance and increase during his lifetime. Thus he made every effort
to persuade the Board of the Institute of Advanced Studies to use different funds to purchase his
collections, but without result. He did not underestimate the hostility that the Institute had shown
him since the beginning, and particularly recently, and when he was invited to sign the contract,
he clearly understood that he was regarded by them as a troublemaker, and that the decision to
pay the price of his collections with the WEBB legacy was merely an expedient devised by the In-
stitute in order to compel him either to lose his prestige as Director or to resign from his office.
Indeed, if BEccarthad sold his collections and kept his office he would have lost his prestige as
a man and as Director, having put his personal interest before his duty. But the Board of the In-
stitute knew that BECCARI was a man of honour and that it would achieve its aim: his resigna-
tion.

Then, as a last attempt, he tried to find some way in which, without going back on his word,
he could withdraw from the compromise of the sale, but without success. Thus, on July 26, 1879,
Beccari resigned as Director of the Botanical Collections and Garden of the Florence Museum.
Only later, on October 31, 1879, as a private citizen, did he sign the contract for the sale of his
own collections. The fight had been lost, but his honour was saved!

Soon after the end of this unhappy and painful experience, on November 16, 1879, BECCARI
left for Ethiopia to stay with his old friend and benefactor, Marquis Giacomo Doria, as members
of an Italian expedition to the Assab Bay on the Red Sea which he had already visited in 1870.
They also spent some days collecting in Aden and returned to Florence on February 26, 1880.

After his return from Ethiopia, BEccARI resumed the study of his collections, which were lo-
cated in a few small rooms on the top floor of the Museum of Natural History. In those modest
and secluded rooms, alone, like a hermit, without any assistant or help, but together with his rich
collections, he worked until his death. There, he wrote his famous works on Malesian plants and
on palm taxonomy, and made the splendid drawings and photographs which adorn his
publications.

The first months there, however, were unfortunately rather hard for him. After his resignation,
in November 1880 TEoDoRO CARUEL was appointed Director of the Botanical Collections and
Garden of the Museum. He was soon instructed by the Institute of Advanced Studies to study the
advisability of removing the Botanic Collections and Garden from the Museum to the Giardino

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Dedication

dei Semplici and adjoining buildings, and eventually to prepare a project for such a removal which
had already been decided, but not realized, before PARLATORE’s death. BECCARI was aware that
CARUEL between 1866 and 1871 had been Director of the Giardino dei Semplici and that he was
in favour of the removal and was preparing the pertinent project. BEccArt had already expressed
his resolute opposition to the removal, chiefly because the buildings near the Giardino dei Semplici
were unsuitable for the collections and library as they were very humid, smaller and worse than
those of the Museum, but also because library, herbaria and garden were well settled in the
Museum and there was no need to remove them, and in doing so waste a large amount of money
which could have been used for their maintenance and growth. Besides, he was strongly convinced
that the great botanical collections and library at the Museum, as a centre of taxonomic research,
had to be kept distinct from the centre of teaching and research on anatomy and physiology at
the Giardino dei Semplici. Thus, in 1880 and 1881 Beccari tried everything, with letters and ar-
ticles in various Italian newspapers, to convince the Faculty of Science, the Board of the Institute
of Advanced Studies, and public opinion that the removal of the botanical collections and garden
would be a great and irreparable mistake. He carried out a referendum against the removal of the
herbaria and library among botanists in Italy and abroad. This was spread far and wide and was
discussed in many Italian and foreign publications. Numerous botanists from every part of the
world, and among them the most eminent taxonomists of the time, declared themselves against
the removal.

In 1881 Beccari published a paper in which the reasons for the protest against the removal of
the botanical department from the Museum and the result of the pertinent referendum were given.
In the same year CARUEL published his study for carrying into effect the project of the removal
of the Botanical Collections and Garden. Despite the opinions of many and outstanding botanists
expressed in the referendum, the Board of the Institute of Advanced Studies decided on the
removal of the Botanical Collections, Library and Garden from the Museum of Natural History,
in Via Romana, to the Giardino dei Semplici and adjoining buildings near San Marco, on the op-
posite side of the river Arno. However, BEccartdid not give in, and he continued to publish other
articles and papers against the removal until 1903. Even if his campaign did not gain its aim, it
greatly contributed to further resolutions of the Institute of Advanced Studies which decided to
enlarge and improve the buildings annexed to the Giardino dei Semplici and later to reserve for
Botany the part of them originally intended for the Zoology department, which remained at the
Museum, at La Specola, where it still is today. These deliberations greatly delayed the removal
of the Botanical Collections, though the living plants of the Garden of the Museum were all moved
to the Giardino dei Semplici during 1883. CARUEL continued to give his botanical lectures at the
Museum until his retirement, in 1896, but the following year, his successor, O. MATTIROLO, under-
took his teaching in the building near the Giardino dei Semplici. When in 1900 he moved to Turin,
P. BAccaRINI succeeded him as Director. At that time, the Library and the Botanical Collections
were still located in the Museum.

In 1901, when the removal was close at hand, BEccaARI was requested to inform the Institute
of Advanced Studies in which rooms of the new botanical building at the Giardino dei Semplici
he wished to have his Malesian collections deposited. He disdainfully replied that he wished his
collections to remain at the Museum in Via Romana in agreement with the contract of their sale,
adding that, if the Institute had decided to move them to the new buildings, he would not follow
them and would give up their study. His Malesian collections remained in the same rooms at the
Museum until the end of his life, but, in 1905, in spite of further protest and particularly after
controversy with P. BACCARINI, the removal of the Library and the Botanical Collections (includ-
ing all the herbaria) from the Museum of Natural History to the new Botanical Institute near the

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FLORA MALESIANA

ancient Giardino dei Semplici, was brought to a conclusion.

However, let us resume the course of BEccart’s life after his sad experience as Director of the
Botanical Collections and Garden of the Museum in 1878—1879, and his hard fight against their
removal started in 1881. These regrettable events marked a turning-point in BECCARI’s career. He
realized that he had lost the chance of making the Florence Herbarium one of the leading centres
for research in plant taxonomy, and decided to retire to private life, devoting himself entirely to
taxonomic research, chiefly to elaborate his own Malesian collections for which he had gathered
a wealth of field observations and drawings.

On January 23, 1882, BECCARI married NELLA GORETTI DE FLAMIN], from a noble family of Ca-
sentino, in the high valley of the Arno. They had four sons: NELLo, Dino, Baccio and RENZO.
The eldest, his devoted son NELLO, became a professor of Comparative Anatomy at Florence Uni-
versity and took great pains in editing some posthumous papers by the father, among them the
book Nuova Guinea, Selebes e Molucche based on the original diaries of his father’s explorations
in eastern Malesia from 1871 to 1876. He also encouraged U. MARTELLI and R.E.G. Picut SER-
MOLLI to revise and edit some works on palm taxonomy which had been left unfinished by his
father.

The years immediately following BECCARI’s marriage, entirely devoted to his family and to
study, were peaceful and fruitful. He set up his home in a villa inherited from his father, the me-
diaeval Castello del Bisarno, near Ripoli in the immediate vicinity of Florence, and he lived there
until his death. According to information obtained from his nephew and from letters to his
friends, we know that he also had another house in the city of Florence where he and his family
spent the week-days, particularly in winter. We also know that he used, as in the years before his
marriage, to spend several weeks, particularly in the summer holidays and during the grape-har-
vest at Radda in Chianti, on the old country estate of his mother’s family. He was very fond of
country life and, following the tradition of the old families of the region, he was particularly inter-
ested in wine-making, in which he attained great experience. He was one of the first producers,
together with Baron B. Ricasott, of that typical wine, well-known in Italy and abroad as ‘Chianti,
Gallo nero’. Nevertheless, during his holidays he did not stop his research, even if he did not work
so actively as in the Florence Museum, where his collections were housed.

However, after a few years his life was troubled by another sad event. In 1877 he had under-
taken the publication of a great work, Malesia, mainly with the intention of embodying in it the
results of the studies dealing with his own collections from the Malesian Archipelago; in addition,
other papers or abstracts of works published elsewhere on plants of that region were also to be
included. The first two volumes were printed in Genoa and BECCARI was greatly helped in editing
them by his faithful friend R. Gestro, the Director of the Civic Museum of Natural History of
Genoa, particularly during his last journey to the East. However, BECcARI undoubtedly corrected
the proofs of all the instalments of Malesia and also those of fascicle 3 of volume 1, issued when
he was in West Sumatra. This is proved by a letter to Gestro from Buitenzorg, now Bogor (dated
May 2, 1878), which accompanied the corrected proofs of that fascicle. BECCARI published the
first two fascicles of volume 1 at his own expense, but later Malesia became a publication of the
Florence Institute of Advanced Studies. However, despite its great interest, the sale of this work
was obviously limited, the text being written entirely in Italian. Consequently the Institute of Ad-
vanced Studies decided that it was not worth continuing its publication and suddenly, in 1887,
stopped all contributions to it while fascicle 3 of volume 3 was not yet complete. Actually, in the
cover of fascicle 3 we find a note which informs us that the publication of Malesia is ended and
explains the reasons for it. However, BECCARI wished to publish at least the text pertinent to the
drawings of the account on Bombacaceae already issued in fascicle 3, but as far as possible, also

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Dedication

other papers and drawings ready for the press. Hence, he was compelled to beg in Italy and abroad
for funds necessary to publish the last two fascicles of volume 3 of Malesia. Fortunately, the Min-
ister of Education, PAOLO BosELLI, and the BENTHAM Trust in England, where he was highly es-
teemed, allowed him the necessary financial support for bringing volume 3 of Malesia to a close.
The last issue appeared in March 1890.

Great was BEccarRt’s disappointment at the unhappy conclusion of the publication of Malesia,
not only because the resolution of the Institute of Advanced Studies represented a slight to him
and to his work, but also because he had lost a safe and certain means of publication for the results
of the study of his collections. He was so much upset by this event that he even thought of visiting
Malesia again. Actually, in the letter to Gestro(April 4, 1890) which accompanied the last fascicle
(‘ultimo definitivo’) of Malesia he asked his friend for information on the departures from Genoa
to Batavia and about the liners. Anyhow, the end of Malesia was another turning-point in his life:
it marked the beginning of a long period of inactivity, after which he never resumed the study of
his own Malesian collections.

Having concluded the studies already undertaken, he published no scientific papers from 1893
to 1902, except some articles and letters protesting against the removal of the Botanical Collec-
tions of the Florence Museum, described above, and the temporary closing of the herbaria and
library in connection with this removal. In these years he was on the point of giving up his
botanical activity entirely and none of his colleagues and friends, not even his devoted pupil UGo-
LINO MaRTELLI, were able to induce him to resume his research. However, another person was to
have the credit for reviving in him the enthusiasm for the country where he had spent the most
fruitful period of his youth.

After his explorations in Sarawak Beccarthad kept alive his friendship with the Rajah and the
Ranee of Sarawak. They liked to spend part of the year in the surroundings of Genoa. It is difficult
to say whether it was by chance or with the definite intention of helping BECcaRI to overcome his
scientific inactivity, but in the early days of May 1897 the Ranee visited Florence and met BECCARI.
A woman of great culture and sensibility, deeply fond of her kingdom of Sarawak, Lady MARGa-
RET BROOKE succeeded in convincing BECCARI to write a book on his fascinating explorations in
Borneo. He soon began his work; the Ranee kindly assisted him in the preparation of the book,
particularly in providing him with the illustrations. Several of them, in fact, area selection from
many fine photographs taken by the Ranee herself in Sarawak; these were assembled in a great
album, still kept in the Florence Botanical Museum, which she presented to BECCARIon June 10,
1897.

The preparation of the book, Nelle foreste di Borneo, led BECcaRI to recover a certain interest
for scientific work and it was not difficult for Prof. OrREsTE MATTIROLO, the Director of the
Botanical Department of the Museum, to complete the Ranee’s enterprise and convince him to
resume botanical research after the publication of his book. However, BEccarthad already real-
ized that the removal of the library and the herbaria from the Museum to the Giardino dei Semplici
was close at hand, and that without them the elaboration of his own Malesian collections would
be rather difficult. Consequently, he did not resume their study, interrupted in 1890 after the pub-
lication of Malesia was stopped, and decided to devote his entire botanical activity to a single
group. This decision was neither easy to take nor satisfying for him, but represented the best solu-
tion for carrying out his research without a big library and a great herbarium at his disposal at
any moment of the day. The selection of the group was easy, as the Palms had intrigued him since
his first visit to Malesia, and he had already done some research on them. On the other hand he
knew that this group offered him a taxonomically nearly unexplored, big field. Thus, BECCARI
started again on his studies with renewed enthusiasm, spending the whole day at the Museum in his

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FLORA MALESIANA

small rooms where he had concentrated everything necessary for his work, including the great
camera he had designed himself specially for making the marvellous photographs which are repro-
duced in the plates of his truly monumental works on Palms. In the years that followed he had
no difficulty in publishing his writings thanks to the great esteem he enjoyed abroad and the
friendship of several Italian botanists. Actually, the publication of his chief work, Asiatic palms,
was made possible by Sir GEorRGE KING, the Director of the Botanic Garden of Calcutta, and other
important papers were published by his close friend, U. MARTELLI, in the periodical Webbia,
which the latter had founded in honour of PHit1p BARKER WEBB. Some interesting works appeared
also in L’Agricoltura Coloniale, a journal edited by the Istituto Agricolo Coloniale, the founda-
tion of which was promoted by Beccarrand other Italian personalities. Several other papers on
palm specimens from all over the world entrusted by their collectors to him for determination,
were issued in various periodicals and books.

Beccariled this last period of his life completely secluded from Italian academic life and nearly
forgotten by most Italian botanists, but he was always overwhelmed by the sympathy and esteem
of foreign botanists. In this period, perhaps more than before, he enjoyed the affection of his old
and devoted friends and particularly of UGOLINO MARTELLI, his only pupil, a very keen botanist
himself, well known for his basic works on the great family of Pandanaceae, whose study he had
undertaken on BEccarr’s advice.

This period, entirely devoted to his family and the palm studies, was serene and creative. In the
last years of his active and eventful life he assembled the materials for a book on his explorations
of eastern Malesia. He had already sorted out a final copy of his diaries and he had also begun
to prepare the illustrations for his book, but unexpectedly death prevented him from accomplish-
ing this last performance.

He died peacefully in the evening of the 25th of October 1920, in Florence, at the age of 77.

ODOARDO BECCARI was a great explorer but at the same time a very clever, many-sided, careful
collector. Indeed he did incredible work in the field. In his long and lonely explorations in Malesia,
BECCARI was attracted by all aspects of nature and human life and assembled an enormous wealth
of botanical, zoological, ethnological and anthropological collections. Not only the number of the
specimens makes his collections really invaluable, but also the fact that these are often accom-
panied by notes and descriptions and sometimes by splendid and detailed drawings made in the
field.

The botanical collections from Malesia are kept in the Herbarium and Museum of the Florence
University. They amount to more than 21,000 sheets, about 2400 flasks of material in alcohol,
about 800 carpological specimens and more than 200 wood samples with the pertinent voucher
specimens (see further information in VAN STEENIS’s Thesaurus Beccarianus). Many collection
numbers are represented in BEccaART’s herbarium by more than one sheet. To these specimens we
must add many duplicates which were distributed to the most important herbaria, among which
those of the British Museum, Kew, Paris, Geneva, Leningrad, Berlin, Leiden, Vienna, Munich,
Stockholm, and Bogor.

The zoological collections consist of several thousand specimens belonging to a very high num-
ber of species, many of which were described as new. Those in the higher groups are represented
by skins, but sometimes also by skeletons, skulls and even heads or other parts of the body pre-
served in alcohol. They are kept in the Civic Museum of Natural History of Genoa, which bears
the name of its founder and Maecenas, Giacomo Doria, the companion of BEccARIon the expedi-
tions to Borneo and the Red Sea. BEccaRI paid attention to special groups and assembled several

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Dedication

collections of particular importance such as that of the great ape, the orang-utan (48 specimens
including a foetus), the set of birds of paradise, one of the best of its kind, and the collections
of fishes, spiders, coleoptera, and ants. These collections have supplied a very rich harvest of study
for many specialists. Indeed 205 works devoted to the study of BEccaArRI’s zoological collections
had been published by 1920 in the Annali del Museo Civico di Storia Naturale of Genoa alone
(see GEsTRO’s biography of BECCARI).

Beccari also collected rich and interesting ethnological collections of great value and beauty.
They are kept in the Anthropological and Ethnological Museum of Florence and some of them
are exhibited in a hall of that Museum. A set of ethnological collections was sent to the Italian
Geographical Society in Rome. Some of the idols, weapons, implements, ornaments, clothing,
etc. were described and illustrated in BEccarr’s books Nelle foreste di Borneo and Nuova Guinea,
Selebes e Molucche for the purpose of trying to establish, with the aid of the anthropological fea-
tures, the origin of some of the peoples of Malesia. The value of these collections was greatly in-
creased by the detailed information on the customs of those peoples which he described in his
books and in the letters to his friends published by E.H. GiGLioL1, G. Cora and the Italian Geo-
graphical Society.

The anthropological collections are kept in the Anthropological and Ethnological Museum of
Florence. They consist of skulls and a few skeletons of natives of Malesia. The most important
is undoubtedly the set of some 200 skulls assembled in Korido in the island of Schouten in north-
western New Guinea. These collections were the base of the first craniological investigations on
Papuans, by P. MANTEGAZzZzA and E. REGALIA.

BEccaRr’s activity in the field also extended to the geographical features of the districts he
visited. During the preparation of his expeditions, he had trained himself in geodetics and topo-
graphy and he had also invented a new instrument (Nuovo orizzonte artificiale. Rivista Marittima
6, 1873, 198—200, f. 1—5) for topographic surveys. During his travels he also made a topographic
survey of several territories, particularly of the northwestern parts of New Guinea. These surveys
later allowed G. Cora to prepare the maps he published in Cosmos (see Appendix 1, C: Maps).
He also discovered a great river, War Samson, in northwestern New Guinea, near Sorong.

Even though Beccarthad succeeded in assembling such enormous and invaluable collections,
his fame is mainly due to his scientific work condensed into more than 150 publications, some of
which consist of monographs of basic importance and those on palms especially still remain stan-
dard works even today.

BEcCaRI’ versatile mind allowed him to devote his attention to problems in different branches
of natural science, but he carried out his activity chiefly in botany. In the first period, during which
he made the great expeditions to Malesia, his botanical activity was essentially applied to the study
of a part of his collections; the result was published mainly in the Nuovo Giornale Botanico Italia-
no which he founded in 1869. Even if some of these papers were written here and there during
his explorations, his scientific output in this period was necessarily small.

With the end of his explorations in Malesia, the second period of BEccarr’s activity begins.
Probably he was unsatisfied when he limited himself to the descriptions of new genera and species,
and in this period he spread his field of research to the monographic or semi-monographic treat-
ment of those families or genera from Malesia which had most attracted him. In this period BEc-
CARI also undertook, at his own expense, the publication of a collection of botanical writings,
Malesia, in order to facilitate the printing of his papers dealing with his Malesian plants and the
reproduction of his splendid drawings. For this purpose he selected the quarto size. In Malesia
he published some of his most outstanding works. The end of its publication in 1890 marks the
end of this second period of his scientific activity and also the end of the study of his own collec-
tions from Malesia.

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FLORA MALESIANA

After about a decade of complete scientific inactivity, due to the sad vicissitudes of his academic
life and the end of Malesia, BEccaripublished Nelle foreste di Borneo in 1902. This year coincides
with the resumption of his scientific studies and marks the beginning of the third period of his
botanical research in which he devoted himself entirely to the study of the family of palms, becom-
ing the best specialist who ever existed.

In the following pages we want to go into more detail about the subjects contained in the mas-
sive oeuvre of the Maestro. Confronted with the multitude of his activities we hope to weave this
into a readable account, with a distinct feeling that our ability for writing falls short of the way
in which Beccari could express himself, as testified by his ‘Wanderings’, which is still a thrilling
guide for exploration in the tropics.

Let us start with this work on Sarawak, the core of his main work in Malesia. He worked under
favourable conditions, having ample equipment and time at his disposal, and the support of the
Tuan-muda, CHARLES BROOKE. His big hut, called ‘Vallombrosa’ on Gunong Mattang, a hill west
of Kuching, was used as a study centre. From there he made excursions and gradually familiarized
himself with the very rich flora of the primary forest. He focussed attention on the big trees (Dipte-
rocarpaceae, Bombacaceae, and others) as well as on the evasive tiny creatures of the saprophytic
Triuridaceae and Burmanniaceae, the parasitic plants, the lianas and so forth, making beautiful
and exemplary complete specimens in a skilled, professional way. This was, especially with un-
wieldy plants such as palms and pandans, gingers and aroids, quite an effort, as every field
botanist must be aware. Perseverance and patience fed by infinite interest must have induced him
to take particular care with these groups.

A special characteristic is that he knew his plants; hardly ever did he make two collections of
the same species. As a scientific collector he was never equalled, and only approached by E.J.H.
Corner and L.J. Brass. What a contrast with most other collectors who, even today, stick to the
disgusting grab-as-grab-can way of collecting on hurried cross-country trips, causing heavy dupli-
cation and absence of vital field notes.

How Beccarimanaged all this at the age of 22, with only a few months training in tropical form
knowledge at Kew, can only be understood if we imagine him as an extraordinarily gifted person
with an intense interest in botany; botany in the widest sense, because he was not satisfied only
with the taxonomy of flowering plants, but collected for example also wood samples and crypto-
gams of all major phyla. His horizon widened to collecting minerals and all sorts of animals, ob-
servations on vegetation types, on edible and horticultural plants, and the way of life of his com-
panions, the Dayak people; in short, he possessed the integrated interest of a born all-round natu-
ralist, whose scope went far beyond the mere plant collecting and description in which he excelled.
BECCARI assembled a great wealth of data on the geographical features of Borneo, on the matter
of useful and horticultural plants, on fibres, rattan, bamboos, resins, camphor, getah percha
yielding trees, and medicinal plants. He was aware of the primitive domestication of species of
Durio (durian), Eugeissona (a palm), Artocarpus (breadfruit), bananas and species of Nephelium,
which he learned from observation of Dayak life. BEccarRi briefly reported on these subjects in
a short summary of his journey in Sarawak to the Italian Geographical Society (1868), and later
he incorporated the complete data in some appendices to his book Nelle foreste di Borneo. Prob-
ably, when gliding easily in canoes on the rivers or proceeding with difficulty on the mysterious
peat of the swamp forests during his long journey in the interior of Sarawak, he ventured on ideas
about the origin of coal in Borneo — in which he was correct. When hunting for orang-utan in
Batang-Lupar or excavating in the limestone caves of Sarawak he made his first guesses about the
origin of man, which he developed in Nelle foreste di Borneo.

He surely must have had a very busy life in his ‘Vallombrosa’, because he could never have con-

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Dedication

ceived the ‘Wanderings’ thirty years later without having copious notes of his field observations
and full diaries. He must also have started in the field to compose botanical papers and prelim-
inary revisions; when still in Sarawak he published some small papers in Italy. BECCARI was much
enchanted with Sarawak, the flora, the country, and the Dayak people, and he even conceived
a project that the Italian government should purchase it as a crown colony.

When Beccarireturned from Borneo to Florence he founded the Nuovo Giornale Botanico Ita-
liano, in which he published freely some papers on spectacular Bornean plants; but before leaving
for the expedition to New Guinea he handed over the journal to T. CARUEL, since it would be diffi-
cult both to edit it and to explore in Malesia. When, loaded with further materials, manuscripts,
field notes and drawings, he returned from New Guinea to Florence, he realized the difficulty of
publishing the botanical results of his explorations in the Nuovo Giornale Botanico Italiano. Thus
he decided to undertake a collection of writings he named Malesia, in which he concentrated near-
ly all his works of that period. The first two instalments of it were published during his one year
interval, spent in Florence, between the last expedition to New Guinea and the journey to Austra-
lia, New Zealand and Sumatra, and the third one appeared during the last mentioned journey.
He undoubtedly must have worked very hard during that period, but he was able to manage it
thanks to his efficient organization en route and the help of his friend R. GesTroin Genoa, where
Malesia was printed.

BeccaRI published in Malesia several works which are important for the subject in itself, but
also various original considerations about some particular subjects, such as evolutionary pro-
cesses, dispersal of seeds, geographical distribution, efc. are dealt with in them. First should be
mentioned the extensive work on the ant plants devoted to the study of the symbiosis between
plants and ants, which occupies the entire second volume. In it, BECCARI gives us his interesting
views on the evolution and the common origin of plants and animals, discussing concepts still valid
and topical today. Likewise of great interest are his considerations on the origin of the insectivor-
ous plants and the distribution of plants in the Malesian archipelago, dealing especially with the
Nepenthaceae. Other important works are the monograph on the genus Phoenix and the account
of the palm genus Pritchardia in which he resumes his considerations on the dispersal of seeds and
fruits and the origin of the flora of the Pacific islands. The three volumes of Malesia contain also
a number of monographic or semi-monographic revisions of families and genera from Malesia,
e.g., Icacinaceae, Menispermaceae, Nepenthes, Bombacaceae, Triuridaceae, Burmanniaceae,
etc., and also a first survey of the palms of New Guinea.

We can conclude that Malesia was intended to embody the botany of Malesia as a repository.
It must have aroused great interest in the botanical world, containing novelties of fascinating
plants with marvellous drawings made by Beccarthimself, a ‘must’ for every botanical institute.
The use of the Italian language, even for monographic contributions from non-Italian collabora-
tors, e.g., on Araceae by ENGLER, was certainly an obstacle to a wide sale, and the edition was
subvented from Italian sources and the third and last volume could only appear thanks to the aid
of the BENTHAM Trustees.

Whether Beccartever intended or hoped to achieve an ultimate incorporation of all the mono-
graphs of Malesian plant families, that is, an attempt towards a true Flora Malesiana, remains
uncertain. The fact that he had certainly intentionally explored all areas of Malesia, except the
Philippines, and that the first instalments of Malesia contained several monographic treatises,
may support this idea. What is certain is that he fully realized that he could never dream of accom-
plishing all this himself. Accordingly he freely entrusted many groups to colleagues in Italy and
to his many correspondents abroad as appears from the bibliography in Appendix 4.

Through the distributed duplicates BECCARI’s material went to various herbaria and later came

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FLORA MALESIANA

into the hands of specialists, but much of his original collection, embodied in the original Herba-
rium Beccarianum, has not been examined by specialists. This original material is separately shelv-
ed in Florence, 400 bundles in 33 cupboards; to it belong a card system with field data and a cover
with drawings; there is a rather large number of unicates or collections of which no duplicates were
distributed. In 1951 vAN STEENIS very roughly sampled a number of families and found that BEc-
CARI Often had indicated and annotated genera in sched. as new, which were later based on other
material, e.g., Koordersiodendron ENGL. (1898), Clavistylus J.J.S. (1910), Neosepicaea DIELS
(1922), Octamyrtus Diets (1922), Haplolobus H.J. Lam (1931), Kjellbergiodendron BURRET
(1936), Macadamia hillebrandii STEEN. (1952), Eriandra v. ROYEN & STEEN. (1952), Whiteoden-
dron STEEN. (1952).

It is a pity that in the past five decades too little initiative has been taken by the curators of the
Florence Herbarium to attract and induce specialists to study the original BEccaRi collections in
Florence. We suppose that it is due to the understaffing of this great Herbarium. Especially the
Herbarium Beccarianum is not a reliquiae, not a closed chapter, but truly a thesaurus, still con-
taining unknown botanical treasures.

As this is not always realized by specialists, we urge them to borrow material of their speciality
from this century-old, inexhaustible source. We sincerely hope that a revival of interest in the Her-
barium Beccarianum is welcome to the future curators of the Florence Herbarium. Its possession
brings with it the scientific obligation of using it, not just in honour of the Maestro, but mainly
for the benefit of scientific botany and as a contribution to the fame of the Florence centre.

After publication of Nelle foreste di Borneo in 1902, BEccaARI decided to concentrate, for the
rest of his life, on the study of one large family on which he possessed more field knowledge than
anyone, before or since, namely the palms. His first contribution to their knowledge dates from
1871, with a provisional account of those of Borneo. Old love never dies! In 1877 he had ac-
counted for the palms of New Guinea, in 1885 for those cultivated in the Botanic Gardens at Bo-
gor, but in about 1890 he spread his wings towards those beyond Malesia, the Indian empire, Indo-
China, and later to Madagascar, Africa, the Pacific islands, etc. Everybody entrusted him with
palm material and from this emanated a massive knowledge of Asiatic palms, embodied partly
in the Records of the Botanical Survey of India, in HooKker’s Flora of India, partly later in Web-
bia, founded by his old pupil and friend U. MarTELLI, but largely in the sumptuous volumes of
the Annals of the Royal Botanic Garden, Calcutta, which also included those of Malesia. For the
large folio plates of these massive plants BECCARI designed a special large camera with suitable
accessories in order to achieve excellent illustrations!. He devoted his attention mainly to the tax-
onomy of palms, but he also studied the cultivated species in some works which appeared in
L’Agricoltura Coloniale, edited by that Institute once named Istituto Agricolo Coloniale, now
Istituto Agronomico per |’Oltremare of which he had solicited the foundation in 1903. Of this big
work a large number of unpublished manuscripts appeared in print after his death, through the
untiring devotion of his pupil, friend, and colleague MARTELLI, who must be given a tribute of
honour for his singularly unselfish efforts. The last of the manuscripts on palms which BECCARI
left unfinished, that of the subfamily Arecoideae, was completed and published by PicHiI SERMOL-
L1in 1955. We should also refer here to Moore’s important and competent evaluation of BECCA-
RI’s massive contribution to the knowledge of the fascinating palm family.

(1) The large camera and other microphotographic cameras designed by BEccarRI are described by LuIGI
PAMPALONI, Apparecchio fotografico universale per laboratorio biologico ideato dal Dottor Beccari. Rend.
Congr. Bot. Naz. Palermo (1903) 164—168, cum fig., and Gli apparecchi microfotografici del Dott. O. Becca-
ri. Bull. Soc. Fotogr. Ital. 14 (1902) 129—145, fig. 1—7.

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Dedication

Dealing with his botanical activity we cannot silently pass over his descriptive work. Also in this
BEccari excelled and showed that he had a remarkable insight into affinities. It appears that his
new genera were always placed in the proper plant family and, moreover, that hardly ever were
new species proposed by him reduced later, stamping him as a most accurate taxonomist. Indeed
he was a taxonomist, but BEccARIShowed his sharp intelligence in other branches of botany often
including his considerations in taxonomical papers. Describing the details of Gnetum led him to
considerations about the ancestry of the flowering plants from the Gymnosperms. The plant geo-
graphy of the palms led him to hypothetical ideas about former landbridges and sunken conti-
nents. His gatherings in Sumatra led him to consider the affinities of its flora with those of South-
east Asia and Java, concluding that the flora of volcanic ranges must be much younger than that
of the more ancient and more stable Sunda lands. Other observations deal with the dissemination
by earthworms; the double dispersal, anemochorous and zoochorous, of the plants of the periodic
swamp forest provided with floating fruits and succulent seeds; pollination by pigeons; the various
colours of flowers of the forest plants, and some others on physiology and ecology.

BEccaRi also left traces of his versatile genius in various writings (papers and letters to his
friends) which lie outside botany but must be mentioned briefly to understand how great he was
as a naturalist. Particularly interesting are the letters to E.H. Gicuiot1and G. Cora in which he
disclosed his views on the origin of the peoples of Malesia, in particular of the Papua-Mafor which
he regarded as derived from a crossing of aboriginals, perhaps descended from Negritos and Hin-
du peoples. Other interesting observations are those on the connections between mosquitos and
malaria which he was one of the first to suppose, those on the agent of the bee pest which he sus-
pected to be due to a protozoon later discovered in America, those on the connections between
flies and cholera and numerous other observations particularly on the customs of animals.

It was in Borneo that BeccarRi perceived the true value of evolution and was primarily fascinat-
ed by the importance of adaptation to environmental conditions. But only later did his views on
the processes of evolution take shape in his mind. It is a fact that the prolonged stays of gifted
naturalists in the tropical wilderness, when their minds are set free from daily minutiae and domes-
tics and solely occupied with the bewildering structural wealth of tropical plants and animals,
allows their minds to open to new, big ideas and syntheses, generating philosophical thought. For
this, one has only to think of von HUMBOLDT, JUNGHUHN, WALLACE, DARWIN, and CorRNER. To
this, BECCARI, with his eager mind and astute power of observation, was no exception.

BECCARI was used to going back from the facts to the causes, and his views on the evolutionary
processes, which arose from the observations he made in nature, were consolidating in his mind
in the course of time. Thus we find his views sketched in some papers and later resumed in others,
whenever he had the chance to develop them on the basis of particular new observations. He did
not supply us, or perhaps he did not want to supply us, with a synthesis of his views on evolution
in an ad hoc publication, perhaps out of humility, since he disliked giving the impression that he
was able to explain the laws of evolution, or perhaps out of honesty, because he perceived that
his ideas had made their way into his mind by intuition and reasoning, without adequate investiga-
tion.

BECCARI was undoubtedly an evolutionist, but he was one in a very original manner. The first
foundation of his theory of ‘plasmation’ was explained in his paper (1876) on the huts and gardens
of Amblyornis inornata, the small bower-bird of paradise which builds a pretty hut with, in front,
a lovely garden of soft moss on which it scatters flowers in shining colours changing them when
they wither. This theory was resumed in the introduction to his work on ant plants (1884) and was
later developed in his paper (1889) on the flowering of Amorphophallus titanum, the gigantic Ara-
cea, and was summarized in his book, Nelle foreste di Borneo.

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FLORA MALESIANA

According to this theory the evolutionary processes of living beings took place, beginning with
the most ancient geological times, fundamentally in two different epochs: a first epoch of plasma-
tion, and a second epoch of conservative heredity, displaying in the course of time the two funda-
mental rules of variability and fixity. In the first epoch, the plasmative strength, unhindered by
heredity, may have given free play to the variability and to the adaptation stimulated by environ-
mental conditions. In that epoch, the organism may have been liable to yield to the stimulus of
external factors and more subject to modelling itself to them, the modifications occurring with
the greatest of ease and even quite suddenly without the offspring necessarily being like the par-
ents. This epoch may have been a period of youth for living beings in which each individual was
allowed to modify itself in conformity with its needs, or rather even according to its wishes, its
vanities, its whims.

This epoch of plasmation, with a maximum of variability and a minimum of fixity, may have
been followed by an epoch of conservative heredity, characterized by that strength which aims at
the conservation of the acquired characters and owing to which the individuals belonging to a spe-
cies transmit to their descendants the characteristics they have inherited from their ancestors. Thus
the plasmative epoch may have been replaced by an epoch with a minimum of variability and a
maximum of fixity. The strength of the conservative heredity becoming stronger in the course of
time may consequently have weakened the faculty to vary, perhaps even to cancel it entirely, thus
impressing the stamp of fixity on all living beings.

He recognized the great influence of environmental factors on the plasmation of living beings
and pointed out several cases of correlation between the morphology of some apparatuses and the
environmental factors. For instance, he was the first to correlate the life form of flood-resistant
plants with the environment and was struck by their similarity in leaf-shape: his ‘stenophyllous
plants’, now called ‘rheophytes’. A still more important correlation amply studied by him was that
of the symbiosis between plants and ants, the ‘piante ospitatrici’, or formicarian plants, to which
he devoted a very large and detailed account (1884). However, he clearly and repeatedly recog-
nized that plasmation could also be stimulated by an interior strength, by the wish of having some
particular functions facilitated, such as defence, pollination, seed dispersal, etc. But BEccARtalso
supposed that plasmation was even influenced, particularly in the animal kingdom, by a psychic
push stimulated by the beauty of the environment, as could have happened in the birds of paradise
desirous of imitating with their feathers the glowing colours of the aurora and dusk of the tropics,
which they greet from the highest trees of the forest with very lively dances.

Though recognizing that the extant being cannot, as a rule, undergo modifications because of
the environment, BECCARI admitted however that even today some changes can take place in the
species on account of a cross between individuals of different species or of the sudden appearance
of hereditary modifications of various, even if unknown, origin. He admitted that these could be
induced by new poisonous substances or by new enzymes arising in the substratum; thus he fore-
saw the existence of mutations and mutagenic substances.

BECCARI was a man of great intelligence, versatility and intuition, who united an exceptional
personality and liberality with uncommon integrity and strength of mind. He was an indefatigable
worker, who devoted hours and hours to his research, without a moment of rest. But he did not
work out of a wish to be praised; prizes and honours did not interest him. He loved his research
studies since he was zealous of the beauty and perfection of nature and only happy when he could
entirely devote himself to investigating its manifold and marvellous aspects. For the same reason
he liked to draw plants and animals and very few excelled him in scientific drawing.

Beccari had an austere and inflexible character, but he was neither obstinate nor autocratic.
His temperament was based on a keen sense of duty. Also in private life, although deeply fond of

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Dedication

his wife, sons and friends, every action was characterized by a clear austerity, and he was greatly
beloved but at the same time he always inspired a certain awe. But this austerity concealed a great
goodness of heart.

He had also a great sense of justice and was a very courageous and stalwart man. These gifts
united with the wish to inspire respect and love rather than fear, were greatly esteemed by native
people and on his dangerous expeditions he never suffered injury from anybody. Undoubtedly
he must have encountered serious danger, but he faced them with resolution, and he recounts them
with indifference and without boasting.

BeEccar! is described as a proud, almost misanthropic spirit and perhaps he was not an easy
character and we understand why some regarded him as a troublesome person. From his youth,
moulded in his long and solitary explorations in Malesia, he was set apart, destined to travel and
to work alone, and he was not afraid of solitude; on the contrary, in his ripe age he found refuge
in it, the better to devote himself to his studies and serve his single purpose and sole end: the
science of nature.

R.E.G. Picut SERMOLLI & C.G.G.J. VAN STEENIS

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FLORA MALESIANA

Appendix 1 — Bibliography of Odoardo Beccari

A — Books and papers
(excluding reviews of books)

1861. Escursione botanica. — L’Araldo Cattolico, Lucca, anno XVIII, 14 agosto 1861, n. 33: 264
(not seen).

1862. Illustrazione dell’ Arnoldia cyathodes Massal. — Comment. Soc. Crittog. Ital. 1: 128—130,
tewle

1868. Descrizione di tre nuove specie di piante Bornensi. — Atti Soc. Ital. Sc. Nat. 11: 197—198.

— Cenno di un viaggio a Borneo. — Boll. Soc. Geogr. Ital. 1: 193-214.

1869. Illustrazione di nuove specie di piante Bornensi. (Balanophoreae, Rafflesiaceae). — Nuovo
Giorn. Bot. Ital. 1: 65—91, t. 2—5S.

—— Varieta e notizie. — Ibid.: 158—160.

— Lamenti del Redattore. Rivista bibliografica. — Ibid.: 222—224.

1870. Illustrazione di nuove specie di piante Bornensi. (Aristolochiaceae). — Nuovo Giorn. Bot.
Ital. 2: 5—8, t. 1.

— Nota di una nuova specie del genere Stenomeris. — Ibid.: 8—12, t. 2.

— Nota sul Trichopodium zeylanicum Thw. — Ibid.: 13-19, t. 3.

— Nota sull’embrione delle Dioscoreaceae. — Ibid.: 149-155, t. 4.

—— Disepalum coronatum nuova specie di Anonacea bornense. — Ibid.: 155—156, t. 5.

—— (O. Antinori & A. Issel). Relazione sommaria del viaggio nel Mar Rosso dei Signori Antinori,
Beccari e Issel. — Boll. Soc. Geogr. Ital. 5(2): 43-60.

1871. Descrizione di due nuove specie di Hydnora d’ Abissinia. — Nuovo Giorn. Bot. Ital. 3: 5—7.

— Petrosavia. Nuovo genere di piante parassite della famiglia delle Melanthaceae. — Ibid.:
(Sle ieeale
— Note sopra alcune palme Bornensi. — Ibid.: 11—30.

— Le Hydrocotyle d’Europa. — Ibid.: 102.

—— Sui generi Bihania ed Eusideroxylon. — Ibid.: 102—103.

— Cattedra di Botanica nell’Universita di Torino. — Ibid.: 103-104.

—— Illustrazione di nuove 0 rare specie di piante Bornensi. (Anonaceae). — Ibid.: 177—193, t.
2-7.

1874. Descrizione di una nuova specie del genere Myrmecodia della famiglia delle Rubiaceae. —
Nuovo Giorn. Bot. Ital. 6: 195—197, t. 6.

1875. Osservazioni supra alcune Rafflesiaceae. — Nuovo Giorn. Bot. Ital. 7: 70—75.

1876. Le Capanne ed i Giardini dell’ Amblyornis inornata. — Ann. Mus. Civ. St. Nat. Genova
9: 382—400, t. 8.

—— Lettera del Dr. A.B. Meyer al Mse. G. Doria (con nota di O. Beccari). — Ann. Mus. Civ.
St. Nat. Genova 8: 333-334.

1877. Relazione intorno alla Baia di Assab. — Cosmos 4: 230—232.

— Della organogenia dei fiori feminei del Gnetum gnemon L. — Nuovo Giorn. Bot. Ital. 9:
91—100, t. 7 (reprinted in Amer. Journ. Sc. 13: 469—471).

—— Sulla Cardiopteris lobata Wall. — Nuovo Giorn. Bot. Ital. 9: 100—108, t. 8.

— Die Flora der Aru-Inseln. — Ausland 50: 759-760.

—— Sul nuovo genere Scorodocarpus e sul genere Ximenia L. della famiglia delle Olacineae. —
Ibid: 273=279 ts 11.

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Dedication

— Della disseminazione delle palme. — Bull. R. Soc. Tosc. Ortic. 2: 167—173.

—— II sagu della Nuova Guinea, Metroxylon rumphii Mart. — Ibid.: 247—249.

— Catalogue of the plants of the Fly River (New Guinea) collected by De Albertis, 1877. —
Journal of Melbourne Logbook 30 Jan. 1877 (not seen).

— Le specie di palme raccolte alla Nuova Guinea da O. Beccari e dal medesimo adesso descritte,
con note sulle specie dei paesi circonvicini. — Malesia 1: 7—96, t. 1 & 2.

— Nuove osservazioni sulle palme della Nuova Guinea. — Ibid.: 97—102.

— Studio monografico sopra le piante della famiglia della Icacineae e delle Menispermaceae sin
qui scoperte nella Malesia e nella Nuova Guinea. — Ibid.: 103—165, t. 3—8.

1877—78. Piante nuove o rare dell’Arcipelago Malese e della Nuova Guinea, raccolte, descritte
ed illustrate da O. Beccari. — Leguminosae-Caesalpinieae, Palmae, Chailletiaceae, Gymno-
spermae papuanae, Coniferae, Gnetaceae, Cycadaceae, Violaceae, Magnoliaceae, Monimia-
ceae, Araliaceae, Ericaceae, Vacciniaceae, Nepenthaceae, Corsia, Burmanniaceae. — Ibid.: 1
(1877) 167—192; contd. (1878) 193—254, t. 9-15.

1878. Sulle piante raccolte alla Nuova Guinea dal Sig. L.M. D’Albertis durante l’anno 1877, con
descrizione di tre nuove specie di Icacineae. — Ibid.: 1 (1878) 255—256; contd. (1883) 257—258
(incl., pp. 257—258: Su di un nuovo genere della famiglia delle Olacineae).

— II Conophallus titanum Beccari. — Bull. R. Soc. Tosc. Ortic. 3: 290—293, f. 32.

1879. Le Nepenthes. — Bull. R. Soc. Tosc. Ortic. 4: 13-19.

— La pit piccola delle Araceae: Microcasia pygmaea Becc. — Ibid.: 179-181, cum fig.

1880. Notes on the plants collected by Sig. L.M. D’Albertis in New Guinea. — In: L.M. D’Alber-
tis, New Guinea: what I did and what I saw. 2: 391—395.

—— Catalogue of the plants of the Fly River, 1877. — Ibid.: 396—400.

— Sul viaggio di O. Beccari in Sumatra. Giugno—Ottobre 1878. — Boll. Soc. Geogr. Ital. ser.
2, 5 (1880) 300—302.

— Lettera di O. Beccari al Preside della Facolta di Scienze naturali del R. Istituto di Studi Supe-
riori di Firenze. Dicembre 1880 (Published by U. Martelli, Webbia 5 (1921) 329—330).

1881. R. Museo di Fisica e Storia Naturale. L’Orto botanico. — Giornale ‘La Nazione’ 6 Gennaio
1881.

—— (& G. Doria) Viaggio ad Assab nel Mar Rosso, dei signori G. Doria ed O. Beccari con il R.
Avviso ‘Esploratore’ dal 16 Novembre 1879 al 26 Febbraio 1880. — Ann. Mus. Civ. St. Nat.
Genova 16: 523—524.

— A proposito del progetto di remozione delle collezioni botaniche dal Museo di Storia Natura-
le di Firenze (not seen).

— Sull’abbandono del museo e del giardino botanico della Specola a Firenze. Protesta dei culto-
ri della botanica in Firenze colle adesioni e le osservazioni dei botanici italiani ed esteri. — Firen-
ze, Tip. Sborgi, in 8°, 72 pp.

— Cenni sopra la Flora di Assab. — Bull. R. Soc. Tosc. Ortic. 6: 108—110.

—— Beitrage zur Pflanzengeographie des malayischen Archipels. — Bot. Jahrb. 1: 25—40 (con-
densed version in German of Malesia 1: 214—238).

1883. Su di un nuovo genere della famiglia delle Olacineae. — Malesia 1: 257—258 (included in:
Sulle piante raccolte alla Nuova Guinea dal Sig. L.M. D’Albertis durante l’anno 1877, con de-
scrizione di tre nuove specie di Icacineae. See 1878).

1884. Piante ospitatrici, ossia piante formicarie della Malesia e della Papuasia descritte ed illustra-
te da O. Beccari. — Ibid.: 2: 5—128, f. 1—9, t. 1-25; contd. (1885) 129—212, f. 10—15, t. 26—54;
contd. (1886) 213—284, f. 16—17, t. 55—65.

—— Veratronia malayana Miq. — Bull. R. Soc. Tosc. Ortic. 9: 70—72, t. 4.

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FLORA MALESIANA

— II The in Italia. — Boll. Notiz. Agrar. Minist. Agric. 6: 279—281; also in L’ Agricolt. meridio-
nale 7: 116-118, and La Natura, Milano, 1: 267—268.

— II colera e le mosche. — La Natura 1884(33): 81-83.

1885. Orto Botanico di Firenze. — Giornale il ‘Corriere’, 14 Maggio 1885.

— Reliquiae Schefferianae. Illustrazione di alcune palme viventi nel Giardino Botanico di Bui-
tenzorg. — Ann. Jard. Bot. Buitenzorg 2: 77—171, t. 1—14.

—— Cyrtosperma (Alocasia Hort.) johnstonii Becc. — Bull. R. Soc. Tosc. Ortic. 10: 5—7.

—— Plantes a fourmis de l’Archipel Indo-Malais et de la Nouvelle Guinée. (Compte rendu de Mr.
E. Levier). — Archiv. Ital. Biol. 6: 305—341.

1886. Rivista delle specie del genere Nepenthes. — Malesia 3: 1—15, t. 1—3.

—— Rivista delle felci e licopodiacee di Borneo e della Nuova Guinea, enumerate o descritte dal
Barone V. Cesati nella memoria che porta il titolo: ‘‘Felci e specie nei gruppi affini raccolte a
Borneo dal Sig. O. Beccari’’ e dell’altra: ‘‘Prospetto delle felci raccolte dal Sig. O.B. nella Poli-
nesia’’. — Ibid.: 16—55, t. 4—S.

— Nota sopra alcune felci raccolte dal Sig. J.E. Teysmann all’isola di Sumba o Sandal-wood
ed in Timor. — Ibid.: 56—57.

— Nuovi studi sulle Palme Asiatiche. — Ibid.: 58—149, t. 6—11.

— Sulla Quercus robur Linn. Nota di botanica legale. Perizia nell’interesse della causa Le Pen-
nec e Monetti contro Vitali, Picard, Charles e Comp.i. — Firenze, Tip. Nicolai, in 4°, 44 pp.

1887. Turgescenza dei petali di Magnolia Yulan. — Malpighia 1: 420.

1887—88. Le palme incluse nel genere Cocos Linn.: Studio preliminare. — Malpighia 1 (1887)
343-354, 441-454, t. 9; 2 (1888) 85—95, 147-156.

1888. Nuove specie di palme recentemente scoperte alla Nuova Guinea. — Nuovo Giorn. Bot. Ital.
20: 177-180.

1889. Nuove palme asiatiche. — Malesia 3: 169—200.

— Le Bombaceae Malesi descritte ed illustrate da O. Beccari. — Ibid.: 201—280, f. 1—16, t.
12—36 (plates 12—35 were published before the text in vol. 3, no. 3, August 1887).

— Fioritura dell’Amorphophallus titanum Becc. — Bull. R. Soc. Tosc. Ortic. 14: 250—253,
266—278, t. 8 (Reprint with separate pagination: 17 pp., 3 plates).

— Palmae. — In: K. Schumann & M. Hollrung, Die Flora von Kaiser Wilhelms Land (Beiheft
zu den Nachrichten tiber Kaiser Wilhelms Land und den Bismarck-Archipel): 15—17.

1890. Le palme del genere Pritchardia. — Malesia 3: 281—317, t. 37—38 (the plates were published
before the text in vol. 3, no. 4, September 1889).

— Le Triuridaceae della Malesia. — Ibid.: 318—344, t. 39—42 (the plates were published before
the text in vol. 3, no. 4, September 1889).

—— Rivista monografica delle specie del genere Phoenix Linn. — Ibid.: 345—416, f. 17, t. 43—44.

1891. Letter to Prof. G. Papasogli. — In: G. Papasogli, La nitrobenzina usata come insetticida.
— Agric. Toscana 9: ?—? [repr. 1—6].

1892—93. (& J.D. Hooker) Palmae. — In: J.D. Hooker, The Flora of British India 6: 402—448,
449-483.

1893. Palmae. — In: G.F. Scott Elliot, New and little known Madagascar plants, collected and
enumerated by G.F. Scott Elliot, M.A., B.Sc., F.L.S. — J. Linn. Soc. London, Bot. 29: 61—62.

1901. Lettera al Presidente della Societa Botanica Italiana. — Bull. Soc. Bot. Ital. (1901) 119—124
(Dated March 23, 1901).

—— Per il Giardino Botanico. — Giornale ‘La Nazione’, 12 Maggio 1901.

— Trasloco delle collezioni botaniche del Museo di Firenze. Lettera al Presidente della Societa
Botanica Italiana. — Bull. Soc. Bot. Ital. (1901) 202—210.

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Dedication

— Sul trasloco delle collezioni botaniche della Malesia dal Museo di Storia Naturale in Via Ro-
mana ne’ nuovi locali presso il Giardino de’ Semplici. — Corrispondenza del D. Odoardo Bec-
cari coll’Istituto di Studi Superiori. — Firenze, Tipogr. S. Landi, 4 pp. (Dated July 2, 1901).

1902. Nelle Foreste di Borneo. Viaggi e ricerche di un naturalista. — Firenze, xvi+667 pp., 81
fig., 4 maps.

—— Systematic enumeration of the species of Calamus and Daemonorops with descriptions of
the new ones. — Rec. Bot. Surv. India 2: 197—230.

1903. L’Istituto di Studi Superiori di Firenze, la chiusura del Museo Botanico e le sue peripezie.
Osservazioni e Critiche. — Rocca San Casciano, Licinio Cappelli, 16 pp., 3 fig.

— Per un Istituto Agricolo Botanico Coloniale. — Giornale ‘Fieramosca’, 21 Marzo 1903 (not
seen).

— L’avvenire agricolo dell’Eritrea. — ‘La Tribuna’, Roma, | Giugno 1903 (not seen).

1904. Palmae. — In: J. Perkins, Fragmenta Florae Philippinae 1: 45—48.

— Wanderings in the great forests of Borneo: travels and researches of a naturalist in Sarawak
(transl. by E.H. Giglioli, rev. and ed. by F.H.H. Guillemard). — Constable, London,
xxiv + 424 pp., 61 fig., 3 maps.

1905. Le palme del genere Trachycarpus. — Webbia 1: 41—68, f. 1—16.

— Note anatomiche sul frutto dei Trachycarpus. — Ibid.: 68—72, f. 17—18.

— Notizie sul Nannorhops ritchieana H. Wendl. — Ibid.: 72—73; also Bull. R. Soc. Tosc. Ortic.
30: 325—326.

— Palme nuove papuane. — Ibid.: 281—313, f. 1-8.

— Le palme delle Isole Filippine. — Ibid.: 315—359.

— Palmae. — In: K. Schumann & C. Lauterbach, Nachtrage zur Flora der Deutschen Schutzge-
biete in der Stidsee: 60—61.

1906. Palmarum madagascariensium synopsis. — Bot. Jahrb. 38, Beibl. 87: 1—41, f. 1.

1907. Le Palme Americane della tribu delle Corypheae. — Webbia 2: 1—343.

— Notes on Philippine palms. I. — Philip. J. Sc. 2: Bot. 219—240.

1908. Asiatic palms — Lepidocaryeae. Part I: The species of Calamus. — Ann. R. Bot. Gard.
Calc. 11 [Fol.]: Letter-press I-VI, 1—518, t. 1—2; Plates t. 1-236 (1—231+25A, 104A, 172A,
188A, 191A).

— Le palme ‘Dum’ od ‘Hyphaene’ piu specialmente quelle dell’ Affrica italiana. — Agric. Co-
lon. 2: 137-183, f. 1—2, t. 1-3.

—— Palmae novae antillanae. — In: Fedde’s Repert. Bot. Syst. 6: 94—96.

— The palms of the Batanes and Babuyanes Islands. — Philip. J. Sc. 3: Bot. 339-342.

1909. New or little known Philippine palms. — Elmer, Leafl. Philip. Bot. 2: 639-650.

— Notes on Philippine palms. II. — Philip. J. Sc. 4: Bot. 601—639, t. 30—31.

—— Palmae. — Nova Guinea 8: 203—222, t. 50—S1.

1910. Glaziova treubiana, nouvelle espece de Cocoinée, avec observations sur le genre Cocos. —
Ann. Jard. Bot. Buitenzorg ser. 2, suppl. 3: 791—806, 2 fig., t. 31.

— Le palme del genere Raphia. — Agric. Colon. 4: 137—170, f. 1—3, t. 1—6 (reprinted with
the original pagination, as an issue of the series ‘Biblioteca Agraria Coloniale’).

—— Studio monografico del genere Raphia. — Webbia 3: 37—130, f. 1—8, t. 1.

—— Palme australasiche nuove 0 poco note. — Ibid.: 131—165, f. 1—6.

—— Descrizione di una nuova specie di Trachycarpus. — Ibid.: 187—190.

— Palmae dell’Indo-China. — Ibid.: 191—245.

— Contributo alla conoscenza delle Lepidocaryeae affricane. — Ibid.: 247—294.

— La Copernicia cerifera in Riviera ed una nuova specie di Livistona. — Ibid.: 295—305, f. 1,
thy.
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FLORA MALESIANA

1911. Asiatic palms — Lepidocaryeae. Part II: The species of Daemonorops. — Ann. R. Gard.
Calc. 12(1) [Fol.]: Letter-press [I—II], 1—237, t. I—II; Plates t. 1—109.

— Classification des palmiers d’Indo-Chine. — Bull. Mus. Hist. Nat. Paris 17: 148—160.

— Le palme che producono fibre di Piassava nel Madagascar. — Agr. Colon. 5: 320—324, f.
A-D.

— The palms of the island of Polillo. — Philip. J. Sc. 6: Bot. 229—230.

1912. Palmae. — In: H. Winkler, Beitrage zur Kenntnis der Flora und Pflanzengeographie von
Borneo. II. — Bot. Jahrb. 48: 89—93.

— Palmae. — In: I. Urban, Symbolae Antillanae 7: 170-172.

1912—13. The palms indigenous to Cuba with appended remarks on the general structure of the
trunk of a palm. — Pomona Coll. J. Econ. Bot. 2 (1912) 253—276, f. 109—118; 351—377, f.
144—153; 3 (1913) 391—417, f. 154-172.

1912—14. Palme del Madagascar. — Firenze, Istituto Micrografico Italiano, [Fol.], [i—iv] +62
pp., 50 fig., 50 tav.

1913. Contributi alla conoscenza delle palme. — Webbia 4: 143—240, f. 1—17.

— Manipolo di palme nuove Polinesiane conservate nell’erbario di Kew. — Firenze, M. Ricci,
October 1913 (preprinted with the same pagination from Webbia 4: 253—291, f. 1—31, 1914).

— Studio sui ‘Borassus’ e descrizione di un genere nuovo Asiatico di ‘Borasseae’. — Firenze,
M. Ricci, November 1913 (preprinted with the same pagination from Webbia 4: 293—385, f.
32-42, 1914).

1914. Asiatic palms — Lepidocaryeae. Supplement to Part I: The species of Calamus. — Ann.
R. Bot. Gard. Calc. 11 (App.) [Fol.]: Letter-press i—viii, 1—142; Plates t. 1—83.

— Contributo alla conoscenza delle palme a olio (Elaeis guineensis). — Agric. Colon. 8: 5—37,
t. 1—5; 108-118, t. 7-14; 201—212, t. 24-26; 255—270, t. 40—41 (reprinted with separate pagi-
nation: 80 pp., 18 tav., as an issue of the series ‘Biblioteca Agraria Coloniale’).

— Neue Palmen Mikronesiens. — In: G. Volkens, Beitrage zur Flora von Mikronesien. — Bot.
Jahrb. 52: 4.

— Neue Palmen Papuasiens. — In: C. Lauterbach, Beitrage zur Flora von Papuasien iv: 26B.
— Bot. Jahrb. 52: 23—39.

— Palmae. — In: K. Rechinger, Botanische und zoologische Ergebnisse einer wissenschaftli-
chen Forschungsreise nach den Samoainseln, dem Neuguinea-Archipel und den Salomonsinseln
von Marz bis Dezember, 1905. V. Teil. — Denkschr. Kaiserl. Akad. Wiss. Math.-Naturwiss.
Kl. 89 (‘1913’) 502—S09, f. 5—12.

1916. Palmae. — In: J.F. Rock, Palmyra Island with a description of its flora. — Coll. Hawaii
Publ. Bull. 4: 44—48, t. 17—18 (reprinted with the title ‘Palmae. Cocos nucifera L. forma pal-
myrensis. Characteristics of the Coconuts of the Palmyra Islands’ and different pagination: p.
1—5, t. 18-19).

—— Il genere Cocos e le palme affini. — Agric. Colon. 10: 435—471, t. 1—4; 489-532, t. 5—13;
585—623, t. 14—15 (reprinted with separate pagination: 128 pp., 15 tav., as issue of the series
‘Biblioteca Agraria Coloniale’).

— Palmae. — In: E. Chiovenda, Resultati scientifici della Missione Stefanini-Paoli nella Soma-
lia Italiana. Volume I. Le collezioni botaniche. Appendice: Le raccolte di Mangano, Scassellati,
Mazzocchi e Provenzale in Somalia. — Firenze, Pubbl. R. Ist. Stud. Super. Mus. ed Erb. Co-
lon. p. 176—177, 230.

1917. The origin and dispersal of Cocos nucifera. — Philip. J. Sc. 12: Bot. 27—43 (reprinted in
Principes 7: 57—69, 1963).

— A new species of Calamus from Amboina. — Ibid.: 81.

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Dedication

— Ona new South Polynesian palm, with notes on the genus Rhopalostylis Wendl. et Drude.
— Trans. & Proc. New Zeal. Inst. Bot. 49: 47—50.

— Palmae. — In: L.S. Gibbs, Dutch N.W. New Guinea. A contribution to the phytogeography
and flora of the Arfak Mountains, &c. p. 91—98, 200.

— Pelagodoxa henriana Becc. — In: D. Bois, Pelagodoxa henriana. Palmier nouveau des Isles
Marquises. — Rev. Hort. 89: 302—304, f. 76-79.

1918—1921. Asiatic palms — Lepidocaryeae. Part III. The species of the genera Ceratolobus, Ca-
lospatha, Plectocomia, Plectocomiopsis, Myrialepis, Zalacca, Pigafetta, Korthalsia, Metroxy-
lon, Eugeissona. — Ann. R. Bot. Gard. Calc. 12(2) [Fol.]: Letter-press [I—II], 1-231, t. i—vi;
Plates t. 1-120 (1—118+50A, 63A).

1919. Palms of the Philippine Islands, collected and distributed by A.D.E. Elmer. — Elmer,
Leafl. Philip. Bot. 8: 2997—3067.

—— The palms of the Philippine Islands. — Philip. J. Sc. 14: 295—362, t. 1—3.

1920. Palmae novae antillanae. II. — In: Fedde, Repert. Spec. Nov. Regni Veg. 16: 436—437.

— Recensione delle palme del vecchio mondo appartenenti alla tribu delle Corypheae con de-
scrizione delle specie e varieta nuove che vi appartengono. — Firenze, M. Ricci, 10 December
1920 (preprinted with the same pagination from Webbia 5: 5—70, 1921).

— Le palme della Nuova Caledonia. — Firenze, M. Ricci, 128 pp., 13 tav., 10 December 1920
(preprinted with separate pagination from Webbia 5: 71—197, t. 1—13, 1921).

1921. Nelle foreste di Borneo. Viaggi e ricerche di un naturalista. Ed. 2. — Firenze, xvi+ 469 pp.,
72 tav., 11 fig., 4 maps, 1 portr. (published posthumously by Beccari’s son Nello Beccari).
1923. Neue Palmen Papuasiens II. — In: C. Lauterbach, Beitrage zur Flora von Papuasien. X.

— Bot. Jahrb. 58: 441—462.

— Note botaniche e botanico industriali bornensi. — Webbia 5: 451—581, f. 1—6 (reprint of
Appendix, Nelle foreste di Borneo, 1902).

1924. Neue Palmen Mikronesiens. — In: L. Diels, Beitrage zur Flora von Mikronesien und Poly-
nesien. III. — Bot. Jahrb. 59: 11—16.

— Nuova Guinea, Selebes e Molucche; diarii di viaggio ordinati dal figlio Nello Beccari, con
introduzione e note del Prof. Luigi Buscalioni. — Firenze, Soc. Anon. Edit. ‘La Voce’,
XXXViii +468 pp., 53 fig., 22 tav., 1 map (Diaries published by Beccari’s son Nello Beccari).

— Palme della trib’ Borasseae. Ed. U. Martelli, G. Passeri, Firenze, 56 pp., 18 fig., 45 tav.

1927. Odoardo Beccari nel Mar Rosso e tra i Bogos. Frammenti di diario inediti, trascritti e ordi-
nati dal figlio prof. Nello Beccari. — Boll. R. Soc. Geogr. Ital. ser. 6, 4: 625—646.

1930. Odoardo Beccari in Sumatra e la scoperta dell’ ‘Amorphophallus Titanum’. (Frammenti di
diario inediti ed ordinati). — Boll. R. Soc. Geogr. Ital. ser. 6, 7: 569—595, f. 1—6, 1 map. (The
author of the paper is given as Nello Beccari).

1933. Asiatic palms — Corypheae. The species of the genera Corypha, Nannorhops, Sabal, Co-
pernicia, Serenoa, Brahea, Acoelorhaphe, Washingtonia, Pritchardia, Erythea, Livistona, Li-
cuala, Pritchardiopsis, Phlolidocarpus, Teysmannia, Rhaphis, Chamaerops, Trachycarpus,
Rhaphidophyllum, Trithrinax, Acanthorhiza, Hemithrinax, Thrinax, Coccothrinax, Cryso-
phyla. (Posthumous work by Dr. Odoardo Beccari. Revised and edited by Prof. Ugolino Mar-
telli). — Ann. R. Bot. Gard. Calc. 13: Letter-press [Qu.] [i—x], 1—356; Plates [Fol.] t. 1-102
(1—99+ 59 bis, 68 bis, 89 bis), (1931’).

1934. Generi, specie e varieta nuove di palme gerontogee della trib Arecaceae lasciate inedite dal
Dr. Od. Beccari ed ordinate a cura di U. Martelli. — Atti Soc. Tosc. Sc. Nat. Pisa. Mem. 44:
114-176. [repr. pp. 1—65].

1935. (By U. Martelli from Beccari manuscripts). I generi e le specie delle palme gerontogee della

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FLORA MALESIANA

tribu delle Arecaceae. Esposizione geografica secondo la monografia inedita del Dr. O. Beccari.
— Nuovo Giorn. Bot. Ital. ser. 2, 41: 693—723 (‘1934’).

—— (By U. Martelli from Beccari manuscripts). La sinonimia delle palme gerontogee della tribu
delle Areceae. — Ibid. 42: 17-88.

1955. (& R.E.G. Pichi Sermolli) Subfamiliae Arecoidearum palmae gerontogeae. Tribuum et ge-
nerum conspectus. — Webbia 11: 1—187, f. 1—47.

B — Letters by Odoardo Beccari

During his travels BECCARI wrote several letters to his friends in Italy, chiefly to G. Doria, E.H.
GIGLIOLI, G. CorA, T. SALVADORI, R. GESTRO, and O. ANTINORI. They contain a wealth of very
interesting observations and comments on the botanical, zoological, ethnological and other natu-
ralistic aspects of Malesia. These letters or fragments of them were published in various Italian
periodicals, usually accompanied by information and comments on BEccarr’s scientific discover-
ies, and on the itineraries and the main events of his adventurous travels.

The bibliographic citations of the papers in which these letters are published are given below
together with an indication of the name of the friend to whom the letter was addressed, and the
date and place in which it was written. In order to facilitate and render more systematic the consul-
tation of these letters, they are quoted according to the periodicals in which they were published.

Letters published in the Nuova Antologia

BEcCARI’s travels in Malesia, Assab and the country of Bogos between 1865 and 1876 were de-
scribed by Enrico H. GIGLIOLI in various instalments published in the Nuova Antologia with the
general title of ‘Odoardo Beccari ed i suoi viaggi’. They were also reprinted, with independent
pagination, and assembled in a special book (Firenze, Le Monnier, 309 pp., 9 fig., 2 small maps,
1872—76) with the title ‘I viaggi del Dott. Beccari da Firenze tracciati e commentati’. There Gi-
GLIOLI published several letters or fragments of letters by BECCARI to his friends and also some
passages from his original diaries. All of them are quoted verbatim between GIGLIOLI’s descrip-
tions and comments.

Borneo. 1865—1868. — Nuova Antologia 21 (1872) 119—160 (Passages of the original diaries).

Samhara e Bogor. 1870. — Ibid.: 22 (1873) 658—668.

Malesia, Molucche e Papuasia. 1871—72—73. — Ibid.: 22 (1873) 668—709. (Letters to G. Doria,
from the Red Sea, December 10—11, 1871; to E.H. Giglioli, from Batavia, February 8; to G.
Doria, from Makassar, February 23; Ceram, March 13; Ambon, March 21; Kapaor, April 21,
with a small map; to E.H. Giglioli, from Sorong, May 3; to G. Doria, from Sorong, June 21,
1872).

Papuasia: Sorong-Mansinam-Andai (Monti Arfak). (Giugno 1872 al Gennaio 1873). — Ibid.: 23
(1873) 194—225, 2 fig. (Letters to G. Doria, from Andai, August 30; Andai, September 8—9,
1872; Ambon, January 2; Ambon, February 2—7, 1873).

Papuasia — Le isole Aru e Kei (Febbraio—Settembre 1873). — Ibid.: 24 (1873) 835—866, 1 map;
25 (1874) 163—192, 1 fig. (Letters to C. Correnti, from Ambon, January 3; to O. Antinori, from
Ambon, January 3; Ambon, February 5; to G. Doria, from Dobbo (Aru Is.), February 24; Wo-
kan (Aru Is.), March 10; Wokan, July 3—4; Tual (Kei Is.), August 27, 1873).

I. Macassar — Kandari (Celebes). II. I Papua (Dicembre 1873, Giugno 1874). — Ibid.: 27 (1874)
420—463, 5 fig. (Letters to E.H. Giglioli, from Makassar, December 4; to R. Gestro, from Ma-

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Dedication

kassar, December 1; to G. Doria, from Makassar, December 4; Makassar, December 26, 1873;
Makassar, January 15; to T. Salvadori, from Makassar, January 14; to R. Gestro, from Makas-
sar, January 15; to G. Doria, from Kandary, April 23, and May 2 and 11, 1874).

Celebes, Giava, Ternate, Amboina (Giugno 1874—Gennaio 1875). — Ibid.: ser. 2, 2 (1876)
802—822. (Letters to G. Doria, from Kandari, June 4; Makassar, August 30; to O. Antinori,
from Makassar, August 28; to T. Salvadori, from Makassar, August 30; to Baron Podesta, the
Mayor of Genoa, from Makassar, end August; to G. Doria, from Ambon, December 7, 1874;
to E.H. Giglioli, from Ambon, January 7; to ??, from Ambon, January 15, 1875).

Seconda esplorazione della Nuova Guinea. La baia di Geelvink (Febbraio—Novembre 1875). —
Ibid.: ser. 2, 3 (1876) 147-163. (Letters to G. Doria, from Dorei, June 5; Andai, June 15;
Hatam, June 21; to T. Salvadori, from Ternate, August 4, 1875).

Terza esplorazione della Nuova Guinea. La baia di Humboldt (Novembre 1875—Marzo 1876). —
Ibid.: ser. 2, 3 (1876) 333—363, 1 fig. (Letters to G. Cora, from Ternate, March 11; to E.H.
Giglioli, from Ternate, March 6, 1876).

Letters published in the Bollettino della Societa Geografica Italiana

Estratti e frammenti di lettere dirette al marchese Giacomo Doria dal naturalista botanico Odoar-
do Beccari, durante il suo viaggio alla Nuova Guinea. — Boll. Soc. Geogr. Ital. 8 (1872)
148—151. (Letters to G. Doria, from Wahaai, March 13; Ambon, March 21; Sorong, May 3,
1872).

Odoardo Beccari nella Nuova Guinea e nelle isole Aru. — Ibid.: 9 (1873) 145—158. (Letters to
G. Doria, from Andai, August 30, 1872; to C. Correnti, from Ambon, January 3; to O. Antino-
ri, from Ambon, January 4; to G. Doria, from Ambon, February 2, 5, 7; to O. Antinori, from
Ambon, February 5; to G. Doria, from Dobbo, Aru Is., February 24; Wokan, Aru Is., March
10, 1873).

Lettera di O. Beccari al marchese Doria. — Ibid. 10: 4—5 (1873) 66—69. (Letter from Wokan, Aru
Is., July 3, 1873).

Frammento di lettera di O. Beccari a Doria da Makassar, il 18 Novembre 1873. — Ibid.: 10: 6
(1873) 38.

Odoardo Beccari alle isole Key. — Ibid.: 87-89. (Letter to G. Doria, from Tual (Small Kei), Au-
gust 27, 1873).

Ultime notizie di O. Beccari. — Ibid.: 11 (1874) 78-81. (Letters to R. Gestro, from Makassar,
December 1; to G. Doria, from Makassar, December 4; to O. Antinori, from Makassar, Decem-
ber 7; to G. Doria, from Makassar, December 26, 1873).

Corrispondenze del dott. Odoardo Beccari. — Ibid.: 276—282. (Letters to T. Salvadori, from Ma-
kassar, January 14; to R. Gestro, from Makassar, January 15; to G. Doria, from Makassar,
January 15, 1874).

Lettere di Odoardo Beccari. — Ibid.: 480—488, carta della parte sud-est di Celebes. (Letters to
G. Doria, from Kandari, April 23 and June 4, 1874).

Nota sui Papua e sulla Nuova Guinea. — Ibid.: 652—659. (Letter to O. Antinori, from Makassar,
August 28, 1874).

Lettere di Odoardo Beccari. — Ibid.: 660—667. (Letters to the Mayor of Genoa, from Makassar,
without dates; to T. Salvadori, from Makassar, August 30; to G. Doria, from Kandari, June
4, 1874).

Lettera di O. Beccari. — Ibid.: 12 (1875) 117—122. (Letter to G. Doria, from Ambon, January
4, 1875).

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FLORA MALESIANA

La Nuova Guinea Olandese. — Ibid.: ser. 2. 1 (1876) 5SO—557. (Letter to G. Cora, from Ternate,
March 11, 1876).

Letters published in Cosmos

The editor of the journal Cosmos of Turin, Gurpo Cora, gave ample information on BECCARIs
travels in Malesia. He published several letters by BEccarito his friends in Italy, which are listed
below. Furthermore, Cora gave various reports on the different stages of BECCARI’s journeys.
The references to the latter are given in Appendix 3 dealing with BEccarrs itineraries.

Recenti spedizioni alla Nuova Guinea. Spedizione di Beccari e D’Albertis:

1. Da Singapore ad Amboina. — Cosmos | (1873—74) 11—15. (Letters to G. Doria, from Ba-
tavia, February 8; Makassar, February 23, Wahai(N. Ceram), March 13; Amboina, March
2 1872):

2. Da Amboina a Sorong. — Ibid.: 15—20. (Letters to G. Doria, from Kapaor, April 21; So-
rong, May 3 and June 21, 1872).

Recenti spedizioni alla Nuova Guinea. Esplorazioni di Odoardo Beccari:

1. Ricerche geografiche nella Nuova Guinea. — Cosmos 2 (1874—75) 7—9. (Letter to G. Cora,
from Makassar, December 1, 1873).

2. Note sulle Isole Kei. — Ibid.: 9-10. (Letter to G. Cora, from Makassar, December 1,
1873).

3. Da Makassar a Kandari. — Ibid.: 92—96. (Letters to G. Doria, from Kandari, May 2 and
11, 1874).

4. Escursioni intorno a Kandari. Ritorno a Makassar. — Ibid.: 203—207. (Letters to G. Doria,
from Kandari, June 4; Makassar, August 30, 1874).

5. Notizie sull’ornitologia di Celebes. — Ibid.: 207—208. (Letter to T. Salvadori, from Makas-
sar, August 30, 1874).

6. Appunti etnografici sui Papua. — Ibid.: 400—404. (Letter to O. Antinori, from Makassar,
August 28, 1874).

7. Soggiorno a Ternate. Da Ternate ad Amboina. Preparativi pel terzo viaggio alla Nuova
Guinea. — Cosmos 3 (1875—76) 83—88. (Letters to G. Doria, from Amboina, January 4,
8, 9, 1875).

8. Da Amboina a Dorei, per Soron e Wakkaré. Scoperta del fiume Wa Samson. Esplorazione
della baia di Geelvink, determinazione della sua vera ampiezza. — Ibid.: 88—92. (Letter
to G. Doria, from Dorei, June 5, 1875).

9. Esplorazione dei Monti Arfak. Ritorno a Ternate per la via di Salvatti, Batanta, Koffiao.
— Ibid.: 92—95. (Letters to G. Doria, from Andai, June 15; Hatam, June 21; to T. Salva-
dori, from Ternate, August 4, 1875).

10. Viaggio a bordo del trasporto olandese ‘Soerabaja’. Itinerario progettato. Da Ternate a
Dorei, per Salvatti. Visita ad Ansus. Lavori idrografici. — Ibid.: 220—221. (Letters to G.
Doria, from Ternate, November 7; Dorei, November 26, 28, 1875).

11. Viaggio a bordo del trasporto olandese ‘Soerabaja’. Da Dorei alla baia di Humboldt per
la baia Vandamen, l’isola Run, il sud di Jobi, le foci dell’ Ambermo. — Ibid.: 349-352.
(Letter to G. Cora, from Ternate, March 11, 1876).

12. Saggio statistico sulla Nuova Guinea Olandese. Popolazione, Commercio, Climatologia,
Nomenclatura. — Ibid.: 352—360. (Letter to G. Cora, from Ternate, March 11, 1876).

13. Viaggio a bordo del trasporto olandese ‘Soerabaja’. La Baia di Humboldt edi suoi abitanti.

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Dedication

La Baia del Disinganno, il Vulcano Ciclope e le isole Arimoa. — Ibid.: 364—372. (Letter
to E.H. Giglioli, from Ternate, March 6, 1876).

14. Viaggio a bordo del trasporto olandese ‘Soerabaja’. Dalla Baia di Humboldt a Ternate pel
nord di Jobi, Dorei, Waigheu, Misol, il golfo Mac Cluer, la baia Gouns, Ghesser, Amboi-
na. — Ibid.: 372—374. (Letter to E.H. Giglioli, from Ternate, March 6, 1876).

15. Questioni etnologiche sui Papua. — Ibid.: 375—379. (Letter to E.H. Giglioli, from Ternate,
March 6, 1876).

Letters published in various journals

Il viaggio di O. Beccari alla Nuova Guinea. — Nuov. Giorn. Bot. Ital. 4 (1872) 208—212. (Letters
to a friend of Florence, from Wahai, N. Ceram, March 13; Ambon, March 21, 1872).

Il viaggio di O. Beccari alla Nuova Guinea. — Ibid.: 291—294. (Letters to G. Doria, from Kapaor,
April 21; Sorong, June 21, 1872).

Lettera di O. Beccari dalle isole Aru. — Ibid. 5 (1873) 330. (Letter to [G. Doria], July 3—4, 1873).

Brano di lettera di O. Beccari da Makassar in data 1° Dicembre 1873. — Ibid. 6 (1874) 205—206.

Una pianta maravigliosa. — Bull. R. Soc. Tosc. Ortic. 3 (1878) 270—271. (Letter to R. Corsi Sal-
viati, from Sumatra, without date. Comment by E.O. Fenzi).

Lettera ornitologica di O. Beccari intorno agli uccelli osservati durante un recente viaggio alla
Nuova Guinea. — Ann. Mus. Civ. St. Nat. Genova 7 (1875) 704—720. (Letter to T. Salvadori,
from Ternate, August 4, 1875. Introduction by T. Salvadori).

Lettera del Prof. Odoardo Beccari a Giacomo Doria. — Ibid. 13 (1878) 451—455. (From Kajt Ta-
nam, Sumatra, September 8, 1878).

C — Maps

In the period of preparation for his travels BEccarRI carefully trained himself also in geodetics and
topography. He also invented a new instrument (Nuovo orizzonte artificiale) for topographic
survey.

During his explorations BEccarI devoted great attention to the topography of the places he vis-
ited and his surveys allowed him to draw some maps which greatly contributed to the delimitation
of the coasts of certain areas of the Malesian Archipelago.

Some of these maps were published by BEccartr himself, others were utilized by G. Cora, to-
gether with surveys of other explorers, to elaborate some of the maps published in his periodical
Cosmos.

BeEccarr’s itineraries are traced on all maps listed below.

Carta originale del viaggio di Beccari e d’Albertis nella Nuova Guinea ovest (Aprile 1872), costrut-
ta e disegnata da Guido Cora. Scala 1:700.000. — Cosmos (G. Cora) 1 (1873-74) Tav. I. —
Explanatory notes in Cora G. Recenti spedizioni alla Nuova Guinea. Memoria sulla Tavola I.
— Ibid.: 22—24.

Carta originale del viaggio di O. Beccari nel Sud-est di Celebes (Maggio— Agosto 1874), costrutta
e disegnata da Guido Cora. Scala 1:1.200.000. — Ibid. 2 (1874-75) Tav. V. — Explanatory
notes in Cora G. Viaggio di O. Beccari nel Sud-est di Celebes. Note sulla Tavola V. — Ibid.:
201-202.

Carta originale della Nuova Guinea N.O. e delle isole Salvatti, Batanta, William, ecc. secondo
i rilievi originali di Lovera, Cerruti, Beccari e le esplorazioni anteriori costrutta e disegnata da

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FLORA MALESIANA

Guido Cora. Scala 1:1.000.000. — Cosmos (G. Cora) 3 (1875) Tav. II. — Explanatory notes
in Cora G. Recenti spedizioni alla Nuova Guinea. Note sulle Tavole II e III. — Ibid.: 81—83.

Carta originale della Baia di Geelvink e del littorale N.O. della Nuova Guinea secondo i rilievi
di Odoardo Beccari, 1875 del Geelvink, di Duperrey, Dumont D’Urville, ecc. costrutta e dise-
gnata da Guido Cora. Scala 1:1.800.000. — Including an inset map: Tracciato comparativo
della Baia di Geelvink secondo la carta dell’ Amm. Ingl. N° 2759a ed i rilievi di O. Beccari. Scala
1:4.800.000. — Ibid.: Tav. III. — Explanatory notes in Cora G. Recenti spedizioni alla Nuova
Guinea. Note sulle Tavole II e III. — Ibid.: 81—83.

Carta originale della Nuova Guinea Nord dai Monti Arfak alla Baia d’ Humboldt secondo i rilievi
di Odoardo Beccari, 1875—76, della nave oland. ‘Soerabaja’, cap. Swaan e le esplorazioni ante-
riori, costrutta e disegnata da Guido Cora. Scala 1:2.000.000. — Including an inset map: I Mon-
ti Arfak. Scala 1:800.000. — Ibid. 3 (1876) Tav. X. — Explanatory notes in Cora G. Recenti
spedizioni alla Nuova Guinea. Note sulla Tavola X. — Ibid.: 347-349.

Piano della Baia d’ Humboldt (Telokh Lintciu) secondo i rilievi della nave oland.e ‘Etna’ 1858 e
le ricerche di O. Beccari, 1875, di Guido Cora. Scala 1:70.000. — Ibid.: Tav. XI.

Carta originale della Provincia di Sarawak, compilata sopra vari documenti e secondo le osserva-
zioni dell’autore. — In O. Beccari, Nelle foreste di Borneo (1902) fig. 36 (p. 187).

Abbozzo di carta (originale) del fiume Bintulu e suoi affluenti. — Ibid.: fig. 58 (p. 351).

Carta del sistema idrografico e delle attuali divisioni politiche di Borneo, compilata sopra i docu-
menti pili recenti e le osservazioni dell’autore. — Ibid.: fig. 66 (p. 407).

Carta orginale degli itinerari dell’autore in Sarawak. — Ibid.: fig. 75 (facing page 504).

Carta speciale della Nuova Guinea Ovest cogli itinerari di O. Beccari e L.M. D’Albertis
(1872—1876) costrutta e disegnata da Guido Cora. — Including an inset map: Carta originale
del viaggio di O. Beccari nel Sud Est di Celebes. 1874. Disegnata da G. Cora. — In O. Beccari,
Nuova Guinea, Selebes e Molucche (1924) at the end of the book.

Carta delle Isole Aru secondo i rilievi di Odoardo Beccari. Luglio 1873. — In O. Beccari, Lettera
di O. Beccari al Marchese Doria. — Boll. Soc. Geogr. Ital. 10: 4—5 (1873) 64—65, at the end
of the paper.

Carta della Parte Sud-est delle Celebes secondo la relazione di O. Beccari e traccia del suo viaggio
da Makassar a Kandari, Febbraio 1874. — Ibid.: 11 (1874) 480—488, at the end of the paper.

Appendix 2 — Biographies of Odoardo Beccari

BALDASSERONI, V. & D. CaRAzzi. L’opera biologica di Odoardo Beccari. Rassegna Sci. Biol. 3
(1921) 84-88.

BARGAGLI PETRUCCI, G. L’ opera biologica di Odoardo Beccari. Pubbl. Ist. Stud. Sup. Firenze,
Sez. Sci. Fis. Nat. In memoria di Odoardo Beccari, pp. 5—16. 1921.

BeccarI, N. Enciclopedia Italiana di Scienze, Lettere ed Arti. Milano, Ist. G. Treccani, 6 (1930)
462.

—— Brief Obituary, Itineraries and Bibliography — Manuscript compiled by Beccari’s son Nello
for Flora Malesiana, Oct. 1947 (in Library Rijksherbarium, Leiden).

BeEGuINoT, A. Boll. R. Soc. Geogr. Ital. ser. 5, 12 (1923) 194—209, portr.

— Rivista Biol. 3 (1921) 118—121, portr.

BERTACCHI, C. Geografi ed esploratori italiani contemporanei. Milano, De Agostini Ed. 1929, pp.
368—372, portr.

BurkiLL, I.H. & J.C. MouLton. Journ. Str. Br. R. Asiat. Soc. 83 (1921) 166-173.

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Dedication

BUSCALIONI, L. La vita e l’opera di Odoardo Beccari. In: O. Beccari, Nuova Guinea, Selebes e
Molucche. Firenze, 1924, pp. xxiii—Xxxviii.

CavaRA, F. Bull. Soc. Afr. Ital. 40 (1921) 46—S51.

CHIOVENDA, E. Notizie biografiche. Odoardo Beccari. Annuario R. Ist. Stud. Sup. Firenze
1920-21 (1921) 155—156.

— Nuovo Giorn. Bot. Ital. ser. 2, 28 (1921) 5—35, portr., bibl. (in historical sequence).

Cora, G. Recenti spedizioni alla Nuova Guinea. I. Spedizione di Beccari e d’Albertis. (Studi e
viaggi di Beccari dalla sua adolescenza al 1872). Cosmos (G. Cora) 1 (1873-74) 8-11.

Dori, G. I naturalisti italiani alla Nuova Guinea e specialmente delle loro scoperte zoologiche.
Boll. Soc. Geogr. Ital. ser. 2, 3 (1878) 154-169.

Gestro, R. Ann. Mus. Civ. St. Nat. Genova, ser. 3, 9 (1921) 242—297, 11 fig. (10 portr.).

GUBERNATIS, A. DE. Dizionario biografico degli scrittori contemporanei. Firenze, 1879, 1:
117-118, portr.

— Dictionnaire international des écrivains du jour. Florence, 1888, 1: 220—221.

— Dictionnaire international des écrivains du monde latin. Suppl. et Ind. Florence, 1906, 30*.

Jackson, B.D. (‘B.D.J.’). Proc. Linn. Soc. London 134 (1922) 37—40.

LEONE E. DE & M. A.ippI CAPPELLETTI. Dizionario biografico degli Italiani. Roma, 1965, 7:
440-444.

Lo Priore, G. Ber. Deut. Bot. Ges. 39 (1922). Gen. Vers. Heft (56)—(87), portr. bibl. (alphabeti-
cal according to titles).

MarRTELLI, U. Webbia 5 (1921) 295-353, portr., 3 maps, bibl. (alphabetical according to titles).

Moore Jr., H.E. Odoardo Beccari (1843—1920). Principes 25 (1981) 29—35, portr., bibl. (on
palms).

NissEN, CL. Die botanische Buchillustration. 2 (1951) 11 (on his drawings).

PALADINI, C. La morte di Odoardo Beccari. ‘Il Nuovo Giornale’ 27 Ottobre 1920.

PAMPANINI, R. Agricoltura Coloniale 14 (1920) 449—453.

Po.iaccl, G. Atti Ist. Bot. Univ. Pavia. ser. 6, 6 (1935), i—xiii, bibl. (in historical sequence).

Puccion1, N. L’opera etnografica ed antropologica di Odoardo Beccari. Pubbl. Ist. Stud. Sup.
Firenze, Sez. Sci. Fis. Nat. In memoria di Odoardo Beccari. pp. 17—26, 1921.

Ropotico, F. Naturalisti esploratori dell’Ottocento Italiano. Firenze, Le Monnier, 1967, pp.
191—222, t. 4 (with reprint of original passages).

Roster, G. Bull. R. Soc. Tosc. Ortic. 46 (1921) 33—36, portr.

Saccarbo, P.A. La Botanica in Italia. Materiali per la storia di questa scienza. Mem. Ist. Veneto
Sci. Lett. Arti. 25:4 (1895) 25; 26:6 (1901) 16.

S.A.S. Kew Bull. (1920) 369—370.

STEENIS, C.G.G.J. vAN. Thesaurus Beccarianus. Webbia 8 (1952) 427—436.

STEENIS-KRUSEMAN, M.J. vAN. Flora Malesiana I, 1 (1950) 43—46, portr.

—— Flora Malesiana I, 8 (1974) xiii (brief addition).

Wittrock, V.B. Catalogus illustratus Iconothecae Botanicae Horti Bergiani Stockholmiensis.
Acta Horti Berg. 3:2 (1903) 160, t. 31 (portr.); 3:3 (1905) 175.

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FLORA MALESIANA

Appendix 3 — Accounts of Odoardo Beccari’s itineraries
in Malesia and Ethiopia
(see also Letters and Maps)

Beccarl, N. Brief Obituary, Itineraries and Bibliography. — Manuscript compiled by Beccari’s
son Nello for Flora Malesiana, Oct. 1947. (in Library Rijksherbarium, Leiden).

Burtt, B.L. Beccari’s ascent of ‘Mount Poi’. Flora Males. Bull. 19 (1964) 1131—1132. (Pointing
out that Mt Poi (Poe, Pueh) in Southwest Sarawak, mentioned in the ‘Wanderings’, a famous
type locality, is not what is nowadays on maps indicated as Mt Poi, but a more southeasterly
peak in the Poi Range, now known as Gunung Berumput, or Gunung Rumput).

Cora, G. Spedizione italiana alla Nuova Guinea. Roma, Stab. Civelli, 1872, 39 pp.

— Recenti spedizioni alla Nuova Guinea. — Cosmos (G. Cora) 1 (1873-74) 7-8.

— Recenti spedizioni alla Nuova Guinea. Spedizione di Beccari e D’Albertis. — Ibid.: 8-11,
141-143.

— Recenti spedizioni alla Nuova Guinea. Odoardo Beccari. — Ibid.: 215-218, 265; 2
(1874-75) 2—4, 86; 3 (1875-76) 75—76.

— Viaggio di O. Beccari nel Sud-est di Celebes. — Ibid.: 2 (1874—75) 200—202, t. V (map).

— Recenti spedizioni alla Nuova Guinea. Secondo viaggio della ‘Vettor Pisani’. — Ibid.: 3
(1875—76) 77-78.

— Recenti spedizioni alla Nuova Guinea. Quarto viaggio di O. Beccari alla Nuova Guinea
(1875—1876) — Ibid.: 217.

— Recenti spedizioni alla Nuova Guinea. I Monti Arfak. — Ibid.: 217-218.

— Carta speciale della Nuova Guinea Ovest cogl’itinerari di O. Beccari e L.M. D’Albertis
(1872—1876) costrutta e disegnata da Guido Cora. — Ann. Mus. Civ. St. Nat. Genova 12
(1878).

— (see also Appendix 1 — Letters by O. Beccari). Recenti spedizioni alla Nuova Guinea. —
Cosmos 1 (1873—74) 11—15, 15—20; 2 (1874-75) 7-9, 9-10, 92—96, 203—207, 207—208,
400—404; 3 (1875-76) 83-88, 88-92, 92-95, 220-221, 349-352, 352-360, 364-372,
372—374, 375-379.

Gestro, R. Ricordo biografico di Giacomo Doria. Ann. Mus. Civ. St. Nat. Genova, ser. 3, 10
(1921) 1—78, portr. (information on Beccari’s travels).

GiGuio1!, E.H. (see also Appendix 1 — Letters by O. Beccari). Odoardo Beccari ed i suoi viaggi.
— Nuova Antologia 21 (1872) 119-160; 22 (1873) 658—668; 668—709; 23 (1873) 194—225; 24
(1873) 835—866; 25 (1874) 163-192; 27 (1874) 420—463; ser. 2, 2 (1876) 802—822; 3 (1876)
147-163, 333-363.

Maaistris, L.F. DE. Biografie di Geografi e di Esploratori contemporanei. IV. Giacomo Doria.
Novara, Ist. Geogr. De Agostini, 1917, 18 pp., portr. (information on Beccari’s travels).

MaRTELLI, U. Odoardo Beccari. Webbia 5 (1921) 295—353, 3 maps.

STEENIS-KRUSEMAN, M.J. vAN. Flora Malesiana I, 1 (1950) 43—45.

Vink, W. Nova Guinea, Bot. n. 22 (1965) 479—481, f. 6. (on itinerary in New Guinea).

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Dedication

Appendix 4 — Studies based on Odoardo Beccari’s botanical collections
(incomplete)

ARCANGELI, G. L’Amorphophallus titanum Beccari, illustrato da G. Arcangeli. — Nuovo Giorn.
Bot. Ital. 11 (1879) 217—223, cum fig.

ASCHERSON, P. Plantae phanerogamae marinae, quas Cl. Eduardus Beccari in Archipelago Indico
annis 1866 et 1867, et in Mari Rubro anno 1870 collegit, enumeratae. — Nuovo Giorn. Bot.
Ital. 3 (1871) 299-302.

BAGLIETTO, F. Lichenes in regione Bogos Abissiniae septentrionalis lecti ab O. Beccari; illustravit
F. Baglietto. — Nuovo Giorn. Bot. Ital. 7 (1875) 239-254.

BaILLon, H. Stirpes exoticae novae. — Adansonia 11 (1873—76) and 12 (1876—79). — Referring
to several Bornean plants collected by Beccari.

BAKER, J.G. On a collection of ferns made by Dr. Beccari in western Sumatra. — Journ. Bot.
18 (1880) 209-217.

BARGAGLI-PETRUCCI, G. Sulla struttura dei legnami raccolti in Borneo dal dott. O. Beccari. —
Malpighia 17 (1903) 280—371, t. 4—15.

CESATI, V. Felci e specie nei gruppi affini raccolte a Borneo dal Signor Odoardo Beccari. — Atti
Accad. Sci. Fis. Mat. Napoli 7:8 (1876) 1—37, t. 1-4.

—— Prospetto delle Felci raccolte dal Sign. O. Beccari nella Polinesia, durante il suo secondo
viaggio d’esplorazione in quei mari. — Rend. Accad. Sci. Fis. Mat. Napoli 16 (1877) 23—31.

— Mycetum in itinere Borneensi lectorum a cl. Od. Beccari enumeratio. — Atti Accad. Sci. Fis.
Mat. Napoli 8:3 (1879) 1—28, t. 1—4.

CHRISTENSEN, C. Revision of the Bornean and New Guinean ferns collected by O. Beccari and
described by V. Cesati & J.G. Baker. — Dansk Bot. Arkiv 9 (1937) 33-52.

DuBarpD, M. Description de quelques types nouveaux ou peu connus de Sapotacées (Illipées)
d’aprés les documents de L. Pierre. — Bull. Mus. Hist. Nat. Paris 14 (1908) 405—409.

ENGLER, A. Araceae, specialmente Bornensi e Papuane raccolte da O. Beccari. — Bull. R. Soc.
Tosc. Ortic. 4 (1879) 265—271; 295—302. (with an introduction, pp. 265—266, by O. Beccari).

—— Araceae della Malesia e della Papuasia raccolte da Beccari, determinate ed illustrate. —
Malesia 1 (1883) 259—304, t. 16—28.

GEHEEB, A. Weitere Beitrage zur Moosflora von Neu Guinea. I. Uber die Laubmoose, welche Dr.
O. Beccari in den Jahren 1872—73 und 1875 auf Neu Guinea, besonders dem Arfak-Gebirge,
sammelte. II. Uber einige Moose von westlichen Borneo. — Bibl. Bot. Heft 44 (1898) 29 pp.,
Qetave

— Musci frondosi in monte Pangerango insulae Javae a Dr. O. Beccari annis 1872 et 1874 lecti.
— Rev. Bryol. 21 (1894) 81-85.

Hampg, E. Musci frondosi in insulis Ceylon et Borneo a Dr. Od. Beccari lecti. — Nuovo Giorn.
Bot. Ital. 4 (1872) 273-291.

Hem, F. Diptérocarpacées de Borneo. — Bull. Mens. Soc. Linn. Paris 120 (1891) 954—958; 122
(1891) 970—976.

— Deux Richetia nouveaux. — Bull. Mens. Soc. Linn. Paris 123 (1891) 979-981.

IRMSCHER, E. Neue Begoniaceen von O. Beccari in Malesien gesammelt. — Webbia 9 (1954)
469—509, f. 1-8.

KRANZLIN, F. On orchids collected by Beccari, in Pflanzenreich; corrections on it by R. Schlechter
in Fedde, Repert. 9 (1911) 286—287.

KREMPELHUBER, A. VON. Lichenes foliicoli quos legit O. Beccari annis 1866—1867 in insula Bor-
neo. — Miinchen, 1874.

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FLORA MALESIANA

—— Lichenes quos legit O. Beccari in insulis Borneo et Singapore annis 1866 et 1867. — Nuovo
Giorn. Bot. Ital. 7 (1875) 5—67, t. 1-2.

MaRTELLI, U. Le Composte raccolte dal Dottore O. Beccari nell’ Arcipelago Malese e nella Papua-
sia. — Nuovo Giorn. Bot. Ital. 15 (1883) 281—305.

— Le Dilleniaceae Malesi e Papuane delle collezioni Beccari. — Malesia 3 (1886) 150—167.

— Florula Bogosensis. Enumerazione delle piante dei Bogos raccolte dal Dott. O. Beccari
nell’anno 1870, con descrizione delle specie nuovo 0 poco note. — Firenze, 1886. vii+ 169 pp.,
1 tav.

— Contribuzione alla flora di Massaua. — Nuovo Giorn. Bot. Ital. 20 (1888) 359—371.

MERRILL, E.D. A brief survey of the present status of Bornean botany. — Webbia 7 (1950)
309-324.

Notaris, C. DE. Epatiche di Borneo raccolte dal Dre O. Beccari nel Ragiato di Sarawak durante
gli anni 1865—66—67. — Torino, Stamperia Reale Paravia, 1874, 44 pp., 35 tav. (preprinted
from Mem. Acad. Sci. Torino 18 (1876) 267—308, t. 1—35; reprinted also in Nuovo Giorn. Bot.
Ital. 8 (1876) 217-251).

PASSERINI, G. Funghi raccolti in Abissinia dal Signor O. Beccari. — Nuovo Giorn. Bot. Ital. 7
(1875) 180—192, t. 4—S.

PicHi SERMOLLI, R. Rapporti tra parassita ed ospite nella Rafflesia tuan-mudae Becc. e Cissus sp.
— Nuovo Giorn. Bot. Ital. ser. 2, 44 (1937) 385—421, t. 8—10, f. 1-16.

REICHENBACH, H.G. Odoardo Beccari novitiae orchidaceae papuanae describuntur. — Bot.
Centr. Bl. 28 (1886) 343—346.

SCHLECHTER, R. Orchidaceae novae Beccarianae. — Notizbl. Berl. Dahl. 8 (1921) 14—20.

SCHUMANN, K. Sterculiaceae Beccarianae. — Bot. Jahrb. 24 (1897), Beibl. 58: 14—21.

Soitms-LAUBACH, H. Uber die von Beccari auf seiner Reise nach Celebes und Neu Guinea gesam-
melten Pandanaceae. — Ann. Jard. Bot. Buitenzorg 3 (1883) 89—104, t. 16.

STEENIS, C.G.G.J. vAN. Thesaurus Beccarianus (including ‘Malaysian sea-grasses collected by
Beccari’ and ‘Some records of Malaysian plants’). Webbia 8 (1952) 427—436.

— & M.J. vAN STEENIS-KRUSEMAN. Numberlists of Beccari: Sumatra, Borneo, New Guinea.
1951—1953. Typed copies in Leiden, Bogor, Florence and the Arnold Arboretum, the Leiden
copy kept up to date by the first author. Piante Sumatrane, PS 1—979, Piante Borneensi, PB
1—4069, Piante Papuane, PP 1—986. These are his collecting numbers; they do not correspond
with the herbarium ‘sheet numbers’ as given in the ‘Herbarium Beccarianum’ at Florence.

STEENIS-KRUSEMAN, M.J. VAN. Flora Malesiana I, 5 (1958) cclvii. — Explaining that Beccari later
renumbered his collections originally named ‘Plantae Beccarianae’ in three series, each starting
with number 1, viz the PS series (Piante Sumatrane), PB (Piante Borneensi) and PP (Piante Pa-
puane).

VENTURI, G. DE. Muschi raccolti dal Signor Odoardo Beccari nella terra dei Bogos in Abissinia.
— Nuovo Giorn. Bot. Ital. 4 (1872) 7—22.

ZANARDINI, J. Phycearum indicarum pugillus, a cl. Eduardo Beccari ad Borneum, Sincapore et
Ceylanum annis MDCCCLXV—VI-—VII collectarum. — Mem. R. Ist. Veneto Sci. Lett. Arti
17 (1872) 109—170, t. 1-12.

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ABBREVIATIONS AND SIGNS

acc. = according

Ak. Bis. = Aklan Bisaya (Philip. language)
Alf. Cel. = Alfurese Celebes (language)
alt. = altitude

Anat. = Anatomy

Ap. = Apayao (Philip. language)

app. =appendix, appendices

appr. = approximate

Apr. = April
Arch. = Archipelago
atl. = atlas

auct. div. =auctores diversi; various authors

auct(t). mal. = auctores malayenses; authors dealing
with Malesian flora

auct(t). plur. = auctores plures; several authors

Aug. = August

Bag. = Bagobo (Philip. language)

basionym = original name of the type specimen; its
epithet remains permanently attached to the taxon
which is typified by it provided it is of the same
rank.

Bg. = Buginese (language)

Bik. = Bikol (Philip. language)

Bil. = Bila-an (Philip. language)

Bill. = Billiton

Bis. = Bisaya (Philip. language)

Bon. = Bontok (Philip. language)

Born. = Borneo

Bt= Bukit; mountain

Bug. = Buginese (language)

Buk. = Bukidnon (Philip. language)

c. =circiter; about

C. Bis. = Cebu Bisaya (Philip. language)

cf. =confer; compare

Chab. = Chabecano (Philip. language)

citations = see references

cm = centimetre

c.n. =see comb. nov.

comb. nov. =combinatio nova; new combination

CS = cross-section or transversal section of an organ

c.s. =cum suis; with collaboration

cum fig. =including the figure

cur. = curante; edited by

D (after a vernacular name) = Dutch

Daj. = Dyak (language)

d.b.h. =diameter at breast height

D.E.I. = Dutch East Indies

descr. added behind a reference=means that this
contains a valid description

diam. = diameter

Distr. (as an item) = Distribution

Distr. (with a geographical name) = District

ditto =the same, see do

Div. = Division, or Divide

div. = diversus (masc.); various

do = ditto (Ital.); the same

Dum. = Dumagat (Philip. language)

dupl. = duplicate

E=east (after degrees: eastern longitude)

E (after a vernacular name) = English

Ecol. = Ecology

ed. = edited; edition; editor

e.g. =exempli gratia; for example

elab. = elaboravit; revised

em(end). = emendavit; emended

em(erg). ed. =emergency edition

Engl. = English

etc., &c. =et cetera; and (the) other things

ex auctt. =ex auctores; according to authors

excl. = exclusus (masc.); excluding, exclusive of

ex descr. =known to the author only from the de-
scription

f. (before a plant name) = forma; form

f. (after a personal name) = filius; the son

f. (in citations) = figure

fam. = family

Feb(r). = February

fide =according to

fig. = figure

fl. =flore, floret (floruit); (with) flower, flowering

For. Serv. = Forest Service

fr. =fructu, fructescit; (with) fruit, fruiting

Fr. (after a vernacular name) = French

G.= Gunung (Malay); mountain

Gad. = Gaddang (Philip. language)

gen. = genus; genus

genus delendum = genus to be rejected

Germ. = German

geront. =Old World

haud = not, not at all

holotype=the specimen on which the original de-
scription was actually based or so designated by the
original author

homonym =a name which duplicates the name of an
earlier described taxon (of the same rank) but
which is based on a different type species or type
specimen; all later homonyms are nomenclaturally
illegitimate, unless conserved

I. = Island

ib(id). =ibidem; the same, in the same place

Ibn. = Ibanag (Philip. language)

ic. =icon, icones; plate, plates

ic. inedit.=icon ineditum, icones inedita; inedited
plate(s)

id. =idem; the same

i.e. =id est; that is

If. =Ifugao (Philip. language)

Ig. =Igorot (Philip. language)

Ilg. =Ilongot (Philip. language)

Ilk. = Iloko (Philip. language)

in adnot. =in adnotatione; in note, in annotation

incl. = inclusus (masc.); including, inclusive(ly)

indet. = indetermined

Indr. = Indragiri (in Central Sumatra)

inedit. =ineditus (masc.); inedited

in herb. =in herbario; in the herbarium

in litt. =in litteris; communicated by letter

in sched. =in schedula; on a herbarium sheet

in sicc. =in sicco; in a dried state

in syn. =in synonymis; in synonymy

Is. = Islands

Is. (after a vernacular name) =Isinai (Philip. lan-
guage)

Ism. = Isamal (Philip. language)

isotype=a duplicate of the holotype; in arboreous
plants isotypes have often been collected from a
single tree, shrub, or liana from which the holotype
was also derived

Iv. =Ivatan (Philip. language)

J(av). = Javanese (language)

Jan. = January

Jr = Junior

Klg. = Kalinga (Philip. language)

Kul. =Kulaman (Philip. language)

Kuy. = Kuyonon (Philip. language)

Lamp. = Lampong Districts (in S. Sumatra)

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FLORA MALESIANA

Lan. = Lanao (Philip. language)

lang. = language

l.c. =loco citato; compare reference

lectotype=the specimen selected a posteriori from
the authentic elements on which the taxon was
based when no holotype was designated or when
the holotype is lost

livr. =livraison, part

IKeex= ex (plurs)

LS = longitudinal or lengthwise section of an organ

m= metre

M = Malay (language)

Mag. = Magindando (Philip. language)

Mak. = Makassar, Macassar (in SW. Celebes)

Mal. = Malay(an)

Mal. Pen. = Malay Peninsula

Mand. = Mandaya (Philip. language)

Mang. = Mangyan (Philip. language)

Mar. = March

Mbo = Manobo (Philip. language)

Md. = Madurese (language)

Minangk. = Minangkabau (a Sumatran language)

min. part. =pro minore parte; for the smaller part

mm = milimetre

Mng. = Mangguangan (Philip. language)

Morph. = Morphology

ms(c), MS(S) = manuscript(s)

Mt(s) = Mount(ains)

n. =numero; number

N=North (after degrees: northern latitude); or New
(e.g. in N. Guinea)

NE. = northeast

nec=not

neerl. = Netherlands, Netherlands edition

Neg. = Negrito (Philip. language)

N.E.I. = Netherlands East Indies

neotype=the specimen designated to serve as no-
menclatural type when no authentic specimens
have existed or when they have been lost; a neotype
retains its status as the new type as long as no auth-
entic elements are recovered and as long as it can
be shown to be satisfactory in accordance with the
original description or figure of the taxon

N.G. = New Guinea

N.I. = Netherlands Indies

no =numero; number

nom. = nomen; name (only)=nomen nudum

nom. al.=nomen aliorum; name used by other
authors

nom. alt(ern).=nomen_ alternativum;
name

nom. cons(erv).=nomen conservandum, nomina
conservanda; generic name(s) conserved by the In-
ternational Rules of Botanical Nomenclature

nom. fam. cons.=nomen familiarum conservan-
dum; conserved family name

nom. gen. cons. =see nomen conservandum

nom. gen. cons. prop.=nomen genericum conser-
vandum propositum; generic name proposed for
conservation

nom. _ illeg(it). =nomen
name

nom. leg(it). =nomen legitimum; legitimate name

nom. nov. =nomen novum; new name

nom, nud. = nomen nudum; name published without
description and without reference to previous pub-
lications

alternative

illegitimum; illegitimate

(46)

nom. rej(ic.) = nomen rejiciendum; name rejected by
the International Rules of Botanical Nomenclature

nom. seminudum=a name which is provided with
some unessential notes or details which cannot be
considered to represent a sufficient description
which is, according to the International Rules of
Botanical Nomenclature, compulsory for valid
publication of the name of a taxon

nom. subnudum = nomen seminudum

nom. superfl. =a name superfluous when it was pub-
lished; in most cases it is a name based on the same
type as an other earlier specific name

non followed by author’s name and year, not placed
in parentheses, and put at the end of a citation =
means that this author has published the same
name mentioned in the citation independently.
These names (combinations) are therefore homo-
nyms.

Compare 56b line 5—4 from bottom. The same can
happen with generic names.

(non followed by abbreviation of author’s name) be-
fore a reference (citation) headed by an other
author’s name = means that the second author has
misinterpreted the taxon of the first author.
Compare p. 419a under species 47 the synonym H.
celebica. DIELS misapplied the name H. celebica as
earlier described by BURCK.

non al. =non aliorum; not of other authors

non vidi=not seen by the author

nov. =nova (femin.); new (species, variety, efc.)

Nov. = November

n.s. =new series

n. sp. =nova species; new species

n. (sp.) prov.=nomen (specificum) provisorium;
provisional new (specific) name

n.v. =non vidi; not seen

NW. =northwest

Oct. = October

op.cit. = opere citato; in the work cited

Pp. =pagina; page

P.=Pulau, Pulu (in Malay); Island

Pal(emb.) = Palembang

Pamp. = Pampangan (Philip. language)

Pang. = Pangasinan (Philip. language)

paratype = a specimen cited with the original descrip-
tion other than the holotype

part. alt. =for the other part

P. Bis. = Panay Bisaya (Philip. language)

P.I. = Philippine Islands

pl. =plate

plurim., = plurimus; most

D.p. = pro parte; partly

pr. max. p. = pro maxima parte; for the greater part

pro=as far as is concerned

prob. = probabiliter; probably

prop. = propositus; proposed

Prov. = Province

pr.p. = pro parte; partly

pt = part

quae est =which is

quoad basionym, syn., specimina, efc. =as far as the
basionym, synonym(s), specimen(s), efc. are con-
cerned

references =see for abbreviations the list in vol. 5,
pp. cxlv—clxv

Res. = Residency or Reserve

resp. = respective(ly)

Abbreviations and signs

S=south (after degrees: southern latitude)

S (after a vernacular name) = Sundanese (language)

Sbl. =Sambali (Philip. language)

SE. = southeast

sec. =secus; according to

sect. =sectio; section

sens. ampl. (ampliss.)=sensu amplo (amplissimo);
in a wider sense, in the widest sense

sens. lat. =sensu lato; in a wide sense

sens. str. (Sstrictiss.)=sensu stricto (strictissimo); in
the narrow sense, in the narrowest sense

Sept. = September

seq., seqq. =sequens, sequentia; the following

ser. = series

s.1. =sensu lato; in a wide sense

S.-L. Bis. =Samar-Leyte Bisaya (Philip. language)

Sml. = Samal (Philip. language)

s.n.=sine numero; (specimen) without the collec-
tor’s number

Sp. =Spanish (language)

sp(ec). = species; species

specim. = specimen(s)

sphalm. = sphalmate; by error, erroneous

Spp. = species; species (plural)

Sr = Senior

S.S. =see sens. Str.

ssp. = subspecies; subspecies

s.str. =see sens. Str.

stat. nov. =Status nova; proposed in a new rank

Sub. = Subanum (Philip. language)

subg(en). =subgenus; subgenus

subsect. = subsectio; subsection

subsp. = subspecies; subspecies

Sul. = Sulu (Philip. language)

Sum. E.C. = Sumatra East Coast

Sum. W.C.=Sumatra West Coast

Suppl. = Supplement

SW. =southwest

syn. =synonymum; synonym

synonyms =the names of taxa which have been re-
ferred to an earlier described taxon of the same
rank and with which they have been united on
taxonomical grounds or which are bound together
nomenclaturally

syntypes = the specimens used by the original author
when no holotype was designed or more specimens
were simultaneously designated as type

t. = tabula; plate

Tag. = Tagalog (Philip. language)

Tagb. = Tagbanua (Philip. language)

Tagk. = Tagaka-olo (Philip. language)

Tapan. = Tapanuli (in NW. Sumatra)

taxon=each entity throughout the hierarchic ranks
of the plant kingdom which can be described and
discriminated from other taxa of the same rank

Taxon. = Taxonomy

Tg =Tandjung (Malay); cape

Ting. = Tinggian (Philip. language)

Tir. = Tirurai (Philip. language)

transl. = translated

type = each taxon above the rank of a species is typi-
fied by a type belonging to a lower rank, for in-
stance a family by a genus, a genus in its turn by
a species; a species or infraspecific taxon is typified
by aspecimen. The name of a taxon is nomenclatu-
rally permanently attached to its type; from this it
cannot be inferred that the type always represents
botanically the most typical or average structure
found in the circumscription of the taxon.

type specimen=the specimen or other element to
which the name of a species or infraspecific taxon
is (nomenclaturally) permanently attached; botan-
ically a type specimen is a random specimen on
which the name was based by description. There-
fore, it does not need to represent the average or
most typical representative of a population. See
holotype, isotype, lectotype, syntype, paratype,
and neotype

typ. excl. = typo excluso; type excluded

typ. incl. =typo incluso; type included

typus =see type and type specimen

var. = varietas; variety

var. nov. =varietas nova; new variety

Vern. = Vernacular

vide = see

viz. = videlicet; namely

vol. = volume

W =west (after degrees: western longitude)

Yak. = Yakan (Philip. language)

+ =about

) = male (flower, efc.)
2 =female (flower, efc.)

(

)=monoecious with unisexual flowers

)= polygamous

) = polygamous

c’= many

> =more than (in size, number, efc.)

< =less than (size, number, efc.)

xX 2/5=2/5 of natural size

Xx montana= means that the epithet montana is that
of a hybrid

(47)

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Pils @! Eiicwws! “am: cure we <p fp a inner
nat s* AE® Myre 7] mm Pat / i

shoe Sgenve wt Pe i) Pe -—--
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vrs

ARALIACEAE—I (W. R. Philipson, Christchurch)!

Trees, shrubs, lianas, woody epiphytes or (extra-Mal.) more rarely herbs. Branches
usually stout with leaves clustered at their ends; armed or unarmed; glabrous or
with a tomentum of stellate or simple hairs; buds either covered by the stipular
sheaths of leaves or by cataphylls. Leaves spiral or rarely opposite or in whorls;
petiole usually clasping the stem; stipules either distinct or united into a ligule or
absent (in Osmoxylon the petiole bears + elaborate crests around its base); lamina
digitately compound or pinnate, sometimes to the second or third degree, or simple,
when either entire or pinnately or palmately lobed, margin entire or dentate.
Inflorescence terminal or more rarely lateral; either simple or compound racemes or
spikes, or more commonly of umbels or capitula, either solitary or arranged in
compound umbels or panicles; bracts usually small and caducous; pedicel either
articulated with the flower or continuous with it. Flowers hermaphrodite or hetero-
sexual, sometimes dioecious; actinomorphic. Calyx lobes small, or reduced to a
rim, or rarely absent. Petals 3 to numerous, often 5, sometimes fused into a calyptra,
or forming a tube with spreading lobes (Osmoxylon), valvate or imbricate in bud,
usually with a broad base but rarely narrowed below. Stamens usually as many as
the petals and alternating with them, or twice as many, or indefinite; filaments
inserted at the edge of the disk; anthers dorsifixed, introrse, pollen sacs 4 or rarely 8.
Ovary inferior, half inferior, or very rarely (extra-Mal.) superior, 1- to many-celled,
the top of the ovary usually a fleshy disk; styles and stigmas as many as the cells,
either connate or wholly or partially free. Ovules solitary, pendulous, anatropous,
with the raphe ventral. Fruit baccate or drupaceous, exocarp usually fleshy,
endocarp forming cartilaginous or membranaceous pyrenes around the seeds.
Seeds one per pyrene, with a small embryo within smooth or ruminate endosperm.

Distribution. About 50 genera with a roughly estimated 1150 species, ranging mainly in the
warmer parts of both hemispheres (especially in montane zones), a small number in or extending
to cool-temperate regions. With the exception of SE. Asia, the family and its centres of distribu-
tion are largely found within the land masses derived from ancient Gondwanaland. In Malesia
17 genera with a total (excluding Schefflera) of 117 species in 16 genera. (The largest genus,
Schefflera, with an estimated 250 species for the region, is omitted from this account.)

Three genera are endemic to Malesia (or nearly so). One of these, Anakasia (related to
Polyscias) is of very local distribution in West New Guinea; the two others, Harmsiopanax (Java,
Lesser Sunda Is., Celebes, to New Guinea) and Aralidium (Malay Peninsula, Sumatra and
Borneo, with an extension to Thailand) are more widely distributed.

Five other genera have their main centres of distribution within Malesia. Two of these extend
further eastwards: Mackinlaya ranges from the Philippines and Celebes through New Guinea
to the Solomon Is. and NE. Australia; Osmoxylon occurs from Botel Tobago (Taiwan) and the
Marianas to Borneo, the Philippines, Celebes, the Moluccas, New Guinea to the Solomon Is. and
the New Hebrides. A third, Trevesia, is confined to western Malesia (Lesser Sunda Is., Java,
Borneo, Sumatra, and Malaya), with an extension into the Asian mainland. Wider ranges are
recorded for Gastonia (widespread in Malesia to the Solomon Is.) with an additional range in the
Seychelles, Mascarenes, and Madagascar (but not East Africa as previously reported) and
Arthrophyllum (all over Malesia with extensions to Thailand, Laos, the Nicobar Islands, and
New Caledonia.

A significant element in the Malesian representation of the family consists of six genera which
occur mainly in SE.-E. Asia, two of which further extending to the Americas, viz Aralia (in

(1) The genus Scheffiera is omitted and will be treated separately by Dr D. G. Frodin (University of
Papua New Guinea). I enjoyed his assistance in drawing the general chapters.

(1)

9) FLORA MALESIANA [ser. I, vol. 9!

America south to Mexico) throughout Malesia, and Dendropanax (tropical America) in West
Malesia (Sumatra, Malay Peninsula, Borneo). The remaining four genera range in Malesia as
follows: Macropanax and Brassaiopsis in Malay Peninsula, Sumatra and Java, Acanthopanax in
Malay Peninsula, Sumatra and the Philippines, and Pentapanax only in East Java.

Only one genus, Delarbrea, has its main centre of distribution to the east of Malesia (Melanesia,
mostly New Caledonia) and Queensland, and extends through New Guinea to the Moluccas and
Lesser Sunda Is. as far west as Timor.

Two other genera have wider distributions. Schefflera, including several segregate genera, is
pantropical although with but few representatives (13) on the African mainland. However, the
several sections of Schefflera as redefined by FRODIN (1970, 1975) often show distinctive regional
distributions paralleling those of several of the other genera. The type section ranges from the
New Hebrides to Samoa and New Zealand, including Fiji and New Caledonia.

Polyscias is widely dispersed in the Old World tropics from the African mainland eastwards to
the Society Is. and Australia, but with only a few species in SE. Asia and western Malesia. As in
Schefflera, the diverse series of species comprising the genus show distinctive regional distribution
patterns, with the type series being mainly Melanesian and East Malesian (in West Malesia
it is only cultivated or adventive).

As said above several Malesian genera extend into, or have their main centres in, mainland
Asia, but only Polyscias (in Sri Lanka) and Schefflera are shared with Africa and the same genera
(with Gastonia) with Madagascar. Only Polyscias and Gastonia occur in the Mascarene Islands
and Gastonia and Schefflera in the Seychelles. Two other genera are restricted to mainland Africa.
Hedera of temperate Eurasia extends to the Canary Islands. The Americas have two (or three)
distinctive endemic genera with recognizable affinities, as well as disjunct groups of the Austral-
asian Pseudopanax, the Asiatic Pentapanax and Dendropanax and (in North America) Oplopanax,
Aralia and Panax, all very closely related to those in East Asia; there are also distinctive sections
of Scheffiera in the neotropics which are nearer those in Africa than in Asia. Many distinctive
endemic genera (or parts of the larger genera) occur in Oceania, the New Zealand region and
Australia, some of them taxonomically very isolated.

The ratio of species : genus is nere estimated as 23 : 1, but if the very large genus Schefflera is
not accounted for this reduces to 13 : 1. Some 30 genera have five or fewer species; in view of the
considerable insular endemism at generic or infrageneric level, weak intercontinental links, and
great distinctiveness of many genera (even though small), the family is surely of great antiquity,
although much of the available palaeobotanical evidence requires re-evaluation (cf. DILCHER &
Do pn, 1970). Many fossils previously referred to Oreopanax must now be placed in Platanaceae
(DoyLeE, pers. comm.). Dendropanax has reliably been reported from Tertiary deposits in both
Europe and North America where it is now absent (DILCHER & DOLPH, /.c.).

References: DiLCHER & DoLpH, Amer. J. Bot. 57 (1970) 153-160; FRopIN, The complex of
Cephaloschefflera in Schefflera (Araliaceae), Thesis, Cambridge, U.K. (1970); J. Arn. Arb. 56
(1975) 427-448.

Ecology. Malesian Araliaceae are usually small trees, shrubs, or lianas, with a number being
sometimes or always epiphytic (especially in Schefflera), and where terrestrial usually in the under-
growth or lower stories of rain-forest, seldom reaching over 20 m. A remarkable exception is
provided by the two species of Gastonia; of these G. spectabilis (HARMS) PHILIPSON of Papuasia
can attain the great height of 40 m with a stem of 1.75 m @;; it is the largest araliad known and is
of very striking appearance. Fig. 27.

Araliads are almost always found scattered in forest and other vegetation, at least in Malesia.
However, a notable exception is provided by the tree Schefflera rugosa (BL.) HARMs in Java;
LAM (1924) recorded its gregarious occurrence on the volcanic cone of Mt Slamet in Central Java
where it is co-dominant with the pyrophilous Albizia lophantha Brn. in elfin forest between 2500—
3050 m, above which it gives way to the open, rocky, treeless slopes below the summit (3428 m).
It is also gregarious on Mt Tjeremai in West Java, where Lam (1925) noted that above some
2650 m a low forest dominated by this species replaces the high forest of Dacrycarpus imbricatus ;
this low forest extends to c. 3000 m. On the Gedeh-Pangrango complex above Puntjak Pass in
West Java Schefflera rugosa is frequent in forest borders. On Mt Ulu Kali in Malaya, east of
Kuala Lumpur, S. nervosa (KING) VIG. is common in young regrowth along the main road

1979] ARALIACEAE—I (Philipson) 3

below the Genting Highlands hotel/casino complex from 1300-1600 m. This pioneering tendency
is shared by a number of other species in the genus, especially in montane parts of Papuasia.
Many of these species are terrestrial as pioneers or in secondary formations, but epiphytic in
closed forest (and then often much more scattered).

Gregarious occurrence, usually in pioneering situations or in forest borders, has also been
observed in the various species of Harmsiopanax ; STEUP (1938) observed H. aculeatus (BL.) WARB.
ex BOERL. as a characteristic pioneer in grass thickets on hills in SW. Celebes. H. harmsii K. Scu.
behaves similarly around Wau, Bulolo, and Sogeri (Rouna) in Papua New Guinea, especially in
narrow intermontane valleys, while H. ingens PHILIPSON can be locally abundant in open situa-
tions in the highlands of New Guinea. Gastonia spectabilis (HARMS) PHILIPSON can be locally
frequent as a pioneer in hill areas, e.g. around Bulolo and on the Madang-Ramu Divide; while
Polyscias elegans (C. MOorRE & F.v.M.) HArMs is frequently seen in stable monsoon scrub and
forest borders in the Port Moresby region and P. /Jedermannii HARMS can be an exceedingly com-
mon regrowth tree in cut-over montane forest subject to frequent cloudiness and rain.

In the subalpine zone of New Guinea Brass (1941) found two species of Schefflera conspicuous
in the stunted forest of Xanthomyrtus-Vaccinium-Papuacedrus-Phyllocladus between 3200-3800 m
on Mt Wilhelmina. From Lake Habbema upwards, the characteristic overtopping tree is S. alti-
gena FRODIN (sect. Brassaia), with its large foliage contrasting sharply with the surrounding
microphyllous vegetation; at higher altitudes it gives way to S. pagiophylla Harms (S. falcata
PHILIPSON), a species of uncertain affinities. Scheffiera chimbuensis FRODIN and S. straminea
FRODIN are likewise often seen in forest borders on Mt Wilhelm and Mt Giluwe respectively in
Papua New Guinea.

In contrast to the large and conspicuous species of Scheffiera, many others are more or less
shade-loving epiphytes or vines of lower storeys within the forest, although they may occur in
sunnier situations where clouding is frequent (S. singularis B. C. STONE on Mt Ulu Kali). One
group of species in New Guinea is almost limited to perhumid moist or wet rain-forests and
usually are small epiphytic shrubs or even herbs (S. gemma FRODIN). A few species are con-
spicuous rosette-trees of the forest understorey (S. stahliana (WARB.) FRODIN).

Climate. In Malesia Araliaceae for the most part shun regions subject to a seasonal climate; all
species are evergreen. A few exceptions include Schefflera thaumasiantha Harms from open
savanna in the hill zone in SE. New Guinea and S. actinophylla (ENDL.) HARMS as a gallery tree
in seasonal parts of southern New Guinea, but especially Harmsiopanax aculeatus whose range
is for the greater part subject to an annual drought period; in addition to SW. Celebes, it is also
frequent on old lava-streams on Mt Idjen in East Java together with Wightia, Casuarina junghuhn-
iana, Dodonaea, and Wendlandia, and is one of the few araliads in the Lesser Sunda Islands.

With respect to altitude, most Araliaceae occur in the lowland, hill and montane zone below
2300 m. The only genera of which all Malesian representatives occur in the hills and mountains
above 1000 m are Pentapanax and Dendropanax, but even these do not exceed 3000 m. Only
certain species of Schefflera and Harmsiopanax continue upwards much higher, especially in
New Guinea where the highest known record belongs to S. pagiophylla HARMS on the Carstensz
complex, where F. J. WissEL found it in 1936 at 3900 m.

Flower biology. Little has been recorded about the floral biology of the family in Malesia, but
BECCARI’s account (1878) of the ‘false fruits’ of Osmoxylon (including Boerlagiodendron) serving
to attract doves which are assumed to effect pollination has become a classic description and
example of ornithophily.

Heterosexual flowers, usually involving combinations of perfect and male flowers, occur
frequently but understanding will require intensive study in the living state.

Though flowers are not generally showy and often veritably inconspicuous, the disk produces
abundant nectar. They also may spread a rather disagreeable scent, somewhat spermatic, that in
Schefflera rugosa resembling that of Ligustrum. For this species DoCTERS VAN LEEUWEN (1933)
observed on Mt Gedeh, West Java, only rare visits (notably by Diptera), but assumed that cross-
pollination will be the rule. Flowers (at least those of Fatsia) will also be visited by Hymenoptera.
In Schefflera stahliana (WARB.) FRODIN, the thick fleshy flowers, numerous stamens, and position
of the inflorescence below the rosette of leaves all point to bat pollination.

Dispersal in the family takes place generally by fruit-fall; but as fruits are baccate or (more

4 FLORA MALESIANA [ser. I, vol. 91

usually) drupaceous, they will also be eaten by birds (for the most part) and bats, as recorded by
RiD.Ley (1930) for Aralia, Hedera, and Schefflera. The black fruits of Schefflera sect. Brassaia in
New Guinea and Australia are especially popular with birds. The hooked mericarps of Harmsio-
panax are exceptional.

Seed germination is most likely after the seed having passed the gut of a bird or after mastica-
tion of the fruit by a bat.

References: BECCARI, Malesia 1 (1878) 193-198; Brass, J. Arn. Arb. 22 (1941) 271-342, esp.
318, 320, 323, 327; DocTERS VAN LEEUWEN, Verh. Kon. Ak. Wet. A’dam sect. 2, 31 (1933) 195;
H. J. Lam, Trop. Natuur 13 (1924) 20; ibid. 14 (1925) 6; RipLey, Disp. (1930); Steup, Trop.
Natuur 27 (1938) 142.

Morphology. Most erect Araliaceae are sparingly branched or even unbranched; their
limbs are upright and the often massive, easily broken twigs generally bear rosettes of big, long-
stalked leaves which leave large scars when they fall. Few form a true crown (4rthrophyllum,
Gastonia, some species of Schefflera and Polyscias), while the others often look like elongated and
grotesque shrubs with a candelabrum-like framework. Branches in most Araliaceae are entirely
orthotropic; generally speaking, the plants would fit into the so-called Holttum, Corner, Tomlin-
son, Chamberlain and Leeuwenberg models of HALLE c.s. (1970, 1978), or their intermediates.

Aralia scandens (MERR.) HA is a true climber. Epiphytic species only occur in the genus
Schefflera; it is not yet recorded that any of them may appear to turn into a hemi-epiphytic habit.

Hedera (not native in Malesia) is almost the only genus with differentiated shoots and marked
vegetative dimorphism.

A preliminary account of shoot-morphology in the family has been given by PHILIPSON
(1978), but much further field work is required in this area. Vegetative buds may be either
proleptic or sylleptic; in the resting phase such buds may be covered by the clasping bases of
foliage leaves (Osmoxylon, Schefflera) or (more rarely) specialized cataphylls (Acanthopanax).
In both types of bud the primordia and young leaves may be covered by exudations of resinous
slime.

There is so far little recorded evidence of the changes in leaf shape and configuration during
the somatic phase of the life cycle known in many araliads in othe: parts of the world; however,
recent observations in New Guinea made by FRODIN suggest that heteroblastism does occur,
although its manifestation is not constant for a given species. Distinctive juvenile and intermediate
foliage has been found in Schefflera eriocephala Harms and (to a lesser extent) in S. stolleana
Harms. Other examples are seen in Brassaiopsis, Trevesia and Schefflera subg. Agalma (S. aroma-
tica (BL.) HARMS; S. nervosa (KING) VIG.); juvenile leaves of some of these are preserved in
Herbarium Bogoriense. In Harmsiopanax, the configuration of the leaves changes abruptly just
below the inflorescence, a phenomenon paralleled in some other genera although less dramati-
cally. In Mackinlaya celebica (HARMS) PHILIPSON and M. schlechteri (HARMS) PHILIPSON, leaf
polymorphism is very marked with the result that in the past several ‘paper species’ have been
described on too limited a range of material; in this revision many reductions have been made.

References: HALLE & OLDEMAN, Essai sur l’architecture et la dynamique de croissance des
arbres tropicaux, Paris (1970); HALLE, OLDEMAN & TOMLINSON, Tropical trees and forests: an
architectural analysis, Berlin (1978); PHILIPSON in Tomlinson & Zimmermann (eds.), Tropical
trees as living systems (1978) 269-284.

Anatomy. General accounts of the vegetative anatomy of the ivy family are given by GUssow
(1900), ViGuiER (1906, 1909) and METCALFE & CHALK (1950). Secretory canals are characteristic
of the stems and leaves, but are absent from Aralidium. A comparison of the wood anatomy of
Araliaceae and Cornaceae is made by PHILIPSON (1967), and an extensive account of vegetative
anatomy in the context of woody Umbellales was provided by RODRIGUEZ (1957, 1971). The
xylem of the former family is characterized by fibres with small, simple pits, and the presence
of scalariform and reticulate perforation plates in the vessel elements (as opposed to simple
perforations) is thought to be a less advanced feature. Recent special reports bearing on taxonomy
include: on stomatal development (INAMDAR c.s., 1969); on sievetube plastids (BEHNKE, 1972), and
on epidermal papillae (Bul, 1974).

Floral anatomy is discussed by BAUMANN-BODENHEIM (1955), PHILIPSON (1967, 1970) and
especially by EypE & TsENG (1971). Embryological characteristics of the family have been re-

1979] ARALIACEAE—I (Philipson) 5

viewed by Davis (1966) as well as by RAO (1972). The single pendulous anatropous ovule has the
funiculus and ovular vascular bundle axial (PHILIPSON, 1970). The embryo is small in a mass of
endosperm (MARTIN, 1946; GRUSHVITZKY, 1967).

References: BAUMANN-BODENHEIM, Bull. Soc. Bot. Suisse 65 (1955) 481-510; BEHNKE, Bot.
Rev. 38 (1972) 155-197; But NGoc-SAnu, Bull. Mus. Hist. Nat. Paris III, Bot. 18 (1974) 85-91
(whole no 271); Davis, Systematic embryology of the angiosperms, New York (1966); EypE &
TsENG, J. Arn. Arb. 52 (1971) 205-239; GRUSHVITZKY, Proc. Int. Symp. Physiol. Ecol. & Bio-
chem. of Germination (ed. H. Borriss) (1967); GUssow, Beitrage zur vergleichende Anatomie der
Araliaceae, Thesis, Breslau (Wroclaw) (1900) 67 pp., illus.; INAMDAR, GOPAL & CHOHAN, Ann.
Bot. n.s. 33 (1969) 67-73; MARTIN, Amer. Midl. Nat. 36 (1946) 513-660; METCALFE & CHALK,
Anatomy of the dicotyledons II, Oxford (1950); PHiLipson, New Zeal. J. Bot. 5 (1967) 134-165;
in Robson, Cutler & Gregory (eds.), New research in plant anatomy, London (1970) 87-100;
Rao, Phytomorphology 22 (1972) 75-87; RoDRIGUEZ, Univ. Calif. Publ. Bot. 29 (1957) 145-318;
in Heywood (ed.), The biology and chemistry of the Umbelliferae, London (1971) 63-91;
VicuiER, Ann. Sci. Nat. Bot. IX, 4 (1906) 1-209; ibid. TX, 9 (1909) 305-405.

Palynology. Palynological studies of Malesian Araliaceae include: on Gastonia (TsENG,
1971); on Tupidanthus and Plerandra (= Schefflera) (TSENG, 1973); on Osmoxylon (as Boer-
lagiodendron) (TSENG, 1974) and on Schefflera (TSENG & SHOUP, 1978). A detailed consideration
of the relations of Araliaceous pollen to those in other orders is given by HIDEAUX & FERGUSON
(1976), and of the affinities of Klotzschia (Umbelliferae/Hydrocotyloideae) to Araliaceae by SHOUP
& TSENG (1977). To date, much useful new evidence has been made available, but better correla-
tion with other classes of attributes is required.

References: HIDEAUX & FERGUSON in Ferguson & Muller (eds.), The evolutionary significance
of the exine, London (1976); SHourp & TsENG, Amer. J. Bot. 64 (1977) 461-463; TsENG, Amer.
J. Bot. 58 (1971) 505-516; Grana 13 (1973) 51-56; Amer. J. Bot. 61 (1974) 717-721; TsENG &
SHoup, Amer. J. Bot. 65 (1978) 384-394.

Chromosome numbers. Lists of chromosome numbers for members of Araliaceae are
given by DARLINGTON & WYLIE (1955), SHARMA & CHATTERJI (1964) and BOLKOVSKIKH c.5S.
(1969). The family shows considerable constancy of the basic number of x = 12, although x = 11
has been recorded for one non-Malesian group of Schefflera. The implications of chromosome
data on the wider relationships of the Araliaceae are discussed by Moore (1971).

References: BOLKOVSKIKH c.s., Chromosome numbers of flowering plants, Leningrad (1969);
DARLINGTON & WyLIE, Chromosome atlas of flowering plants, ed. 2, London (1955); Moore in
Heywood (ed.), The biology and chemistry of the Umbelliferae, London (1971) 233-255;
SHARMA & CHATTERJI, Cytologia 29 (1964) 1-12.

Phytochemistry. Information on the chemistry of the Araliaceae should be sought in
HEGNAUER (1964, 1978) where references to original sources are given. The family is characterized
by the occurrence of essential oils and resins in canals and by the presence of polyacetylenic
compounds (especially falcarinone-type), triterpenic sapogenins of the oleanene-, ursene- and
dammarene-types, seed oils with petroselinic acid, and by the absence of true tannins. The
chemistry of the family fully confirms its close relationship with the Umbelliferae and also more
distantly with the Pittosporaceae and the Compositae (HEGNAUER, 1969, 1971; BOHLMANN,
1971). The rareness of flavones and the predominance of flavonols in Araliaceae suggests closer
relationship with two of the three subfamilies of the Umbelliferae, flavones having not yet been
found in Hydrocotyloideae and Saniculoideae (HARBORNE, 1971). The absence of iridoid sub-
stances and true tannins and the presence of polyenes, petroselinic acid and isoprenylated
coumarins in the Umbellales contrasts with the Cornales (HEGNAUER, 1969; JENSEN c.s., 1975),
and this led to a suggestion that the Umbellales and Cornales had to be separated (BATE-SMITH
c.s., 1975), an argument with increasing support from other lines of inquiry. However, insufficient
evidence appears to be yet available for the detection of possible lines of relationship within the
Araliaceae.

References: BATE-SMITH c.s. Biochem. Syst. Ecol. 3 (1975) 79-89; BOHLMANN in Heywood (ed.),
The biology and chemistry of the Umbelliferae (1971) 279-291; HARBORNE, /.c. 293-314;
HEGNAUER, Chemotaxonomie der Pflanzen 3 (1964); in Harborne & Swain (eds.), Perspectives
in phytochemistry (1969) 121-138; in Heywood (ed.), The biology and chemistry of the Umbelli-

6 FLORA MALESIANA [ser. I, vol. 91

ferae (1971) 267-277; in Cauwet-Marc & Carbonnier (eds.), Les Ombelliféres. Contributions
pluridisciplinaires a la systématique. Perpignan (1978) 335-363; JENSEN, NIELSEN & DAHLGREN,
Bot. Notis. 128 (1975) 148-180.

Taxonomy. The Araliaceae are on all grounds closely connected with the Umbelliferae, a
very large but mainly temperate and tropical-montane group comprising mostly aromatic herbs
with a restricted floral scheme. It seems likely that the ancestors of the ivy family were the woody
tropical stock from which the herbaceous Umbelliferae evolved under the rigours of cooler
climates (CORNER, 1940) with the arborescent Heteromorpha of upland Africa and Myrrhidendron
in Central and South America, both in the subfamily Apioideae, perhaps representing relics of
the transition on account of the presence of a number of attributes primitive for the family
(RODRIGUEZ, 1957); on the other hand, the umbellifers may have originated as megaherbs on
tropical mountains (PHILIPSON, 1978). A few Araliaceous genera exhibit some features charac-
teristic of Umbelliferae, such as Harmsiopanax, Mackinlaya, and especially Myodocarpus (from
New Caledonia) and Stilbocarpa (from southern New Zealand and associated ‘subantarctic’
islands).

This led HARMs (1898) to show three different lines leading from Araliaceae to Umbelliferae,
suggesting that considerable overlap between the families existed; more recently RODRIGUEZ
(1971) again called attention to this phenomenon and THoRNE (1968, 1973) has gone further by
merging Umbelliferae into Araliaceae (a step also advocated by HALLIER f. in 1905) and assuming
the three subfamilies of the former to have arisen separately, perhaps in different parts of the
world at different times, from proto-Araliaceous ancestors. However both he and RODRIGUEZ
have concluded in agreement with CORNER (/.c.) and BAUMANN-BODENHEIM (1946) that the
Araliaceae sensu stricto, ““‘because of their greater evolutionary breadth and their retention of
many more primitive features, would seem closest to the proto-araliad stock’’ (THORNE, 1973).

Nevertheless, the concept of Araliaceae as a separate natural family appears to serve a useful
purpose and has been retained for this Flora. An isolated, doubtfully included genus is the West
Malesian Aralidium; it is the only genus lacking resin ducts but it would be equally anomalous
in the Corndceae to which it has also been referred. It shows some resemblance with the New
Zealand genus Griselinia, usually relegated to the Cornaceae, be it as a marginal member
(PHILIPSON, 1967).

Subdivision. Since the first significant family monograph by SEEMANN (1868), several systems
have been proposed which, taken together, are notable for their lack of consistency. This results
from a lack of agreement on the relative importance of the comparatively minor structural and
gross anatomical features of reproductive parts traditionally used and by conflicting claims on
the relative ‘antiquity’ of polymery versus pentamery. Until recently, there has been for a priori
reasons (cf. EYDE, 1975) little recognition of the potential value of vegetative features and their
acceptance as valid evidence for a system; and information from wood anatomy, floral histology,
palynology, karyology, phytochemistry, and other areas is only beginning to be utilized. While
woodiness is generally accepted as a primitive feature in Araliaceae, the impact of the work of
CORNER and others on tree structure dnd growth rhythms (summarized in HALLE, OLDEMAN &
TOMLINSON, 1978; see also BORCHERT, 1969; HLADIK, 1970; PHILIPSON, 1978) has still to be fully
assimilated. Much more work is also required on inflorescences, although FRoDIN (1970),
PHILIPSON (1970b) and others have made a beginning. The monothetic interpretation of most
attribute states usual in systems of the family was first challenged by BAUMANN-BODENHEIM
(l.c.) who considered that phyletic changes could have taken place in parallel; and EypE & TsENG
(1969) showed that at least some supposedly unidirectional sequences were reversible. This has
tended to reduce the supposed importance of many of the traditional attributes, with a conse-
quent reduction in the number of genera.

Sufficient evidence is not yet available, however, for the construction of a more balanced,
polythetically based system of the family, and the long-standing subdivision proposed by HARMS
(1898) into three tribes, Schefflereae, Aralieae, and Mackinlayeae, based monothetically on the
structure and aestivation of the perianth, is retained. In spite of its now recognized imperfections,
it is more valid than the systems of VIGUIER (1906) and HUTCHINSON (1967) and remains the most
widely accepted. Some steps towards the formulation of a new system on polythetic principles
have been taken by EYDE & TsENG (1971: 221) who make a fundamental distinction based on

1979] ARALIACEAE—I (Philipson) |

basic leaf-organization (either pinnate or palmate) and recognize the heterogeneity of HARMS’
Aralieae; however, this system was deliberately not completely developed.

Generic delimitation. Generic limits within the Araliaceae have long been unstable. As in the
Umbelliferae, the flower conforms to a simple and relatively uniform pattern throughout most
of the family and systematists have resorted to small technical differences to delimit genera.
HARMS recognized 51 genera, while ViGuIER, little more than a decade later, recognized 80;
HUTCHINSON distinguished 84 genera but with criteria very differently weighted as compared with
VicuiER. Nevertheless, in Malesia as elsewhere there are several distinctive and very natural
genera, including Harmsiopanax, Aralidium, Osmoxylon, Anakasia, Mackinlaya, Arthrophyllum,
Delarbrea, and Trevesia. Many other genera in the family, including those Malesian ones not
noted above, are mutually less distinctive and delimitation is based on various combinations of a
number of attributes which have been subject to many different standards of weighting and inter-
pretation. These include: petals valvate or imbricate; pedicel articulated or not; leaves digitately
compound, pinnate, or simple; stamen and locule number and the relationship of these numbers;
style free or connate; endosperm ruminate or not; and thorns present or not.

The present treatment is characterized by a number of generic reductions or exclusions; only
one genus described since 1900 has been retained (Anakasia). Hederopsis is united with Macro-
panax, a genus with a very similar facies and (in part) overlapping range and which was separated
merely on the number of cells in the ovary and variations in inflorescence structure. Wardenia
has been united with Brassaiopsis because apart from its simple palmately veined leaves there are
no differences; better material collected in recent years has provided evidence that the ovary is in
fact 2-locular, but that one ovule aborts and the fruit is as a result 1-seeded. Acanthophora
differs from Aralia only in habit (VAN STEENIS, 1948). PHILIPSON (1951) already regarded Anomo-
panax as insufficiently distinct from the older Mackinlaya, a union retained here. The same author
(1973) combined Boerlagiodendron with the earlier described Osmoxylon as a number of species
intermediate between the two genera had come to light. The Malesian species formerly included
in Tetraplasandra as well as Peekeliopanax were likewise by PHILIPSON (1970a) reduced to two
species of Gastonia; but it should be noted that in Malesia as on the SW. Indian Ocean islands
this genus is very close to Polyscias, particularly the very similarly ranging sect. Eupteron
(P. nodosa, P. ledermannii, etc.).

The two largest Malesian genera are herein both treated in a wider sense. Polyscias includes as
its type section a distinctive Melanesian/Micronesian group of species which in the wild state
extends into eastern Malesia and the Philippines, and in cultivation further west. The remainder
of the genus as represented in Malesia comprises some rather distinctive species, several of which
have been given generic rank (Eupteron, Kissodendron, and Palmervandenbroekia). If the name
Polyscias were to be confined to the type-section of the genus it would be necessary to recognize a
considerable number of small genera. Retention of a wide concept for the genus therefore appears
to be preferable.

Schefflera, by far the largest genus of the family in Malesia, is likewise more broadly conceived
than in the past, although in our region only a small number of species from segregate genera are
involved (i.e. those formerly in Brassaia, Plerandra, Scheffleropsis, and Tupidanthus). On a
world-wide basis, FRODIN (1975) recommended the reduction of 12 segregate genera; improved
knowledge of the genus required that, as in Polyscias, this step be taken or have the genus split
into a number of differently organized smaller genera with more serious nomenclatural conse-
quences. A broad concept of Schefflera has therefore been adopted. It may be noted here that the
flowers in both the former genera Plerandra and Tupidanthus, characterized by numerous
stamens, are thought by FRopDIN to represent a secondary development related to bat pollination,
and the large fruits for dispersal by bats or larger birds; this is supported by the position of the
inflorescences, which are beneath the leafy rosettes as a result of retarded development although
the axes remain sympodial and the shoot units mostly orthotropic. As divisions of the larger
genus Schefflera, they are not at all closely related, conforming to the views of TsENG (1974) on
the pollen morphology and contrasting with the views of HARMs (1898) and EyDE & TsENG (1971),
who regarded them (as genera) as of close affinity and relatively primitive within the family. In
fact, rather different levels of specialization are represented overall by the two taxa, not just in the
pollen morphology (TsENG, /.c.).

8 FLORA MALESIANA [ser. I, vol. 9!

Specific delimitation. This has offered many problems, but fortunately the very abundant
material now available from many areas has revealed that many of the species described up to
1951 are in fact conspecific. Especially was the great plasticity in vegetative and inflorescence
development not sufficiently appreciated in the past, partly due to imperfect field knowledge of
the plants. This has led to considerable reductions in Arthrophyllum, Aralia, Osmoxylon,
Gastonia, and parts of Schefflera and Polyscias. Other species have been reduced through a
regional approach to the genera. However, in Osmoxylon and Schefflera, many species from the
Philippines, Celebes, western New Guinea, and to a lesser extent Borneo and Sumatra are still
known only from very few or even only a single collection; this has meant a rather tentative
treatment in many cases. The same applies in a more limited way in some of the other genera.
Certain species are rather polymorphic, and at least in Schefflera several ‘species-complexes’
have been discerned.

References: BAUMANN-BODENHEIM, Bull. Soc. Bot. Suisse 56 (1946) 13-112; BoRCHERT, Amer.
J. Bot. 56 (1969) 1033-1041; CorNER, Ways. Trees Malaya (1940) 153; Eypge, Amer. Sci. 63
(1975) 430-437; EYDE & TSENG, Science 166 (1969) 506-508; J. Arn. Arb. 52 (1971) 205-239;
FRODIN, The complex of Cephaloschefflera in Schefflera (Araliaceae), Thesis, Cambridge, U.K.
(1970); J. Arn. Arb. 56 (1975) 427-448; HALLE, OLDEMAN & TOMLINSON, Tropical trees and
forests: an architectural analysis, Berlin (1978); HALLIER f. New Phytol. 4 (1905) 151-162;
Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1898) 1-62; HLapik, Adansonia 10 (1970) 383-407;
HUTCHINSON, Gen. FI. Pl. 2 (1967) 52-81; PHILIPSON, Bull. Brit. Mus. Nat. Hist. Bot. 1 (1951)
3-20; New Zeal. J. Bot. 5 (1967) 134-165; Blumea 18 (1970a) 490-495; ibid. 18 (1970b) 497-505;
ibid. 21 (1973) 81-89; in Tomlinson & Zimmermann (eds.), Tropical trees as living systems
(1978) 269-284; RopRIGUEZ, Univ. Calif. Publ. Bot. 29 (1957) 145-318; in Heywood (ed.), The
biology and chemistry of the Umbelliferae (1971) 63-91; SEEMANN, Revision of the natural order
of Hederaceae, repr. from J. Bot. London (1868); VAN STEENIS, Bull. Bot. Gard. Btzg III, 17
(1948) 390-391; THORNE, Aliso 6 (1968) 57-66; Notes R. Bot. Gard. Edinb. 32 (1973) 161-165;
TsENG, Amer. J. Bot. 61 (1974) 717-721; Vicuter, Ann. Sci. Nat. Bot. IX, 4 (1906) 1-209.

Uses. A variety of minor local uses are reported by BURKILL (1966), HEYNE (1927), and
OcHSE & BAKHUIZEN VAN DEN BRINK (1931) as well as in the notes under individual species in
this Flora. The only species that form articles of trade are the taxa of Polyscias sect. Polyscias
popularly grown as foliage and hedge plants; however, many other species in a variety of genera
are of actual or potential ornamental worth, with Schefflera actinophylla (ENDL.) HARMS and
S. longifolia (BL.) Vic. being particularly widely used. In Papua New Guinea, Gastonia spectabilis
is cut for timber and the wood used for light carpentry, boxes, etc.

Monkeys are fond of the flush of some aromatic species of Schefflera, as observed in West
Java and North Sumatra.

References: BURKILL, Dict. rev. ed. (1966); HEYNE, Nutt. Pl. (1927); OcHsE & BAKHUIZEN
VAN DEN BRINK, Veget. D.E.I. (1931).

Notes. Nomina nuda and invalidly published names have only been quoted if they have been
cited in Index Kewensis.

Notes for collectors. Many araliads present problems to collectors because of the size of their
leaves and inflorescences. It is often advisable to select leaves of medium size, but the maximum
size of leaves should be recorded on the label. It is important to preserve the junction of leaf and
stem and also sufficient of the leaf to allow reconstruction of the whole. Likewise, with inflores-
cences the base, main axis and some primary branches should be preserved so that the whole can
be visualized — ultimate branches alone are insufficient. Fruiting material is as useful as a flower-
ing specimen. Collectors should be alert to note the existence of vegetative heteroblasty and
floral dimorphism and document these with specimens and notes; the form in immature plants
is important. Rapid drying is essential or all parts will disarticulate and very fragmentary
specimens result.

KEY TO THE GENERA
1. Petals imbricate. Tribe ARALIEAE.
2. Leaves simple, palmately or pinnately lobed, or entire.
Ss EeaVvesspalmatelvalObeGaitOMment OSC mm uriiCinrsince cnn mnie a on 1. Harmsiopanax
3, Leaves pinnarciy lobed on entire, glabrous’ =). 3 << 2) eet Renee ee 2. Aralidium

1979] ARALIACEAE—I (Philipson) 9

2. Leaves pinnately compound (or bi- or tripinnate).

Amibeavesmwice (Or more) pinnate (eo ee ac sk ke sk we tw ee 3. Aralia
4. Leaves once pinnate.
Seebenheroumany csi: ©. seen Lae ROE ew. Stes. sla ao oe aes 4. Delarbrea
Sra HerNE OTC WOR 8, oe Re eM ed can Ban 055) odd ios outa 5. Pentapanax
1. Petals valvate.
6. Petals with a narrow base, or claw. Tribe MACKINLAYEAE ........... 6. Mackinlaya

6. Petals with a broad base. Tribe SCHEFFLEREAE.
7. Inflorescence rays trifid: central branch shorter with ‘false fruits’, the two lateral longer with normal

TOG: 9: Sahara iment Meme ys Bent ee er en A 7. Osmoxylon
7. Inflorescence branches not as above.
fo. LO EER Sod a NVoy 1 ite] | Oe aan SP ee De et ee a 8. Arthrophyllum

8. Ovary with more than one cell.
9. Leaves pinnate.
lOfPedicel not articulated below the flower.5..> 44 ne eee 9. Gastonia
OM Pedicellarticulatedibelow;themowel cme ae en eee een oe ae: 10. Polyscias
9. Leaves not pinnate.
11. Pedicel articulated below the flower.
12. Leaves digitately compound (or rarely unifoliolate) (Malay Peninsula, Sumatra, Java)
11. Macropanax
12. Leaves simple, not articulated with the petiole (West New Guinea) . . . . 12. Anakasia
11. Pedicel not articulated below the flower.
13. Leaf simple (or unifoliolate) or palmately lobed.
14. Leaf palmately lobed.

BO VARY 2-COMEG: (yes a4 oc si o Sehst a os Ste S o 13. Brassaiopsis
IS Ovary,.10=orsmore-celledvose-s.... oachaaet wud) Bile Soo Gickbs OR Seoue 14. Trevesia
14. Leaf simple (or unifoliolate).
16. Articulation present between petiole and blade. ........2.2.2.2.. Schefflera
16. No articulation between petiole and blade.
AV ROVATYP2-COLCA) graye cers. oy Bn sk She Te Rs By GED tsuki, Sieies dates ess 13. Brassaiopsis
ieeOvary’4-.or;more-celled o..) 0.546" ¢ os EA ee thes Cl NS at db es 15. Dendropanax
13. Leaf digitately compound.
18. Petiolules joined together by a web of tissue. . .........2.2.4.-. 14. Trevesia

18. No such web of tissue present.
19. Styles or stigmas 2.

Boe Otyle DING teri Woes 6 We sb earane 6 Feb. oxen? ecaeey WA Moa aee” 16. Acanthopanax

20 estylestunited intola columnysure-ges eee lee) eRe ee nome) oe lee 13. Brassaiopsis

ios Styles or stigmas more: than.2. oi Spill nls Shen sin eine Nelsen iene Schefflera
1. HARMSIOPANAX

Wars. in E. & P. Nat. Pfl. Fam. Nachtr. 1 (1897) 166; HArRms, Bot. Jahrb. 56
(1921) 413; Hutcu. Gen. FI. Pl. 2 (1967) 62; PHiLipson, Blumea 21 (1973) 81. —
Schubertia BL. Bijdr. (1826) 884, nom. illeg., non MirRB. 1812. — Horsfieldia BL. ex
DC. Prod. 4 (1830) 87, non WILLD. 1805; Bru. in B. & H. Gen. Pl. 1 (1865) 937;
BoerL. Hand. 1 (1890) 633; HARMs in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 60. —
Fig. 1-3.

Sparingly branched or single-trunked often monocarpic trees, up to 18 m. Stems
stout, bearing terminal clusters of large, palmately lobed, often peltate, exstipulate
leaves. Trunk, petioles, and sometimes the blades spiny. Flowers in very large,
repeatedly branched, terminal panicles which develop after the leaves have fallen.
Umbellules arranged racemosely on the ultimate branchlets, sessile or peduncled,
each consisting of a few to many pedicelled flowers. Pedicels not jointed, subtended
by a bract and bearing two subulate bracteoles. Flowers hermaphrodite or with
hermaphrodite flowers on terminal and male flowers on basal branches. Calyx a

, Rh SN a SS SH litte:
y YY a ea a a Sana 2) REREeeah es Seen
+) ,—,, NAN, . = ON * 4 r. i, o ayiNatal BP une > NY See
Nea pa era SRO SN AS RY ACO Ke -—.
Brea Ne vo Ss r ‘a rl (3 AR Sy a p = > =,
SORUN AALS OS. Ueraaaia SS | aay, Sg AN oa, ie ete TA ba

TR RS eS mame ie i

MS

[ser. I, vol. 91

FLORA MALESIANA

10

Seep Nd
als
‘ Bs a

lule, x 4, d. developing fruit, x 12 (a NGF 36901, b-d Puiipson 3483).

Fig. 1. Harmsiopanax ingens PHILIPSON ssp. ingens. a. Leaf, x 4/., b. part of inflorescence, x 1/3, c. umbel-

1979] ARALIACEAE—I (Philipson) 11

minute rim. Petals.5, free, valvate with a broad base. Stamens 5, dorsifixed, versa-
tile, introrse. Ovary inferior, narrowly obconic, densely bristly; cells 2; disk
conical, deeply cleft between the two subulate styles. Fruit consisting of 2 dry
mericarps, each 3-ribbed and bearing a persistent slightly hooked style.

Distr. Malesia: 3 spp. from Java, the Lesser Sunda Is., Celebes, and New Guinea.

Ecol. Montane and mossy forest and in regrowth on grassy hillsides.

Notes. Harmsiopanax is a small structurally isolated genus confined to Malesia. The three species are
uniform both in their vegetative and their reproductive features. It has long been recognized that some
of the characters of this genus are anomalous within Araliaceae and a return to its earlier position within
Umbelliferae would have something in its favour. The monocarpic habit is unknown elsewhere in Aralia-
ceae, but is not uncommon in Umbelliferae. The character of the fruit, which splits into two dry mericarps,
closely approaches the fruit structure of Umbelliferae, and the vascularization of the gynoecium is also
characteristic of that family. However, the structure of the leaf-base, the woody habit, and the shape of
the petals all incline towards Araliaceae.

KEY TO THE SPECIES

1. Upper surface of leaves uniformly setulose.
PemUuMIDEIUIESISESSIIC.. 5 6 es He
2 Wmbellules peduncled. . ..... .4 4.

1. Upper surface of leaves with many (or rarely few) larger spines among the setulose hairs

1. Harmsiopanax aculeatus (BL.) WARB. ex BOERL.
Hand. 3 (1900) 88; Koorp. Exk. Fl. Java 2 (1912)
719; Atlas 4 (1916) f. 668 & 669; BakH. f. &
OoststR. in Back. Bekn. Fl. Java (em. ed.) 7
(1946) fam. 159, p. 19; Back. & BAKH. f. Fl. Java 2
(1965) 171; STEEN. Mt. FI. Java (1972) pl. 3-2;
PHILIPSON, Blumea 21 (1973) 82. — Schubertia
aculeata BL. Bijdr. (1826) 885. — Horsfieldia
aculeata (BL.) DC. Prod. 4 (1830) 87; BENN. PI.
Jav. Rar. (1840) 123, t. 26; K. & V. Bijdr. 7 (1900)
57; BoeRL. Handl. 1 (1890) 647. — Horsfieldia
peltata BTuH. in B. & H. Gen. Pl. 1 (1862) 937. —
Fig. 2.

Tree up to 4 m, with a slender spiny trunk.
Young stems covered more or less densely with
woolly hairs, bristles, and spines with bulbous
bases, the spines enlarging on older stems. Leaves
rounded, variable in size, often 60 cm or more in @,
deeply palmately lobed, usually peltate in mature
leaves, sinuses between the lobes broad or narrow,
lobes 7-10, usually sharply and irregularly incised
and toothed, apex acute, upper surface rather
sparsely covered with evenly-spaced, appressed,
sometimes branched hairs (denser on the main
veins), underside densely clothed with a soft,
woolly tomentum, often with some bristles on the
main veins; petiole c. 60 cm, 1 cm @ at base,
terete with clasping base, densely covered with
woolley hairs, bristles, and some spines. Inflores-
cence up to c. 70 cm long, main branches rather
sparsely covered with a short tomentum and, when
young, bearing numerous bracts similar to the
leaves but smaller, not peltate, and often 3-lobed
or entire; ultimate branchlets slender and often
woolly-tomentose, bearing minute linear bracts
which subtend the sessile umbellules. Umbellules

si Yous 46: JeMve® “eLie se, im. a5 ea) Pw) ie, Se

about 4 mm @ in flower, the broadly ovate outer
bracts forming a more or less distinct involucre.
Flowers hermaphrodite or male, either mixed in an
inflorescence, or separate, c. 10-15 per umbellule,
each subtended by a lanceolate receptacular bract
c. 2 mm long. Pedicel c. !/, mm long. Calyx rim
fringed. Petals strap-shaped, c. 11/, mm long at
anthesis. Filaments c. 2 mm; anthers c. 0.3 mm
long, orbicular. Ovary covered with cilia which
lengthen as the fruit ripens. Mericarps long-ciliate,
crowned with the divergent styles.

Distr. Malesia: Java, Lesser Sunda Is. (Bali,
Lombok, Sumbawa, Flores, Timor), southern half
of Celebes. There is a single KoRTHALS sheet in L,
ticketed from Central Sumatra, but this is presum-
ably wrongly localized.

Ecol. Usually in rather dry, open localities, but
also in forest, in secondary forest, also pioneering
on rocks, in grasslands and on lava-streams,
300-1800 m. FI. fr. April-Nov. ScHMutTz found it
in Flores flowering in October, but leaves had
fallen.

Vern. Java: djankurang, d. tjutjuk, djogloran-
grang, S, gabus, garang, g. lanang, gungrang,
udulan laki, J.

Note. In this species lateral shoots usually
appear below the infructescences so that the trees
are not normally monocarpic.

2. Harmsiopanax harmsii K. Scu. in K. Sch. &
Laut. Nachtr. (1905) 329; Harms, Bot. Jahrb. 56
(1921) 413.

Tree up to 7 m, with a slender trunk. Young
stem covered with bristles, hairs, and spines, older
stems with smooth bark with small rounded lenti-
cels and numerous spines. Monocarpic. Leaves

12 FLORA MALESIANA

[ser. I, vol. 9!

Fig. 2. Habit of Harmsiopanax aculeatus (BL.) WARB. ex BOERL. Coarse shrub on old lava-streams in
E. Java (Mt Idjen) at c. 900 m altitude (Photogr. VAN STEENIS).

rounded, up to 30 by 40 cm, deeply palmately
lobed, cordate at base, lobes 5—9 with broad sinuses
between them, margin unevenly and sharply den-
tate, apex acute, upper surface densely covered with
evenly spaced bristles of varying size (larger on the
main veins), appressed and directed towards the
leaf margin, often with woolly hairs inserted on
their enlarged bases, the underside very densely
woolly and with many bristles, usually bearing
crisped hairs on their enlarged bases; petiole

50 cm, !/, cm @ at base, terete with clasping base,
densely covered with bristles, woolly hairs, and
spines. Panicle at first with numerous leaf-like
bracts, the principal branches with some spines,
rather sparsely covered with bristles and hairs,
ultimate branches slender and tomentose, bearing
linear bracts c. 4 mm long subtending peduncled
umbellules; peduncles up to 5 mm, slender, tomen-
tose, bearing 2 minute bracts. Umbellules spheri-
cal, c. 4-5 mm @ in flower, outer bracts not form-

1979]

ARALIACEAE—I (Philipson) 1)

ing a distinct involucre. Flowers hermaphrodite,
maturing in basipetal succession, the lower bracts
of a branch either with sterile umbellules or lacking
flowers; up to 60 in an umbellule, each subtended
by a lanceolate ciliolate bract c. 1 mm long, and
borne on a glabrous pedicel c. 1'/, mm long.
Calyx rim fringed with many lacerate filaments.
Petals ovate, c. 1 mm long. Filaments c. 1 mm;
anthers c. '/, mm long. Ovary covered with cilia
which lengthen as the fruit ripens. Mericarps with
rounded ribs, long-ciliate, crowned by the diver-
gent styles.

Distr. Malesia: Papua New Guinea (Madang
Distr., Western Highlands, Morobe Distr. &
Central Distr.).

Ecol. Forested hills, grassy slopes, and road-
sides, 100-1800 m.

Vern. Opme, Ganja, Mt Hagen, mafiong,
Sattelberg, Morobe Distr.

Note. Information about the habit is inade-
quate. The stalked spherical umbellules are very
distinctive.

%

3
.
4
3
®
.

3. Harmsiopanax ingens PHILIPSON, Blumea 21
(1973) 84.

ssp. ingens. — Fig. 1, 3.

Unbranched tree up to 18 m witha thick or some-
times slender trunk densely covered, except
towards the base of mature specimens, with long,
sharp, upwardly directed spines and marked with
leaf-scars. Monocarpic. Leaves usually peltate,
rounded, up to 1 m @, deeply palmately lobed,
lobes usually with minor lobes and coarsely
dentate, apex acute, upper surface bearing few to
many long spines, especially on the midrib and
principal veins between which the surface is often
rugose and glabrous except for the remains of a
tomentum of branched hairs, or with many bristles
often with woolly hairs on their bases, the under-
surface also with few to many long spines and
usually clothed with a fawn or greyish woolly
tomentum of branched hairs, or densely furnished
with bristles usually with woolly hairs on their
bases, or occasionally glabrous between the spines

Fig. 3. Harmsiopanax ingens PHILIPSON. Left: apex of leafy stem; right: the large inflorescence (Photogr.
FRODIN, Murmur Pass, 1971).

14 FLORA MALESIANA

[ser. I, vol. 91

except for a few bristles; petiole up to 1 m and
3 cm @, terete with clasping base, covered with
woolly hairs and bearing many spines. Panicle up
to 5 m long and 5 m wide, leafless or with lobed
bracts c. 10-20 cm long, principal branches spiny
especially below, ultimate branches slender,
tomentose, bearing linear bracts c. 1 cm long sub-
tending peduncled or sessile umbellules; peduncles
elongating as the fruit ripens, up to 4 mm, rather
stout, tomentose, bearing 1 or 2 minute bracts.
Umbellules bowl-shaped, c. 6-10 mm @ in flower,
enlarging slightly in fruit, with an involucre of
about 8 ovate bracts, 2-4 mm long and ciliolate
distally. Flowers hermaphrodite, maturing in
basipetal succession, terminal branches bearing
maturing fruit while lower branches bear flowers or
unopened buds; usually c. 12-16 (8-20) in an
umbellule each subtended by an involucral bract
or a narrower receptacular bract and borne on a
glabrous pedicel 1-2 mm long. Calyx rim fringed
with many lacerate filaments. Petals ovate, 1-2 mm
long. Filaments 2-31/, mm; anthers 4/,—3/, mm
long. Ovary covered with cilia which lengthen as
the fruit ripens. Mericarps with rounded ribs, long-
ciliate, crowned by the divergent styles.

Distr. Malesia: New Guinea (NW. Irian and
extending along the central mountains from the
Orion Mts to the Owen Stanley Range, Murray
Pass).

Ecol. Montane and mossy forest and second-
growth forest, 2000-3600 m, occasionally rather
lower.

Vern. Papua: mauku, Huli; Mandated Terr.:
Sepik Distr., kKamul, Hindenburg Ra.; Western
Highlands: murri, Hagen, tolsan, Minj, mauri,
Melpa, mai, Mendi, kinogore, makua, makw, Enga;
Eastern Highlands: kimu, Ka, ollu, Chimbu.

Notes. A striking, single-trunked, monocarpic
tree bearing immense inflorescences. The bark is
described as grey brown and the wood white witha
wide pith. The inflorescence has the appearance of
bearing female flowers above and male flowers
below, but this is evidently due to a basipetal
sequence of anthesis. The terminal flowers have

stamens when freshly opened and all those on
lower branches bear styles. Apparently, the female
organs of the lower branches are functional because
branches from mature inflorescences bear fruit
uniformly. Nevertheless, herbarium specimens
cannot adequately represent such a large inflores-
cence so that the possibility remains that some
female-sterile flowers occur in this species.

Variation occurs in both tomentum and in-
florescence characters. For example, most speci-
mens from West Irian have small umbellules and
fewer leaf-spines. In the eastern part of the Eastern
Highlands District a number of gatherings display
a series of variations: the under-leaves give the
appearance of being glabrous between bristles, the
inflorescence branches bear small leafy bracts, the
umbellules are sessile, with rather numerous
{c. 18-21) small flowers subtended by rather broad
bracts. Specimens from Mt Otto show all these
features combined, but other specimens from this
region diverge from the typical state in only some of
these characters. No specimens of this subspecies
are known from the Finisterre Range and only one
from the Owen Stanley Range.

ssp. moniliformis PHILIPSON, Blumea 21 (1973) 86.

Umbellules disposed irregularly along the
branches, singly or in small groups, with bare spaces
intervening, sessile; flowers usually c. 20-30 per
umbellule, floral parts smaller than in ssp. ingens;
fruiting heads rather small (c. 5 mm @).

Distr. Malesia: Papua New Guinea (districts
bordering on the Huon Gulf).

Vern. Morobe Distr.: mobian, Finschhafen.

Note. This subspecies occurs at lower altitudes
than is usual for ssp. ingens {1500-2000 m). No
specimens of either subspecies have been collected
from higher altitudes in the mountains north of the
Markham River and the Huon Gulf. At higher
altitudes in the Owen Stanley Range ssp. ingens is
known from one gathering. The most south-easterly
gathering at present known (CARR 13603) has a
distinctive appearance due to the straight rigid in-
florescence branches with small sessile umbellules.

2. ARALIDIUM

Mia. Pl. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1 (1856) 762, t. 13; Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 25; Bru. in B. & H. Gen. Pl. 1 (1865) 936; HEmsL. in Hook. Ic.
Pl. 16 (1886) t. 1549; Bogert. Handl. 1 (1890) 631; Harms in E. & P. Nat. Pf.
Fam. 3, 8 (1894) 60; Hutcu. Gen. FI. Pl. 2 (1967) 59. — Fig. 4.

Unarmed tree or shrub with simple, exstipulate pinnately lobed, irregularly
incised, or entire leaves. Inflorescence a large panicle, with cymules of small
flowers arranged racemosely on the branches. Pedicels articulated below the ovary.
Dioecious. Male flowers: calyx 5; petals 5, imbricate; stamens 5, anthers dorsifixed.
Female flowers: calyx and corolla similar to male; staminodes 5; ovary with 3

1979] ARALIACEAE—I (Philipson) 15

Fig. 4. Aralidium pinnatifidum (JUNGH. & DE VriesE) Mia. a. Habit, x 2/;, b. 3 flower and bud, c. 2 flower
and bud, both x 8, d. fruit, seed, and CS, slightly enlarged (a VAN BALGOOY 2185, b CocKBURN FRI
8376, c SINCLAIR 9884, d fresh material). Drawn by W. R. PHILIPSON.

16 FLORA MALESIANA [ser. I, vol. 91

locules (2 abortive) and 1 ovule, styles 3-4, tapering from broad bases, stigmas
terminal. Fruit drupe-like. Seed solitary, pendulous from a thickened funicle,
4-5-grooved; endosperm deeply ruminate.

Distr. Monotypic. Peninsular Thailand and Malesia: Malay Peninsula, Sumatra, and Borneo.

The record from Java by Mique (Ann. Mus. Bot. Lugd.-Bat. 1, 1863, 25) is erroneous (see K. & V.
Bijdr. 7, 1900, 2).

Ecol. Primary and secondary forest, from sea-level to c. 1250 m.

Notes. The single species forms a genus with several unique features, namely the large, simple, charac-
teristically pinnately lobed leaves, the absence of resin ducts, the diffuse panicles of male or female
flowers, the 3-carpellate ovary with a single surviving loculus resulting in a single-seeded fruit, the dorsal
raphe, the deeply ruminate endosperm, and the enlarged funicle.

The genus is treated here as a member of the Araliaceae mainly as a matter of convenience. Sometimes
it has been placed in the Cornaceae (e.g. RiDL. Fl. Mal. Pen. 1, 1922, 894, and Vicuter, Ann. Sc. Nat.
Bot. 4, 1906, 171), and on full investigation it may well prove to be better placed in that family. The absence
of resin ducts and the dorsal raphe strongly support a relationship with the Cornaceae and its immediate
allies, though the absence of borders to the pits of the xylary fibres is characteristic of Araliaceae.

Several genera formerly placed in the Cornaceae have now been elevated to the rank of family. If this
course is followed then Aralidium should also be segregated. Many of the features of Aralidium approach
those of Griselinia (segregated as Griseliniaceae) and possibly these two genera should be united as a

single family.

1. Aralidium pinnatifidum (JUNGH. & DE VRIESE
Mia. FI. Ind. Bat. 1, 1 (1856) 763; HEMsL. in Hook.
Ic. Pl. 16 (1886) t. 1549; BoerL. Handl. 1 (1890)
631; Ripv. Fl. Mal. Pen. 1 (1922) 895; Puiieson,
J. Bot. 78 (1940) 118. — Aralia pinnatifida JUNGH.
& DE VRIESE, Ned. Kruidk. Arch. 1 (1846) 15; Ann.
Sc. Nat. III, 6 (1846) 115. — A. dentatum Mia.
Sum. (1861) 340. — A. integrifolium Herne in
Fedde, Rep. 54 (1951) 245. — Fig. 4.

Shrub or small tree up to c. 10 m, rarely reaching
20 m and 25 cm @, glabrous in its vegetative parts;
buds enclosed in long sheathing leaf-bases. Leaves
spaced with distinct internodes, usually c. 30 by
22 cm or more, + regularly pinnately incised,
frequently as deep as the midrib, lobes oblong-
acuminate and decurrent on the midrib, c. 2!/, cm
wide or more, the lobing sometimes irregular, and
occasionally the blade entire and broadly ovate
(up to 25 by 20 cm) or rarely lanceolate, leaf mar-
gin either entire or coarsely dentate, especially on
the terminal lobe; petioles 5-12 cm, broadly
channelled above, clasping the stem with a slightly
dilated base, exstipulate. Panicles terminal, or
occasionally in the upper axils, to 50 cm long,
pendulous, puberulous; main bracts caducous, but
the minute bracteoles often persisting until
anthesis. Flowers numerous, small (buds c. 2!/, mm
long), fragrant, creamy or red-tinged, ovary, calyx
lobes and petals densely covered in a minute but
coarse puberulence. Male flowers with the corolla
persistent during anthesis, petals c. 1!/, mm long,
strap-shaped, spreading, stamens c. 1 mm with
flattened filaments and round anthers; stylopodium
a succulent disk with a concave centre; styles
absent, the ovary 11/, mm long, narrowly turbinate,

without a loculus. Female flowers with the corolla
caducous at anthesis together with the staminodes,
styles divergent from their gibbous bases, ovary
ovate with a single loculus (two abort early); ovule
pendulous. Fruit usually obliquely ellipsoid,
tapering to the apex and c. 3-4'/, cm long, but
rarely subspherical, white when immature, ripening
to purplish or black, juicy; exocarp fleshy, endocarp
chartaceous. Seed broadly ellipsoid, 2—2'/, cm long,
with the surface patterned with deep ruminations.

Distr. Peninsular Thailand; in Malesia: Malay
Peninsula (from Kedah southwards common;
Singapore), throughout Sumatra (also in Simalur
I.), Anambas Is. (Siantan) and throughout Borneo.

Ecol. Frequent in evergreen primary rain-
forest, also in open bamboo forest and secondary
growths, from sea-level to c. 1250 m, in Borneo up
to 1500-1800 m. FI. fr. Jan.—Dec.

Uses. The only use, once mentioned, is from
Brunei, as “leaves make good ghost medicine’’.

Vern. Malay Peninsula: /émpédu buaya, (poko)
balai, pungar, sahalat, sébalai tingal, sibilai,
tébalai, M; Sumatra: (kayu) attarodan, Asahan,
Batak lang., ségéntut, Gajo, médung, M, maneél
silai, mannel dotan, sukun dotan, M, Simalur;
Anambas Is.: ballok, M, Siantan; Borneo: daun
tutchol antu, Brunei, Iban lang.

Note. Entire leaves are not infrequent through-
out the range of the species, so that the recognition
of a second species using this character is not
justified. Coarsely dentate leaf-margins were also
employed as a specific character but are merely a
minor variation. Some specimens from Mt Kina-
balu have rather small globose fruits, but the typical
form of fruit also occurs on that mountain.

1979] ARALIACEAE—I (Philipson) 17

3. ARALIA

LinnE, Gen. Pl. ed. 5 (1754) 134; Sp. Pl. (1753) 273; DC. Prod. 4 (1830) 257; Mia.
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 6; BTH. in B. & H. Gen. Pl. 1 (1865) 936;
BoeRL. Handl. 1 (1890) 629; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 56;
STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 391; Hutcu. Gen. FI. Pl. 2 (1967) 63;
STONE, Gard. Bull. Sing. 30 (1977) 134; PHiLipson, /.c. 97. — Acanthophora MERR.
Philip. J. Sc. 13 (1918) Bot. 316, non LAMOUREUX, 1813 (Algae); STEEN. Bull. Bot.
Gard. Btzg III, 17 (1948) 390. — Fig. 5, 6.

Sparingly branched shrubs or small trees, or climbing, rarely (extra-Mal.)
herbaceous, glabrous or hairy, often prickly. Leaves pinnate to tripinnate, usually
with leaflets at the insertion of the lateral pinnae; leaflets serrate; petiole with a
sheathing base. /nflorescence a terminal panicle; flowers sessile or pedicelled, with
an articulation below the flower; calyx with 5—6 teeth; petals 5—6, imbricate; ovary
2-6-celled; styles 2-6 free or shortly connate below. Fruit a fleshy drupe; pyrenes
cartilaginous, compressed; endosperm uniform.

Distr. More than 30 spp. in North America (S. to Mexico) and East Asia, 6 spp. in Malesia: Sumatra,
Malay Peninsula, Java, Lesser Sunda Is. (Sumba), Borneo, Celebes, Philippines, and West New Guinea.

Ecol. Usually on scrubby hillsides and in secondary growth, often in ravines or near streams, or in
thickets near or above the limit of tree-growth, at low altitude (100 m), but usually in the montane zone,
up to 3000 m.

Note. For a discussion of specific distinctions see VAN STEENIS, /.c. 391. Hut-Liw Li in Sargentia 2
(1942) 101, treated some species that extend into Malesia. MERRILL considered that the climbing habit and
recurved spines of Acanthophora justified its separation as a distinct genus, but more recent authors have
not agreed.

KEY TO THE SPECIES

1. Flowers sessile (capitate) or very shortly pedicelled.
2. Flowers sessile, underside of leaf + densely tomentose, hairs of the branches and inflorescence +
appressed and felted, bracts around the capitula enveloped in hairs 1. A. dasyphylla
2. Flowers short-pedicelled, underside of leaf sparsely tomentose, hairs of the branches and inflorescence
+ patent, bracts around the capitula less densely tomentose 2. A. javanica
1. Flowers +/,-11/, cm pedicelled (umbellate).
3. Climbing or scrambling liana, spines curved
3. Erect shrubs or small trees, spines straight.
4. Leaflets glaucous beneath, margins with few crenations. Fruit small (c. 3 mm long)
4. A. bipinnata
4. Leaflets green (or with fawn pubescence) beneath, margins serrate. Fruit rather larger (4-6 mm long).
5. Young parts and undersurface of leaves glabrous (but with small spines) 5. A. ferox
5. Young parts and undersurface of leaves pubescent

Pe oR ee 2 eS ee te ce ee eA

1. Aralia dasyphylla Mia. FI. Ind. Bat. 1, 1 (1856)
751; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9, incl.
var. strigosa Mia. et var. latifolia MiQ.; BOERL.
Handl. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 53;
Koorp. Exk. Fl. Java 2 (1912) 718; Atlas 4 (1916)
f. 673 A-K; Hut-Lin Lt, Sargentia 2 (1942) 20;
BAKH. f. Blumea 6 (1947) 367, incl. var. urticifolia
(BL. ex Mia.) BAKH. f.; BAKH. f. & OosTsTR. in
Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159,

p. 18; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948)
391; NGoc-SANH Bul, Adansonia 4 (1964) 464;
Back. & BAKH. f. Fl. Java 2 (1965) 170; STEEN.
Mt. FI. Java (1972) pl. 3-1; Puitrrson, Gard. Bull.
Sing. 30 (1977) 98; Y.-R. Lina, Acta Phytotax. Sin.
15 (1977) 86. — A. chinensis (non L.) BL. Bijdr.
(1826) 870. — A. urticifolia BL. ex Mia. Ann. Mus.
Bot. Lugd.-Bat. 1 (1863) 9; BoerL. Handi. 1 (1890)
646; K. & V. Bijdr. 7 (1900) 55; Koorp. Atlas 4

18 FLORA MALESIANA [ser. I, vol. 91

@
(9 \, i)

2S

9 08

= od “N

2 @ re Z
Ves)

Fig. 5. Aralia bipinnata BLANCO. a. Upper branches of inflorescence, b. pinna, c. base of petiole, all x 2/.,
d. flower bud, e. fruit, both x 3 (JAcoss 7017). Drawn by W. R. PHILIPSON.

1979]

ARALIACEAE—I (Philipson) 19

(1916) f. 673 L-N. — A. beccarii Riu. J. Mal.
Br. R. As. Soc. 1 (1923) 64. — Fig. 6.

Prickly shrub or small tree, often unbranched,
to c. 5 m; young parts densely brown pubescent.
Leaves forming large rosettes at the summit of the
stems, c. 1 m long (or more), bi- or tripinnate, with
a pair of leaflets (occasionally pinnate) at each
division of the rachis, the petiole, rachis and lateral
rachides prickly or unarmed, densely pubescent;
leaflets subsessile or petiolule c. 5 mm long (or
longer), usually densely pubescent on the lower
surface, less dense above, ovate to oblong-ovate,
c. 5—14(-18) by 3—5(—10) cm, base rounded to sub-
cordate, apex acuminate, margin finely or some-
times coarsely serrulate; petiole c. 40 cm, with an
elongated sheathing base and a small ligule.
Inflorescence a large terminal panicle, 70 cm or
more long, densely brown pubescent, rachis
bearing several secondary branches c. 30-40 cm
long, with ultimate branches arranged racemosely,
bracts ligulate, ending in heads of several sessile
flowers, surrounded by an involucre of small usually
densely pubescent bracts. Calyx with 5 usually
obtuse teeth; petals 5, c. 11/, mm long, glabrous;
stamens 5; ovary c. 2 mm long, glabrous, 5-celled;
styles 5, slightly connate below. Fruit globose,
c. 31/, mm @, ribbed when dry.

Distr. Northwards to southern China; in
Malesia: Sumatra, Malay Peninsula, West and
Central Java.

Ecol. Primary forest and secondary growths in
deep ravines or open hillsides, from low altitude
(c. 100 m) to 2500 m.

Vern. Sumatra: kaju burle lasét, k. sépak-
sipang, k. si marsuga-suga, k. sipang-sipang, sami-
mpadan, M; Java: gorang, osangsing, J, pangang
tjutjuk, S.

Note. The capitulate flowers are characteristic
(see also under A. javanica). The presence of this
species in the Malay Peninsula has often been
overlooked, though it extends into southern China.
Its variability was discussed by VAN STEENIS (1948,
Ec):

2. Aralia javanica Mia. Pl. Jungh. 3 (1855) 420;
Fl. Ind. Bat. 1, 1 (1856) 749; Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 9; BoERL. Handl. 1 (1890) 646;
K. & V. Bijdr. 7 (1900) 55; Koorp. Exk. Fl. Java 2
(1912) 718; Atlas 4 (1916) f. 670; Baku. f. &
OoststrR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 159, p. 17; Back. & BAKH. f. Fl. Java 2
(1965) 170; Puiirson, Gard. Bull. Sing. 30 (1977)
98.

A shrub or small tree, often unbranched, young
parts covered with brown pubescence which per-
sists on the stems and rachides of the inflorescence
as + patent hairs. Leaves tufted at the ends of the
branches, bipinnate with a pair of leaflets at the
divisions of the rachis; leaflets variable in size,
subsessile or the petiolules up to c. 2 cm, blade
ovate to elliptic, up to 18 by 8 cm (usually smaller),

both surfaces with sparse, short, appressed, bristly
hairs, base cuneate to truncate, apex acuminate,
margin finely and unevenly serrulate; petiole
20-30 cm. Inflorescence a large terminal panicle,
rachis bearing several secondary branches c. 30-40
cm long, with the ultimate branches bearing
heads (or subumbellules) of c. 10 flowers, sur-
rounded by an involucre of small linear bracts.
Calyx with 5 small teeth; petals 5; stamens 5;
ovary c. 2 mm long, glabrous, 5-celled; styles erect
at anthesis. Fruit ovoid, c. 5 mm long, ribbed when
dry, with the persistent styles recurved.

Distr. Malesia: West and Central Java (Mts
Papandayan, Malabar, Diéng, Surakarta).

Ecol. Mountain forests, 2000-3000 m.

Note. This imperfectly known species may prove
to be a form of the widespread A. dasyphylla, from
which it appears to differ in the shortly pedicelled
flowers, the sparser leaf-tomentum, the more
patent hairs on the inflorescence branches and the
bracts of the umbellules less thickly enveloped in
hairs.

3. Aralia scandens (MERR.) HA, Nov. Sist. Vyssh.
Rast. 11 (1974) 229; Stone, Gard. Bull. Sing. 30
(1977) 276, f. 1; Puiirson, /.c. 99. — Acantho-
Phora scandens MeErRR. Philip. J. Sc. 13 (1918) Bot.
316; En. Philip. 3 (1923) 236; STEEN. Bull. Bot.
Gard. Btzg III, 17 (1948) 390. — A. ferox (MiaQ.)
Kina, J. As. Soc. Beng. 67, ii (1898) 45; Koorp.
Minah. (1898) 498; MeErRR. En. Born. (1921) 458;
Rip.. Fl. Mal. Pen. 1 (1922) 872; Masam. En.
Phan. Born. (1942) 564.

Prickly scandent shrub, glabrous, reaching a
height of 10 m or more, stems c. 2!/, cm @. Leaves
dispersed (c. 30 cm apart), up to 11/, m long, tri- or
quadripinnate with a pair of leaflets at each
division of the rachis, prickly on the petiole,
rachides, and sometimes on the leaf veins; leaflets
ovate to elliptic-ovate or ovate-lanceolate, petio-
lules 3-10 mm, blade 5-14 by 2!/,-5 cm, base
rounded or subcordate, apex acuminate, margins
finely spinulose-denticulate; petiole to 35 cm, with
an elongated sheathing base and a small ligule.
Inflorescence a large terminal spiny panicle, the
main rachis to c. 60 cm, bearing secondary
branches singly or in whorls, up to 50 cm; ultimate
branches 1-4 cm, subtended by lanceolate bracts,
racemosely arranged, ending in umbellules;
umbellules c. 10—20-flowered, pedicels slender,
10-12 mm, with lanceolate bracts 2-3 mm long,
articulated below the flower. Calyx with 5-6 short
acute teeth; petals 5-6, with a broad base, imbri-
cate; stamens 5-6, filaments c. 4 mm long, anthers
c. 1 mm long; ovary turbinate c. 2'!/, mm long,
5-6-celled, styles 5-6, free or only slightly connate
below, at first erect. Fruit ellipsoidal, c. 5 mm long,
purple to blue-black, deeply furrowed when dry,
crowned by the persistent radiating styles.

Distr. Malesia: Malay Peninsula (Perak, Selan-
gor, Pahang), Sabah (Mt Kinabalu), Philippines

[ser. I, vol. 9!

FLORA MALESIANA

20

Fig. 6. Aralia dasyphylla Mig. Habit, Tjibodas, on slope of Mt Gedeh, W. Java, c. 1400 m altitude

(Photogr. PHILIPson, 1973).

1979]

ARALIACEAE—I (Philipson) 21

(Luzon, Panay, Catanduanes, Mindanao), Celebes
(Menado, Buton I., S. Celebes).

Ecol. Thickets on slopes and mountainsides,
often near streams, or among secondary growths,
180-1550 m.

Uses. In Mindanao is reported that scrapings
of the bark are applied to wounds and a decoction
of the boiled bark is drunk to releave internal
pain.

Vern. Philippines:
simbar, Bag.

Note. The only species with the habit of a liana,
with spaced leaves, and recurved spines. The
flowers are whitish or yellowish, and slightly fra-
grant and are visited by numerous small bees. The
fruit is purple and fleshy.

cwangayan, Mindanao,

4. Aralia bipinnata BLANCO, FI. Filip. (1837) 222;
Me_rkR. Sp. Blanc. (1918) 294; En. Philip. 3 (1923)
235; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948)
392, incl. f. inermis STEEN.; PHILIPSON, Gard. Bull.
Sing. 30 (1977) 99. — A. hypoleuca PREsL, Epim.
(1851) 250; Mia. FI. Ind. Bat. 1, 1 (1856) 751;
F.-VILL. Nov. App. (1880) 101; VmAL, Phan.
Cuming. Philip. (1885) 117; Rev. Pl. Vasc. Filip.
(1886) 144; Harms, Bot. Jahrb. 23 (1896) 18; MERR.
Philip. J. Sc. 5 (1910) Bot. 369; En. Philip. 3
(1923) 235. — A. javanica (non Mi.) F.-VILL. Nov.
App. (1880) 101. — A. glauca Merr. Philip. J.
Sc. 2 (1907) Bot. 291; En. Philip. 3 (1923) 236. —
A. apoensis ELMER, Leafl. Philip. Bot. 7 (1914)
2325; MERR. En Philip. 3 (1923) 235. — Fig. 5.

A shrub or small, sparsely branched tree to 7 m,
with prickly stems. Leaves to 1'/, m or more long,
forming large crowns at the ends of the branches,
bipinnate, with a pair of pinnae at each division of
the rachis, with some prickles, especially on the
petiole or unarmed, the rachis swollen and articu-
lated at the nodes; leaflets sessile or with a short
petiolule, ovate to lanceolate, usually 4-5 by
2-2'/, cm, but variable in size, apex acute or acumi-
nate, base rounded to cordate, usually markedly
oblique in lateral leaflets, margin conspicuously
crenate, upper surface green, glabrous, lower sur-
face glaucous, pubescent along the veins and some-
times sparingly on the mesophyl!, sometimes only
in the angles of the lower veins, or almost glabrous
throughout, primary and secondary veins con-
spicuous; petiole to 30 cm, base long sheathing and
slightly ligulate. Inflorescence a large terminal
panicle 30-70 cm long (or more), peduncle and
also usually the main branches prickly, the whole
either almost glabrous or pubescent; peduncle
5-18 cm long, stout; main rays c. 5-10, mostly
clustered at the apex of the rachis, 25-65 cm long,
bearing many short tertiary branches along their
length; tertiary branches usually 5-10 cm long,
ending in umbellules, and bearing a small number
of lateral umbellules, or branches, minute lanceo-
late bracts subtending the branches of the third or
higher orders; umbellules with c. 20-30 radiating

pedicels ; pedicels 5-10 mm. Calyx lobes 5, rounded,
1/, mm long; petals 5, 1'/, mm long; stamens 5;
ovary 5-celled, styles subulate, free. Fruit spheroi-
dal, c. 3 by 4 mm, strongly 5-ribbed when dry,
persistent styles spreading.

Distr. Taiwan; in Malesia: Philippines (Luzon,
Leyte, Negros, Mindoro, Mindanao) and West
New Guinea (Vogelkop Peninsula, possibly also in
Swart Valley).

Ecol. In rather open forests, ravines, and in
thickets and secondary growths, (700-)1000-
2450 m.

Vern. Philippines: badbaranai, C.Bis., dasanat,
Neg., karugi, Buk., magkasau, Bis., mara-bauya,
Bag., papang, Bon., sugsuga, Ig.

Note. VAN STEENIS /.c. discussed the vari-
ability in pubescence and the development of
spines.

5. Aralia ferox Mia. Fl. Ind. Bat. 1, 1 (1856) 750;
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9; BoeRL.
Handl. 1 (1890) 629; K. & V. Bijdr. 7 (1900) 49;
Koorop. Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916)
f. 671; Fl. Tjib. 2 (1923) 229; Baku. f. & OoststrR.
in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159,
p. 18; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948)
394; Back. & BAKH. f. Fl. Java 2 (1965) 170;
PHILIPSON, Gard. Bull. Sing. 30 (1977) 99. —
A. filicifolia R1pL. J. Fed. Mal. St. Mus. 8 (1917)
42, non C. Moore, 1876.

Spiny shrub or small tree, usually unbranched,
to c. 10 m. Leaves forming a large rosette at the
summit of the stem, up to c. 1 m long, or shorter
below the inflorescence, bi- or tripinnate, with a
pair of leaflets (often pinnate) at each division of
the rachis, prickly on the petiole, main rachis, and
often on the lateral rachides; leaflets sessile or
petiolules to c. 5 mm, ovate or ovate-oblong,
usually c. 3 by 1°/, cm, but variable in size, base
truncate to rounded or cuneate, apex acute acumi-
nate, margin sharply serrate, both surfaces with
small bristle-like spines, especially on the veins,
sometimes with small spines on the underside of
the midrib; petiole to c. 25 cm, with an elongated
sheathing base and a small ligule. Inflorescence a
large terminal panicle, 25-50 cm long, glabrous,
the main rachis rather short, bearing a few lateral
or a terminal cluster of branches c. 15-25 cm long;
tertiary branches disposed singly or in subverticils,
ending in umbellules and bearing a variable
number of lateral umbellules. Flowers c. 10-12 per
umbellule; pedicels c. 5-6 mm, articulated below
the ovary; calyx a rim bearing 5 narrow or trian-
gular teeth; petals and stamens 5; ovary turbinate
c.2 mm long; styles 5, erect at first, soon spreading,
scarcely united at the base. Fruit spheroidal, c. 6 by
5 mm, deeply furrowed when dry, the persistent
styles radiating.

Distr. Malesia: Central W. Sumatra (Mt
Kerintji), W. Java (Mts Gedeh, Patuha and
Tangkuban Prahu).

22

FLORA MALESIANA

[ser. I, vol. 91

Ecol. In montane scrub and among scattered
trees, 1900-2900 m.

Vern. Pabong, pangang njirvan, panggang-
tjérmé, S.

forma nana STEEN. Bull. Bot. Gard. Btzg III, 17
(1948) 394, f. 1.

Smaller, probably +/,-1 m, leaves tripinnate,
30 cm long, spiny all over; leaflets 4-13 by 2-7 mm,
rachides of the ultimate pinnae winged.

Distr. Malesia: Central W. Sumatra (Mt
Talang).

Ecol. Growing about 2500 m.

Note. VAN STEENIS considered this interesting
dwarf form to represent the extreme of a series in
size variability.

6. Aralia montana BL. Bijdr. (1826) 870; Mia. FI.
Ind. Bat. 1, 1 (1856) 750; Ann. Mus. Bot. Lugd.-
Bat. 1 (1863) 9, incl. var. acutata MiQ.; BOERL.
Handl. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 51;
Koorp. Exk. Fl. Java 2 (1912) 718; Atlas 4 (1916)
f. 672; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948)
391; BAKH. f. & OosTstrR. in Back. Bekn. FI. Java
(em. ed.) 7 (1948) fam. 159, p. 17; BAKH. f. Blumea
6 (1950) 367, incl. var. crassifolia BAKH. f.; BACK. &
BAKH. f. Fl. Java 2 (1965) 170; Puitirson, Gard.
Bull. Sing. 30 (1977) 100. — A. bipinnata REINW.
ex BL. Cat. (1823) 43, nomen; ex DE VRIESE, PI. Ind.
Bat. Or. (1857) 84, nomen in synon. — Panax
armatus WALL. [Cat. (1832) n. 4933, nomen] ex
G. Don, Gen. Syst. 3 (1834) 386. — A. decomposita
REINW. ex DE VRIESE, Pl. Ind. Bat. Or. (1857) 84,
nom. illeg. altern. — A. armata (WALL.) SEEM. J.
Bot. 6 (1868) 134; CLarkE, Fl. Br. Ind. 2 (1879)
723; KING, J. As. Soc. Beng. 67, ii (1898) 44; RIDL.
Fl. Mal. Pen. 1 (1922) 873; Hur-Lin Lt, Sargentia 2
(1942) 106. — A. thomsonii SEEM. J. Bot. 6 (1868)
134; CLARKE, FI. Br. Ind. 2 (1879) 723; Kine, J.
As. Soc. Beng. 67, ii (1898) 44; RipL. Fl. Mal.
Pen. 1 (1922) 873; Hut-Lin Li, Sargentia 2 (1942)
i»

Shrub or small tree, frequently unbranched, with
prickly stems, occasionally attaining a height of
12 m. Leaves to 1 m or more long, forming large
crowns at the ends of the branches, bipinnate, with
a pair of simple or occasionally pinnate leaflets at
each division of the rachis, usually with some
prickles, especially on the petiole, or unarmed, the
rachis constricted at the joints; leaflets sessile or
with a short petiolule, ovate, up to 14 by 7 cm,
apex acute to acuminate, base truncate or rounded,
oblique in lateral leaflets, margin sharply serrate,
upper surface with the remains of a strigose tomen-
tum, often + rugose, lower surface often with a +
velvety tomentum, or with more harsh hairs +
confined to the veins; petiole to 30 cm, its base
sheathing and ligulate. Jnflorescence a large ter-
minal panicle, peduncle and branches tomentose,
prickles, if any, confined to the peduncle and main

rachis, small usually persistent linear or ovate
bracts c. 11/, cm long subtending the branches and
also spaced along the peduncle; bracts of tertiary
branches similar but smaller; secondary branches
at intervals along the main rachis, c. 35 cm long,
bearing numerous tertiary branches along their
length; tertiary branches usually c. 6 cm long,
ending in umbellules, and often bearing a number
of lateral umbellules; umbellules with c. 20-30
radiating pedicels; pedicels usually 12-15 mm,
occasionally shorter, pubescent. Flowers herm-
aphrodite; calyx lobes 5, triangular or rounded;
petals 5, c. 2 mm long; stamens 5; ovary 5-celled,
glabrous; styles subulate, connate below, free and
spreading above. Fruit spheroidal, up to c. 4 by
4 mm, strongly 5-ribbed when dry, surmounted by
the reflexed styles.

Distr. Malesia: Malay Peninsula, Sumatra,
Java, Borneo (Sarawak, Sabah), Celebes, Lesser
Sunda Is. (Sumba).

Ecol. Primary and secondary forest, bamboo
forest and low-lying moist ground, from near sea-
level to 2600 m.

Vern. Gorang, panggang tjutjuk, S; Malay
Peninsula: poko dulang-dulang; Sarawak: tepa
paluk.

Notes. This species is considered to include all
West Malesian examples with pedicelled flowers
and pubescent leaves. This broad concept is
contrary to former treatments which have recog-
nized several species (A. thomsonii, A. armata).
The alliance with A. chinensis L. and A. decaisneana
HANCE is also very close. The application of names
to this and other Javanese species has been very
confused. The position is ably discussed by
VALETON (in K. & V. Bijdr.) and by VAN STEENIS
(/.c.). In most specimens the lower leaf surface and
the pedicels are densely tomentose, but there are
specimens in which the leaf is only sparsely hairy
and the pedicels may be glabrous. A. armata
appears to be within the range of variation of the
complex although this plant is very spiny, its
leaves and inflorescences always being provided
with numerous short spines. The leaflets also are
thinner and smoother and, like the pedicels, are less
densely pubescent (see NGoc-SANH Bul, Adan-
sonia 9, 1969, 461). However, A. armata (if distinct)
has been collected only very rarely in the Malay
Peninsula and only in the extreme north. The most
aberrant specimens are those with glabrous umbel-
lules which are mostly from Sumatra but also from
Java: they may indicate that A. foliolosa SEEM.
should also be included in this complex.

Excluded

Aralia capitulata JUNGH. & DE VRIESE, Ned.
Kruidk. Arch. 1 (1846) 17; Ann. Sc. Nat. III, 7
(1846) 116 is, cf. SLEUMER, FI. Males. I, 7 (1971)
24 = Gomphandra capitulata (JUNGH. & DE
VRIESE) BECC. (Icacinaceae).

1979] ARALIACEAE—I (Philipson) 23

Fig. 7. Delarbrea collina VIEL. a. Habit, x */s, b. flower bud, c. flower and ditto in LS, d. petal, e. stamen,
f. flower after anthesis, x 13, g. fruit and ditto in CS, x 3 (ae, g RIDSDALE NGF 36736, f SOEKMA $.n.).
Drawn by HELENE MULDER.

24 FLORA MALESIANA [ser. I, vol. 92

4. DELARBREA

VIEILL. Bull. Soc. Linn. Norm. 9 (1865) 342, 393; Bru. in B. & H. Gen. Pl. 1 (1865)
935; BRITTEN in Forbes, Nat. Wand. (1885) 506 (see also p. 354); HARMs in E. & P.
Nat. Pfi. Fam. 3, 8 (1894) 61; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 485; Hutcu.
Gen. FI. Pl. 2 (1967) 63. — Fig. 7.

Glabrous unarmed shrubs or small trees. Leaves large, imparipinnate, with a
stipular sheath; leaflets alternate or opposite, entire or indistinctly dentate. Flowers
in umbellules grouped in large terminal panicles; pedicels articulated below the
flower. Calyx lobes 5. Petals 5, imbricate, obovate, narrowed towards the base.
Stamens 5, filaments stout, anthers dorsifixed. Ovary inferior, 2-celled, disk fleshy,
obconic, crowned by two erect styles with clavate stigmas. Fruit ovoid, crowned
by the small calyx lobes and the recurved style arms (which eventually fall);
exocarp thin, fleshy, with peripheral oil vesicles; endocarp papery; endosperm
with shallow longitudinal grooves not ruminate.

Distr. Queensland, Melanesia and East Malesia, 3 or 4 spp., from the Lesser Sunda Is. eastwards to
New Guinea (also New Britain), Queensland, Solomons, New Caledonia, and New Hebrides. In Malesia
1 sp.

Ecol. Lowland to montane forest.

Note. The corolla is distinctive, the petals being strongly imbricate and narrowed towards their
insertion. The fruit also has a characteristic appearance, since the calyx and stylopodium, although
persistent, do not enlarge as in most other araliads. The fruit, therefore, is a smooth ellipsoid berry without
a prominent rim around the apex.

1. Delarbrea collina VIEILL. Bull. Soc. Bot. Norm.
9 (1865) 342; PuiLipson, Bull. Br. Mus. Nat. Hist.
Bot. 1 (1951) 18. — D. sp. HEMSL. Rep. Challenger,
Bot. 1, pt 3 (1885) 155. — D. paradoxa (non
VIEILL.) BRITTEN in Forbes, Nat. Wand. (1885)
506. — D. lauterbachii HARMs in K. Sch. & Laut.
Fl. Schutzgeb. (1900) 485.

Sparsely branched shrub to 5 m high, with the
multijugate leaves clustered at the ends of the
branches. Leaves c. 70-100 by 30-40 cm; rachis not
articulated; leaflets alternate or in pairs, c. 7 on
each side; petiolules c. 1 cm long; lamina c. 17-20
by 4-6!/, cm, lanceolate, ovate, oblong or elliptic,
gradually tapered to an acute apex, base truncate,
rounded or cuneate, usually oblique, margin
entire; petiole c. 17-20 cm, terete, lenticellate, with
a heavily lenticellate clasping base with mem-
branous margins. Inflorescence a terminal panicle
of umbellules, rachis up to 60 cm long, bearing
well-spaced secondary branches 6-25 cm long,
bracts caducous; tertiary branches c. 2-8 cm long,
terminating in a circlet of broadly ovate bracts
(mostly caducous) surrounding the umbellules,
sometimes with smaller (male) lateral umbellules;
umbellules c. 2 cm @ at anthesis, with c. 30-40
flowers. Pedicels c. 5 mm (elongating to 10-15 mm
in fruit), pustulate. Calyx lobes 5, obtuse, united
below into a tube. Petals 5, c. 11/, by 3/, mm, keeled
within. Stamens 5, 1 mm long. Ovary sometimes
prominently ribbed when dry, c. 2 mm long; disk

and styles c. 1 mm high at anthesis. Fruit 16 by
10 mm, purplish black when mature.

Distr. Solomon Is. to New Caledonia and
Queensland; in Malesia: Lesser Sunda Is. (Timor,
Wetar, Babar), Moluccas (Tenimber Is., Banda),
New Guinea (Aru Is., Kar Kar Is., Madang, New
Britain). Fig. 8.

Ecol. Rain-forest, from sea-level to 1000 m.

Vern. Don, Madang.

Notes. The most wide-ranging of any Malesian
araliad. It was collected in Malesia by ForBEs in

Fig. 8. Range of Delarbrea collina VIEILL., localities
in Malesia dotted.

1979]

ARALIACEAE—I (Philipson) 25

Timor in 1882, when it was incorrectly identified
as the New Caledonian species D. paradoxa
VIEILL. Eight years earlier it had been collected in
the Aru Is. during the Challenger Expedition
(HEMSLEY, /.c.) and also been referred as close to
D. paradoxa. Nearly twenty years later LAUTER-
BACH collected it in the Moluccas, when HARMS
described it as a new species. The statement by
HaArMs that the genus was known previously only
from New Caledonia cannot be reconciled with his
note in the Pflanzenfamilien in which he recorded

BRITTEN’s report of it in Timor. Solomon Islands
collections were identified as D. collina VIEILL. by
PHILIPSON in 1951. The arrangement of the um-
bellules differs in the panicles of D. collina and
D. paradoxa. All the material from Malesia con-
forms to the characters of D. collina. The plant is
evidently rare, few collections having been made
in spite of its wide distribution.

It was formerly cultivated in the Botanic Garden
at Bogor until about 1958, having been introduced
from Banda.

5. PENTAPANAX

SEEM. J. Bot. 2 (1868) 294; HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 55; Koorp.
Bull. Jard. Bot. Btzg III, 1 (1919) 181; Hurcu. Gen. FI. Pl. 2 (1967) 63; NGoc-SANH
Bul, Adansonia 9 (1969) 389; PHILIPsoN, Austrobaileya 1 (1977) 23. — Fig. 9.

Trees or shrubs, often scandent, unarmed. Leaves imparipinnate, exstipulate,
glabrous. Flowers in racemes or umbels, which are arranged in panicles or com-
pound umbels, pedicels articulated below the ovary. Calyx 5. Petals 5, imbricate in
the bud. Stamens 5. Ovary inferior, 5-celled, disk + conical, surmounted by the
styles which are united their whole length or become free down to half their length.
Fruit globose; exocarp leathery, enclosing crustaceous pyrenes. Seeds compressed,
endosperm smooth.

Distr. About 14 spp. in India, Thailand, Vietnam, Ceylon, Burma, southern China, Taiwan, in Malesia:
1 sp. locally in E. Java.
South American species formerly included are best excluded, and the 2 Queensland spp. are now

referred to Polyscias.
Ecol. Forest and scrub.

1. Pentapanax elegans Koorpb. Bull. Jard. Bot.
Btzg III, 1 (1919) 182, pl. 16 & 17; Baku. f. &
Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 159, p. 17; BAcK. & BAKH. f. Fl. Java 2 (1965)
169. — Fig. 9.

var. elegans.

Epiphytic scrambler or terrestrial shrub up to
10 m, with unarmed branches, leaf and flower
buds separate, enclosed in + persistent imbricated
cataphylls. Leaves disposed along the branches;
petiole c. 6-10 cm, flattened above, base scarcely
dilated, and sometimes minutely fimbriated, articu-
lated with the rachis, and the rachis articulated with
the petiolules, articulations minutely fimbriate,
rachis to 5 cm, petiolules of lateral leaflets up to
5 mm, of terminal leaflet to 20 mm, leaflets 5 or
fewer, ovate to oblong-elliptic, the lateral some-
times oblique, up to 10 by 7 cm, usually c. 5 by
2'/, cm, apex acute, base rounded or cuneate,
margin entire or with subulate teeth, glaucous
beneath. Inflorescence terminal with persistent
cataphylls at the base of the main axis, umbels
solitary or 1-4 smaller (apparently male) lateral

umbels arising from the axils of minute bracts on
the rachis; rachis 8-16 cm, slender, glabrous;
terminal umbel 3-6 cm @, many-flowered; pedicels
1/,-3 cm, filiform, glabrous, with minute brac-
teoles surrounding the articulation below the
flower. Calyx lobes ligulate, obtuse, c. 1 mm long;
petals triangular to ligulate c. 2 mm long; filaments
yellow, 3 mm, anthers purple, 1/, mm long. Ovary
broadly obconic, surmounted by a stylar column,
2 mm long. Fruit globose, 3-4 mm @, disk broadly
conical, crowned by the persistent calyx lobes and
an awl-shaped stylar column which may divide at
apex.

Distr. Malesia: E. Java (Mts Ardjuno and
Tengger), the variety in Thailand.

Ecol. Uncommon, in light forest or scrub,
including Casuarina junghuhniana forest, 1700-
2600 m.

Note. A remarkably isolated species of a genus
otherwise unrepresented in Malesia.

var. pubescens Koorb. Bull. Jard. Bot. Btzg III, 1
(1919) 183 (as var. puberula in f. 17); NGoc-SANH
Bul, Adansonia 9 (1969) 389.

26 FLORA MALESIANA [ser. I, vol. 91

WY © & A
‘ %
iY .
aN
6
as ae

Fig. 9. Pentapanax elegans Koorv. a. Habit, x 1/, b. flower bud, c. flower in anthesis, d. ovary in CS,
enlarged (VAN STEENIS 10879). Drawn by P. PRENDERGAST.

1979] ARALIACEAE—I (Philipson) 27

Peduncle and pedicels tomentose. Note. Since all specimens from Mts Tengger

Distr. Thailand; in Malesia: E. Java (Mt Jang). and Ardjuno lack pubescence, the retention of the

Ecol. In Casuarina junghuhniana forest, scat- variety appears justified. However, more collections
tered, 1900-2300 m. are required from all localities.

6. MACKINLAYA

F.v.M. Fragm. 4 (1864) 119; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 62; Bot.
Jahrb. 56 (1921) 413; PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 3; Hutcu.
Gen. FI. Pl. 2 (1967) 65. — Anomopanax Harms [in Dalla Torre & Harms, Gen.
Siph. (1903) 364, nomen;] Ann. Jard. Bot. Btzg 19 (1904) 13; in K. Sch. & Laut.
Nachtr. (1905) 332; in E. & P. Nat. Pfl. Fam. Nachtr. 3 (1908) 255; Bot. Jahrb. 56
(1921) 414; Hutcu. Gen. FI. Pl. 2 (1967) 59. — Fig. 10.

Glabrous unarmed shrubs, often unbranched (sympodial). Leaves with a petiole
having a dilated sheath encircling the stem and (in dried material) a constriction at
the apex, and with a leaf-blade either unifoliolate or digitately compound, the
central leaflet, or the three central leaflets, sometimes digitately lobed or compound.
Inflorescence terminal (but sympodium often continued by axillary branching), the
peduncle bearing umbellately arranged branches which terminate either in umbel-
lules or in cymes. Flowers male or hermaphrodite, the male flowers either in
distinct inflorescences or towards the periphery of mixed inflorescences. Pedicel
articulated below the flower. Calyx lobes 5—6, triangular or lanceolate. Petals 5-6,
narrowed below into a distinct claw, and above into a long incurved process.
Stamens 5-6; anthers subglobose. Ovary inferior, with two uni-ovulate cells.
Disk prominent, with a crenulate margin. Styles 2, subulate, free, recurved in fruit.
Fruit strongly compressed, 2-seeded (or one aborted), with a longitudinal furrow
between the seeds; exocarp leathery, endocarp cartilaginous. Endosperm smooth.

Distr. 5 spp., Solomon Is., Queensland, and in Malesia (3 spp.): Philippines, Celebes and throughout
New Guinea.

Ecol. Understorey of rain-forest and montane forest, or epiphytic. Also in secondary growth.

Notes. The leaves are palmately divided or they may be reduced to a single leaflet, especially on the
upper branches of M. schlechteri. The central leaflet, or the three central leaflets, are either lobed or com-
pound, a character rarely found in other genera of this family. Another foliar character rare in the family
is the insertion of the leaf-sheath round the whole circumference of the stem. This character occurs also
in a section of Polyscias, but is more typical of Umbelliferae. The narrow base of the petal is also very rare
in the family, but is characteristic of Umbelliferae. The constantly 2-celled ovary is also typical of Umbel-

liferae, but other characters of the fruit appear to justify the retention of these plants in the Araliaceae.
Reasons for regarding Anomopanax as congeneric with Mackinlaya are given by PHILIPSON (i.c.).

KEY TO THE SPECIES

1. Ultimate branches of the inflorescence in irregularcymes ............ 1. M. celebica
1. Ultimate branches of the inflorescence in umbellules.
22 Primary, fays of the inflorescence many (30-50) < <= sss). 2 2 = eee ee 2. M. radiata
> Prumary rays.of the inflorescence ‘¢.,15 ordewer 2) a0-e 4 ee eee 3. M. schlechteri

1. Mackinlaya celebica (HARMS) PHILIPSON, Bull. (1904) 15. — Anomopanax warburgii Harms, /.c.
Br. Mus. Nat. Hist. Bot. 1 (1951) 8. — Anomopanax 15. — M. amplifolia Hemst. Kew Bull. (1909)
celebicus Harms, Ann. Jard. Bot. Btzg 19 (1904) 260; Harms, Bot. Jahrb. 56 (1920) 413. — Anomo-
14; Ic. Bog. 2 (1906) t. 176 & 177. — Anomopanax panax arfakensis Gripes, Arfak (1917) 163. —
philippinensis Harms, Ann. Jard. Bot. Btzg 19 Anomopanax digitata Merr. Philip. J. Sc. 17

28 FLORA MALESIANA [ser. I, vol. 92

Fig. 10. Mackinlaya celebica (HARMS) PHILIPSON. a. Part of inflorescence, x ?/., b. leaf, c. leaflet, xis
d. flower cluster, e. flower, x 7, f. petal, x 13, g. fruit, x 5/; (BRAss 28056). Drawn by HELENE MULDER.

1979]

(1920) 301. — Polyscias cibaria WHITE & FRANCIS
ex LANE-POOLE, For. Res. (1925) 129, descr. angl.
minim. — Anomopanax variifolius C. T. WHITE,
J. Arn. Arb. 10 (1929) 256. — M. digitata (MERR.)
PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951)
7. — M. warburgii (HARMS) PuHiuipson, /.c. 8. —
Fig. 10.

Shrub or small sparsely branched tree to 6 m.
Leaves very variable in size and complexity; petiole
up to 52 cm, 1'/, cm wide, terete, striate, with a
membranous base ensheathing the stem; leaflets 5
(rarely 3) or the central petiolule (or the central
3 petiolules) frequently dividing to bear three, or
more rarely 5 leaflets; lateral petioles short (c. 1-2
cm), the three central longer (up to 12 cm); lamina
elliptic or ovate, up to 48 by 22 cm, base abruptly
attenuated into the petiolule or subcordate, apex
acuminate or gradually narrowed, acute, margin
entire, denticulate or coarsely serrate, especially
towards the apex, membranous, lamina of the
terminal leaflet (and less frequently of the central
three leafiets) sometimes deeply 3-lobed or with
3-5 separate leaflets, of which the lateral are
markedly oblique at the base. Inflorescence a
terminal compound umbel, very variable in size,
either entirely of male flowers or with male and
hermaphrodite fiowers, often overtopped by sym-
podial growth; peduncle terete, striate, stout, up to
30(-45) cm, 3/, cm 2, bearing lanceolate bracts
below the rays; primary rays c. 9-18, 10-20 cm,
striate, with distal small linear bracts; secondary
tays about 5-10, 3-6 cm, dividing again (often
repeatedly) either umbellately or in an irregular
cymose manner, the central ray frequently more
strongly developed. Calyx lobes 5, triangular,
c. 1 mm long. Petals 5, obovate c. 11/, mm long.
Filaments c. 11/, mm, anthers small. Ovary obconic,
1-2 mm long, narrowly turbinate in male flowers,
ovoid and quickly swelling in female flowers. Fruit
up to 2!/, by 3 cm, compressed, rotund, constricted
in the mid-axis, the two halves frequently unevenly
developed.

Distr. Solomon Is.; in Malesia: New Guinea
(incl. New Britain and Aru Is.), Celebes, and
Central & S. Philippines.

Ecol. Rain-forest, open hill forest, and montane
forest, also in secondary growths, from sea-level to
1450 m.

Uses. LANE-POOLE (/.c.) recorded that at Mt
Obree leaves and flowers are cooked with coconut
oil and put in armlets in dances.

Vern. Philippines: binlaon, C.Bis., pararau,
Bag., tagima, Sub., lumot-lumot, Mindanao; New
Guinea: bugini, wale, yam bonga, Sepik Distr.,
lak-lak, W. Highlands, po’undo, S. Highlands,
Papua, nere, Central Distr., Papua, narona, New
Britain.

Notes. The leaves have a strong parsley-like
odour. The flowers are creamy white, and the
fruits blue to purple with a glaucous bloom. Salt is
said to be obtained from the ashes of the leaves.

ARALIACEAE—I (Philipson)

29

Although collected frequently in the Philippines
and New Guinea, this species is unrecorded for the
Moluccas. There is considerable variation in the
size of the leaves and of the inflorescence. A few
New Guinea specimens are intermediate in charac-
ter between this and the equally common M.
schlechteri, and are interpreted as hybrids.

2. Mackinlaya radiata PHiipson, Bull. Br. Mus.
Nat. Hist. Bot. 1 (1951) 6.

Slender shrub to 5 m. Petiole c. 20 cm, terete and
finely striate, base ensheathing the stem, mem-
branous. Leaflets 5, or the central petiolule bearing
three leaflets, the two lateral petiolules short (1-
11/, cm), the three central longer (6-8 cm, or the
midpetiolule to 11 cm); lamina of the lateral
leaflets elliptic or ovate, up to 20 by 12 cm, base
abruptly attenuated into the petiolule, apex
gradually narrowed, acute, margin entire or
minutely denticulate towards the apex, mem-
branous; lamina of the central leaflet similar or
deeply 3-lobed or with 3 separate leaflets of which
the lateral are strongly oblique at the base. Jn-
florescence a terminal compound umbel; peduncle
terete, striate, stout, from 20 cm to considerably
longer, 4-6 mm @, bearing lanceolate bracts below
the rays; primary rays numerous (c. 30-50),
9-18 cm, slender, striate, with distal minute, linear,
caducous bracts; secondary rays (pedicels)
numerous (35-130), filiform, 1-2 cm; outer
flowers male, central hermaphrodite. Calyx lobes
5, narrowly triangular, c. '/, mm long. Petals 5,
obovate, c. 1 mm long. Ovary narrowly obconic
in male flowers, ovoid in hermaphrodite flowers,
c. 0.7 mm long. Fruit (immature) ovate, com-

pressed.
Distr. Malesia: New Guinea (NW. Irian;
Sepik Distr.).

Ecol. Montane rain-forest and mossy forest,
900-1200 m.

Vern. Apiyetimber, Sepik Distr.

Note. Flowers creamy white.

3. Mackinlaya schlechteri (HARMS) PHILIPSON,
Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 7. — Anomo-
panax schlechteri Harms in K. Sch. & Laut.
Nachtr. (1905) 332, t. 13. — Anomopanax versteegii
Harms, Nova Guinea 8 (1910) 276. — M. versteegii
(HARMS) PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot.
1 (1951) 7. — M. brassii Puitipson, I.c. 6. —
M. klossii Putrrson, I.c. 6. — M. subulata
PHILIPSON, /.c. 7.

Slender shrub to 6 m. Leaves 1-3{rarely 4-)folio-
late. Petiole usually less than 10 cm, but occasion-
ally longer (to 20 cm), especially in compound
leaves, terete and finely striate, base ensheathing
the stem, membranous. Lamina elliptic obovate, or
oblong, occasionally irregularly lobed, up to 28 by
12 cm but usually considerably smaller, base
cuneate or truncate (of lateral leaflets often

30 FLORA MALESIANA

[ser. I, vol. 9?

On nw
y ch W 3 4 iF (\ fd
ke IZ e iy
“1 ‘ae
PF
{ D i : Yale Y
50 0 aS aigtZ XV D .
0 Soy aoa 4:
At Wy * Wa
a ‘ my * sf 2
eo" ys \ » Art 1A /
rY bes eh | Y
*. v QQ §
SIG GS pr } \
WW SS}
| g 5 % ° Z
| £ NN \ f

Fig. 11. Osmoxylon novoguineense (ScuerF.) Becc. a. Leaf half, b. base of petiole, both x 2/;, c. inflores-
cence, x */;, d. flower and CS of ovary, x

4, e. false fruit and ditto in CS (CRAVEN & SCHODDE 789).
Drawn by W. R. PHILIPSON.

1979]

ARALIACEAE—I (Philipson) 31

oblique), apex shortly acuminate, acute, margin
entire or dentate towards the apex, membranous or
chartaceous. Inflorescence a terminal compound
umbel, often overtopped by a lateral branch at its
base; peduncle terete, striate, c. 10-20 cm, bearing
small lanceolate bracts below the rays; primary
rays 6-15, c. 4-7 cm, striate, with minute distal
caducous bracts; secondary rays (pedicels) c. 10-20,
filiform or rather rigid, usually 5-12 mm; male
flowers towards the outside of the umbellules.
Calyx lobes 5, triangular to subulate, '/,-1 mm
long. Petals 5, obovate. Ovary narrowly turbinate,
in male flowers obconic or ovoid, c. */,; mm long in
hermaphrodite. Fruit large, 15 by 22 mm, com-
pressed, rotund, constricted above and below on
the central axis; styles persistent, recurved.

Distr. Malesia: New Guinea (along the Central
Ranges, from the Star Mts east to Meyamya), also
in New Britain.

Ecol. Rain-forest and montane forest, 600—
2300 m.

Uses. The cut stem exudes a viscous sap which
is an irritant. The leaves are aromatic. The plant is
reported to be poisonous and to have a number of
medicinal uses. The boiled leaves are eaten to

reduce fever and to relieve ‘korima’. Pieces of leaf
placed in a cavity relieve toothache. The leaves are
wrapped around taro at planting to encourage
growth.

Vern. Dako, Wissel Lakes, kolobang, kulbang,
Sepik Distr., auke, kenata, muklofo, E. Highlands,
narona, New Britain.

Notes. The flowers are white and the ripe fruit
mauve to purple with a glaucous bloom.

A large number of collections made in recent
years throughout New Guinea all have regularly
compound umbels with the flowers borne on
branches of the third degree in the form of strict
umbels. In two of the earliest gatherings (SCHLECH-
TER 14365 and VERSTEEG 1419) the third degree
branches frequently divide again either umbellately
or cymosely. These two specimens were described
as species by HARMS. PHILIPSON later (1951) kept
the forms with regular umbellules separate (de-
scribing three species). All these five entities are now
considered conspecific, the SCHLECHTER and
VERSTEEG specimens being regarded as rare
anomalies in a widespread and abundant species.
It is possible that the VERSTEEG plant is a hybrid
with M. celebica.

7. OSMOXYLON

Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; BTH. in B. & H. Gen. Pl. 1 (1865) 944;
Becc. Malesia | (1878) 193; BoeRL. Ann. Jard. Bot. Btzg 6 (1887) 123; O. K. Rev.
Gen. Pl. 1 (1891) 645; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; Bot. Jahrb.
56 (1920) 384; Hutcu. Gen. FI. Pl. 2 (1967) 73; PHILIPSON, Blumea 23 (1976) 99. —
Eschweileria Zipp. ex BOERL. Ann. Jard. Bot. Btzg 6 (1887) 112, non Eschweilera
Mart. 1828; Handl. 1 (1890) 640. — Pseudosandalum O. K. Rev. Gen. Pl. 1 (1891)
271 (‘Pseudosantalum’), nom. illeg. — Boerlagiodendron HARMS in E. & P. Nat.
Pfi. Fam. 3, 8 (1894) 31; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484; Bot. Jahrb.
56 (1920) 377; Hutcu. Gen. FI. Pl. 2 (1967) 72. — Fig. 11, 13-16.

Unarmed, glabrous or tomentose shrubs or trees. Leaves palmately lobed or
simple, rarely digitately compound; stipules forming a ligule, and the base of the
petiole furnished with one to several spiral or transversal crests or collars (very
rarely absent). Inflorescence a terminal compound umbel; peduncle short; primary
rays each terminating into three branches; the central branch bearing a head or
umbellule of almost always sterile bacciform flowers (‘pseudo-fruits’); the two
lateral branches each bearing a head or umbellule of hermaphrodite flowers. Calyx
an obsolete rim or 0. Corolla with few to many lobes above, tubular below.
Stamens 4-30, filaments thick, anthers oblong, exserted. Ovary inferior, not
articulated with the pedicel, cells 1-many; disk flat with a central raised boss
bearing the pustulate stigmas. Fruit subglobose (ribbed when dry); exocarp fleshy,
endocarp crustaceous. Seeds compressed, endosperm smooth or wrinkled.

Distr. About 50 spp., of which 40 occur in Malesia, extending from Borneo and the Philippines east-

wards through Celebes and Moluccas to New Guinea, the remainder lying further north and east in
Taiwan, Micronesia, Melanesia and the New Hebrides. Fig. 12.

a2 FLORA MALESIANA [ser. I, vol. 9!

Ecol. Mainly understorey trees in primary rain-forest, also in second growth forest, usually at low
altitudes, especially in shaded situations and near rivers, 15. O. borneense a characteristic rheophyte.

Notes. The foliage, inflorescence and flowers of this genus are all unique within the family. The base
of the petiole often bears a spiral crest, or this may form a simple collar; the blade varies from simple to
elaborately compound; the inflorescence is composed of trifid rays, the central branch bearing sterile
bacciform flowers; the corolla is tubular.

The central bacciform flowers (pseudo-fruits) are sterile, except in 12. O. yatesii, in which apparently
mature seed was once found.

By exception a specimen of 40. O. /uzoniense had apparently fertile flowers on the central branches of
the inflorescence.

KEY TO THE SPECIES
1. Leaves simple, without lobes.

Dweetioleybase without crestsi(dNewaGuinea)) sas. carsueeneee ec ese ee 1. O. miquelii
2. Petiole base with 1 or more crests.
Ss Petioleibaseawithiseveralcrests (Philippines)ia = —p ence eee Geen 8. O. oblongifolium

3. Petiole base with a single collar-like crest.
4. Fertile flowers pedicelled (Philippines).

5. Leaf obovate to oblanceolate, petiole 3cmorless........... . 2. O. dinagatense
SS elWeatdelliptics petiole icmiormmoressaewacs 200s ees cl cence ee ree 3. O. simplicifolium
4. Fertile flowers sessile (or subsessile).
6s Weafibroadly obovate (Solomons) masseuse) | ee ae ete 4. O. spathipedunculatum
6. Leaf lanceolate or narrowly obovate.
deePseudo-truits;pedicelled’ (Moluccas) Ream .ct suche cant eee ee 5. O. articulatum
7. Pseudo-fruits sessile.
$=inflorescenceses60:cmyG (Moluccas), 20 net se eee een een 6. O. umbelliferum
82 Inflorescencee: 30%em) @ (New ireland) ag. e405 eee cee eee nee 7. O. lanceolatum

1. Leaves lobed or digitately compound.
9. Leaves digitately compound.
10 Ovary 10=16-celled\((New Guinea) = 0. 4 616 see ee ee 13. O. geelvinkianum
10. Ovary 4—5-celled.
11. Leaflets lobed (Philippines).

12. Primary rays of intiorescence'c; 1Ojor fewer... <a. cs 2 20s & 9. O. catanduanense
12. Primary rays of inflorescence more than 20.
iS eintlorescencerc: 16=201emU Dh cams ts ee ee RE Coon eames See eee 10. O. caudatum
IS ssiniorescence:cs.9=6/CM tem ae ene a een Ione 11. O. heterophyllum
11. Leaflets entire.
14. Leaflets elliptic-oblong to ovate (Philippines). . ............. 12. O. yatesii
14. Leaflets linear-lanceolate.
15 Reafiets-4—7-spetiolei4—6 cm (Philippines) 5) <1. a ee ae 14. O. lineare
15. Leaflets 9 or more, petiole longer than 10 cm (Borneo). ........ 15. O. borneense

9. Leaves lobed (sometimes smaller simple leaves below inflorescence).
16. Pseudo-fruits sessile.
17. Petiolar crests long-pectinate. Bracts of the peduncle heavily setose (Philippines)
16. O. pulcherrimum
17. Petiolar crests fimbriate or undulate. Bracts of the peduncle not setose (Moluccas).
18. Lateral inflorescence branches with opposite, persistent bracts close to the base 17. O. soelaense
18. Lateral inflorescence branches with scars of opposite caducous bracts near the middle
18. O. globulare
16. Pseudo-fruits pedicelled.
19. Lateral inflorescence branches without an articulation.
20. Petiolar base with a single collar-like crest (Bormeo) ......... 19. O. kostermansii
20. Petiolar base with several crests.
21. Petiolar crests long-pectinate.
22. Fertile flowers pedicelled. Stamens at least 7 (KeyIs.)......... 20. O. barbatum
22. Fertile flowers sessile (or subsessile). Stamens 5 (Bismarck Arch.). .... . 21. O. pfeilii
21. Petiolar crests fimbriate or undulate.
23. Stamens and ovary cells more than 5. Flowers usually pedicelled (Moluccas) 22. O. palmatum
23. Stamens and ovary cells 4. Flowers sessile or subsessile (Philippines). . . . 23. O. ramosii
19. Lateral inflorescence branches with an articulation (2 bracts or bract-scars).

1979] ARALIACEAE—I (Philipson) 33

24. Fertile flowers pedicelled.
25. Pedicels of fertile flowers longer than 7 mm (New Guinea) ..... 24. O. novoguineense
25. Pedicels of fertile flowers 5 mm or shorter.
26. Ovary cells 7 or more.

Piet eat lobes elliptic((Celebesjit Geer. « 6) ci. Sie sae es . . . 25. O. teysmannii
Dieteath lobes pinnatilobed (Batian)iece ~ = Sas Gok is Esse. ee 41. O. insigne
26. Ovary cells 4-5.
Pe Petiolar crests=- entire (Philippmesy .. 29=4. (ees be 2 ae ee 26. O. humile
28. Petiolar crests with long setae.
29. Leaves coriaceous, margin thickened, teeth obtuse (Philippines). . . .27. O. pectinatum

29. Leaves membranaceous, margin not thickened, teeth setulose (Celebes)
28. O. masarangense
24. Fertile flowers sessile (or if subsessile bracteoles longer than the very short pedicels).
30. Ovary cells 10 or more (New Guinea).
31. Flower buds large (c. 9-12 mm long). Primary rays of inflorescence very strong
29. O. insidiator
31. Flower buds smaller (c. 4-6 mm long). Primary rays of inflorescence more tenuous.
32. Heads of fertile flowers + discoid at anthesis ............ 30. O. boerlagei
Secads OF tertile flowers! globose. Sect =. & 208 Ryde See ee 31. O. sessiliflorum
30. Ovary cells 6 or fewer.
33. Leaf lobes lanceolate.
34. Leaf lobes 4-7; petiole 4-6 cm (Philippines) ............. 14. O. lineare
34. Leaf lobes 9 or more; petiole longer than 10cm(Borneo) ...... 15. O. borneense
33. Leaf lobes broader.
35. Leaf surfaces retaining some trace of setulose tomentum. Umbellules of pseudo-fruits small
(Simi oor less)'€New Guinea) =. . Shae ee Se Se eee 38. O. micranthum
35. Leaf surfaces glabrous at maturity. Umbellules of pseudo-fruits larger (1 cm @ or more).
36. Ovary cells 3 (Philippines).
375 Petiolar erests; entire: Czas «4 = Rb ees ee 32. O. camiguinense
aa betiolarierests long-pecunate: GE vr5, .° Scions ewes Lee oe 33. O. fenicis
36. Ovary cells 4 or more.
38. Umbellules of pseudo-fruits large (3-5 cm @) (Philippines).
39. Primary rays of the inflorescence 9-12 cm long. Leaves with strong radiating veins, usually

mores thant. goa aoe tk Ses Os arich See Eee. ce dees See 34. O. eminens
39. Primary rays of the inflorescence 4-5 cm long. Leaf-veins less strongly developed, usually
fewermthanstl ge ks ee ct. SO ee eee 35. O. serratifolium

38. Umbellules of pseudo-fruits smaller (2'/, cm @ or less).
40. Leaf with a small triangular lobe below the middle of each side of the blade (not strictly

paimately;lobed) (Philippines) >... se) ee ee 8. O. oblongifolium
40. Leaf palmately lobed.
41. Petiolar crests long-pectinate (Celebes) . . ......-..... 36. O. celebicum

41. Petiolar crests fimbriate, entire, or undulate.
42. Articulation of lateral branches of umbels close to the base (Talaud Is.)
37. O. talaudense
42. Articulation of lateral branches of umbels near the middle.
43. Inflorescence over 20 cm @ (New Guinea) ......... 31. O. sessiliflorum
43. Inflorescence under 20 cm @ (Philippines).
44. Leaf usually 3-lobed. Inflorescence rays delicate, indistinctly setose to glabrous
39. O. trilobatum
44. Leaf usually 5—7-lobed. Inflorescence rays sturdy, markedly setose 40. O. luzoniense

1. Osmoxylon miquelii BorrLt. Ann. Jard. Bot. Sparsely branched tree, 15 m. Leaves glabrous,
Btzg 6 (1887) 125, t. 16; Harms, Bot. Jahrb. 56 simple, subcoriaceous; stipules small forming a
(1920) 384; Puitipson, Blumea 23 (1976) 103. — __ bicuspid ligule; petioles long (to 19 cm), swollen
O. amboinense Mia. Ann. Mus. Bot. Lugd.-Bat. 1 distally; blade oblong-elliptic, 22-36 by 9-12 cm,
(1863) 6, p.p.; Becc. Malesia 1 (1878) 194, p.p.— subrounded at base and apex or mucronulate,

Gastonia simplicifolia Zier. ex SEEM. J. Bot. 3 midrib prominent below, secondary veins arched-
(1865) 75, nomen in synon.; ex BoERL. Ann. Jard. ascending and uniting, c. 1-2 cm apart, margin
Bot. Btzg 6 (1887) 125, nom. inval. in synon. —_ entire to undulate. Umbel terminal, sessile, with
Pseudosandalum miquelii (BOERL.) O. K. Rev. Gen. many (28-32) radiating rigid, angular, trifid
Pl. 1 (1891) 271. branches c. 7 cm long to first joint. Central

34 FLORA MALESIANA

[ser. I, vol. 91

=o *

\
x.
DS a
Ce
i =|o

pee N aye 1

—— — —— —

Fig. 12. Species density of Osmoxylon Mia. in
Malesia; above the hyphen the number of endemic
species, below it the non-endemics.

branches unknown. Lateral branches c. 5 cm long,
articulate near the base. Flowers 20—30, sessile on
the expanded ends of the inflorescence branches.
Corolla and stamens unknown. Drupes crowded,
subrotund, c. 4 mm @ (dry), c. 8-10 ribbed when
dry, crowned by a semiglobose entire stigma,
8-10-celled. Seeds with slightly ruminate endo-
sperm.

Distr. Malesia: West New Guinea. Only known
from the type (coll. ZIPPELIUs).

2. Osmoxylon dinagatense (MERR.) PHILIPSON,
Blumea 23 (1976) 103. — Boerlagiodendron
dinagatense MeERR. Philip. J. Sc. 17 (1920) 301;
Me_rr. En. Philip. 3 (1923) 222.

Glabrous shrub, c. 2 m. Leaves crowded at the
ends of the branches; petiole 2-3 cm, channelled
above, with a small triangular base, bearing a short
stipular ligule (2-3 mm long) and extending around
the base of the petiole as a single narrow collar;
blade obovate to oblanceolate, to 23 by 8'/, cm;
base narrowed into the petiole, apex rounded with
a very short apiculum, margin slightly revolute,
entire to obscurely undulate with minute teeth,
coriaceous. Inflorescence a terminal compound
umbel; peduncle c. 1!/, cm, with 1-few bracts
(reduced leaf-bases) with small triangular bracts
(3 mm long) among the primary rays; primary rays
about 15, 8-10 mm long, flattened, bearing opposite
bracts (2 mm long) at the apex, each bearing three
branches; central branch c. 4 mm long, bearing
a head of sessile, bacciform flowers c. 2!/, mm @;
lateral branches 1'/,-2 cm long with two opposite
small bracts about the middle and ending in an
involucre of minute rounded bracts around a
terminal umbellule of c. 7-10 flowers; pedicels
11/,-2'/, mm long. Calyx a minute rim. Corolla
and stamens unknown. Ovary 3-4-celled. Ripe
fruit unknown.

Distr. Malesia: Philippines (Dinagat I.).

Note. A species clearly demonstrating the con-
generity of Osmoxylon and Boerlagiodendron.

3. Osmoxylon simplicifolium (ELMER) PHILIPSON,
Blumea 23 (1976) 103. — Boerlagiodendron
simplicifolia ELMER, Leafi. Philip. Bot. 7 (1914)
2329; MERR. En. Philip. 3 (1923) 224.

Glabrous shrub, to 5 m, with numerous /eaves
clustered near the ends of the branches; petiole
5-7 cm, 2 mm wide, terete, with a small clasping
base, an inconspicuous stipular ligule, and a single
broad disk-like crest around the lower part of
the petiole; blade simple, elliptic, base broadly
cuneate, apex acute to apiculate, to 20 by 6!/, cm,
coriaceous, margin thickened, coarsely dentate,
midrib prominent, principal nerves c. 8-10 mm
apart. Inflorescence a terminal compound umbel,
spherical, c. 7 cm @; peduncle 11/,-2 cm; primary
rays c. 25-30, c. 1 cm long with two small obtuse
bracts at the apex, ending in three branches; the
central branch c. 6 mm long, bearing a subglobose
umbel of c. 10-12 sterile bacciform flowers (2 mm
@), 3-4 mm pedicelled; lateral branches c. 2—2!/,cm
long, articulated about the middle, terminating in
an umbel of c. 10-20 flowers; pedicels c. 2!/, mm.
Calyx rim obsolete. Corolla 3—4-lobed, tubular
below, 2 mm long. Stamens 3-4, exserted, 3 mm
long. Ovary subcylindric, 24-celled, 1 mm long.
Drupe spherical, c. 5mm @ (dry), 2-4-ribbed when
dry; surface of endosperm shallowly wrinkled.

Distr. Malesia: Philippines (Mindanao: Agusan
Prov., Cabadbaran).

Ecol. On wind-swept ridge at 1750 m, on moss-
covered soil with stones.

Vern. Bolauanon, Mbo.

Note. A wide-spreading shrub. Bark thick,
yellowish, becoming grey. Wood soft, yellowish.
Twigs repeatedly branched, the leafy portion sub-
erect, leaves mostly ascending, rigidly coriaceous.
Inflorescence branches green. Flowers orange,
odourless. Berries becoming purple-black.

4. Osmoxylon spathipedunculatum (PHILIPSON)
PHILIPSON, Blumea 23 (1976) 103. — Meryta
spathipedunculata PHILIPSON, Bull. Br. Mus. Nat.
Hist. Bot. 1 (1951) 12.

Glabrous tree, to 20 m, with spreading branches.
Leaves crowded at the ends of the branches; petiole
to 14 cm with a small clasping base bearing a
stipular ligule and a rim-like collar around the base
of the petiole; blade obovate 17-30 by 10-15 cm,
attenuate at the base, apex obtuse, margin entire,
midrib prominent, secondary veins arched and
uniting, 1!/,-2 cm apart. Inflorescence a terminal
compound umbel; primary rays c. 12, stout, com-
pressed 9-17 cm long, bearing three branches at the
apex; central branches and sterile flowers unf-
known; the two lateral branches 9-14 cm long with
an articulation c. 1-2 cm from base, bearing
helmet-shaped bracts which fall to reveal the
terminal head of c. 12 flowers sessile on an ex-
panded receptacle with an involucral rimc.1cm @.
Calyx rim 0. Corolla split into 5 lobes above,
tubular below. Stamens 5. Ovary subcylindr:c, +

1979]

ARALIACEAE—I (Philipson) 3D

9-celled; disk raised in the centre to the pustulate
stigmas. Drupes in a spherical head, globose, c. 12
mm @, c. 9-ribbed when dry, crowned by the
prominent, persistent, confluent stigmas.

Distr. Solomon Islands (Bougainville and
Guadalcanal).

Ecol. Rain-forest, 800-1200 m.

5. Osmoxylon articulatum PHILIPSON, Blumea 23
(1976) 103.

Tree with stout branches, glabrous. Leaves well
spaced towards the ends of the branches; petiole
10-13 cm, 2 mm broad, narrowly channelled above,
with a small triangular base, bearing a short stipu-
lar ligule (c. 2 mm long) and extending around the
base of the petiole as a collar; blade obovate, to
27 by 9 cm, base narrowly cuneate, apex rounded or
acute and shortly apiculate, margin thickened,
remotely dentate towards the apex, midrib promi-
nent, principal lateral veins c. 11/,-11/, cm apart.
Inflorescence a terminal compound umbel, almost
sessile, saucer-shaped bracts caducous; primary
rays c. 24, 8-11 cm long, flattened, c. 4 mm broad,
bearing three branches at the apex; central branch
4-6'/, cm long, the apex expanded and bearing an
umbel of c. 10 sterile bacciform flowers c. 6 by
6 mm (when dry) apparently 1-celled, c. 9-14 mm
pedicelled; the two lateral branches c. 7 cm long at
anthesis with an articulation c. 8-10 mm above the
base, bearing helmet-shaped bracts which fall to
reveal the terminal head of c. 15—18 flowers, sessile
on an expanded receptacle with an involucral rim
c.6mm @. Calyx rim 0. Corolla split into c. 4 lobes
above, tubular below, c. 2!/, mm long. Stamens 5,
exserted. Ovary subcylindric, c. 11/, mm long,
7-8-celled, disk with a pustulate central stigmatic
boss. Drupes in a spherical head c. 2 cm @ (when
dry), strongly 7—8-ribbed (when dry), c. 8 mm @;
stigmas persistent, prominent.

Distr. Malesia: Moluccas (Halmaheira: Ake
Mumar to upper reaches of the Kakatua-matawe).
Fr. Sept.

Note. This plant is evidently similar to 6.
O. umbelliferum described by RUMPHIUS. However,
a number of differences make it unlikely that it is
the same species. The diameter of the inflorescence
of the Halmaheira plant is only about half that
given by RuMPHIUs; the lateral rays of the inflores-
cence are distinctly articulated near the base, a
feature now shown in Rumpuius’ figure; and the
sterile bacciform flowers are long-pedicelled, where-
as RuMPHIus described and figured his as borne in
capitula.

6. Osmoxylon umbelliferum (LAMK) MEerR. Int.
Rumph. (1917) 406; Puitirson, Blumea 23 (1976)
104. — Pseudo-Sandalum amboinense RUMPH.
Herb. Amb. 2: 54, t. 12. — Aralia umbellifera
LAMK, Encycl. 1 (1783) 225. — Hedera umbelli-
ferum (LAMx) DC. Prod. 4 (1830) 262. — Gilibertia
saururoides DC. l.c. 256. — Gastonia saururoides

Roxs. [Hort. Beng. (1814) 90, nomen;] FI. Ind.
ed. Carey 2 (1832) 408 (‘sasuroides’). — O. amboi-
nense Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 6,
p.p. — Pseudosandalum umbelliferum (LAMK)
O. K. Rev. Gen. Pl. 1 (1891) 271.

According to RUMPHIUs: Tree with stout trunk,
the branches marked with prominent round leaf-
scars. Leaves clustered at the ends of the branches,
glabrous; petioles long; blades simple, lanceolate
(30-36 by 10-12 cm), base rounded, apex acute,
margin dentate. Flowers in large spreading umbels,
the radiating branches tripartite, c. 30 cm long,
each ending in a capitulum.

Distr. Malesia: Moluccas. Infrequent on Am-
bon, but said to be more numerous in Ceram and
the Sula Islands. Only known from Rumputus’
excellent plate and description; not yet re-
collected in Ambon.

Ecol. Evidently in forest in the hills of Ambon,
and also planted at the time of RUMPHIUS.

Vern. Sasuru, Leytimor, tonokuku, Hitu.

Notes. Valued for the perfume of its wood and
foliage.

Since this plant is known only from a description
and a figure, some uncertainties remain as to its
specific characters. In the description it is stated
that the young leaves possess a few small teeth of
which some signs remain on the older leaves. It is
not clear whether the teeth are best developed on
distinctive juvenile foliage, or whether the newly
expanded normal foliage is intended. The leaves in
the figure have prominent teeth, but as they are
not shown associated with the inflorescence, they
may be from a juvenile shoot. The description of
the size of the flower buds is confusing, and it seems
likely that sterile pseudo-fruits were mistaken for
flower buds. Nevertheless, most characters of the
plant are adequately portrayed and there can be no
doubt that this species is distinct from the other
simple-leaved species with a single, collar-like,
petiolar crest.

7. Osmoxylon lanceolatum PHILIPSON, Blumea 23
(1976) 104. — Fig. 13.

Small tree with few branches, up to 16 m,
glabrous. Many /eaves clustered towards the ends
of the branches; petiole 8-15 cm, terete, with a
small triangular base, bearing a short stipular
ligule (c. 2mm long) and extending around the
base of the petiole as a collar; blade oblanceolate,
to 33 by 7'/, cm, base narrowly cuneate, apex acute
or slightly apiculate, margin entire, midrib promi-
nent, lateral veins arched ascending, c. 2-3 cm
apart. Inflorescence a terminal compound umbel,
almost sessile, saucer-shaped; bracts caducous;
primary rays c. 15, c. 10 cm long, flattened,
c. 4-5 mm broad, bearing three branches at the
apex; central branch 5-6 cm long, the apex
expanded and bearing a spherical head of c. 8-12
sessile, sterile, bacciform flowers c. 5 by 5 mm
(when dry), 1-2-celled; the two lateral branches

36 FLORA MALESIANA [ser. I, vol. 92

Fig. 13. Osmoxylon lanceolatum PutILipson. Above: habit of inflorescence and leaves; below, left: twig
showing collar-like crests at the base of the petioles; below, right: the trifid branches of the inflorescence
(New Ireland, SANDs 795).

1979]

ARALIACEAE—I (Philipson) a7

c. 5 cm long at anthesis, with an articulation
immediately above the base, bearing two helmet-
shaped bracts which fall to reveal the terminal
head of c. 8-10 flowers, sessile on an expanded
receptacle with an involucral rim c. 7 mm @.
Calyx rim 0. Corolla known only in bud, c. 2!/, mm
long. Stamens c. 5. Ovary gibbous, c. 1'/, mm high,
4-celled. Fruit unknown.

Distr. Malesia: New Ireland (Namatanai
Subdistr., Danfu R. area, inland from Manga).

Ecol. Understorey tree in ridge top forest on
limestone, 750-850 m.

Note. The bark is pale grey, + smooth with
fine cracks. The twigs and cut branches are strongly
aromatic. The wood is soft and dark straw-
coloured. The central branches of the inflorescence
rays are held + horizontally or depressed and come
to maturity before the lateral branches which are
held erect.

8. Osmoxylon oblongifolium PHILipson, Blumea 23
(1976) 105.

Shrub c. 2 m, glabrous when mature, setulose on
young parts. Leaves clustered at the ends of the
branches; petiole to 16 cm, channelled above, 4 mm
broad, with a clasping base prolonged upwards as a
stipular ligule 11/,-2 cm long, and with entire or
fimbriate crests encircling the lower part of the
petiole; blade simple, oblong-ovate, occasionally
with a small triangular lobe on each side below the
middle, to 46 by 17(-24) cm, base rounded to
truncate, apex shortly acuminate, margin serrate,
midrib prominent, principal lateral veins c. 3-4 cm
apart (at broadest part of leaf). Inflorescence a
terminal compound umbel, hemispherical, c. 13 cm
@; peduncle 3-4 cm, c. 6 mm wide, bearing
fimbriate, lanceolate bracts (to 2 cm long) along its
length and around and among the primary rays;
primary rays c. 20, rather short and stout (16-20
by 3-4 mm) with small opposite caducous bracts at
apex, each ray ending in three branches; the
central branch c. 4 mm long; sterile flowers un-
known; the two lateral branches 31/,-4 cm long,
articulated below the middle, terminating in a head
of c. 20-30 sessile or subsessile flowers. Flowers
unknown. Fruits crowded in a spherical head,
drupes c. 5mm @ (when dry) on pedicels c. 1 mm,
4-ribbed, pyrenes 4, cartilaginous; endosperm with
faint reticulate ridging.

Distr. Malesia; Philippines (Samar).

Ecol. In dipterocarp forest, along creek bank, at
200 m.

9. Osmoxylon catanduanense (MERR.) PHILIPSON,
Blumea 23 (1976) 105. — Boerlagiodendron
catanduanense MERR. Philip. J. Sc. 13 (1918) Bot.
318; En. Philip. 3 (1923) 222.

Shrub c. | m, glabrous except for parts of the
inflorescence. Leaves clustered at the ends of the
branches; petiole c. 30 cm, terete, 3 mm wide,
base with a short ligule, and inconspicuous

recurved crests; blade digitately compound,
leaflets 7, chartaceous to subcoriaceous; petiolules
4-5 cm, the lateral shorter; leaflets lanceolate,
mid-leaflet to 26 cm long, base cuneate, apex +
caudate; irregularly lyrately lobed, the sinuses
reaching to within c. 8 mm of the midrib, lobes
2-4 cm long, patent, margin slightly thickened,
entire or obscurely dentate. Inflorescence a terminal
compound umbel, c. 10 cm @, peduncle stout,
bearing ovate bracts c. 1 mm long; primary rays
c. 7-10, 2-21/, cm long, to 2 mm wide, minutely
pubescent, with 2 broadly ovate bracts 4-5 mm
long at apex; central branch 3 mm or less, bearing
a head of sterile flowers; lateral branches 2!/, cm
long, bearing 2 broad bracts near the middle, and
ending in a spherical head of c. 15 sessile flowers.
Calyx an obsolete rim. Petals and stamens un-
known. Ovary 4-celled. Fruit ovoid, 4-ridged when
dry, 7 by 5 mm.

Distr. Malesia: Philippines (Catanduanes).

Ecol. On forested slopes, at c. 350 m.

Note. Together with 10. O. caudatum and 11.
O. heterophyllum this species forms a small group
with leaves composed of lyrate leaflets. Although
the foliage is similar (though not identical), the
inflorescences are distinctive. In O. heterophyllum
the rays are short, resulting in a compact compound
umbel; in O. caudatum there are few rays (10 or
fewer); in O. catanduanense there are many,
relatively long rays, resulting in a large, diffuse
compound umbel. Since O. catanduanense and
O. caudatum are known only from the type collec-
tions, the range of variation of these species is not
known, but the inflorescence differences justify the
retention of all three species.

10. Osmoxylon caudatum (MERR.) PHILIPSON,
Blumea 23 (1976) 105. — Boerlagiodendron
caudatum MeErR. Philip. J. Sc. 14 (1919) 440; En.
Philip. 3 (1923) 222.

Erect shrub, c. 2 m, becoming glabrous. Leaves
clustered towards the ends of the branches; petiole
c. 45 cm, terete, striate, 4-5 mm wide, base with a
stipular ligule c. 11/, cm long, and 2-3 fimbriate or
pectinate crests; blade digitately compound,
leaflets S—9, subcoriaceous; petiolules of the central
leaflets to 5 cm long, lateral leaflets + sessile;
leaflets elliptic-lanceolate, mid-leaflet to c. 25 cm
long, base decurrent on the slightly winged petio-
lule, apex caudate-acuminate, the larger leaflets
with 1-3 pairs of lyrate lobes reaching almost to
the midrib, lobes ascending, margin slightly
revolute with often prominent usually incurved
teeth. Inflorescence a terminal compound umbel,
c. 18 cm @; peduncle stout, c. 2 cm, bearing few
ligulate bracts c. 11/,-2 cm long, rough, with short
setae on the back; primary rays c. 25, c. 4-5 cm
long, glabrous and striate, subtended by ligulate
bracts, similar opposite bracts at the apex, c. 1 cm
long; central branch 1-1'/, cm long, glabrous,
ending in a whorl of obtuse rough coriaceous bracts

38 FLORA MALESIANA

[ser. I, vol. 91

c. 3 mm long and a head (c. 1 cm @) of sterile
bacciform flowers c. 3 mm @, on pedicels 3-8 mm;
lateral branches 4-5 cm long, with opposite
fimbriate bracts near the middle, bearing an
ellipsoid head of c. 20 sessile flowers, each flower
subtended and + enclosed by an ovate fimbriate
bract 3-4 mm long. Calyx an obsolete rim. Petals
and stamens unknown. Ovary 2-2!/, mm long,
turbinate, obscurely angled, 4-celled. Fruit
unknown.

Distr. Malesia: Philippines (Luzon: Ilocos
Norte Prov., Mt Palimlin).

Ecol. On forested slopes near the summit, at
c. 1000 m.

Note. Apparently never re-collected. For a dis-
cussion of distinctive features see under 9.
O. catanduanense.

11. Osmoxylon heterophyllum (MERR.) PHILIPSON,
Blumea 23 (1976) 106. — Boerlagiodendron
heterophyllum Merr. Philip. J. Sc. 9 (1914) Bot.
329; En. Philip. 3 (1923) 223.

Erect tree, c. 5 m, glabrous. Leaves clustered at
the ends of the branches; petiole to 25 cm, base
clasping, prolonged as a stipular ligule to 21/, cm
long, and bearing several pectinate crests (becom-
ing recurved when the bristles may be obscured
or shed); blade to 35 cm long, digitately com-
pound (or some leaflets imperfectly separated);
leaflets 3-7, unequal in size, oblong-ovate to
broadly oblong-oblanceolate, acuminate, lyrately
lobed and irregularly dentate, the base gradually
narrowed to the petiolule; petiolule up to 7 cm.
Inflorescence a dense terminal compound umbel;
peduncle stout, c. 2 cm, bearing many lanceolate
bracts 2-3 cm long; primary rays c. 30, 10-15 mm
long, subtended by lanceolate c. 2!/, cm long
bracts, sometimes with bristles on the back, and
bearing similar opposite terminal bracts 8-10 mm
long, each ending in three branches; central
branch short (not seen fully developed) terminating
in an umbellule of c. 15 sterile bacciform flowers
(c. 3 mm long), pedicels 3-4 mm; lateral branches
8-10 mm long (? fully developed), articulation
present ending in heads (c. 1 cm @) of numerous
flowers. Calyx rim obsolete. Corolla lobes 4, 2 mm
long. Stamens 4. Ovary 4-celled. Fruit unknown.

Distr. Malesia: Philippines (Samar, Biliran and
Mindanao).

Ecol. Primary forest, under shade near creek,
100-550 m.

Vern. Arafas, Bis., kayuang, Mbo, magusayag,
C.Bis.

Note. The description is partially based on the
original publication as I have seen only immature
inflorescences. For a discussion of distinctive
features, see under 9. O. catanduanense.

12. Osmoxylon yatesii (MERR.) PHILIPSON, Blumea
23 (1976) 106. — Boerlagiodendron yatesii MERR.

Philip. J. Sc. 13 (1918) Bot. 44; En. Philip. 3 (1923)
225%

Shrub, 1 m, glabrous, except for the inflores-
cence. Leaves clustered towards the ends of the
stout branches; petiole to 38 cm, channelled above,
5 mm @, base with a stipular ligule 1 cm long, and
1-2 inconspicuous non-fimbriate crests around the
back of the petiole; blade digitately compound,
leaflets 5-7; petiolule 2-7 cm (the lateral shorter);
blade elliptic-oblong to ovate, mid-leaflet to 23 by
8 cm, base gradually tapered, apex acuminate-
caudate, margin dentate or somewhat undulate.
Inflorescence a terminal compound umbel, sub-
sessile or peduncle stout 1-3 cm, bearing few ovate
bracts 1 cm long; primary rays 5-10, tomentose,
c. 3 cm long and 3 mm wide, subtended by ovate
bracts 6 mm long, similar opposite bracts at apex;
central branch 2-3 mm long, pubescent, ending
in a whorl of blunt bracts (3 mm long) and an
umbellule of c. 10 sterile flowers 4 mm @, 2-4-
celled, pedicels 3-8 mm; lateral branches 2—21/, cm
long, pubescent, with small opposite bracts about
the middle, bearing a head of 10-15 sessile
flowers, subtended by ovate ciliate bracts. Calyx
rim obsolete, sometimes fimbriate. Corolla 4—5-
lobed, tubular below, 11/, mm long in bud. Stamens
4-5. Ovary 4-celled. Young fruit (MERRILL, /.c.)
shortly pedicelled (2-3 mm).

Distr. Malesia: Philippines (Luzon and Catan-
duanes).

Ecol. In rain-forest and mossy forest, from low
altitude to 1250 m.

Vern. Magalayag, Dinagat.

Note. The leaves are unlike any other Osmoxy-
lon, resembling those of Macropanax or Schefflera.
The flowers are described as yellow, and the fruit
black. The only instance of a pseudo-fruit con-
taining apparently normal seeds occurred in this
species.

13. Osmoxylon geelvinkianum BeEcc. Malesia 1
(1878) 196; PHILipson, Blumea 23 (1976) 106. —
Eschweileria geelvinkiana (BECC.) BOERL. Ann.
Jard. Bot. Btzg 6 (1886) 120. — Trevesia geelvin-
kiana (BEcc.) O. K. Rev. Gen. PI. 1 (1891) 272. —
Boerlagiodendron geelvinkianum (BECC.) HARMS in
E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. —
Eschweiler(i)a elegans RIDL. Trans. Linn. Soc. II, 9
(1916) 63. — Boerlagiodendron elegans (RIDL.)
Hars, Bot. Jahrb. 56 (1920) 380. — Boerlagioden-
dron stenolobum HARMS, /.c. 382, f. 1 kt.
Glabrous shrub with few slender branches, up to
3 m high. Leaves clustered near the ends of the
branches; petiole up to 22 cm, usually shorter,
2-3!/, mm wide, narrowly channelled above,
with a sheathing base prolonged as a membranous
stipular ligule up to 4 cm or longer and with
fimbriate or + entire crests encircling the lower
part of the petiole; blade up to 30 cm @ (usually
20 cm or less) very deeply 5—11-lobed, or with
distinct digitately arranged leaflets, the lobes or

1979)

ARALIACEAE—I (Philipson) 39

leaflets linear-lanceolate to lanceolate-obovate,
entire or irregularly pinnatifid with narrow finely-
tapering lobes, base gradually narrowed, apex
narrowly caudate, margin serrate; leaves below the
inflorescence sometimes reduced to a single leaflet.
Inflorescence terminal hemispherical, c. 12-20 cm
@; peduncle 1 cm or less, with caducous lanceolate
bracts mostly clustered below the primary rays,
1-2 cm long; primary rays rather few, spaced,
2'/.-6 cm long, slender, bearing two caducous
lanceolate bracts at the apex, up to 1'/, cm long,
each ray ending in three branches; the central
branch c. 4-6 mm long bearing a whorl of lanceo-
late caducous bracts and an umbel of c. 7-12
sterile, globose or ovoid bacciform flowers (c. 7 mm
@ when dry) with pedicels c. 5 mm long and 6-9-
celled; the two lateral branches c. 3-4 cm long at
anthesis, articulated about the middle, terminating
in a small head of 10-20 sessile or subsessile
flowers. Calyx rim obsolete; corolla splitting into
c. 4 irregular lobes above, tubular below, c. 21/, mm
long. Stamens 10-14, exserted, 3 mm long, anthers
small. Ovary cylindric, c. 2 mm long, 10—16-celled;
disk with a central raised boss formed by the
pustulate stigmas. Fruit globose, fleshy (ribbed
when dry), c. 10 mm @.

Distr. Malesia: New Guinea (Irian Jaya, to
Sepik and Fly R. areas).

Ecol. Primary forest, along creeks and river
banks, flood-resistant, from near sea-level to 850 m.

Vern. Amamutapu, Kamora, korinki, Orne,
ida’ pforpforsami, Kutubu.

Note. The narrow leaf segments, almost or
quite separated at their bases, are characteristic,
even though variable in outline. The flowers are
described as orange or reddish, and the soft fleshy
fruits as dark purple, dark blue, or black.

14. Osmoxylon lineare (MERR.) PHILIPSON, Blumea
23 (1976) 106. — Boerlagiodendron lineare MERR.
Philip. J. Sc. 3 (1908) Bot. 253; En. Philip. 3 (1923)
2233

Glabrous, erect shrub, c. 3 m. Leaves crowded
near the ends of the branches; petiole 4-6 cm, the
base with a small stipular ligule (5 mm long) and a
few fimbriate crests; blade to 20 cm @, digitately
compound with 4-7 leaflets (or very deeply
divided into as many lobes); leaflets linear-
lanceolate, c. 1—1!/, cm wide, the base decurrent on
the winged petiolule, apex attenuated, margin
thickened, denticulate especially above. Jnflores-
cence a terminal compound umbel; peduncle short
(c. 1 cm), bracteate; primary rays c. 10, 2-23/, cm
long, each ending in three branches; the central
branch 4-5 mm long bearing a spherical head of
numerous fimbriate bracts (sterile flowers fallen);
the lateral branches 3-3!/, cm long with a pair of
minute fimbriate bracts about the middle, ending in
a capitulum of c. 20 sessile flowers subtended by
small fimbriate bracts, c. 7 mm @. Calyx rim
minute. Corolla and stamens unknown. Ovary

5-celled. Fruit ovoid c. 3 mm long, 5-ridged (when
dry).

Distr. Malesia: Philippines (Luzon).

Note. Apparently never re-collected. The
original specimen is without field information. The
foliage is similar to that of 15. O. horneense but
with shorter petioles and fewer leaflets.

15. Osmoxylon borneense SEEM. J. Bot. 6 (1868)
141; PuiLirson, Blumea 23 (1976) 107. —
O. helleborinum Becc. Malesia 1 (1878) 198. —
Eschweileria helleborina (BECC.) BOERL. Ann. Jard.
Bot. Btzg 6 (1887) t. 13. — Trevesia helleborina
(BECC) 2O Ke Rev Gen! Ply 1.113891)? 272. —
Boerlagiodendron helleborinum (BEcc.) HARMS in
E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31. — Boerlagio-
dendron borneense (SEEM.) MERR. En. Born. (1921)
456.

Glabrous, spreading shrub, up to 3 m. Leaves
clustered near the ends of the branches; petiole up
to 24 cm, narrowly channelled above, 2-3 (or 5) mm
wide, with a sheathing base prolonged as a mem-
branous stipular ligule up to 2!/, cm long, and
usually with fimbriate, + entire, or more rarely
long-setose crests encircling the lower part of the
petiole; blade up to 20 cm @ digitately compound
(or the bases of the leaflets joined by a very short
web of tissue); leaflets 9-13, linear-lanceolate to
lanceolate, gradually narrowed to the base and
apex, up to 20 by 3 cm, usually much narrower,
margin serrate, principle veins numerous, c. 5—
10 mm apart. Inflorescence terminal, hemispherical,
c. 6-13 cm @; peduncle 1-2 cm or shorter, with
caducous lanceolate-ovate entire or fimbriate
bracts mostly clustered below the primary rays,
1-11/, cm long; primary rays rather few (5-12),
2'/.-4 cm long, with 2 ovate bracts at the apex,
c. 8 mm long, each ray ending in three branches;
central branch c. 5-12 mm, bearing an umbel of
c. 6-16 sterile, globose, bacciform flowers, c. 3—
5 mm @ (when dry) with pedicels 4-5 mm long and
5-celled; the two lateral branches 1'/,—3 cm long at
anthesis, articulated about the middle, terminating
in a small head of c. 20-25 sessile flowers with
minute rounded bracts. Calyx rim obsolete; corolla
splitting into few irregular lobes above, tubular
below, c. 11/,-2 mm long; stamens 5-6, slightly
exserted; ovary turbinate, angled, c. 1 mm long,
5-celled, disk with a central raised boss formed by
the pustulate stigmas. Fruit globose, fleshy (ribbed
when dry) c.5 mm @.

Distr. Malesia: Borneo (Sarawak, Sabah and
Kalimantan).

Ecol. Characteristic of rocky river banks, not
beyond flood-level, often in deep shade, from near
sea-level to 950 m.

Vern. Medong, Kayan, empasia abor, Iban,
kayan, Tamang, koung, Kinabalu, bungor, Murut
Bokan, salimpangaya, Murut Kalabakai.

Notes. The leaves of some specimens of 13.
O. geelvinkianum (New Guinea) resemble this

40

FLORA MALESIANA

[ser. I, vol. 91

species closely. This species can be distinguished by
the more compact and smaller fertile flowers, and
by its leaflets being uniformly unlobed.

Both species are characteristic of river banks, and
O. borneense has a low spreading habit, with the
branches often rooting, resulting in extensive
patches of this low shrub. It is a characteristic
rheophyte confined to below flood-level.

The flowers are described as greenish white or
cream and the inflorescence branches are frequently
dark purple.

16. Osmoxylon pulcherrimum VIDAL ex F.-VILL.
Novy. App. (1880) 102; Sinopsis Atlas (1883) 28,
t. 55, f. B; PHmirson, Blumea 23 (1976) 107. —
Eschweileria pulcherrima (VIDAL) BOERL. Ann.
Jard. Bot. Btzg 6 (1887) 123. — Trevesia pulcher-
rima (VIDAL) O. K. Rev. Gen. Pl. 1 (1891) 272. —
Boerlagiodendron pulcherrimum (VIDAL) HARMS in
E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; MerrR.
Philip. J. Sc. 3 (1908) Bot. 254; En. Philip. 3 (1923)
224. — Boerlagiodendron sibuyanense ELMER,
Leafi. Philip. Bot. 7 (1914) 2328; Merr. En. Philip.
3 (1923) 224.

Erect, sparsely branched tree, up to 10 m,
glabrous when mature, except for the inflorescence.
Leaves crowded at the ends of the branches; petiole
to 1 m, channelled above, clasping base heavily
lenticellate, prolonged as-a broad stipular ligule
c. 2 cm long, usually with strong bristles on the
back, and with strong long-pectinate crests
encircling the base of the petiole; blade coriaceous,
fan-shaped, c. 40 cm long, base broadly cuneate to
truncate, palmately 7-11-lobed, lobes extending to
within c. 12 cm from the base, lanceolate, coarsely
serrate, sometimes irregularly lobulate, slightly
narrowed towards the base, apex acute, sinuses
rounded. Inflorescence a terminal compound
umbel, c. 18 cm @; peduncle very short, bearing
heavily setose bracts; primary rays 15-20, c. 34cm
long, 3-4 mm broad, setulose, at the apex bearing
opposite, ovate-lanceolate bracts 10-15 mm long,
each ending in three branches; central branch
c. 15-20 mm long, terminating in a globular head
(c. 12 mm @) of c. 20-30, sessile, sterile, bacciform
flowers (3-4 mm @) 3-celled, subtended by small
ovate-lanceolate bracts; lateral branches c. 5!/, cm
long (7'/, cm in fruit), with opposite bracts
(c. 3-4 mm long) near the middle, terminating in a
globose head of c. 40-50 sessile flowers, bracts
between the flowers very small, setulose. Calyx rim
obsolete. Corolla 4-lobed, tubular below, 2 mm
long in bud. Stamens 4. Ovary 4-celled. Fruit
globose c. 6-8 mm long, 4-ribbed (dry).

Distr. Malesia: Philippines (Luzon, Mindoro
and Sibuyan), recorded also from Formosa and
Micronesia (Palau), cf. KANEHIRA, En. Micron.
Pl. (1935) 384.

Ecol. Damp primary forests, 225-800 m.

Vern. Cf. MERRILL: paladukai, Bik., salapak,
Neg.; cf. ELMER: palad-amok, Vis.

Notes. The fan-shaped leaves with several nar-
tow lobes and prominent main veins resemble those
of 34. O. eminens but are less strikingly developed.
The inflorescence is considerably smaller with the
pseudo-fruits forming a compact head borne on a
comparatively long peduncle. The heads of true
flowers, and of the fruits, are considerably smaller
than those of O. eminens.

Although VIDAL’s material is no longer avail-
able, the figure and description relate well to later
collections.

The specimens on which ELMER based his
Boerlagiodendron sibuyanense have the lobes of the
leaf rather simpler in outline than is usual, but the
fragments of young inflorescence are quite typical
of the taxon and the name is reduced to synonymy.

17. Osmoxylon soelaense PHILIPSON, Blumea 23
(1976) 108.

A glabrous shrub or small tree. Petiole c. 32 cm,
broadly channelled above, clasping base prolonged
as a stipular ligule 11/, cm long, and bearing
c. 3 fimbriate crests; blade c. 40 cm long, broadly
cuneate at the base, palmately 7-lobed to within
c. 12 cm from the base, lobes narrowly ovate to
oblong-elliptic, slightly narrowed towards the
rounded sinuses, apiculate, margin denticulate.
Inflorescence a terminal compound umbel;
peduncle c. 2 cm, bearing lanceolate bracts c. 2 cm
long (similar smaller bracts subtend the primary
rays); primary rays c. 1'/, cm long, 5 mm wide,
flattened, bearing opposite, terminal, persistent
bracts 10-12 mm long, with lenticels and branched
bristles on the back, ending in three branches;
central branch c. 10 mm long terminating in a
head 1 cm @ of 10-15 sessile sterile bacciform
flowers (4 by 3 mm when dry; 3-celled) surrounded
by an involucre of ovate bracts (3 mm long) and
with minute bracts interspersed; lateral branches
4'/,-5 cm long, bearing opposite ovate bracts
(4 mm long) c. 8 mm above the base, terminating
in a dense head c. 1 cm @ of 25-30 sessile flowers
interspersed with inconspicuous obtuse fimbriate
bracts. Calyx rim obsolete. Corolla 5-lobed above,
tubular below. Stamens 5. Ovary 4~5-celled. Fruit
unknown.

Distr. Malesia: Moluccas (Sula Is.: Taliabu
and Sulabesi).

Note. Fora discussion of the distinctive features,
see under 31. O. sessiliflorum.

18. Osmoxylon globulare PHiLirson, Blumea 23
(1976) 108.

Shrub to 4 m, furfuraceous on the young parts.
Petiole to 55 cm long, broadly channelled above,
c. 1 cm wide, clasping base heavily lenticellate,
prolonged as a stipular ligule 4 cm long sometimes
scaly on the back, and bearing numerous irregular
undulate crests on the base of the petiole often
continued up the petiole, as rough fascicles of
bristles as far as the blade; blade 45 cm long, base

1979]

ARALIACEAE—I (Philipson) 4]

cordate or emarginate, with some bristles under-
neath, palmately 7-lobed to within c. 10-15 cm
from the base, lobes narrowly ovate to oblong-
elliptic, slightly narrowed towards the broadly
rounded sinuses, apex acute, margin serrate.
Inflorescence a terminal compound umbel, spheri-
cal, c. 15 cm @; peduncle stout, 2-3 cm, bracts
together with those among the primary rays
caducous; primary rays 30-40, rigid only slightly
flattened (subterete), 2-4 cm long, 2—2'/, mm wide,
bearing opposite bract-scars at the apex, ending in
three branches; central branch 8-10 mm long,
terminating in a head c. 13 mm @ of c. 20 sub-
sessile sterile bacciform flowers (5 by 4 mm when
dry, 2-3-celled); pedicels to 2 mm interspersed
with minute bracts; lateral branches 3-4'/, cm
long, with the scars of opposite bracts near the
middle, terminating in a dense head 1-11/, cm @
(in bud) of 20-30 sessile flowers interspersed with
inconspicuous bracts. Calyx rim obsolete. Corolla
6-8-lobed above, tubular below, 2'/, mm long (in
bud). Stamens 6-8. Ovary turbinate, obscurely
ribbed, 5—8-celled (varying on the same plant).
Fruit unknown.

Distr. Malesia: Moluccas (Halmaheira, Moro-
tai).

Ecol. In forest from sea-level to 800 m. Said to
be rare in Halmaheira but common in Morotai.

Vern. Bungan-gutu, saha-sasate, Djailolo.

Note. For a discussion of the distinctive features
see under 31. O. sessilifiorum.

19. Osmoxylon kostermansii PHILIPSON, Blumea 23
(1976) 108.

Glabrous, small tree, 8 m. Leaves clustered near
the ends of the branches; petiole up to 35 cm,
narrowly channelled above, c. 3 mm broad, witha
sheathing base prolonged as a stipular ligule c.2 cm
long, continued around the back of the leaf-base
as a single wide crest with an entire recurved
margin; blade up to 30 cm @, base cordate, deeply
5—7-lobed, lobes elliptic, slightly narrowed towards
the sinuses and with a short acute apiculum,
Margin minutely serrate and sometimes with small
sub-lobes, sinuses rounded. Inflorescence a terminal
compound umbel, hemispherical, c. 14 cm @ at
anthesis; peduncle c. 1 cm long, bearing small
lanceolate bracts (c. 3 mm long) below and among
the numerous (c. 20-24) primary rays; primary rays
4-5 cm long and 1 mm broad, with opposite bracts
(2 mm long) at the apex, each ending in three
branches; the central branch 5S—6 mm long, bearing
a spherical umbel of c. 20 small, sterile, bacciform
flowers (2 mm @) on pedicels c. 5-7 mm long,
2-celled; the two lateral branches c. 31/,4 cm long,
with no articulation or bracts except for a minute
involucre around the terminal umbellule of
c. 10-14 flowers; pedicels c. 2-3 mm. Calyx rim
obsolete; corolla 4-lobed, 2 mm long in bud;
stamens 4; ovary subcylindric, angled, c. 1 mm
long, 5—7-celled, disk with a central stylar boss.

Fruit spherical, c. 6 mm long, strongly ribbed when

Distr. Malesia: Borneo (Kalimantan: Sang-
kulirang Distr., Mt Medadam).

Ecol. On limestone at 450 m.

Note. The foliage is similar to that of 22. O. pal-
matum, except for the distinctive petiolar crest.
The inflorescence is also similar to O. palmatum,
especially in the lack of an articulation on the rays
below the umbellules.

20. Osmoxylon barbatum Becc. Malesia 1 (1878)
197; PHiLtipson, Blumea 23 (1976) 109. — Esch-
weileria barbata (BECC.) BoERL. Ann. Jard. Bot.
Btzg 6 (1886) 117. —Trevesia barbata (BEcc.)
O. K. Rev. Gen. Pl. 1 (1891) 272. — Boerlagio-
dendron barbatum (BEcc.) HARMS in E. & P. Nat.
Pfi. Fam. 3, 8 (1894) 31.

Small, glabrous tree. Leaves clustered near the
ends of the branches; petiole to 32 cm, channelled
above, 4-5 mm broad, with a sheathing base
prolonged as a stipular ligule 2-3 cm long, and
with several long setose crests encircling the lower
part of the petiole; blade up to 43 cm @, base
cordate to truncate; deeply 5-7-lobed; lobes
elliptic-lanceolate, narrowed towards the sinuses
and tapered to an acuminate apex, margin minutely
serrate, sinuses broadly rounded. Inflorescence a
terminal compound umbel, hemispherical, to
12 cm @; peduncle 1-2 cm; primary rays c. 20,
3—5 cm long, with three branches at the apex; the
central branch c. 1 cm long, bearing a subglobose
umbel of c. 15-20, sterile, bacciform flowers
(3-4 mm @) on pedicels S—7 mm long, 2-4-celled;
the two lateral branches about 4 cm long with no
clear articulation but 1 or 2 obsolescent bracts,
terminating in an umbellule of c. 15-20 flowers;
pedicels c. 34 mm. Calyx rim obsolete; corolla
4-lobed above, tubular below, c. 4 mm long;
stamens 7 or more, rarely fewer, exserted; ovary
subcylindric c. 3 mm long, angled, 7-celled; disk
with a central stigmatic boss. Fruit unknown.

Distr. Malesia: SE. Moluccas (Key Is.).

Note. Distinguished from the closely related
22. O. palmatum by the long-setose petiolar crests.
For discrimination from 21. O. pfeilii see that

species.

21. Osmoxylon pfeilii (WARB.) PHILIPSON, Blumea
23 (1976) 109. — Eschweileria pfeilii WARB. Bot.
Jahrb. 13 (1891) 396. — Boerlagiodendron pfeilii
(Wars.) Harms in E. & P. Nat. Pfil. Fam. 3, 8
(1894) 32.

A tree developing a crown when mature, up to
16 m, glabrous when mature, young parts slightly
setulose. Leaves in terminal clusters; petiole up to
60 cm, channelled above, c. 6 mm broad, with a
sheathing base prolonged as a membranous
stipular ligule 24 cm long, and with many pro-
nounced, long-setose crests encircling the lower
part of the petiole; blade up to 75 cm @, deeply

42 FLORA MALESIANA

[ser. I, vol. 91

7-11-lobed, base cordate or emarginate; lobes
lanceolate to narrowly elliptic-oblong, narrowed
towards the sinuses and tapered to an attenuate
apex, margin sharply and remotely serrate, sinuses
broadly rounded. Inflorescence a terminal com-
pound umbel, hemispherical, to 20 cm @; peduncle
short (2-3 cm); primary rays numerous (c. 30-40),
4-5 cm long, glabrous, with obsolete bracts at the
apex, each ending in three branches; central
branch c. 10 mm long, bearing a subglobose umbel
of c. 20, small, sterile, bacciform flowers (1—-11/, mm
@) on pedicels c. 3-4 mm long, 2—5-celled; the two
lateral branches c. 3-4 cm long, with no clear
articulation but 1 or 2 obsolescent bracts, termi-
nating in a head of c. 12-16 subsessile flowers
(pedicel c. 1 mm, becoming longer in fruit) sur-
rounded by an inconspicuous involucral rim. Calyx
rim obsolete; corolla 5-lobed, 11/, mm long in bud;
stamens 5; ovary subcylindric, angled, 11/, mm
long at anthesis, 5—16-celled, disk with a central
boss formed by the united pustulate stigmas. Fruit
spherical, fleshy, c. 8 mm @, ribbed when dry, the
stigmatic boss persistent and prominent; pyrenes
cartilaginous.

Distr. Malesia: Bismarck Archipelago (New
Britain, Duke of York Group and New Ireland).

Ecol. Primary rain-forest, from near sea-level
to 600 m.

Vern. Sare, sasare, sare a lauvolau, New Britain,
Pomio; a ibalur, New Ireland.

Notes. The bark is grey-brown and pustular,
the wood straw-coloured and soft. The flowers are
orange, and the ripe fruit dark red-violet.

In the original description the ovary is recorded
to possess 10-14 cells. However, some other
specimens have as few as 5 cells in the ovary, but
in other respects agree with specimens with the
large number of seeds. Since the inflorescence,
leaf-shape, and especially the nature of the petiolar
crests, as well as the distribution, are all highly
distinctive within the genus, all the specimens can
be accepted as examples of ome species with a
highly variable number of carpels.

This species is very close to 20. O. barbatum of
the Key Islands. The original diagnostic character
of the number of cells in the ovary has been found
to be unreliable. However, since the primary rays
in the inflorescence are more numerous and the
pedicels of the fertile and sterile flowers are shorter
this geographically distinct species is maintained.

22. Osmoxylon palmatum (LAMK) PHILIPSON, comb.
nov. — Folium polypi mas (et femina?) RUMPH.
Herb. Amb. 4: 101, t. 43. — Aralia palmata LaMK,
Encycl. 1 (1783) 224, type, non Lour. 1790, nec
R. & S. 1820. — Trevesia moluccana Mia. FI. Ind.
Bat. 1, 1 (1856) 748; Bonplandia 4 (1856) 137. —
Trevesia zippeliana Mia. Ann. Mus. Bot. Lugd.-
Bat. 1 (1863) 11. — Unjala bifida REINW. ex DE
VRIESE, Pl. Ind. Or. (1867) 83, nomen in synon.; ex
Boert. Ann. Jard. Bot. Btzg 6 (1887) 166, in

synon. — O. moluccanum (MiqQ.) Becc. Malesia 1
(1878) 195; PHILIPSON, Blumea 23 (1976) 109. —
O. zippelianum (MiQ.) BEcc. Malesia 1 (1878) 195.
— Eschweileria palmata Zipp. ex BOERL. Ann.
Jard. Bot. Btzg 6 (1887) 116, t. 14. — Boerlagio-
dendron palmatum (Zipp. ex BOERL.) HARMS in
E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31; Merr. Int.
Rumph. (1917) 407. — Boerlagiodendron moluc-
canum (Miq.) BAKH. f. & Ooststr. in Back. Bekn.
Fl. Java (em. ed.) 7 (1948) fam. 159, p. 3; Blumea 6
(1950) 367; BACK. & BAKH. f. Fl. Java 2 (1965) 163.

Small, glabrous tree to 15 m. Leaves clustered
near the ends of the branches; petiole up to 40 cm,
channelled above, 4-5 mm broad, with a sheathing
base prolonged as a stipular ligule up to 5 cm long,
and with several fimbriate crests encircling the
lower part of the petiole; blade up to 45 cm @, base
cordate, deeply S5S—9-lobed, lobes elliptic, slightly
narrower towards the sinuses, acuminate, margin
serrate and sometimes with small sub-lobes, sinuses
rounded. Inflorescence a terminal, compound
umbel, subspherical, to c. 20 cm @ at anthesis;
peduncle c. 1-2 cm, with small caducous bracts
below and among the numerous (20-60) primary
rays; primary rays 3-5 cm long, 2-3 mm broad,
with two opposite caducous bracts at the apex,
each ending in three branches; central branch
6-10 mm long, bearing a spherical umbel of
c. 20-30 small, sterile, bacciform, flowers (2-4 mm
@) on pedicels 4-6 mm long, 4~7-celled; two lateral
branches c. 2!/,-5 cm long, rigid and straight, with
no articulation (indistinct scars of bracts may
occur below the involucre), bearing a minute
involucre around the terminal umbellule of
c. 12-20 flowers; pedicels to c. 5 mm (occasionally
flowers subsessile). Calyx rim obsolete. Corolla
irregularly 4-5-lobed above, tubular below,
c. 5 mm long. Stamens 6-9, exserted, filaments
stout. Ovary subcylindric, angled, c. 2 mm long,
6-9-celled, disk with a raised central stigmatic
boss. Fruit globose, fleshy, c. 10 mm @, strongly
ribbed when dry.

Distr. Malesia: Celebes (once, not localized)
and Moluccas (Buru, Ceram, Ambon, Banda,
Tenimber Is.).

Also cultivated in the Bogor Botanic Garden.

Ecol. An understorey tree in primary rain-
forest.

Uses. The leaves are used for culinary and medi-
cinal purposes (against gonorrhoea).

Vern. Daun gurita, pelenda darat, saha-saha,
Moluccas, fumala-alas, Tenimber Is.

Note. The spherical inflorescence is characteris-
tic, having straight rigid rays with no articulation
on the secondary branches, and the pseudo-fruits
are well separated from the true flowers. MIQUEL
distinguished Trevesia zippeliana because the
collector noted that its ovary was 4-celled. Possibly
this number related to the sterile flowers. In two
gatherings from Ambon (Waai, TEYSMANN;
G. Salhutu, BoERLAGE 179) the fertile flowers are

1979]

ARALIACEAE—I (Philipson) 43

subsessile in heads, but otherwise conform to the
characters of this species. The only record of this
genus from the Tenimber Is. consists of leaves only,
but their characters conform to this species.

23. Osmoxylon ramosii (MERR.) PHILIPSON,
Blumea 23 (1976) 110. — Boerlagiodendron
ramosii MERR. Philip. J. Sc. 11 (1916) Bot. 27;
Me_rr. En. Philip. 3 (1923) 224.

Erect, unbranched, or sparingly branched, 4 m
high shrub, becoming glabrous. Petiole to 40 cm,
the clasping base prolonged as a stipular ligule
c. 2 cm long, and with few to several prominent
recurved, obscurely fimbriate, crests surrounding
the lower part of the petiole; blade to 30 cm long,
base emarginate, palmately 3-7-lobed, lobes
extending to within 3-8 cm from the base, oblong-
ovate, margin serrate, sometimes lyrately lobulate,
apex acuminate, sinuses broadly rounded. Jnflores-
cence a terminal compound umbel, 10-15 cm @;
peduncle stout, with lanceolate bristle-bearing
bracts; primary rays c. 15, 2-3 cm long, 2-3 mm
wide, flattened, subtended by lanceolate bracts
c. 2 cm long, with bristles on the back and bearing
similar opposite terminal bracts c. 1 cm long, each
terminating in three branches; central branch
c. 4mm long, slightly pubescent, terminating in an
umbellule (c. 11/, cm @) of c. 10-15 sterile, bacci-
form flowers 4-5 mm @, 2-celled, pedicels 5-8 mm
long, subtended by caducous bracts; lateral
branches 2!/, cm long (slightly longer as fruits

form), without any articulation or bracts except
for a caducous small involucre around the terminal
head of c. 25-35 sessile or very short-pedicelled
flowers, c. 1 cm 2, bracts among the flowers
obscure. Calyx rim obsolete. Corolla 4-lobed
above, tubular below. Stamens 4, exserted,
filaments broad. Ovary subcylindric, obscurely
angled, 4-celled. Fruit spherical 9 mm @, 4-ribbed
when dry.

Distr. Malesia: Philippines (Luzon).

Ecol. On low-lying, wet ground, in forest, or on
forested slopes, 700-800 m.

Notes. The flat-topped inflorescence is c. 15 cm
@ with much-reduced leaves below it. The in-
florescence rays are dark purplish and the flowers
orange-yellow. The bark is grey and the wood soft.

This is the only species in the Philippines without
opposite bracts on the lateral branches of the
inflorescence rays. In this respect it resembles 22.
O. palmatum and a few other species.

24. Osmoxylon novoguineense (SCHEFF.) BECC.
Malesia 1 (1878) 197; PHiLipson, Blumea 23 (1976)
110. — Trevesia novo-guineensis SCHEFF. Ann.
Jard. Bot. Btzg 1 (1876) 26. — Eschweileria
novoguineensis (SCHEFF.) BOERL. ibid. 6 (1886) 118.
— Boerlagiodendron novoguineense (SCHEFF.)

Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31;
Back. & BAKH. f. Fl. Java 2 (1965) 163. —
Boerlagiodendron lauterbachii HARMS in K. Sch. &
Laut. Fl. Schutzgeb. (1900) 484. — Fig. 11, 14.

Fig. 14. Osmoxylon novoguineense (SCHEFF.) BECC. Petiolar base with ligule and collar-like crests (Photogr.
PHILIPSON, Wantoat, 1968).

44 FLORA MALESIANA

[ser. I, vol. 9!

Tree or shrub, unbranched or sparingly
branched, up to 16 m, the young parts rufous-
furfuraceous, glabrescent. Large leaves forming
terminal crowns; petiole up to 1 m, stout (1-2 cm
broad), flattened above, with a sheathing base
prolonged as a strong stipular ligule up to 7 cm
long, and with fimbriate crests encircling the lower
part of the petiole; blade up to 1.20 m @, with 5-7
strong ribs radiating from the top of the petiole,
deeply lobed almost to the base of these ribs, the
lobes in turn deeply lobed and incised, the central
lobes especially being strongly pinnatisect or
digitately tripartite, apices acute, margins serrate;
upper leaves associated with inflorescences may be
smaller, more simply lobed, or entire. Inflorescence
terminal, a large compound umbel, bowl-shaped,
up to 35 cm @; peduncle up to 10 cm, stout, with
lanceolate caducous bracts (c. 4 cm long) below
and among the numerous (c. 50-70) primary rays;
primary rays c. 12-15 cm long at anthesis, c. 3 mm
@, bearing two caducous bracts (1 cm long) at the
apex, each ray ending in three branches; central
branch c. 2 cm long, bearing an umbel of c. 20-40,
sterile, bacciform flowers (c. 6 mm @ when dry) on
pedicels c. 10 mm, and 2-6-celled; the two lateral
branches c. 4-6 cm, with two opposite or sub-
opposite bracts about the middle, terminating in a
subspherical umbel 21/,-3 cm @ of 30-50 flowers
on pedicels c. 8-10 mm long. Calyx rim obsolete,
undulate. Petals with irregular erect lobes, tubular
below. Stamens 6-10 exserted. Ovary turbinate
somewhat angled; glabrous, 6—-14-celled; disk flat
with a central double row of pustulate stigmas.
Fruits on stiff radiating pedicels, ovoid or spherical,
fleshy, ribbed when dry.

Distr. Solomon Is.; in Malesia: throughout
New Guinea and in the Bismarck Archipelago.

Ecol. Primary and second-growth forest, from
sea-level to 1600 m.

Vern. Lebe, Mooi, teresakui, Manikiong,
akriek, Biak, hoppung, Hottam, uger, Wagu,
faliifalii, Tifal, ap gan dandam, aimaini, Mamig,
ida’ pfopforsami, Kutubu, pulaka, Gazelle Peninsula.

Notes. The foliage is similar to that of 30.
O. boerlagei, but the pedicelled flowers of O. novo-
guineense distinguish it readily from that species.
The ripe fruits are usually ovoid, but in the Solomon
Is. they are characteristically spheroidal, and this
feature recurs in some specimens from the Bis-
marck Archipelago and the adjacent coast of New
Guinea.

The fawn bark is pustulate with many lenticels.
An orange exudate flows from the cut stems. The
wood is soft and straw-coloured. The inflorescence
branches are dark purple, the corolla and stamens
usually deep red, and the ripe fruit shining purple
or blue-black.

25. Osmoxylon teysmannii (BOERL.) PHILIPSON,
Blumea 23 (1976) 111. — Eschweileria teysmannii
Borrt. Ann. Jard. Bot. Btzg 6 (1887) 119. —

Trevesia teysmannii (BOERL.) O. K. Rev. Gen. Pl. 1
(1891) 272. — Boerlagiodendron teysmannii
(BOERL.) HARMS in E. & P. Nat. Pfl. Fam. 3, 8
(1894) 31.

A small, glabrous tree, 6 m. Leaves clustered at
the ends of the branches; petiole to 40 cm, chan-
nelled above, 4 mm broad, with a sheathing base
prolonged as a stipular ligule 2-2'/, cm long, and
with several fimbriate or entire crests on the lower
part of the petiole; blade c. 30 cm @, membranous,
cordate at the base, deeply 7-lobed, lobes elliptic,
slightly narrowed to the broadly rounded sinuses,
narrowed to a fine apiculum at the apex, margin
finely serrulate. Inflorescence a terminal compound
umbel, c. 10cm @; peduncle c. 1 cm, bearing ovate
bracts (ligules of reduced leaves) and terminating
in a cluster of bracts (c. 10 mm long) below and
among the primary rays; primary rays c. 12-15,
c. 3-4 cm long, with a pair of lanceolate bracts at
the apex (c. 1 cm long); central branch c. 1 cm long,
terminating in an umbellule of c. 5-8 sterile bacci-
form flowers (c. 4 mm @ when dry) on pedicels
6-9 mm long interspersed with linear bracts 5 mm
long; two lateral branches c. 3!/, cm long, articu-
lated about the middle, terminating in an umbellule
surrounded by caducous linear bracts (leaving a
rim-like involucre); flowers c. 8-12, pedicels
c. 2-3 mm. Calyx rim obsolete. Corolla 3 mm long,
with 7-8 lobes above, tubular below. Stamens
7-8, filaments stout, anthers exserted. Ovary sub-
cylindric, angled, c. 11/, mm long, 7—8-celled; disk
flat, with a central stigmatic boss. Fruit unknown.

Distr. Malesia: SW. Celebes (Tjamba, Kosali-
Porema) and NW. Central Celebes (Palu-Parigi
and Mt Nokilalaki).

Ecol. In rain-forest, 800-1000 m.,

26. Osmoxylon humile (ELMER) PHILIPSON, Blumea
23 (1976) 111. — Boerlagiodendron humilis ELMER,
Leafl. Philip. Bot. 7 (1914) 2327; Merr. En.
Philip. 3 (1923) 223.

Erect, small, sparsely branched shrub, up to
11/, m. Leaves clustered at the ends of the branches;
petiole to 25 cm, terete, base prolonged as a
stipular ligule c. 1 cm long, and with c. 3 entire
crests at the base; blade palmately 5-lobed, 24 cm
long, base truncate or cordate, lobes reaching to
within 3-6 cm from the base, elliptic, 4-6 cm wide,
narrowed towards the broadly rounded sinuses,
tapered to an acute apiculum, margin serrate in the
upper part, the outer lobes with a lobule on the
lower edge. Inflorescence a terminal compound
umbel, 9 cm @, subtended by a few foliaceous
bracts; peduncle stout, 2-3 cm, with furfuraceous,
oblong bracts; primary branches crowded, numer-
ous, 21/, cm, furfuraceous, flattened, striate with
opposite minute bracts at the apex; central branch
c. 3 mm, bearing an umbellule of sterile, bacciform
flowers 31/, mm @, pedicels 4'/, mm; lateral
branches 3-31/, cm, articulated about the middle,
terminating in an umbellule of c. 15 flowers, bracts

1979]

ARALIACEAE—I (Philipson) 45

inconspicuous, fimbriate, pedicels 11/,-2 mm.
Calyx an obsolete rim. Corolla and stamens not
known. Ovary 1'/,-2 mm, 4-celled, with a flat
disk and a raised central stigmatic boss, 4-celled.
Fruit 6 by 4 mm (dry) 4-ribbed; pyrenes crusta-
ceous; endosperm rugose.

Distr. Malesia: Philippines (Mindanao).

Ecol. Damp fertile ground in dense forest, on
south side of Baruring R., at 1000 m.

Vern. Sardng-ka-mdno, Bag.

27. Osmoxylon pectinatum (MERR.) PHILIPSON,
Blumea 23 (1976) 111. — Boerlagiodendron
pectinatum MeErkR. Philip. J. Sc. 3 (1908) Bot. 253,
424; En. Philip. 3 (1923) 224; KANEHIRA, Form.
Trees rev. ed. (1936) 520, f. 480; Hur-Lin Li,
Woody FI. Taiwan (1963) 666, f. 273.

Shrub or small glabrous tree up to 8 m. Leaves
clustered at the ends of the branches; petiole to
18 cm, with a clasping base prolonged as a short
acute stipular ligule, and with several basal crests
fringed with 1-2 cm long bristles; blade to 25cm @,
base truncate to broadly cuneate, palmately 5-7-
lobed, lobes reaching to about the middle of the
lamina, sinuses narrow-rounded, lobes oblong-
elliptic, usually slightly narrowed below, obtuse to
acute, margin thickened, coarsely dentate, coria-
ceous. Inflorescence a terminal compound umbel;
peduncle c. 1 cm, with small bracts (3 mm) below
and among the primary rays; primary rays c. 25-
35, c. 2-3 cm long, with opposite ovate caducous
bracts at the apex, each terminating in three
branches; central branch c. 8-11 mm long, ending
in an involucre of minute bracts (1 mm) surround-
ing an umbellule of c. 15—20 ovoid sterile flowers
(c. 3 mm long, 3-celled), pedicels 5-6 mm long;
lateral branches c. 2!/, cm long at anthesis, with an
articulation about the middle, ending in an umbel-
lule c. 1 cm @ with minute fimbriate bracts;
flowers c. 30, pedicels c. 11/, mm (elongating
slightly in fruit). Calyx rim obsolete. Corolla lobes
4-5, tubular below, 2 mm long. Stamens 4-5.
Ovary turbinate, 11/, mm long, 4—6-celled. Fruits
globose, 5 by 5 mm (dry), 4—6-ribbed when dry.

Distr. Taiwan (Botel Tobago and Lutao I.,
east off Taiwan proper); in Malesia: N. Philippines
(Batan I.).

Ecol. Forested slopes at 650 m.

Vern. Narapan, Iv.

28. Osmoxylon masarangense PHILIPSON, Blumea
23 (1976) 111.

Small tree, 5 m, the young parts setulose,
becoming + glabrous. Leaves in terminal clusters;
petiole c. 17 cm, rather narrow (1'/.-2 mm @),
channelled above, with a small clasping base,
prolonged as a stipular ligule, 1-11/, cm long,
setulose on the back, and with a number of long-
setulose crests encircling the lower part of the
petiole; blade c. 18 by 22 cm, deeply 3—5-lobed (or
below the inflorescence sometimes simple), the

base truncate or emarginate, lobes oblong or
elliptic, slightly narrowed to the broadly rounded
sinuses, apex with a short apiculum, membranous,
margin finely setulose-serrate, sinuses c. 6 cm frob-
base of the blade. Inflorescence a terminal sum
sessile compound umbel; primary rays c. 10,
setulose, 1°/,-2 cm long, each ray ending in three
branches; central branch 4-5 mm long, ending in
an umbellule, pedicels 6 mm, sterile flowers not
known; two lateral branches 2-2'/, cm long, 1 mm
broad, with two bract scars about the middle but
usually not opposite, terminating in an umbellule
with c. 10 pedicels 3-5 mm long (in fruit). Flowers
unknown. Fruit (when dry) ovoid, 6 by 4 mm,
5-seeded.

Distr. Malesia: N. Celebes (Minahasa, Tomo-
hon, Mt Masarang).

Ecol. Secondary forest at edge of crater lake, at
1200 m.

Note. This species is similar in aspect to the
Philippine 39. O. trilobatum, but the petiolar
crests are distinctive.

29. Osmoxylon insidiator BEcc. Malesia 1 (1878)
195; PHILIPSON, Blumea 23 (1976) 112. — O. carpo-
phagarum BeEcc. Malesia 1 (1878) 196. — Esch-
weileria insidiatrix (BECC.) BOERL. Ann. Jard. Bot.
Btzg 6 (1886) 120. — Eschweileria carpophagarum
(BEcc.) BOERL. /.c. 121, t. 15. — Trevesia insidiator
(BEcc.) O. K. Rev. Gen. Pl. 1 (1891) 272. —
Trevesia carpophagarum (BeEcc.) O. K. lc. —
Boerlagiodendron insidiator (BECC.) HARMS in E. &
P. Nat. Pfl. Fam. 3, 8 (1894) 32. — Boerlagiodendron
carpophagarum (BEcc.) Harms, /.c. — Boerlagio-
dendron pachycephalum Harms, Nova Guinea 8
(1910) 271.

Small tree to 12 m, young parts with uniform
scurfy tomentum. Large /eaves forming terminal
crowns; petiole up to 80 cm, stout (1-2 cm 9),
broadly channelled above, with a sheathing base
prolonged as a strong stipular ligule c. 9 cm long,
and with moderately developed irregular (not
fimbriate) crests encircling the lower part of the
petiole; blade up to 85 cm @, with 5-7 strong ribs
radiating from the top of the petiole, deeply lobed
almost to the base of these ribs, lobes in turn deeply
lobed and incised, the median often digitately
tripartite, apices long acuminate, margin irregu-
larly and remotely serrate, subglabrous when
mature or showing remnants of the tomentum.
Inflorescence a terminal hemispherical compound
umbel, c. 15 cm high by 30 cm wide; peduncle
short, stout (11/.-2 cm @), with lanceolate bracts
4-6 cm long below and among the numerous
(15-20) primary rays; primary rays 6-10 cm long,
c. 5-12 mm wide, rigid, bearing 2 lanceolate bracts
(2-3 cm long) at the apex, each ray ending in three
branches; central branch c. 11/, cm long, bearing
an umbel of c. 30 sterile bacciform flowers (c. 5-
12 mm @ when dry), the pseudo-fruits and their
pedicels + rufous tomentose, pedicels 10-12 mm,

46

FLORA MALESIANA

[ser. I, vol. 92

and 6-celled, surrounded by an involucre of short
ovate bracts (3-8 mm long); two lateral branches
c. 6 cm at anthesis, rigid, slightly flattened, to
8 mm broad, bearing a pair of bracts (c. 11/, cm
long) about the middle, terminating in a subglobose
head 31/,4 cm @ of c. 30-40 sessile flowers, and
surrounded by an involucre of ovate bracts
c. 10-14 mm long. Calyx rim fimbriate. Petals
irregularly 4-S-lobed, 7-8 mm long, connate
below to form a fleshy tube, pubescent on the
outer surface. Stamens c. 15-26, filaments strap-
like, projecting beyond the corolla, anthers c.4mm
long. Ovary shortly turbinate, 24 mm long,
angled, furfuraceous, c. 13—25-celled; disk flat,
with a central boss formed of the pustulate stigmas.
Fruits in a compact spherical head, the individual
drupes angled by mutual pressure, and bearing the
persistent stigmas on the exposed face, c. 10-14 mm
long, the numerous pyrenes compressed and flat;
cartilaginous.

Distr. Malesia: throughout New Guinea, but
local; also Waigeo I.

Ecol. Primary rain-forest and regrowths,
frequently beside streams, from sea-level to 350 m.
Vern. Angit, kangit, Waigeo, pennifogo,

Orakawa, Papua.
Notes. The bark is greyish brown, slightly
fissured with many lenticels. The wood is soft and

% ae xe

white. The flowers are reddish-brown or purple,
with orange-red filaments and the fruit purple.

BEcCaRI provided a detailed description of the
living plant, and noted that the fruits are eaten by
various species of pigeon.

Boerlagiodendron pachycephalum Harms has
very strongly developed umbels and leaves, but
apart from size, it does not differ from this species.
Since a range in stature is shown by the several
gatherings now available, the whole is best re-
garded as a single species.

Similarly, the very short pedicels of O. carpo-
Phagarum, which BEccarI used to distinguish it
from O. insidiator, can in fact be matched on
several specimens of that species.

30. Osmoxylon boerlagei (WARB.) PHILIPSON,
Blumea 23 (1976) 112. — Eschweileria boerlagei
Wars. Bot. Jahrb. 13 (1891) 395. — Boerlagioden-
dron warburgii HARMS in E. & P. Nat. Pfl. Fam. 3,
8 (1894) 32, nom. illeg. superfl.; in K. Sch. & Laut.
Fl. Schutzgeb. (1900) 484. — Boerlagiodendron
boerlagei (WARB.) HARMS, Bot. Jahrb. 56 (1920)
382. — Fig. 15.

Small to fairly large tree, unbranched or
sparingly branched, up to 24 m, glabrous, at least
when mature. Large Jeaves forming terminal
crowns; petiole up to 1 m, stout (1-2 cm @),

” *e,"

Fig. 15. Osmoxylon boerlagei (WARB.) PHILIPSON. Showing that each ray ends into a central umbel of
bacciform flowers and two lateral umbels with normal flowers (Photogr. PHILIPSON, Kassam Pass, E. New
Guinea, 1968).

1979]

ARALIACEAE—I (Philipson) 47

broadly channelled above, with a_ sheathing,
heavily lenticellate base prolonged as a strong
stipular ligule up to 7 cm long, and with lacerate
crests encircling the lower part of the petiole;
blade up to 1.15 m @, with 5—7 strong ribs radiating
from the top of the petiole, deeply lobed almost to
the base of these ribs, lobes in turn deeply lobed and
incised, the central lobes especially being strongly
pinnatisect or digitately tripartite, apices acute,
margin undulate or indistinctly serrate. Jnflores-
cence terminal (or overtopped by a lateral leafy
branch), a large compound umbel, bowl-shaped
with a slightly convex top, up to 60 cm @; peduncle
c. 10 cm, stout, with lanceolate bracts below and
among the very numerous radiating primary rays;
outer primary rays c. 20 cm long at anthesis
(elongating in fruit), inner rather shorter, woody,
bearing two caducous bracts at the apex, each ray
ending in three branches; central branch c. 4 cm,
bearing an umbel of c. 20 sterile bacciform flowers
(c. 8 mm @ when dry) with rigid pedicels c. 11/, cm
long, and 5—6-celled; two lateral branches c. 9 cm
long at anthesis, articulated about the middle,
terminating in a button-like head of c. 20-30
sessile flowers and surrounded by ovate bracts
which soon fall leaving a bowl-shaped involucre,
c. 11/, cm @. Calyx rim obsolete. Petals c. 13, bud
flat-topped, angled, minutely pubescent, apparently
falling as a calyptra. Stamens 8-13. Ovary shortly
turbinate, angled, glabrous, 10-14-celled; disk
flat, with a central double row (or ellipse) of
pustulate stigmas. Fruits spreading to form a
+ spherical head obscuring the involucre, each
c. 9 by 7 mm (when dry) with prominent persistent
stigmas.

Distr. Malesia: throughout New Guinea.

Ecol. Primary forest, and secondary growths on
old cultivations, from near sea-level to 1800 m.

Vern. Eunya, Gimi, apiatambay, Washkuk,
ma-korr-korr, Jal, teresahui, Manikiong.

Note. Bark yellow grey with shallow fissures
and many pale corky pustules; wood straw-
coloured, fibrous. The large terminal inflorescence
is shallowly convex on top and is surrounded by
several large leaves. Flowers reddish brown. Fruit
purplish black and succulent.

31. Osmoxylon sessiliflorum (LAUT.) PHILIPSON,
Blumea 23 (1976) 113. — Boerlagiodendron
sessiliflorum Laut. Nova Guinea 8 (1910) 272.
Small tree, up to 18 m, glabrous when mature, or
tomentum persistent on the inflorescence. Leaves
crowded at the ends of the branches; petiole to
60 cm, with the clasping base prolonged as a stipu-
lar ligule up to 6 cm long, and with few to several
strong or weak crests around the base of the petiole,
margin undulate or fimbriate; blade to 50 cm long,
base cordate, palmately 5—9-lobed, lobes extending
to near the base, elliptic, coarsely serrate, often
irregularly lobulate, apex acute, sinuses rounded.
Inflorescence a terminal compound umbel;

peduncle to c. 4 cm, bearing lanceolate bracts to
2/2 cm long, caducous or persistent, occasionally
with some bristles on the back, primary rays
c. 20-30, c. 9 cm long, pubescent, bearing
opposite caducous or rarely persistent bracts at the
apex; central branch variable in length (2-18 mm),
bearing an umbellule or head 1-2'/, cm @ of
sterile bacciform flowers (4-5 mm @), 4-8-celled,
subtended by minute bracts, pedicels variable in
length (5-18 mm); lateral branches 3-7 cm, articu-
lated near the middle, terminating in a globose
head of c. 20-30 sessile flowers (or pedicels 1'/, mm
long), bracts between the flowers very small. Calyx
rim obsolete. Corolla few- to many-lobed, tubular
below, 1'/.4 mm long in bud. Stamens 6-17.
Ovary 5—18-celled. Fruit a globose head of drupes;
drupes c. 10 by 6 mm, obovoid, ribbed when dry.

Distr. Malesia: throughout New Guinea.

Ecol. Rain-forest, especially along the muddy
banks of rivers, from sea-level to 100 m.

Vern. Akriek, Biak, korinki, Orne, kwita-kwita,
Milne Bay, sapi-ai, Jense, terrasahui, Manikiong.

Notes. Unbranched or sparsely branched with
crowns of large leaves. The bark is light brown and
the wood cream. The inflorescence branches are
purple, the flowers red, and the succulent ripe
fruits black.

The variation in the numbers of floral parts is
considerable. Most specimens have more than 10
stamens and the same number of cells in the ovary,
or more. Three gatherings have from 5—7 stamens
and cells. These may possibly require to be
segregated as a distinct species, but other evidence
to support this course is lacking. An even more dis-
tinctive gathering has central branches to 3 cm
long with the pseudo-fruits on short pedicels (less
than 5 mm) and flowers with 17 stamens and 25
cells in the ovary. These characters have not been
included in the specific description as this specimen
is only very tentatively referred to this species.

This species forms an eastward extension of a
complex of species, represented in the Moluccas by
37. O. talaudense, 17. O. soelaense and 18. O. glo-
bulare. ‘Several similar species occur in the Philip-
pines. They are characterized by dense spherical
heads of flowers.

Osmoxylon talaudense resembles some specimens
of O. sessiliflorum rather closely, and the difficulty
of preserving the characters of these large-leaved
plants in an herbarium probably obscures several
good diagnostic features. The most reliable charac-
ter to distinguish these two species is the position
of the articulation on the lateral branches of the
inflorescence rays. In specimens from the Talaud
Is. this is close to the base (below the apex of the
central umbellule of pseudo-fruits) and the bracts
are persistent, whereas in the New Guinea material
it is near (or above) the middle, and is usually
marked by two inconspicuous scars which fre-
quently are not opposite. In both species the
pseudo-fruits are pedicelled, whereas in the other

48 FLORA MALESIANA

[ser. I, vol. 91

two Moluccan species the pseudo-fruits are sessile
or subsessile forming spherical heads. In O. globu-
lare (from Morotai and Halmaheira) the lateral
branches are rigid and only slightly flattened with
the articulation near the middle and the bracts
caducous. In O. soelaense the articulation is much
nearer the base, the bracts are persistent, and the
branches are broader and much flattened.

32. Osmoxylon camiguinense (MERR.) PHILIPSON,
Blumea 23 (1976) 113. — Boerlagiodendron
camiguinense MERR. Philip. J. Sc. 3 (1908) Bot. 252;
En. Philip. 3 (1923) 222.

Shrub to 2 m, glabrous except for the inflores-
cence. Leaves clustered at the ends of the branches;
petiole to 18 cm, with a sheathing base prolonged
as a stipular ligule 1 cm long, and with several
entire crests encircling the lower part of the
petiole; blade 20 by 22 cm, base truncate, 3—-5-lobed
to about the middle, lobes oblong, scarcely nar-
rowed towards the base, apex acuminate, sinuses
broad, rounded, margin dentate, coriaceous.
Inflorescence a terminal compound umbel; primary
rays c. 2—2!/, cm, pubescent with opposite lanceo-
late bracts (c. 12 mm long) at the apex; central
branch c. 4 mm long, bearing a globose umbellule
(c. 12 mm @) of sterile bacciform flowers (c. 2 mm
@), pedicels c. 3 mm, subtended by numerous
ligulate bracts; lateral branches c. 3 cm long, with
opposite lanceolate bracts 4 mm long, ending in a
head of c. 20-30 sessile flowers. Corolla and
stamens not seen, described by MERRILL as
3-merous. Fruit globose 6 mm @ (dry), 3-seeded.

Distr. Malesia: Philippines (Babuyan Is.:
Camiguin I.).

Ecol. On slopes in forest, at 500 m.

33. Osmoxylon fenicis (MERR.) PHILIPSON, Blumea
23 (1976) 114. — Boerlagiodendron fenicis MERR.
Philip. J. Sc. 13 (1918) Bot. 44; En. Philip. 3 (1923)
223. — Boerlagiodendron tayabense MERR. Philip.
J. Sc. 13 (1918) Bot. 45; En. Philip. 3 (1923) 224.
Erect shrub or treelet a few m high, glabrous
except for the inflorescence. Petiole to 45 cm,
clasping base prolonged as a broad stipular ligule,
1!/,-3 cm long, several prominent long pectinate
petiolar crests extending up the petiole as oblique
groups of bristles; blade to 30 cm long, base
cordate or truncate, deeply palmately 3—7-lobed,
lobes extending to within c. 7 cm of the base,
broadly elliptic to oblong, entire or with subsidiary
lobes, somewhat narrowed towards the sinuses,
apex abruptly apiculate, margin remotely denticu-
late or serrate, sinuses very broadly rounded.
Inflorescence a terminal, compound, furfuraceous
umbel, hemispherical, c. 10-12 cm @; peduncle
2-3 cm, densely enclosed in broadly ovate bracts
c. 2 cm long, bearing dense fascicles of strong
bristles on their blades; primary rays 15-30,
c. 21/,-3 cm long, 2'/, mm wide, pubescent, sub-
tended by large bristly bracts c. 1'/, cm long,

bearing opposite terminal usually bristly bracts
6-15 mm long, each ending in 3 branches; central
branch c. 4-10 mm, pubescent, bearing a terminal
umbellule of c. 10-20 sterile bacciform flowers
2-3 mm @, 2-celled, pedicels 3-4 mm tomentose
sometimes with a ruff of hairs around the pseudo-
fruits, subtended by early-caducous small fimbriate
bracts; lateral branches 2!/,-3 cm long, with two
opposite bracts (2-3 mm) about the middle,
bearing a terminal head (8 mm @ without corollas)
of c. 15-30 sessile flowers, subtended by incon-
spicuous ovate tomentose bracts. Calyx an
obsolete rim. Corolla in bud c. 1 mm long. Stamens
3. Ovary 3-celled. Fruiting head 10-12 mm @;
drupes c. 6 mm long, crowded, sessile, 3-angled;
seeds 3.

Distr. Malesia: Philippines (Luzon).

Ecol. In primary dipterocarp forest, on rocky
slopes near streams, 300—400 m.

Note. Closely allied to 40. O. luzoniense and 27.
O. pectinatum, but the combination of long-
fringed petiolar crests, pubescent inflorescence
branches, excessively bristly bracts and 3-merous
flowers is distinctive.

34. Osmoxylon eminens (BULL) PHILIPSON, Blumea
23 (1976) 114. — Trevesia eminens BULL, Cat. New
Plants (1884) 17; Retail List (1885) 64, fig. —
Boerlagiodendron mindanaense MerRR. Philip. J.
Sc. 3 (1908) Bot. 154. — Boerlagiodendron eminens
(BULL) Merr. En. Philip. 3 (1923) 223.

Small tree, up to 10 m, with few stout branches.
Leaves large, forming terminal crowns, glabrous
when mature, young inflorescence densely fur-
furaceous; petiole to 1 m, stout (to 2 cm @),
flattened above, base clasping the stem, heavily
lenticellate, prolonged as a stipular ligule 2 cm
long, bicuspid, often with scales or bristles on the
back, and bearing few to several entire, fimbriate
or occasionally long setose crests; blade to 60 cm
long, palmately 10—-19-lobed, base cordate, lobes
reaching to near the base, lanceolate to oblong, up
to 15 cm wide, in outline either strap-shaped or
irregularly pinnatisect, or the central lobe occa-
sionally distinctly tripartite, margin coarsely and
irregularly dentate, apex acuminate. Inflorescence
a terminal compound umbel c. 40 cm @; peduncle
stout c. 6 cm, 1!/, cm wide, bearing many lanceo-
late scaly bracts 3-6 cm long; primary rays
numerous, rigid, flattened, 9-12 cm long, 6-10 mm
broad, bearing opposite oblong scaly bracts
(2-3 cm long) at the apex, each ending in three
branches; central branch !/,—1 cm long, terminating
in an umbellule (3-5 cm @) of c. 20-40 sterile
bacciform flowers, 7 mm @, 2-3-celled, pedicels
1-2 cm long, surrounded by an involucre of small
bracts (to 8 mm long); lateral branches c. 12 cm
long, with opposite bracts (6-10 mm long) near
the middle, terminating in a head of c. 50-60
sessile flowers, 1!/,-2 cm @ (with corollas), heads
spherical at anthesis, ovoid after corollas absciss;

1979]

ARALIACEAE—I (Philipson) 49

bracts between the flowers very small. Calyx rim
obsolete. Corolla 5-6-lobed, tubular below,
4-5 mm long. Stamens 4-6, filaments 7 mm long,
anthers 1'/, mm long. Ovary 2-3 mm long (at
anthesis), 5—6-celled. Fruits crowded in dense ovoid
heads 3-4 by 2!/.-3 cm, drupes c. 9 by 5 mm,
5-6-angled by mutual pressure, narrowed to the
base, crowned by the persistent stigmatic boss;
pyrenes crustaceous; endosperm wrinkled.

Distr. Micronesia: Carolines; in Malesia:
throughout the Philippines.

Ecol. In primary forest from low altitude
(100 m) to ridge forest and mossy forest at 950 m,
often in shady ravines.

Vern. Cf. MERRILL: apalong or apulong, Bis.,
bunglui-babde, pina-pina, Sul., mangunpulun, Bag.,
palad-ulot, S.L.Bis., ulo-ulo, C.Bis.; in addition:
lolobongan, Lan.

Notes. This is the most widespread and most
frequently collected species in the Philippines. It is
also the most striking. Its large, many-lobed, fan-
shaped leaves and the strong inflorescences, with
globular flower-heads and large clusters of pseudo-
fruits are distinctive. Only 16. O. pulcherrimum
resembles it somewhat in its leaf characters, but the
central branches of the inflorescence rays of that
species are much longer and its pseudo-fruits are
sessile.

The inflorescence branches are described as dull
reddish brown, the flowers as light orange, and the
fruits as indigo-black.

35. Osmoxylon serratifolium (ELMER) PHILIPSON,
Blumea 23 (1976) 114. — Boerlagiodendron
serratifolium ELMER, Leafl. Philip. Bot. 2 (1908)
505; Merr. En. Philip. 3 (1923) 224.

Sparingly branched shrub to 5 m. Petiole to
50 cm long, channelled above, clasping base pro-
longed as an obtuse stipular ligule, and with few
narrow + fimbriate crests around the base of the
petiole; blade to 50 cm long, base cordate, pal-
mately lobed (up to 11 lobes), lobes extending to
within about '/,; from the base, narrowly elliptic,
margin serrate (or slightly lobulate), apex acumi-
nate, sinuses narrowly rounded. Inflorescence a
terminal compound umbel c. 30 cm @; peduncle
stout, bracteate; primary rays 20-30, 4-5 cm long,
5-6 mm wide, flattened, subtended by lanceolate
bracts 3-5 cm long, sometimes with bristles on the
back, and bearing similar opposite terminal bracts
21/2 cm long, each terminating in three branches;
central branch 12-15 mm long, terminating in an
umbellule (3-4 cm @) of c. 20-25 sterile bacciform
flowers 5-6 mm @, 3-4-celled, pedicels to 10 mm,
interspersed with persistent small bracts; lateral
branches c. 9 cm long, with opposite bracts
(c. 6 mm long) 2-3 cm from the base, terminating
in a spherical head of c. 30 sessile flowers c. 11/, cm
@ (in bud), bracts between the flowers very small,
obtuse, fimbriate. Calyx rim obsolete. Corolla
5-7-lobed above, tubular below, 5 mm long.

Stamens 5-6, exserted, filaments 7 mm long,
anthers 2 mm long. Ovary 3 mm long, 5-celled.
Fruit unknown.

Distr. Malesia: Philippines (Leyte, Camiguin,
Panay).

Ecol. ELMer noted that this species was rare in
the low hills of Leyte.

Notes. The flowers are orange-yellow (salmon),
the fruits dark purple.

The inflorescence is very similar to that of 34.
O. eminens, but the leaves lack the many strong
fan-like ribs of that species.

36. Osmoxylon celebicum PxHiLieson, Blumea 23
(1976) 115. — Boerlagiodendron celebicum HARMS
ex Koorpb. Minah. (1898) 489, nomen.

A small, sparsely branched tree, 6 m high. Large
leaves forming terminal crowns; petiole 50 cm by
8 mm, flattened above, with a sheathing base
prolonged as a strong stipular ligule 3 cm long
with branched fibrous setae on the outer surface,
and with several crests bearing similar setae on the
lower part of the petiole; blade 50 cm @, base
emarginate, deeply 9-lobed, lobes narrowly elliptic
to lanceolate, narrowed towards the sinuses, apex
broadly cuneate, margin minutely and remotely
serrate, sinuses broadly rounded. Inflorescence a
terminal compound subspherical umbel c. 20cm @;
peduncle short, stout (15 mm wide) with large
setose bracts (c. 4 cm long) below and among the
primary rays; primary rays c. 15, c. 6 cm long,
5 mm broad, with a pair of large setose bracts (22
by 10 mm) at the apex, each ending in three
branches; central branch c. 6 by 2 mm, terminating
in an involucre of setose ovate bracts (c. 4 mm long)
and an umbel of c. 20-30 sterile bacciform flowers
(c. 3 mm @ when dry, 2-3-celled, on pedicels
6-10 mm long) interspersed with bracts covered
with crisp reddish-brown setulae; the two lateral
branches c. 2 cm long, with opposite setulose bracts
(c. 1 cm long) below the middle, terminating in a
spherical head of c. 30-40 sessile flowers each sub-
tended by a reddish brown setulose cymbiform
bract. Calyx rim obsolete. Corolla c. 2'/, mm long
in bud (not seen in open condition). Stamens 5.
Ovary subcylindric, c. 1 mm long in bud, 5-celled;
disk with a central stigmatic boss. Fruit unknown.

Distr. Malesia: Celebes (Minahasa, Manado).

Ecol. On rich volcanic sand, at 10 m.

Vern. Sinomaha.

Note. The flower buds are orange and the fruits
deep purple.

37. Osmoxylon talaudense PHILIPSON, Blumea 23
(1976) 115.

Shrub or small tree, to 6 m, glabrous. Leaves at
the ends of the stout branches; petiole to 60 cm,
broadly channelled above, clasping base prolonged
as a stipular ligule c. 2 cm long, and bearing 2-3
fimbriate crests; blade c. 50 cm long truncate to
cordate at the base, palmately 7—-11-lobed to within

50 FLORA MALESIANA [ser. I, vol. 91

Fig. 16. Osmoxylon micranthum (HARMS) PHILIPSON. a. Habit, x 1/,, b. flower, c. false fruit and ditto in
CS, x 15, d. CS of fruit, x 6 (a-c KANIs 1384, d PULLEN 428). Drawn by W. R. PHILIPSON.

1979]

ARALIACEAE—I (Philipson) 51

+ 1/4 of the base, lobes elliptic oblong slightly
narrowed towards the rounded sinuses, apiculate,
margin denticulate to undulate. Inflorescence a
terminal compound umbel, peduncle 1-2 cm,
bearing broad ovate bracts 2-3 cm long, with
bristles on the back, (similar persistent bracts
subtend the primary rays); primary rays c. 15,
c. 3-4 cm long, 4 mm wide, flattened, bearing
opposite terminal persistent bracts 11/,-2 cm long,
sometimes with a few bristles on the back, ending
in three branches; central branch 8-10 mm long,
terminating in an umbellule 2 cm @ of 15-20 sterile
bacciform flowers (4 mm @, 2-celled) surrounded
by an involucre of obtuse bracts 1-2 mm long,
pedicels 5—7 mm; lateral branches 4'/,-5 cm long,
bearing opposite ovate persistent bracts (3-7 mm
long) c. 5-10 mm above the base, terminating in a
dense head c. 1 cm @ of 30-40 sessile flowers inter-
spersed with inconspicuous obtuse bracts. Calyx
rim obsolete. Corolla 5-lobed. Stamens 5. Ovary
turbinate, 11/, mm long, 5-celled. Fruit in spherical
heads 2 cm @ (when dry); drupes c. 9 by 6 mm,
obovoid, 5-ribbed.

Distr. Malesia: N. Moluccas (Talaud Is.:
Karekelong and Salebabu).

Ecol. Common in forest, besides streams, from
near sea-level to 100 m.

Vern. Laripatu, Talaud.

Note. The flower is yellow-orange and the fruit
dark purple. For a discussion of the distinctive
features, see under 31. O. sessiliflorum.

38. Osmoxylon micranthum (HARMS) PHILIPSON,
Blumea 23 (1976) 115. — Boerlagiodendron
micranthum Harms, Bot. Jahrb. 56 (1920) 379. —
Boerlagiodendron sayeri HARMS, /.c. 379, f. 1 a—j. —
Eschweileria gawadensis BAKER f. J. Bot. 61 (1923)
22. — Boerlagiodendron tricolor PHILIPSON, Bull.
Br. Mus. Nat. Hist. Bot. 1 (1951) 11. — Fig. 16.
A sparsely branched shrub to 8 m, sometimes
trailing or semi-scandent, young parts uniformly
setulose, buds without cataphylls. Leaves in
terminal clusters; petiole up to 30 cm, rather
narrow (2-4 mm @), becoming sparsely setulose,
channelled above, with a sheathing base prolonged
as a membranous stipular ligule up to 3 cm long,
and with a number of lacerate crests encircling the
lower part of the petiole; blade deeply 3-5- or
more rarely 7-lobed, or below the inflorescence
sometimes simple, base cordate or emarginate, the
central lobe up to 30 cm long, the lobes oblong,
lanceolate or broadly elliptic, entire or irregularly
lobed or incised, or with small sub-lobes, apices
long cuspidate, acute, margin serrate, sinuses
between the lobes broad and rounded, surfaces
become sparsely setulose to subglabrous. /nflores-
cence a terminal compound umbel, often appearing
subterminal by growth of a leafy brafich at the
base of the peduncle; peduncle short (1-2 cm),
heavily setulose, occasionally with a flowering ray
arising from the axils of bracts on or below the

peduncle, bearing distally many lanceolate bracts
5-10 mm long; primary rays 12-18, 10-20 mm
long, setulose, with two lanceolate bracts at the
apex, each ray ending in three branches; the central
branch very short (2-3 mm) bearing a subglobose
umbel of many (c. 40) small sterile bacciform
flowers (c. 1'/. by 1 mm) with filamentous pedicels
c. 2 mm long, and 1-2 cells each with 1 abortive
ovule; the two lateral branches c. 2 cm long, with
two minute bracts about their middle, setulose,
terminating in a head of c. 20 sessile flowers sur-
rounded by an involucre of small rounded bracts.
Calyx rim obsolete. Corolla 4(—5)-lobed, united
below, c. 2 mm long. Stamens 4(—5), filaments
ribbon-like elongating beyond the corolla tube at
anthesis, 3-4 mm, anthers small. Ovary shortly
subcylindric, c. 1 mm high, faintly angled, glabrous,
1-5-, usually 4-celled, disk fleshy, rising slightly to
the central stigmas. Fruit an ellipsoid drupe with
1-5 cartilaginous pyrenes; seeds with smooth
endosperm.

Distr. Malesia: New Guinea (Irian Jaya:
Idenburg R. to Milne Bay Distr.).

Ecol. In primary forest from the foothills to the
montane mossy forest, often in swampy or deeply
shaded situations, 700-2400 m.

Vern. Diande, Chimbu, kenata, Okapa.

Note. The inflorescence branches are often red
or purple, and the flowers either orange or reddish
with yellow anthers. The ripe fruits are deep purple
or black. The shape and size of the leaf can vary
greatly, even on the same plant. The ovary usually
has 4 cells, but plants with 3, 2 and 1 occur. Since
these are alike in other respects they have been
treated as a single species. Although the type of
Boerlagiodendron tricolor has an ovary with 5 cells
and is from much further west than other gather-
ings, it is not considered to be specifically distinct.

39. Osmoxylon trilobatum (MERR.) PHILIPSON,
Blumea 23 (1976) 116. — O. cumingii SEEM. J. Bot.
6 (1868) 141, nomen. — Boerlagiodendron trilo-
batum Merk. Philip. J. Sc. 2 (1907) Bot. 289; En.
Philip. 3 (1923) 224.

Slender shrub or small tree up to 5 m, becoming
glabrous except for slight pubescence on the
inflorescence. Leaves clustered near the ends of the
branches; petioles to 25 cm, c. 3 mm wide, clasping
base prolonged as a broad stipular ligule c. 1 cm
long, and with 2-3 entire, or obscurely fimbriate,
often recurved crests surrounding the base of the
petiole; blade 3- or occasionally 5-lobed (leaves
below the inflorescence sometimes simple), to 30
by 28 cm, base broadly cuneate, rounded or trun-
cate (emarginate in 5-lobed leaves), lobes about
1/,*/, of the blade, narrowly or broadly oblong,
often slightly narrowed below and sharply acumi-
nate to caudate, margin serrate. Inflorescence a
terminal compound spherical umbel, 7-15 cm 2,
either rather compact or branches lax; peduncle
2-3 cm with broad ovate bracts; primary rays 8-20

52 FLORA MALESIANA

[ser. I, vol. 9

or more, slightly pubescent, 11/.-4 cm long, sub-
tended by ovate bracts 5-10 mm long, opposite
ovate bracts at the apex, 1-3 mm long; central
branch 11/,-6 mm long, pubescent, bearing an
umbel (1—11/, cm @) of sterile bacciform flowers up
to 5 mm @, 1-4-celled, pedicels 2-6 mm long,
subtended by ovate bracts 1-3 mm long; lateral
branches 1'/,-3 cm with opposite small bracts
about the middle, bearing a terminal head, c. 1 cm
@ of c. 8-20 flowers, surrounded by an involucre of
small rounded pubescent bracts, pedicels c. 1 mm
or less (up to 3 mm in fruit). Calyx rim obsolete.
Corolla 4—5-lobed above, tubular below, 2-3 mm
long. Stamens 4-5, exserted. Ovary subcylindric,
4-5-celled. Fruit a spherical drupe (when dry 4—5-
ribbed, 7 by 5 mm).

Distr. Malesia: widespread in the Philippines
(Luzon to Mindanao).

Ecol. In primary forest, frequently beside
streams in damp ravines, from the lowland at
75 m to 750 m.

Vern. Kamay-kamay, Tag., ayum, C.Bis.

Note. Sparingly branched but wide-spreading
slender shrub, with yellowish bark, at first heavily
dotted with brown lenticels. The flowers are white
and the ripe fruit smooth and purple.

40. Osmoxylon luzoniense (MERR.) PHILIPSON,
Blumea 23 (1976) 116. — Boerlagiodendron
luzoniense MerRR. Philip. J. Sc. 3 (1908) Bot. 252;
En. Philip. 3 (1923) 223. — Boerlagiodendron
clementis MERR. Philip. J. Sc. 3 (1908) Bot. 155;
En. Philip. 3 (1923) 222. — Boerlagiodendron
agusanense ELMER, Leafl. Philip. Bot. 7 (1914)
2330; MerrR. En. Philip. 3 (1923) 222. — Boer-
lagiodendron diversifolium MERR. Philip. J. Sc. 10
(1915) Bot. 333; En. Philip. 3 (1923) 223.

Erect, unbranched or sparsely branched shrub or
tree to 8 m, becoming glabrous except for the
inflorescence. Leaves clustered at the ends of the
branches; petiole to 40 cm, with a clasping base
prolonged as a short stipular ligule, and with
several basal entire or shortly fimbriate crests;
blade to 33 cm @, base truncate or cordate, deeply
palmately 3—7-lobed, sinuses broad, rounded, lobes
elliptic, usually narrowed below, entire or with
subsidiary lobes (the central lobe especially often
narrow below and strongly pinnately lobed), apex
acute, margin coarsely serrate, coriaceous; upper-
most leaves often reduced and simple. Inflorescence
a terminal compound, subsessile umbel 10-15 cm
@; primary rays c. 20-30, 2-3 cm long, subtended
by lanceolate furfuraceous and + fimbriate bracts,
furfuraceous villose or + hirsute, with opposite
rounded or lanceolate hirsute bracts at the apex,
each ending in three branches; central branch
c. 2-8 mm long, hirsute, ending in an involucre of
minute bracts (1 mm) surrounding a globose
(2 cm @) umbellule of c. 15-20 sterile flowers
(c. 6 by 6 mm, 2-3-celled), pedicels 2-3 mm, hirsute;
lateral branches c. 21/,-31/, cm long at anthesis,

with an articulation about the middle, + hirsute,
ending in a globose head, 2 cm @ (with open
corollas), with ovate tomentose obtuse bracts,
c. 2 mm long; flowers 30-40, + sessile (rarely
pediceis to 2 mm). Calyx rim obsolete. Corolla
4-5-lobed above, tubular below, 3'/.-4 mm long.
Stamens 4-5, exserted, filament stout, 5 mm, anther
1 mm long. Ovary 4-S-celled. Fruit 6 by 5 mm
(dry), strongly 4—-5-ribbed.

Distr. Malesia: widespread in the Philippines
(Luzon to Mindanao), also in N. Celebes.

Ecol. In forests, often by streams and on ridge
in mossy forest, 280-1650 m.

Vern. Philippines: bolwang hi inalahan, If.,
iyangnok, Mbo, malakapayas,S.L.Bis., molonpolon,
Buk., tafigan-tangan-batu, Buk., tachung, vafiganeg,
Ig.
Notes. Inflorescence yellow to red (salmon),
fruits blue-black or purple.

MERRILL did not liken his Boerlagiodendron
diversifolium (from Mindanao) to this species, no
doubt because he gave importance to the occur-
rence of variable leaves and the 5-merous flowers.
However, specimens from Luzon may possess
simple leaves below the umbel, and both 4- and
5-merous flowers occur in both Luzon and
Mindanao. The greater range of material now
available establishes the identity of the two species.
Similarly, no features seem to distinguish Boer-
lagiodendron clementis, and B. agusanense though a
greater range of collections would be desirable.

The species is treated here in a broad sense. The
inflorescence characters of most specimens are
uniform, being hirsute and with the bracts at the
apex of the primary rays obtuse and short. Some
specimens (CURRAN 5088, ELMER 16762) have
longer lanceolate bracts with some bristles on the
back. The northernmost specimen, from Ilocos
Norte, has finer and less hairy inflorescence rays
(recalling 39. O. trilobatum), but the foliage agrees
with this species. Leaf-shape is more variable, even
on the same specimen. Other species which
resemble O. luzoniense in some respects are: 32.
O. camiguinense with broader, shallower lobing, a
more delicate inflorescence, and _ tri-merous
flowers; 27. O. pectinatum with glabrous inflores-
cence branches and long-pectinate petiolar crests;
and 26. O. humile with pedicelled flowers forming
less dense heads.

A specimen from Surigao Province (BS 83562)
has a most interesting abnormal structure. The
central branches of the inflorescence rays bear
heads of apparently fertile flowers, with lobed
corollas and exserted stamens.

41. Osmoxylon insigne (MIQ.) Becc. Malesia 1
(1878) 195; Puiipson, Blumea 23 (1976) 117. —
Trevesia insignis Mia. Ann. Mus. Bot. Lugd.-Bat. 1
(1863) 222. — Trevesia palmata var. insignis
CLARKE, FI. Br. Ind. 2 (1879) 732, pro nomen. —
Eschweileria insignis (M1Q.) BoERL. Ann. Jard. Bot.

1979]

ARALIACEAE—I (Philipson) 53

Btzg 6 (1887) 122. —. Boerlagiodendron insigne
(Miq.) Harms in E. & P. Nat. Pfl. Fam. 3, 8
(1894) 31.

A glabrous tree. Leaves large, palmately lobed;
petiole to 45 cm, 8-10 mm wide, flattened above,
with a sheathing base prolonged as a stipular
ligule, numerous fimbriate crests around the base
of the petiole, and irregular tufts of bristles along
the whole length of the petiole; blade to 55 cm @,
deeply 5S—7-lobed, the sinuses broadly rounded and
c. 4-5 cm from the base of the blade, lobes
pinnatilobed, with a narrow base and an attenuated
apex, margins serrate. Inflorescence a terminal
compound umbel c. 16 cm @; peduncle stout, with
lanceolate bracts (11/,-2 cm long) subtending the
primary rays; primary rays 25-30, c. 4 cm long,
bearing 2 caducous lanceolate bracts (c. 1 cm long)
at the apex, each ray ending in 3 branches; central
branch c. 12-16 mm long, bearing an umbel
(c. 2 cm @) of c. 20 sterile ovoid bacciform flowers
(c. 4 mm long when dry) with pedicels c. 4-6 mm
long, and 3-celled; two lateral branches c. 31/, cm
long, with an articulation about the middle, termi-
nating in an umbellule (c. 2 cm @) of c. 10-15
flowers on short stout pedicels c. 2 mm long,
umbellules surrounded by a receptacular rim after
caducous bracts have abscissed. Calyx rim minute.
Petals c. 31/, mm long in bud (when dry), with 8-9
lobes above, tubular below. Stamens 8-9, with
stout filaments. Ovary cylindric, 8—-9-celled; disk
with a central double row of pustulate stigmas.
Fruit unknown.

Distr. Malesia: Moluccas (Batjan).

Note. Tufts of bristles along the entire length of

the petiole together with the pinnatifid lobes of the
leaf are distinctive. SEEMANN (J. Bot. 4, 1866, 353)
referred to 5-flowered umbels with 5-angled drupes,
but this probably relates to the New Guinea
specimen which he included under this name.

Insufficiently known

Boerlagiodendron ledermannii HARMS, Bot. Jahrb.
56 (1920) 383; PHILIPSON, Blumea 23 (1976) 117. —
Type: LEDERMANN 12293.

HARMS compared this species with Boerlagio-
dendron geelvinkianum. The size of the foliage and
flowers prevents it from being included within that
species. If it represents a local species, it has not
been re-collected since the original gathering of
LEDERMANN in 1912. The type specimen, which was
incomplete, was destroyed during the war.

Boerlagiodendron monticola HaARMs in K. Sch. &
Laut. Fl. Schutzgeb. Nachtr. (1905) 330; Puitip-
SON, Blumea 23 (1976) 117. — Type: SCHLECHTER
14471.

The incomplete type specimen, gathered by
SCHLECHTER, was destroyed during the war. This
species was evidently similar to 38. Osmoxylon
micranthum, but the ovary was possibly 10-celled.
I have tentatively identified RopBins 1644 as this
species: it is close to O. micranthum but its ovary,
with 8 cells, is outside the range of variation of that
species and the pedicels are longer (in fruit). Its
distribution (foothills of Adalbert Range) is not
dissimilar to that of Boerlagiodendron monticola
(Torricelli Mts).

8. ARTHROPHYLLUM

BL. Bijdr. (1826) 878; DC. Prod. 4 (1830) 266; Mia. Ann. Mus. Bot. Lugd.-Bat. 1
(1863) 27; Bru. in B. & H. Gen. PI. 1 (1865) 944; Harms in E. & P. Nat. Pfi. Fam.
3, 8 (1894) 54; Koorpb. Atlas 4 (1916) f. 675 & 676; HuTcH. Gen. FI. Pl. 2 (1967)
80; STONE, Gard. Bull. Sing. 30 (1977) 276; PHILIPSON, /.c. 299, f. 1-16; Adansonia
17 (1978) 329. — Mormoraphis JACK ex WALL. Cat. (1831) n. 4931, nomen. —
Eremopanax BAILL. Adansonia 12 (1878) 158. — Fig. 17, 19-23.

Unarmed, sparingly branched trees or shrubs. Leaves on vegetative shoots and
lower leaves spirally arranged, imparipinnate, often crowded at the end of the
branches, those on flowering branches often opposite, smaller, or reduced to a
single leaflet; petiole terete; rachis articulated at the insertion of the pinnae and
leaflets; leaflets entire; stipular sheath clasping, small, ligule a mere rim. Jnflores-
cence consisting of compound umbels, either solitary and terminal or more
commonly borne on a cluster of specialized leafy branches arising from the axils
of the uppermost leaves; pedicels not articulated. Flowers bisexual. Calyx an
undulate rim, sometimes with indistinct teeth, persistent. Petals 4-6, valvate in
bud. Stamens 4-6, anthers curved, basifixed. Ovary turbinate, 1-celled; disk

FLORA MALESIANA [ser. I, vol. 91

. 17. Habit of Arthrophyllum diversifolium Bu. (Photogr. HOOGLAND, Bogor).

1979] ARALIACEAE—I (Philipson) $5

fleshy, rising in the.centre to the + sessile capitate stigma. Fruit ovoid or spheroidal,
often oblique; exocarp leathery; endocarp cartilaginous. Seed solitary, pendulous;
endosperm deeply transversely ruminate.

Distr. About 31 spp. (17 in Malesia) extending from the Nicobar Is. and Indo-China to the Philippines,
New Guinea and New Caledonia. Fig. 18.

Notes. The principal distinctive characters of this isolated genus are the single-celled ovary and the
arrangement of the umbels on specialized lateral branches in the majority of the species.

The present treatment, following that which I gave in 1977, /.c., remains tentative until widespread field
studies can be undertaken.

KEY TO THE SPECIES

1. Inflorescence becoming paniculate by the successive development of branches below the umbellules.
Fig. 19a 1. A. proliferum

1. Inflorescence a compound umbel.
2. Inflorescence with four orders of branching.
3. Leaflets lanceolate.
4. Leaflets c. 4-7 cm long. Fig. 20f
4. Leaflets much longer
3. Leaflets broader.
5. Pedicels (at early anthesis) c. 10 mm long.

6. Peduncles of umbellules with bracts or their scars. Fig. 21a
6. Peduncles of umbellules without bracts or their scars. Fig.21b. . . .. .

2. A. ashtonii
3. A. angustifolium

4. A. ahernianum
5. A. engganoense

5. Pedicels (at early anthesis) c. 5 mm long, or shorter.

i eaeeaves associated with. the umbels rotund. Fig. 2le.... . 2 0.2 26 ws ns

6. A. collinum

7. Leaves (or leaflets) associated with the umbels ovate or elliptic.
8. Leaves associated with the umbels ovate, + fleshy, with the lower surface smooth (lateral veins

obscure). Fig. 21f

7. A. crassum

8. Leaves (or leaflets) associated with the umbels + elliptic, coriaceous or chartaceous, veins

visible.
9. Young parts glabrous
9. Young parts with rufous tomentum.

10. Umbels at anthesis with numerous filamentous pedicels. Fig. 21c.
10. Umbels at anthesis with fewer stout pedicels. Fig. 21d

Re E Spt GAG cuee cus 3m (sania date oN Sa ce 8. A. pacificum

9. A. diversifolium
10. A. macranthum

2. Inflorescence with three (or fewer) degrees of branching.

11. Leaflets membranaceous or chartaceous.

12. Mid-leaflets c.16cmlong. .......

12. Mid-leaflets c. 8 cm long or shorter.

C) eter te hela web Me. die lek baie) cel) ce) Mis ais,

11. A. papyraceum

13. Leaflets usually 5—7 (Fig. 20a). Primary inflorescence branches usually short (c. 3-6 cm) and

without articulations

Pernt? “Ope Whe Oe Be ake ao)

12. A. maingayi

13. Leaflets more numerous (Fig. 20d). Primary inflorescence branches longer (10-20 cm), with one

or more leafy nodes
11. Leaflets coriaceous.

14, Leaflets c. 5

14. Leaflets more numerous.

15. Petals and stamens 6
15. Petals and stamens 4 or 5.

16. Leafiets 6 cm long, or longer. Fig. 20b .

16. Leaflets shorter. Fig. 20c

phos Lek. jet itis. le amie

1. Arthrophyllum proliferum PHILIPSON, Gard.
Bull. Sing. 30 (1977) 302, f. 3-4. — Fig. 19a-b.
Medium-sized, glabrous tree. Leaves multijugate,
up to 90 by 24 cm, of the flowering branches smaller
with fewer pinnae or usually simple; petioles up to
28 cm, 5 mm @; petiolules 1-1'/, cm; leaflets
obovate-oblong, c. 16 by 7 cm, chartaceous, margin
slightly revolute, base broadly cuneate to truncate,

es Ce oa ne CARTS ye Or MO. ey) mc

Chet <TC h Tyee Gomet ye mary

© 05 ey 0) ew edule. Me) 6. ems. elk lo bike

oe fe wee le is el Je Biel ee ce ow

oY detense, i eemey <. cee cen el (isl Me exe ela ccle eialek. (al Ble. mie

13. A. kjellbergii

Sn ee ae ren ral Mn’ chal ited 14. A. cenabrei

sometimes oblique, apex acute. Inflorescences
terminating specialized plagiotropic shoots, bear-
ing axillary flowering branches and ending in
umbellules of a few flowers below which pairs or
whorls of branches continue the growth of the
inflorescence to produce an elongated panicle of
umbellules; pedicels 7-8 mm (slightly longer in
fruit). Calyx often with 5 indistinct teeth. Petals 5,

56 FLORA

MALESIANA

[ser. I, vol. 91

———- — a

Fig. 18. Species density of Arthrophyllum BL. in

Malesia; above the hyphen the number of endemic

species, below it the non-endemics. Complete

range of genus encircled; 17 of the total of 31 spp.
occur in Malesia.

3'/, mm long, narrowly triangular. Stamens 5,
filaments c. 2 mm, anthers reniform, c. 1/, mm long.
Ovary obconical, c. 2 mm long; disk fleshy,
cushion-like; stigma capitate + sessile. Fruit
ovoid, fleshy, capped by the calyx and the enlarged
beak-like stylopodium, c. 10 by 5 mm when dry.

Distr. Malesia: E. New Guinea (Morobe Distr.).

Ecol. Mid-mountain rain-forest, reaching the
canopy, on steep slopes, 300-1200 m.

Notes. The thick outer bark is grey-brown,
fissured, and peeling in small flakes. Wood straw-
coloured. Cut stems exude brown latex. The thick
petals are yellow-green.

The flower and fruit are typical of this well-
defined genus, but the branching of the inflores-
cence is unlike that found in all other species.

2. Arthrophyllum ashtonii PHILIPson, Gard. Bull.
Sing. 30 (1977) 303, f. 12. — Fig. 20f.

Slender small tree, to 5 m, with the leaves dis-
persed for some distance from the apex of the
branches, young parts with brown scurfy tomen-
tum which persists on the umbellules. Leaves
multijugate, to c. 30 cm long; of the flower-bearing
branches smaller with fewer pinnae, or simple;
petioles slender, c. 7-9 cm; petiolules c. 4-7 mm;
leaflets lanceolate to broadly lanceolate, 3-7!/, by
1-2 cm, thinly coriaceous, margin revolute, base
broadly cuneate, apex tapered to subcaudate, veins
channelled above, visible beneath. Inflorescence a
terminal cluster of specialized leafy branches; main
rays variable in length in the same inflorescence,
the longest from c. 16-25 cm long with a pair of
opposite simple leaves about the middle (with
flowering branches in their axils) and ending in a
whorl of secondary rays subtended by simple
leaves; secondary rays 4-12 cm long, bearing
simple leaves, usually in an opposite pair and
terminating in compound umbels; umbellules with
c. 8-10 flowers; pedicels 4-10 mm, furfuraceous.
Flower buds c. 2 mm long, calyx a furfuraceous rim.
Petals 5, broadly triangular. Stamens 5, anthers

curved, basifixed. Ovary glabrous, obconical,
c. 1 mm long; stigma on a raised stylopodium at
the centre of a flat disk. Fruit spheroidal, with a
persistent stylopodium, c. 6 mm long when dry.

Distr. Malesia: Borneo (Sarawak and Brunei).

Ecol. Mossy forest on sandstone ridge, and in
kerangas forest, 1000-1550 m.

Note. The small narrow leaflets are very dis-
tinctive. No other species with small leaflets has
inflorescences which branch to the fourth degree.

3. Arthrophyllum angustifolium RID. J. Fed. Mal.
St. Mus. 10 (1920) 136; Fl. Mal. Pen. 1 (1922) 885;
PHILIPSON, Gard. Bull. Sing. 30 (1977) 304.

Shrub or small tree, up to 5 m, young parts
rufous-tomentose, glabrescent. Lower /eaves multi-
jugate, rachis dilated, c. 70-90 by 30-40 cm;
petioles c. 15-22 cm, 3-5 mm wide, ligule a rim
c. 2 mm long; petiolules c. 7-10 mm long; leaflets
coriaceous, lanceolate, c. 15-22 by 11/,-21/, cm,
tapering to an acute or obtuse apex, base cuneate,
margin slightly revolute; upper leaves reduced,
mostly unifoliolate, opposite, broader, with
petioles 2-4 cm long. Jnflorescence a terminal
cluster of specialized leafy branches; main rays
30 cm (or more) long, bearing simple leaves in
opposite pairs with small flowering branches in
their axils, ending in a whorl of c. 10-12 secondary
rays subtended by simple leaves; secondary rays
8-12 cm long, bearing pairs of simple leaves, and
terminating in an umbel of 5-12 tertiary rays
c. 2-3 cm long, with bract scars about the middle,
and ending in an umbellule of c. 8-12 flowers,
pedicels c. 5 mm. Petals 5, 2-3 mm long in bud.
Stamens 4. Ovary turbinate, inconspicuous at
anthesis; disk fleshy; stigma + sessile. Fruit
spheroidal, c. 5 by 5 mm when dry, calyx and
stylopodium small.

Distr. Malesia: Malay Peninsula (Perak) and
Borneo (Brunei).

Ecol. Forest and old regenerated forest on
peat swamp at low altitude or on ridges.

Note. The lanceolate leaflets are unlike those of
any other species. The grey bark is minutely
fissured and bears many small orange lenticels.
The wood is soft and white. The Malayan and
Bornean specimens are similar, except that the
flower buds are larger in the Brunei plant.

4. Arthrophyllum ahernianum Merr. Philip. J.
Sc. 1 (1906) Suppl. 109; En. Philip. 3 (1923) 235;
PHILIPSON, Gard. Bull. Sing. 30 (1977) 304, f. 13. —
A. pinnatum (non CLARKE) F.-VILL. Nov. App.
(1880) 103; VIDAL, Sinopsis Atlas (1883) 28, t. 55
f. c. — Macropanax sp. VIDAL, Rev. Pl. Vasc. Filip.
(1886) 145. — A. sablanense ELMER, Leafl. Philip.
Bot. 1 (1908) 331. — A. borneense MERR. PI. Elm.
Born. (1929) 231, non BAKER, 1896. — A. elmeri
MERR. Webbia 7 (1950) 319. — A. merrilliana
FurTAbo, Gard. Bull. Sing. 19 (1962) 185. —
Fig. 21a.

1979]

ARALIACEAE—I (Philipson) 57

d

Fig. 19. Arthrophyllum proliferum PHILIPSON. a. Part of inflorescence in fruiting state, x 1/2, b. ditto,

showing detail of ultimate branches in flowering stage, nat. size. — A. maingayi PHILIPSON. c. Terminal

inflorescence, < 1/,. A. montanum RIDL. d. Single flowering shoot, « 1/3; (Courtesy Gard. Bull. Sing. 30,
1977).

Tree up to 15 m, young parts with rufous tomen-
tum. Leaves clustered at the ends of the branches,
multijugate, up to 200 by 60 cm; of the flower-
bearing branches smaller with fewer pinnae, or
simple; petiole stout, up to 35 cm; petiolules
5-20 mm; leaflets ovate-oblong, occasionally
oblong-lanceolate, up to 35 by 12 cm, membrana-
ceous or chartaceous, margin revolute, base
cuneate to rounded, usually oblique, apex short
acuminate. Inflorescence a whorl of specialized
leafy branches forming a terminal crown; main rays
up to 150 cm (or more), bearing pinnate leaves
usually in 1-2 opposite pairs, and with flowering

branches in the upper axils, ending in a whorl of
secondary rays subtended by pinnate or more
rarely simple leaves; secondary rays up to 30 cm
bearing simple or pinnate leaves in opposite pairs,
and terminating in compound umbellules; um-
bellules with c. 10-20 flowers c. 3 cm @; peduncles
with 1-2 pairs of small simple, often caducous
leaves; pedicels c. 1 cm (at anthesis) subtended by
minute caducous bracts. Petals 5, 4 mm long.
Stamens 5, anthers curved. Ovary turbinate; disk
fleshy; stigma + sessile. Fruit c. 10 by 7 mm,
ellipsoidal, calyx and stylopodium forming a
prominent beak.

58 FLORA MALESIANA

[ser. I, vol. 91

:

i?)

Fig. 20. Foliage leaves of Arthrophyllum spp. a. A. maingayi PHILIPSON, b. A. montanum RIDL., c. A. pulga-
rense ELMER, d. A. kjellbergii PHILIPSON, e. A. alternifolium MAINGAY ex RIDL., f. A. ashtonii PHILIPSON.
All = 1/3; (Courtesy Gard. Bull. Sing. 30, 1977).

Distr. Malesia: N. Borneo and throughout the
Philippines to the northern Moluccas (Talaud,
Ternate).

Ecol. Primary and second-growth forest, from
the lowlands to 1000 m.

Vern. Philippines, cf. MERRILL: alabihig,
dokloi, P.Bis., binalayon, C.Bis., danipo, \g., higin,
Mang., malapapdya, puyga-puygahan, Tag., pama-
latafigen-a-purau, Ik. Additional names: Philip-
pines: /ulpo, Luzon, mayari, Mindoro, bungyo,
Palawan; Moluccas: Jangator’a, Talaud.

Notes. This species replaces the more westerly
9. A. diversifolium which it closely resembles. It is

characteristically larger in all its parts, particularly
in the size of the individual flowers and the length
of their pedicels. There are fewer flowers in an
umbellule.

The distinction between these two species is not
always easy to make, especially when the material
is fragmentary; PHILIPSON, /.c. A few specimens
from the Philippines appear very similar to
A. diversifolium, and it is possible that this species
extends beyond Borneo. I have regarded them as
part of the range of variation of A. ahernianum.
Similarly, at least one specimen from S. Borneo
approaches A. ahernianum in appearance.

1979]

5. Arthrophyllum engganoense PHILIPSON, Gard.
Bull. Sing. 30 (1977) 305, f. 14. — Fig. 21b.

Tree to 21 m high, becoming glabrous. Lower
leaves imparipinnate, multijugate, 60 cm long or
more; petiole 24 cm, 6 mm wide, rachis articulated
at the insertion of the leaflets; petiolules 10-18 mm;
leaflets broadly elliptic to elliptic-oblong, c. 12-15
by 6-7 cm, base rounded with a short asymmetrical
cuneate centre, apex shortly apiculate, margin
entire, often undulate chartaceous. Flowering
branches c. 40 cm; leaves opposite, simple, or
unifoliolate, petiolules c. 5-7 cm, leaflets ovate,
c. 13 by 6 cm, with inflorescence branches in their
axils; ending in a whorl of simple leaves surround-
ing a compound umbel to 30 cm @; secondary rays
c. 8, c. 10-15 cm long at flowering, slender (2 mm @)
and striate, each bearing a pair of small leaves
about the middle (sometimes with inflorescences
in their axils) and ending in an umbel; tertiary
rays c. 8, slender, c. 20-40 mm long, without
bracts; tertiary rays pedicels c. 5-10 per umbellule,
c. 10-15 mm at anthesis. Ca/yx an undulate rim.
Petals 5, c. 2 mm long in bud. Stamens 5, anthers
curved. Ovary turbinate, obscurely ribbed,
c. 2'/, mm long at anthesis. Fruit ellipsoid, c. 10 by
7 mm, with a rather small persistent calyx and
stylopodium.

Distr. Malesia: S. Sumatra (Enggano I.), two
collections.

Ecol. Forest at low altitude, up to c. 100 m.

Vern. Langkapu utan kaauh, Enggano.

Note. The two known collections of this species
are very similar and contrast with the widespread
9. A. diversifolium because of the few-flowered
umbellules with long, spreading pedicels.

6. Arthrophyllum collinum PHILIpsoN, Gard. Bull.
Sing. 30 (1977) 305, f. 17. — Fig. 21e.

Sparingly branched shrub or small tree up to
12 m, all young parts with dense, rufous tomentum
which persists on flowers and inflorescences.
Leaves multijugate, up to 60 (or more) by 32 cm;
of the flowering branches smaller with fewer pinnae
or more frequently unifoliolate, leaflets more
rotund and with longer petioles; petiole up to
20 cm; petiolules c. 1 cm; leaflets oblong, broadly
elliptic or rotund, up to 16 by 7 cm, coriaceous,
margin slightly revolute, base truncate to rounded,
unequal, apex rounded, obtuse, or shortly and
bluntly apiculate, midrib prominent, lateral veins
usually clearly visible below, upper surface fre-
quently rugose. Inflorescence a cluster of specialized
leafy branches forming a terminal crown; main rays
up to 60 cm long, bearing one or more, rarely two,
opposite pairs of usually unifoliolate rotund leaves
often with flowering branches in their axils, and
ending in a whorl of secondary rays, subtended by
usually unifoliolate leaves; secondary rays up to
17 cm, bearing a pair of unifoliolate leaves with
flowering branches in their axils, and terminating
in compound umbellules; tertiary rays c. 5—10,

ARALIACEAE— | (Philipson) 59
Q ol. 9

pots if

Fig. 21. Umbellules of some Arthrophyllum spp.

a. A. ahernianum MEeERR., b. A. engganoense

PHILIPSON, c. A. diversifolium BL., d. A. macranthum

PHILIPSON. All nat. size. — Bracts of two Arthro-

phyllum spp. e. A. collinum PHILIPSON, f. A. crassum

PHILIPSON. Both x !/; (Courtesy Gard. Bull. Sing.
30, 1977).

c. 3-6 cm long, articulated near the middle;
pedicels c. 10, c. 2-3 mm long, rufous tomentose.
Petals 5, broadly triangular, c. 2 mm long. Stamens
5, anthers curved. Ovary turbinate, rufous-tomen-
tose, c. 1!/, mm long at anthesis; disk fleshy, stigma
sessile. Fruit ellipsoidal, c. 9 by 5 mm (when dry),
the calyx and stylopodium prominent.

Distr. Malesia: Borneo (Sarawak and Sabah).

Ecol. Forest and scrub, sometimes growing as
an epiphyte in the crowns of trees, 600-2700 m.

Notes. The bark is grey and smooth, the wood
pale and soft, and the cut stems exude a yellowish
or orange latex.

This species is characterized by the rotund,
coriaceous leaves on the flowering branches.

7. Arthrophyllum crassum PHILIPSON, Gard. Bull.
Sing. 30 (1977) 305, f. 18. — Fig. 21f.

Sparingly branched shrub or small tree up to
8 m, occasionally epiphytic, all young parts with
dense, rufous tomentum which persists on the

60 FLORA MALESIANA

[ser. I, vol. 9!

flowers and inflorescence. Leaves multijugate, up
to 100 (or more) by 50 cm, occasionally with 4
leaflets inserted at some of the lower articulations;
of the flowering branches smaller with fewer pinnae
or more frequently unifoliolate; petiole up to
25 cm; petiolules 10-20 mm; leaflets elliptic,
oblong or lanceolate, up to 24 by 71/, cm, coria-
ceous, margin strongly revolute, base cuneate to
rounded, often oblique, apex narrowed to an acute
often caudate apiculum, midrib prominent, lateral
veins faint to obscure. Inflorescence a cluster of
specialized leafy branches forming a terminal
crown; main rays up to 70 cm, bearing an opposite
pair (or rarely more pairs) of unifoliolate or (less
frequently) pinnate leaves, often with flowering
branches in their axils, and ending in a whorl of
secondary rays, subtended by usually unifoliolate
leaves; secondary rays up to 25 cm, bearing a pair
of usually unifoliolate leaves with flowering
branches in their axils, and terminating in com-
pound umbellules; tertiary rays c. 10, c. 2-3 cm
long, articulated near the middle; pedicels c. 12-14,
c. 3-4 mm long, rufous-tomentose. Petals 5,
broadly triangular, c. 2 mm long. Stamens 5,
anthers curved. Ovary turbinate, rufous-tomentose,
c. 1*/, mm long at anthesis; disk fleshy, stigma
sessile. Fruit spheroidal, c. 6 by 5 mm (when dry),
calyx and stylopodium prominent.

Distr. Malesia: Borneo (Sarawak and Kaliman-
tan).

Ecol. Swampy peat forest and heath woodland,
both primary and disturbed, from sea-level to
c. 150m. Some fragmentary collections from higher
altitude (1000 m) further inland may belong to this
species.

Note. The leaves associated with the in-
florescence are distinctively fleshy, have a strongly
revolute margin and a smooth lower surface with
indistinct lateral veins, and are ovate with rather
acute apex. The leaflets of the pinnate leaves on the
vegetative shoots are also rather leathery with
indistinct lateral venation.

8. Arthrophyllum pacificum PHILIPSON, Gard.
Bull. Sing. 30 (1977) 306.

A slender, glabrous tree to 14 m. Leaves multi-
jugate, c. 60 by 24 cm; of the flower-bearing
branches smaller with fewer pinnae or simple;
petioles c. 22 cm, 3-4 mm @;; petiolules c. 1 cm;
leaflets elliptic, oblong or ovate, c. 8-10 by 4-5 cm,
rather membranaceous, margin entire, very slightly
revolute, base abruptly cuneate, often oblique,
apex obtuse, acute, or slightly apiculate (in a
specimen from Morotai attenuated). Inflorescence
a whorl of specialized leafy branches forming a
terminal crown to the vegetative shoots; main rays
35 cm, bearing small pinnate leaves in opposite
pairs and with flowering branches in the axils,
ending in a whorl of secondary rays, subtended by
a whorl of simple leaves (bracts); secondary rays
10-24 cm long, bearing simple or trifoliolate leaves

usually in opposite pairs, and terminating in
umbellules; umbellules c. 7—-12-flowered; pedicels
c. 5 mm at anthesis, slightly elongating in fruit,
minute bracts caducous. Petals 5, broadly triangu-
lar, 2'/.-3 mm long. Stamens 5, anthers reniform.
Ovary obconical, 1'/, mm long; disk fleshy,
furrowed when dry; stigma + sessile. Fruit
ellipsoidal, fleshy, somewhat oblique, conical beak
with calyx and stigma, c. 9 by 5 mm when dry.

Distr. E. Malesia: Moluccas (Morotai) to the
Bismarck Archipelago.

Ecol. Primary forest on mountain slopes,
attaining Nothofagus mossy forest, 500-2000 m.
Reported as common in most localities.

Vern. Kainsoka, Ambai, Japen I.

Notes. All specimens from islands to the north
of New Guinea are similar, in having more delicate
foliage than 10. A. macranthum from the mainland
of New Guinea. The specimens from Morotai are
sterile, and have narrower more tapering leaflets.

The bark is light brown and the cut branches
exude a clear latex. The flowers are light green with
yellow anthers, and the fruit is black.

9. Arthrophyllum diversifolium BL. Bijdr. (1826)
879; DC. Prod. 3 (1830) 266; Mra. FI. Ind. Bat. 1,
1 (1856) 767; Sum. (1861) 340, incl. var. lanceolata
Miq.; CLARKE, FI. Br. Ind. 2 (1879) 733; K. & V.
Bijdr. 7 (1900) 46; Koorp. Exk. Fl. Java 2 (1912)
717; Atlas 4 (1916) f. 675 & 676; BACK. & BAKH.
f. Fl. Java 2 (1965) 169; Stone, Gard. Bull. Sing. 30
(1977) 135; Putieson, /.c. 306, f. 15. — A. javani-
cum BL. Bijdr. (1826) 879; DC. Prod. 4 (1830) 266;
Back. & BAKH. f. Fl. Java 2 (1965) 169. —
A. ellipticum BL. Bijdr. (1826) 879; DC. Prod. 3
(1830) 266. — Mormoraphis sumatrana JACK ex
WALL. Cat. (1831) n. 4931, nomen. — A. blumeanum
Z. & M. Syst. Verz. (1846) 41, nom. illeg.; Mia.
Fl. Ind. Bat. 1, 1 (1856) 768; Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 27, incl. var. oblongatum Mia.,
var. ellipticum (BL.) Mia. et var. ovalifolium
(JUNGH. & DE VRIESE) Mig. — A. ovatifolium
JUNGH. & DE VRIESE, Ned. Kruidk. Arch. 1 (1846)
19; Ann. Sc. Nat. Paris III, 6 (1846) 117; Mia.
Fl. Ind. Bat. 1, 1 (1856) 768, t. 14 (‘ovalifolium’);
Rip. Fl. Mal. Pen. 1 (1922) 885; Stone, Gard.
Bull. Sing. 30 (1977) 136, f. 2. — A. dilatatum
Mia. FI. Ind. Bat. 1, 1 (1856) 768. — A. borneense
BAKER, Kew Bull. (1896) 23; Merr. En. Born.
(1921) 457. — A. congestum RIDL. J. Fed. Mal.
St. Mus. 10 (1920) 137; Fl. Mal. Pen. 1 (1922) 887;
Stone, Gard. Bull. Sing. 30 (1977) 136. — A. havi-
landii RipL. Kew Bull. (1933) 494. — A. rufosepa-
lum Rip. Kew Bull. (1946) 40. — A. rubiginosum
Ripe. .¢741) is Ad 2ic. 225235

Small tree, up to 14 m, young parts with rufous
tomentum. Leaves clustered at the ends of the
branches, imparipinnate or bipinnate (rarely
tripinnate) with leaflets at the insertion of the
lateral rachides, multijugate, 150 by 45 cm (wider
in bipinnate leaves); usually in opposite pairs on

1979] ARALIACEAE—] (Philipson) 61

KO

: i
E ~~)
Na

RS AxS

S SS ) =D

AS BY &

Ak HK W S

/) g \ & =>

» (WES
jel

AY

She

IN
CE
Wire

tS

A)
AA
Fie We
(iY AY BE
Be AG
ae

AERERe
CHhbhic

ss

Fig. 22. Arthrophyllum diversifolium BL. Diagrammatic sketch of habit, showing vegetative and flowering
branches, 3/3, (Courtesy Gard. Bull. Sing. 30, 1977).

the inflorescence-bearing branches and smaller rarely unifoliolate) leaves mostly in opposite pairs
with fewer pinnae or unifoliolate; petiole up to and usually with flowering branches in the axils of
40 cm; petiolules '/,-11/, cm; leaflets ovate-oblong the upper leaves, ending in an umbel of secondary
or elliptic, up to 24 by 11 cm (leaflets of bipinnate rays subtended by a whorl of pinnate or unifoliolate
leaves usually c. 10 by 5 cm), + coriaceous or leaves; secondary rays up to c. 30 cm, bearing
(especially in bipinnate leaves) somewhat mem- mainly simple leaves in opposite pairs with
branaceous, margin slightly revolute, base truncate, flowering branches in their axils and terminating
rounded, or cuneate, often oblique, apex shortly in compound umbellules each subtended by a
acuminate, veins usually 5—7 pairs. Inflorescence a whorl of often caducous bracts; tertiary rays
cluster of specialized leafy branches forming a (peduncles) c. 5 cm, articulated about the middle;
terminal crown which abscisses after fruiting; umbellules with c. 30-40 flowers, c. 17-20 mm 2;
main rays up to 150 cm, bearing pinnate (or more _ pedicels c. 20, c. 5 mm at anthesis (longer in fruit),

62 FLORA MALESIANA [ser. I, vol. 9!

Fig. 23. Arthrophyllum diversifolium BL. Schematic drawing of a single flowering shoot, with four degrees
of branching, « 1/; (Courtesy Gard. Bull. Sing. 30, 1977).

1979 ]

with an involucre of minute caducous bracts.
Petals 5, c. 2 mm long. Stamens 5, anthers curved.
Ovary turbinate, often inconspicuous at anthesis;
disk fleshy, rising in the centre to a sessile stigma.
Fruit ellipsoidal, c. 9 by 7 mm; calyx and stylo-
podium forming a conspicuous beak.

Distr. Malesia: Sumatra, Malay Peninsula,
Java, Borneo, Celebes.

Ecol. In a wide variety of habitats, on dry
sandy soil to swampy humus, in primary lowland
and montane rain-forest and also in secondary
forest, heath-forest and waste land, from sea-level
to 1600 m.

Uses. Concoctions of the root and bark are
reported to have medicinal properties, including
a remedy for syphilis, and the plant has stupifying
and poisonous properties.

Vern. Sumatra: antjaneudeung uding, bidju,
bolu bolu, bulu, (kayu) abang-abang, k. attu turut,
légung, Riouw Arch., mapang, miu, obang, potah,
silanta, ténjam dakan, Banka, tocrah. Malay
Peninsula: chindangan utan, jolok hantu, lupa
dahan, (pokok) restong, ségan budahan, susun
kélapa, tum bong ninyor, tusum perpah. Java: dajo,
délahan, déleg, délek, djangkorrang, gombong,
(kayu) gompang, kédrja, kidjangkurang, kingom-
pang, ki ompong, kléntjang, krépang, langitj, malas
bérdahan, pongporang, putjangan. Borneo: merje-
meh, Sarawak, karadjungjung, ‘Kalimantan.
Celebes: kKambabah, susangkangan.

Notes. The very widespread A. diversifolium is
variable in many characters, and may comprise a
number of geographic subspecies, but no basis for
this is apparent at present. Most individuals have
the lower leaves simply imparipinnate, whereas
others have bipinnate, or rarely tripinnate, leaves.
The flowers and inflorescences of these forms
appear to be identical, though rapid changes in the
umbellules after flowering produce a deceptively
distinctive appearance in specimens at different
stages of development. Field experience over the
. whole range of the species will be required to
understand this interesting leaf-polymorphism.
In treating all forms as one species I am partly
influenced by the fact that most authors who have
been familiar with the plants in Java (where both
forms occur) have regarded the complex as a single
species (the fact that some authors have recognized
the variant from Mt Salak as a distinct species does
not affect the problem of leaf-polymorphism).

Apart from the strikingly different leaf forms
just discussed, certain local variants may eventually
be shown to justify specific rank. A form growing
on Mt Salak (near Bogor) has often been regarded
as distinct (see, for example, HOCHREUTINER,
Candollea 2, 1925, 481, and BACKER & BAKHUIZEN
VAN DEN BRINK f. FI. Java 2, 1965, 169). Indeed
this form is the basis of the name 4A. diversifolium.
I retain this name in preference to the other two
names published simultaneously by BLUME
because it has been most consistently adopted since

ARALIACEAE—I (Philipson) 63

it was first used in this comprehensive sense by
CLARKE, /.c. On the evidence available I do not
consider the Salak plants any more distinctive than
many other local variants.

It might be considered that 5. A. engganoense is
also no more than another such variant, but its
facies is so marked that specific rank appears
justified.

It is possible that RIDLEY was correct in distin-
guishing A. congestum, but the material is not good
and appears inadequate to confirm specific status.

Five collections from Brunei and a neighbouring
district of Sarawak are all very alike and sufficiently
distinct from both A. diversifolium and 7.
A. crassum to suggest that they represent a
separate taxon, but for the present they are tenta-
tively retained as a form of A. diversifolium.

Similarly, the two collections described by
Rip.ey as A. rubiginosum and A. rufosepalum are
based on collections which are not altogether
typical of A. diversifolium, but which come closest
to that species. In the absence of more supporting
material, it is advisable not to retain them as
species. The first of these names (A. rubiginosum)
has been widely used in identifications of Bornean
specimens, but the specimens concerned are either
typical A. diversifolium or belong to the distinctive
7. A. crassum.

Specimens from Mt Kinabalu described by
Ripcey as A. havilandii have bipinnate leaves, and
appear to conform well with A. diversifolium. This
form was again collected on. Mt Kinabalu by
CLEMENS and is also known from Sarawak.

The smooth bark is whitish to greyish brown
with pustulate lenticels; the wood is cream, with a
colourless aromatic exudate. The flowers are
yellowish with a sickly sweet scent. Seedlings have
simple and trifoliolate leaves.

10. Arthrophyllum macranthum PHILIPSON, Bull.
Br. Mus. Nat. Hist. Bot. 1 (1951) 18; Gard. Bull.
Sing. 30 (1977) 308, f. 16. — A. diversifolium (non
BL.) Harms, Bot. Jahrb. 56 (1920) 413. — Fig. 21d.

Tree up to 25 m, sparsely branched with leaves
crowded at the ends of the branches, all young
parts with dense rufous tomentum which may
persist on the flowers and inflorescence. Leaves
multijugate, up to 100 by 30 cm; of the flower-
bearing branches smaller with fewer pinnae, or
simple; petioles stout, up to 40 cm, clasping base
heavily lenticellate; petiolules '/,-2 cm; leaflets
ovate to oblong, up to 16 by 8 cm, coriaceous, mar-
gin revolute, base rounded, truncate, or cordate,
very rarely cuneate, often oblique, apex obtuse or
bluntly apiculate, principal veins arched-ascending,
reticulations visible especially beneath (pinnae of
leaves on the flowering branches usually elliptic
with a cuneate base). Inflorescence a whorl of
specialized leafy branches forming a terminal
crown; main rays up to 60 cm, bearing pinnate
leaves often in opposite pairs and with flowering

64 FLORA MALESIANA

[ser. I, vol. 91

branches in the upper axils, and ending in a whorl
of secondary rays subtended by a whorl of usually
simple leaves (bracts); the secondary rays c. 10—
20 cm, bearing simple leaves, usually in opposite
pairs, and terminating in compound umbels;
umbellules (c. 2 cm) with c. 10-15 flowers; pedicels
stout, 3-5 mm to 10 mm or more in fruit, sub-
tended by minute bracts. Flower buds 5 mm or
more long; calyx a rim or with 5 indistinct teeth.
Petals 5, triangular, c. 4 mm long, fleshy. Stamens
5. Ovary obconical, c. 2'/, mm long; disk fleshy
(hemispherical in living material, conical and
furrowed when dry); stigma capitate, + sessile.
Fruit ellipsoid, sometimes slightly oblique, c. 12 by
8 mm, the stylopodium forming a conical beak
with the persistent stigma; exocarp fleshy, endo-
carp cartilaginous.

Distr. Malesia; New Guinea (from Irian Jaya
to Milne Bay Distr. and the Bismarck Archipelago).

Ecol. Usually a sub-canopy tree of rain-forest
ranging from the lower montane zone to mossy
subalpine woodland and scrubland, occasionally
in second growth, usually above 1000 m (up to
2700 m), but occurring also on the coastal scarps
of the Astrolabe Range.

Vern. Maguva, W. Sepik, agagwa, agare,
agugwa, alolo, angga, engga, pooli, tipilan, W. High-
lands, arua, hagegoa, wonkurumeh, E. Highlands,
kolom, S. Highlands.

Note. Trees become very different in appearance
when in flower or fruit: the spiral pinnate foliage
leaves are surmounted by tufts of branches which
end in inflorescences and bear much smaller leaves.
The foliage leaves are fleshy, leathery and glossy.
The ripe fruit is purple and shining. The bark is
grey, at first smooth with many leaf-scars and
lenticels, but small longitudinal fissures develop.
The cut branches exude a brown latex and a scent
of celery. The soft wood is white or straw-coloured.

11. Arthrophyllum papyraceum PHILIPSON, Gard.
Bull. Sing. 30 (1977) 308.

Shrub, rufous-tomentose on the young parts.
Leaves alternate, imparipinnate; petiole 15—20 cm;
leaflets c. 7, membranaceous, elliptic, c. 12-24 by
5-10 cm, base broadly cuneate, apex finely
acuminate, margin slightly revolute. /nflorescence
a terminal compound umbel; primary rays few (2),
2-3 cm long, 2 mm wide, without bracts (caducous),
secondary rays few (3), c. 13-18 mm long, articu-
lated about the middle, ending in an umbellule of
c. 10-12 flowers; pedicels 2-3 mm, slightly fur-
furaceous. Petals 5, c. 2 mm long in bud. Stamens
5, anthers curved. Ovary turbinate, glabrous,
obscurely ribbed. Fruit unknown.

Distr. Malesia: E. Sumatra (East Coast Res.
near Aek Sordang), one collection.

Ecol. Primary rain-forest.

Note. Known from a single collection (with no
duplicates) this species resembles 12. A. maingayi
in its simple inflorescence, and the few pinnae of its

foliage leaves. However, the large size of the leaflets
precludes its inclusion in that species.

12. Arthrophyllum maingayi PHILIpson, Gard.
Bull. Sing. 30 (1977) 309, f. 7. — A. pinnatum
CLARKE, Fl. Br. Ind. 2 (1879) 734, excluding
synonyms [see also SEEM. J. Bot. 4 (1866) 294];
RIDL. Fl. Mal. Pen. 1 (1922) 886. — Fig. 19c, 20a.

Low shrub or slender tree, rarely as high as
10 m, rufous-tomentose on the very young parts,
soon becoming glabrous. Leaves tufted at the ends
of the branches, usually with 5—7 leaflets (but up to
15), up to 30 by 18 cm; petiole to 12 cm (usually
shorter), 2 mm @; petiolules c. 0-10 mm; leaflets
elliptic or elliptic-lanceolate (occasionally the
lowermost pair of leaflets is replaced by pinnate
leaf segments), c. 8 by 3!/, cm, rather thin, base
cuneate or rounded, apex apiculate or caudate,
margin slightly revolute, lateral veins faint and
obscure; leaves below the flowering branches
sometimes reduced to 3 or | leaflet(s). Inflorescence
consisting of a number of primary branches
radiating from the end of a leafy shoot (which
forms a longer or shorter peduncle); primary
branches, often rather few, usually 3-6 cm long
and devoid of leaves except for a few terminal
simple or trifoliolate leaves around the compound
umbels, occasionally the branches bear pairs of
opposite leaves when they may be up to 30 cm long;
secondary rays c. 2-3 cm long; pedicels 4-10 cm.
Petals 5, 11/,-2 mm long in bud. Stamens 5. Ovary
turbinate, disk fleshy. Fruit spheroidal, c. 8 by
6 mm, calyx and stylopodium rather inconspicuous.

Distr. Malesia: Central W. Sumatra (Mt
Kerintji), throughout the Malay Peninsula (incl.
Penang) and Borneo.

Vern. Poko minta anak, Kedah, karon baru,
Sumatra.

Notes. The name ‘A. pinnatum’, misapplied by
CLARKE, /.c., has been in general use for this
species. However, this name was based on Panax
pinnatum LAMK, which in turn was based on the
Rumphian name ‘Scutellaria secunda’ ; this is quite
a distinct plant (see under 6. Polyscias cumingiana).

The three species A. maingayi, 16. A. montanum
and 17. A. alternifolium are similar in having
simpler inflorescences than 9. A. diversifolium.

A. montanum can be distinguished from A. main-
gayi by its more leathery leaves with more promi-
nent nervation, and by the more woody and leafy
flower-bearing primary branches of the inflores-
cence.

A. alternifolium is distinguished from both these
species by its small, coriaceous leaflets with incon-
spicuous nervation.

Most specimens can be readily distinguished, but
a few can be assigned to a species only doubtfully,
usually because the material is inadequate. This is
usually due to the junction between the vegetative
(spiral phyllotactic) shoots and the flower-bearing
branches being omitted.

1979 ]

13. Arthrophyllum kjellbergii PHriLIpsoN, Gard.
Bull. Sing. 30 (1977) 309, f. 10. — Fig. 20d.

Small tree, 10 m, branches c. 1 cm @, young parts
covered by rufous tomentum. Leaves clustered
towards the ends of the branches, multijugate,
25-30 by 16 cm; petioles c. 8 cm, 2 mm wide;
petiolules c. 3-8 mm; leaflets elliptic or ovate-
oblong, up to 8 by 3'/,; cm, chartaceous, margin
minutely revolute, base rounded to cuneate often
oblique, apex tapered to a blunt apiculum, or
rounded and mucronate. Inflorescence a terminal
cluster of c. 5—10 specialized branches (primary
rays); primary rays c. 15-20 cm, 1'/,-2 mm wide,
bearing near the middle an opposite pair of simple
leaves or sometimes trifoliolate leaves with short
flowering branches in their axils, and sometimes
with a second pair higher up, and 2-3 similar leaves
below the terminal cluster of secondary rays;
secondary rays c. 12, c. 2'/,-3'/, cm long, each
subtended by a small bract and bearing opposite
caducous bracts near the middle, terminating
in an umbellule of c. 12 flowers surrounded by an
involucre of caducous bracts (1 mm long). Flowers
known only in young bud. Fruit spheroidal,
c. 5 by 4 mm, calyx and stylopodium prominent;
pedicel 5-6 mm.

Distr. Malesia: SE. Celebes (Kendari).

Ecol. Primary rain-forest, 50-150 m.

Note. The small leaves and relatively simple
inflorescences are distinctive.

14. Arthrophyllum cenabrei Merr. Philip. J. Sc. 20
(1922) 417; En. Philip. 3 (1923) 235; PHILIPsoN,
Gard. Bull. Sing. 30 (1977) 309.

Glabrous tree, c. 10 m, ultimate branches c. 5mm
@. Upper leaves pinnate, up to 10 cm long, leaflets
mostly 5, sometimes 3, or the uppermost reduced
to simple leaflets, the rachis and petiole c. 4 cm;
leaflets mostly elliptic, 4'/,-6 by 2!/,-31/, cm,
chartaceous to subcoriaceous, very shortly and
obtusely acuminate, base acute, brownish oliva-
ceous and slightly shining when dry, nerves
3-4 pairs, slender; petiolules 5-10 mm. Peduncles
c. 4 cm, umbellately arranged at the tops of
the branchlets, usually however with solitary
inflorescences in the axils of the uppermost leaves,
thus forming a somewhat leafy inflorescence.
Fruits 5-8 in each umbel, ovoid, c. 7 mm @;
pedicels 8-10 mm.

Distr. Malesia: Philippines (Cebu; FB 28343,
type, not seen).

Ecol. On slopes at 600 m.

Vern. Bingliu, C.Bis.

Note. No specimen of this species has been
located. The above description is taken from
MERRILL’s original account. In placing this species
in the key, it has been assumed that the
inflorescence branching is relatively simple.

15. Arthrophyllum pulgarense ELmer, Leafl. Philip.
Bot. 7 (1915) 2551; Merr. En. Philip. 3 (1923) 235;

ARALIACEAE—I (Philipson) 65

PHILIPSON, Gard. Bull. Sing. 30 (1977) 311, f. 9. —
Fig. 20c.

Small tree, branches c. 1 cm @, young parts with
red tomentum, becoming glabrous except on the
ovaries. Leaves clustered towards the ends of the
branches, leaflets c. 6 pairs, c. 22 by 8 cm; petioles
c. 6 cm, 3 mm @; petiolules 5-6 mm; leaflets
elliptic to rotund, c. 4 by 23/, cm, coriaceous,
margin revolute, rounded to broadly cuneate, apex
rounded or abruptly tapered to a short obtuse
apiculum. Inflorescence a terminal cluster of
specialized branches (primary rays); primary rays
c. 6-10 cm, 3-4 mm @, bearing near the middle an
opposite pair of simple rotund leaves, sometimes
with flowering branches in their axils, and with a
whorl of similar leaves below the terminal cluster
of secondary rays; secondary rays c. 6-8, 2-4 cm
long, articulated about the middle with scars of
bracts or bearing a pair of small simple leaves
terminating in an umbellule of c. 8-12 flowers;
bracts caducous. Calyx with indistinct teeth. Petals
6, 2'/, mm long (in bud). Stamens 6, anthers
curved. Ovary obconical, 2 mm long, furfuraceous.
Fruit ellipsoid, c. 8 by 5 mm, the stylopodium
forming a beak with stigma and calyx; exocarp
fleshy.

Distr.
Pulgar).

Ecol. Common in montane forest on Mt Pulgar.

Note. The coriaceous, small, often rotund leaf-
lets are characteristic.

Malesia: Philippines (Palawan: Mt

16. Arthrophyllum montanum RIDL. J. Fed. Mal.
St. Mus. 4 (1909) 24; Fl. Mal. Pen. 1 (1922) 886;
PuHILIPsoN, Gard. Bull. Sing. 30 (1977) 311, f. 8. —
A. nitidum RipL. J. Fed. Mal. St. Mus. 7 (1916)
42; Fl. Mal. Pen. 1 (1922) 886. — A. ovatum
RIpDL. J. Fed. Mal. St. Mus. 7 (1916) 42; Fl. Mal.
Pen. 1 (1922) 886. — Fig. 19d, 20b.

Shrub or small tree to 6 m, unbranched or
sparingly branched, rufous-tomentose on the
young parts, becoming glabrous. Leaves tufted at
the ends of the branches, multijugate, c. 30-55 by
12-22 cm; petiole 9-21 cm, 3 mm @; petiolules
c. 10-15 mm; leaflets elliptic or oblong, 6-10 by
21/4 cm, coriaceous or chartaceous, base
cuneate, apex with a short blunt apiculum, margin
entire, revolute, the few principal lateral veins
usually rather prominent; the leaves associated
with the umbels usually unifoliolate, broadly ellip-
tic to rotund, with a petiole to 4'/, cm. Inflores-
cences on specialized leafy branches either in
terminal clusters or axillary in the upper leaves;
branches 10-30 cm, leaves mostly simple in
opposite pairs, usually without flowering branches
in their axils, branches ending in a whorl of simple
leaves surrounding a compound umbel; primary
rays c. 5-15(-20), 2-6 cm, with scars of caducous
bracts; pedicels 4-8 mm. Petals (4-)5, 2 mm long
in bud. Stamens (4-)5. Ovary turbinate; disk
fleshy, stigma + sessile. Fruit spheroidal, c. 8 by

66 FLORA MALESIANA [ser. I, vol. 91

Fig. 24. Gastonia serratifolia (MiQ.) PHILIPSON. a. Habit, x */3, b. lower bud, c. flower, d. fruit, both x5
(a—c SCHMUTZ 3612, d VAN ROYEN 4090). Drawn by HELENE MULDER.

1979]

ARALIACEAE—]I (Philipson) 67

5 mm, calyx and stylopodium rather prominent.

Distr. Malesia: Malay Peninsula (Kedah to
Selangor).

Ecol. Primary forest ascending to the montane
zone, and in second-growth, 250-1500 m.

Note. Similar to 12. A. maingayi but dis-
tinguished by the more leathery leaves and by the
leafy inflorescence branches which are usually
absent in A. maingayi.

17. Arthrophyllum alternifolium MAINGAY ex
RIDL. Fl. Mal. Pen. 1 (1922) 886; PHILipson, Gard.
Bull. Sing. 30 (1977) 311, f. 11. — A. pinnatum
CLARKE, FI. Br. Ind. 2 (1879) 734, p.p., excel.
basionym; Kina, J. As. Soc. Beng. 67, ii (1898) 59,
p.p. — A. alternifolium MAINGAY ex CLARKE, FI.
Br. Ind. 2 (1879) 734, nomen in synon. — Fig. 20e.

Slender, sparingly branched shrub to 2 m,
rufous-tomentose on the young parts, becoming
glabrous. Leaves tufted at the ends of the branches,
multijugate, c. 20-25(-30) by 9-12(-15) cm;
petiole terete, c. 3-6(-9) cm, 2-3 mm @; petiolules
c. 2 mm; leaflets ovate, elliptic or lanceolate,
31/,4(-6) by 1-2(-2'/,) cm, coriaceous, base
cuneate, apex acuminate to caudate, obtuse,
margin revolute, veins obscure; leaves associated
with the umbels (if any) reduced, with fewer
leaflets or unifoliolate, sometimes broadly ovate.
Inflorescence usually a terminal compound umbel,
occasionally a whorl of leafy branches (5-14 cm
long) (leaves usually simple in opposite pairs), each

ending in a compound umbel; peduncle 1'/,—
4'/, cm, with one or more usually caducous simple
(or trifoliolate) leaves at the apex; primary rays
c. 5, 3-4'/, cm, with scars of caducous leaves about
the middle, each ending in an umbellule of c. 12-25
flowers, pedicels 5-8 mm. Calyx a rim or minutely
4-5-dentate. Petals 4-5, 2 mm long in bud. Stamens
4-5. Ovary turbinate; disk fleshy, stigma + sessile.
Fruit spheroidal, c. 5 by 5 mm when dry, calyx and
stylopodium small.

Distr. Malesia: Malay Peninsula (Johore: Mt
Ophir; Pahang, Selangor, and Malacca).

Ecol. In shady montane forest, with Rhodo-
dendron and Dacrydium, 900 m and above.

Note. The small, coriaceous, often apiculate
leaflets are characteristic. Although collected most
frequently on Mt Ophir it occurs on other high
ridges in southern Malaya.

Excluded

Arthrophyllum ceylanicum Mia. Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 27 (type in L), is according to
VAN STEENIS, Rec. Trav. Bot. Néerl. 24 (1927) 819
= Oroxylum indicum (L.) Kurz (Bignoniaceae).

Arthrophyllum reticulatum Bui. ex Mia. Ann.
Mus. Bot. Lugd.-Bat. 1 (1863) 27 (type in L), is
according to MIQUEL (/.c. 318) and VAN STEENIS
(vide supra) = Oroxylum indicum (L.) Kurz
(Bignoniaceae).

9. GASTONIA

Comm. ex LAMK, Encycl. 2 (1786) 610; Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1863)
5; HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 43; Hurcu. Gen. FI. Pl. 2 (1967) 68;
PHILIPSON, Blumea 18 (1970) 491, 497, f. 1-10. — Tetraplasandra (non A. GRAY)
Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 4; HARMs in E. & P. Nat. Pfl. Fam. 3, 8
(1894) 29, p.p.; Nachtr. 2 (1900) 253. — Indokingia HEMSL. in Hook. Ic. Pl. (1906)
t. 2805. — Peekeliopanax HARMS, Notizbl. Berl.-Dahl. 9 (1926) 478, fig. — Fig. 24,
26, 27.

Trees unarmed with thick branches, glabrous or tomentose. Leaves large, im-
paripinnate, exstipulate; rachis articulated; leaflets in pairs entire or crenate;
petiole terete, with clasping base. Flowers in umbellules which are arranged race-
mosely, or in verticils, on strong inflorescence branches; pedicels not articulated
below the ovary. Calyx forming a continuous rim with an entire or indistinctly
denticulate margin. Corolla of 5-13 free petals or calyptrate, fleshy, valvate.
Stamens either equal in number to the petals or up to several times as many;
filaments usually rather short and thick; anthers large, often irregularly lobed,
dorsifixed. Ovary inferior, broadly obconic, cells 7-22; disk fleshy with stylar
processes equal in number to the cells arising from its centre. Fruit a spherical berry
(strongly ribbed when dry), with an indistinct calyx rim, a flattened disk, and a

68 FLORA MALESIANA

[ser. I, vol. 9!

prominent stylopodium bearing a ring of radiating stigmatic arms; exocarp fleshy,
endocarp crustaceous. Endosperm with smooth surface.

Distr. About 10 spp. in East Africa, Madagascar, the Seychelles and Mascarenes, Malesia, and the

Solomon Is.

Ecol. Primary and second-growth forest, or in open country, from sea-level to lower montane zone.
Taxon. I have amply analyzed and discussed the affinities of Gastonia (Blumea 18, 1970, 497). I have
come to the conclusion that within its alliances Gastonia is the only genus west of Samoa; in Polynesia

there are three other closely allied genera.

KEY TO THE SPECIES

1. Corolla with free petals. Ovary cells and style arms 6—9(-12). Whole plant glabrous. Leaflets usually
entire or sparsely crenate. Main inflorescence branches borne along an elongated axis with caducous

bracts; peduncles of the umbellules mostly aggregated into pseudo-whorls

1. G. serratifolia

1. Corolla calyptrate. Ovary cells and style arms 12-18(—22). Young parts with scurfy tomentum, per-
sisting on the ovary and bracts. Leaflets strongly crenate. Main inflorescence branches borne sub-
umbellately on a short axis with persistent bracts; peduncles of the umbellules scattered

1. Gastonia serratifolia (MIQ.) PHILIPSON, comb.

nov. — Arthrophyllum serratifolium Mia. Sum.
(1861) 341, type from Sibolga, leg. TEYSMANN
(in U). — G. papuana Mia. Ann. Mus. Bot.

Lugd.-Bat. 1 (1863) 5; Harms, Bot. Jahrb. 56
(1921) 408; PHitipson, Blumea 18 (1970) 492, 500
f. 3. — Tetraplasandra paucidens Mia. Ann. Mus.
Bot. Lugd.-Bat. 1 (1863) 4; Koorb. Minah. (1898)
488. — G. eupteronoides T. & B. Nat. Tijd. N. I. 25
(1863) 416. — Polyscias papuana (MiIQ.) SEEM.
J. Bot. 3 (1865) 181. — Tetraplasandra koordersii
Harms, Ann. Jard. Bot. Btzg 19 (1904) 12; Ic.
Bog. 2 (1906) t. 178. — Tetraplasandra philippinen-
sis Merr. Philip. J. Sc. 1 (1906) Suppl. 219; En.
Philip. 3 (1923) 222. — G. winkleri Harms in
Fedde, Rep. 15 (1917) 20. — Tetraplasandra
solomonensis PHILIPSON, Bull. Br. Mus. Nat. Hist.
Bot. 1 (1951) 11. — Fig. 24.

Shrub or small tree, sometimes epiphytic, up to
27 m, with clear bole to 15 m, dbh 34 cm, crown
sparsely branched with leaves crowded at the ends
of the branches, glabrous. Leaves up to 80 by
20 cm; petiole c. 13 cm; leaflets c. 10 pairs,
petiolules up to 1 cm; blade oblong, ovate or
lanceolate, middle leaflets 8—-14(-18) by 2!/,-
31/,(-8) cm, chartaceous, entire and subrevolute,
or with a few obscure, more rarely several promi-
nent crenations, apex rounded and bluntly apicu-
late or tapering and acute, base broadly cuneate,
midrib prominent. Inflorescence terminal, glab-
rous, with a stout primary axis 15-25 cm long,
bearing scattered or clustered branches along its
length and ending in an umbel of c. 10 branches;
bracts caducous; secondary branches 30-40 cm,
bearing numerous subverticillate peduncles along
their length and ending in an umbel; peduncles
c. 3 cm, somewhat longer in fruit; pedicels
1-21/, cm, forming umbellules of c. 10 flowers.
Flower buds (when dry) c. 7 by 3 mm. Calyx rim

2. G. spectabilis

undulate. Petals 5-9, slightly fleshy, fully separated.
Stamens variable in number, (7—)14—55; filaments
short; anthers broad and irregularly lobed, variable
in size. Ovary glabrous, smoothly rounded below,
slightly constricted below the calyx; cells 6-12,
usually c. 9, disk with a prominent rim and, at
anthesis, a central boss formed by closely appressed
subulate style arms equal in number to the ovary
cells. Fruit c. 9 by 7 mm (without stylopodium),
the flattened stylopodium ending in a ring or
double row of radiating subulate stigmatic arms,
black when ripe, the fleshy exocarp enclosing com-
pressed crustaceous pyrenes.

Distr. Solomon Is.; in Malesia: Central W.
Sumatra (Sibolga, Enggano I.), Malay Peninsula
(Johore), Sunda Straits (islet Dwars in den Weg),
West Java, Lesser Sunda Is. (Sumba, Timor,
Wetar, Flores), W. Borneo (east of Pontianak,
Karimata), N. Borneo (Sabah), Philippines

Fig. 25. Distribution of Gastonia in Malesia.

Known localities of G. serratifolia (M1Q.) PHILIP-

SON: black dots; range of G. spectabilis (HARMS)
PHILIPSON: broken line.

1979] ARALIACEAE—I (Philipson) 69

SEQ NTT RC

Fig. 26. Gastonia spectabilis (HARMS) PHILIPSON, two inflorescences visible below the leaves (Photogr.
PHILIPSON, Kassam Pass, E. New Guinea, 1968).

70 FLORA MALESIANA

[ser. I, vol. 91

(Palawan, Balabac), Celebes (Lepo-Lepo, Luwuk,
Minahassa), N. Moluccas (Talaud Is.), New
Guinea (Vogelkop, NW. & SW. Irian, Schouten
I., Waigeu I.). Fig. 25.

Ecol. Primary and secondary forest, or in open
country, usually at low altitude and often on the
shore or sea-cliffs, but ascending to 1000 m. EyMa
noted it to be a characteristic tree on Mt Tam-
bunan, Luwuk, E. Celebes.

Vern. Bajur talang ékoaho, M, Enggano, jarum,
Sabah, raka, Sumba, kre, wangka, Flores, lampo
pad, bungku, Celebes, bufigio, Palawan, lantora,
Talaud Is., mansnongoree, mantsenongor, Schouten
I., raauwrack, ara-orach, Vogelkop, Maibrat.

Note. The entire or serrate nature of the leaf
margins, the number of ovary cells, and especially
the number of stamens are variable characters but
show no discernable geographical segregation. The
outer bark is described as light brown, with small
oblong brittle scales. Inner bark, leaves and
inflorescences with copious sticky juice. Wood soft
white. The petals are variously described as purple,
light green, and white.

2. Gastonia spectabilis (HARMS) PHILIPSON, Blumea
18 (1970) 494, pl. 1. — Peekeliopanax spectabilis
Harms, Notizbl. Berl.-Dahl. 9 (1926) 478, fig.;
PHILIPSON, Blumea 18 (1970) 500 f. 2. — G. bori-
diana Harms, Bot. Jahrb. 69 (1938) 282. —
Fig. 26, 27.

Tree up to 40 m high, clear bole to 28 m, dbh
1.75 m, crown sparsely branched with the branches
whorled or regularly forked and the leaves crowded
at the ends of the branches, all young parts with
scurfy indumentum more evident in dried material.
Leaves up to 80 by 30 cm; petiole c. 15 cm, with
some tomentum remaining at the joints; leaflets
c. 11 pairs on petiolules c. 2-8 mm, oblong or
elliptic, middle leaflets 10-15 by 4-6 cm, charta-
ceous when dry, prominently crenate, apex nar-
rowed to a short blunt apiculus, base rounded or
truncate. Inflorescence in forks well below the
leaves, when in bud covered with large scurfy
cataphylls, and when mature with radiating
branches forming clusters c. 130 cm @. Primary
axis short (5-10 cm) with persistent bracts;
secondary branches numerous, radiating, up to
65 cm, bearing small, persistent, scurfy bracts and
peduncles scattered along their length and clustered
in a terminal umbel; peduncles c. 1'/,-5 cm;
pedicels 14/,-1'/, cm, forming umbellules of
c. 5-12 flowers. Flower buds c. 8 by 5 mm when dry
(fresh c. 12 by 8 mm). Calyx rim straight. Petals
6-12, very fleshy, incompletely separated (often
splitting into c. 5 lobes). Stamens 25-66, often
c. 35, filaments short; anthers broad and irregularly
lobed, variable in size. Ovary with a short dense
indumentum, smoothly rounded below, cells
usually c. 16, very rarely fewer than 10, as many as
22; disk at anthesis with a prominent rim and a
central boss formed by closely appressed subulate

Fig. 27. Gastonia spectabilis (HARMS) PHILIPSON.

Young tree, free bole 19 m, 47 cm @, crown 10 m,

longest leaves 2 m, not yet flowering (Photogr.
G. PEEKEL, New Ireland, Ugana, 1940).

styles equal in number to the cells. Fruit c. 8 by
10 mm (dry and without stylopodium), the flattened
disk with a prominent stylopodium ending in an
elliptical ring of radiating subulate stigmatic arms;
the fleshy exocarp enclosing compressed crusta-
ceous pyrenes.

Distr. Solomon Is.; in Malesia: New Guinea
(Vogelkop, NW. Irian, Papua New Guinea),
Bismarcks, and New Ireland. Fig. 25.

Ecol. Primary and secondary rain-forest, and in
cultivated areas, 200-2000 m.

Vern. Amoriga, bekuak, djak, ntjier, tuju,
Vogelkop, bohko, boinga, gabiel, jamwa, mestic,
sikoto, tubat, tumbala, Madang Distr., waki,
W. Highlands, aita, E. Highlands, kuhuh, Papua.

Note. Possibly the largest araliad known. The
regular habit of branching results from the vegeta-

1979] ARALIACEAE—I (Philipson) 71

Fig. 28. Polyscias nodosa (BL.) SEEM. a. Leaf and part of inflorescence, x */s, D. leaflet, x 2/3, c. branch
of inflorescence, x 2/3, d. umbellule, e. flower bud and ditto in LS, x 8, f. ovary and ditto in CS, x 8,
g. stamens (VERSTEEGH BW 3868). Drawn by HELENE MULDER.

7/p2 FLORA MALESIANA

[ser. I, vol. 9!

tive shoots springing in pairs or whorls from below
terminal inflorescence buds. Several flushes of
growth occur as an inflorescence bud matures, so
that at anthesis the inflorescences are situated in
forks well below the leafy crown, with a succession

Bole without buttresses or with buttresses 1 m high
and 2 m wide. Outer bark brown with prominent
pustular lenticels and small shallow fissures.
Exudate from cuts abundant, clear and aromatic.
Wood soft. Flowers cream, stamens yellow. Ripe
fruit dark red-brown.

of younger inflorescence buds in higher forks.

10. POLYSCIAS

J. R. & G. Forster, Char. Gen. (1776) 63, t. 32; DC. Prod. 4 (1830) 257; SEEM.
J. Bot. 3 (1865) 179; Bru. in B. & H. Gen. PI. 1 (1876) 941; Harms in E. & P. Nat.
Pfi. Fam. 3, 8 (1894) 43; Koorp. Atlas 4 (1916) f. 677-680; MerR. Int. Rumph.
(1917) 408; Harms, Bot. Jahrb. 56 (1921) 409; Merr. En. Philip. 3 (1923) 233;
Hutcu. Gen. FI. Pl. 2 (1967) 75; BERNARDI, Candollea 26 (1971) 13; PHILIPSON,
Blumea 24 (1978) 169. — Eupteron Mia. Pl. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1
(1856) 762; Hutcu. Gen. FI. Pl. 2 (1967) 68. — Nothopanax Mig. P\. Jungh. 3
(1855) 425; Bonplandia 4 (1856) 139; Fl. Ind. Bat. 1, 1 (1856) 765; SEEM. Fl. Vit.
(1866) 114; Merr. Int. Rumph. (1917) 409; En. Philip. 3 (1923) 233. — Irvingia
F.v.M. Fragm. 5 (1865) 17, non Hook. f. 1860. — Kissodendron SEEM. J. Bot. 3
(1865) 201; ibid. 6 (1868) 129; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45;
Bot. Jahrb. 56 (1921) 412; Hutcu. Gen. Fl. Pl. 2 (1967) 75. — Palmervandenbroekia
Gisss, Arfak (1917) 162; Hutcu. Gen. Fl. Pl. 2 (1967) 75. — Gelibia Hutcu. Gen.
FI. Pl. 2 (1967) 57. — Fig. 28, 30.

Unarmed shrubs or trees, glabrous or furfuraceous. Leaves imparipinnate or
2-3-pinnate (rarely unifoliolate) with an elongated or short sheathing base;
rachis articulated; petiole terete; leaflets in pairs, entire, crenate or dentate.
Inflorescence terminal, often large, a panicle, corymb, or compound rarely simple
umbel. Flowers in umbellules, capitula, or racemose; pedicels articulated below the
ovary. Calyx a rim with an undulate or dentate margin. Petals 4-5(-8 or more),
valvate. Stamens equal in number to the petals; anthers dorsifixed. Ovary inferior,
4-5(-8 or more)-celled; disk fleshy; styles either free and recurved (at least in fruit)
or joined to form a beak-like stylopodium. Fruit a spherical or ovoid drupe,
crowned by the persistent calyx rim and the styles or stylopodium; exocarp fleshy,
endocarp chartaceous. Endosperm with an uneven surface, fissured, or rarely
smooth.

Distr. About 100 spp. throughout the tropics of the Old World (incl. Australia), and the Pacific
Islands; in Malesia 23 spp. of which 3 adventive: rare in West (none native in Sumatra and Malaya), more
common in East, the majority in New Guinea. Fig. 29.

Ecol. Primary or second-growth forest, from sea-level to 2650 m.

Note. Araliads with pinnate leaves and an articulated pedicel are here regarded as forming one genus,
divided into several sections. These sections have formerly been segregated as genera, but I follow

BERNARDI, /.c., in uniting them. The most distinctive is sect. Polyscias, characterized by an elongated leaf-
sheath. Several of the species of this section are cultivated and have a number of cultivars.

KEY TO THE SECTIONS

1. Leaf-sheath elongated, extending along the petiole for about '/, of its length (Spp. 1-7)
1. Sect. Polyscias
1. Leaf-sheath short or obsolete, restricted to the base of the petiole.
2. Style arms spreading, at least in fruit.

1979]

3. Flowers arranged racemosely (Sp.8)... .

3. Flowers arranged in umbellules or capitula (Spp. 9-14)... 2... .
2. Style arms erect, fused, forming a beak in fruit.

4. Inflorescence large, + as long as the leaves (Spp. 15-19)

4. Inflorescence much shorter than the leaves (Spp. 20-23).

ARALIACEAE—I (Philipson) 13

. . . 4 Sect. Kissodendron
5. Sect. Palmervandenbroekia

1. Section Polyscias

PHILIPSON, Blumea 24 (1978) 169.

Aromatic, glabrous shrubs or small trees, often cultivated. Leaf-sheath elon-
gated along the petiole for '/,'/, of its length. Styles spreading, at least in the fruit.

Distr. Polynesia, Queensland, Malesia, and SE. Asia.

KEY TO THE SPECIES

femlbeaves 2—S-pinnate.s im cit, <o.- oe

1. Leaves imparipinnate, or unifoliolate.
2. Ovary 2-celled (or predominantly so).

3. Main inflorescence branches diffusely branched
3. Main inflorescence branches with verticils of short branches

1. P. fruticosa

2. P. macgillivrayi
3. P. verticillata

2. Ovary variable, but many flowers with more than 2 cells.

4. Leaflets orbicular or reniform. Leaves unifoliolate or trifoliolate. ......

4. P. scutellaria

4. Leaflets ovate, oblong, or elliptic (cultivated forms often laciniate or lanceolate). Leaves with 3 or

more pairs of leaflets.

5. Leaf margin sharply serrate (blade often rhomboidal and variegated with light yellow)

5. Leaf margin entire, or obscurely dentate.

5. P. guilfoylei

6. Peduncles of the ultimate umbellules bearing 1 or more pairs of small bracts (reduced flowering

branches sometimes present in their axils)

©. Keema Senge: le ie) alen cee sep uicl ume Kel ge fe.

6. P. cumingiana

6. Peduncles of the ultimate umbellules either without bracts, or with 1 or more small bracts inserted

singly

1. Polyscias fruticosa (L.) HARMs in E. & P. Nat.
Pfi. Fam. 3, 8 (1894) 45; K. & V. Bijdr. 7 (1900) 2;
Harms in K. Sch. & Laut. Fl. Schutzgeb. (1900)
486; Koorp. Exk. Fl. Java 2 (1912) 716; Atlas 4
(1916) f. 680; BAILEY, Rhodora 18 (1916) 153, incl.
var. plumata (Hort.) BAILEY; BACK. & BAKH. f.
FI. Java 2 (1965) 168; PHiLipson, Blumea 24 (1978)
169. — Scutellaria tertia RuMpH. Herb. Amb. 4:
78, t. 33. — Panax fruticosum LINNE, Sp. PI.
ed. 2 (1763) 1513; BL. Bijdr. (1826) 830; DC.
Prod. 4 (1830) 254; BLANCO, FI. Filip. ed. 2 (1845)
156; ed. 3, 1 (1877) 281, t. 78; CLARKE, FI. Br. Ind.
2 (1879) 725; F.-ViLL. Nov. App. (1880) 101;
BoeRL. Handl. 1 (1890) 647; F.v.M. Descr. Pap.
Pl. 9 (1890) 60; Wars. Bot. Jahrb. 13 (1891) 396;
Koorpb. Minah. (1898) 488; MERR. Philip. J. Sc. 3
(1908) Bot. 84. — Panax obtusum BL. Bijdr. (1826)
880; DC. Prod. 4 (1830) 254. — Aralia tripinnata
BLANCO, FI. Filip. (1837) 223, cf. MERR. Sp. Blanc.
(1918) 295. — Nothopanax fruticosum (L.) Mia.
Pl. Jungh. 3 (1855) 425; Fl. Ind. Bat. 1, 1 (1856)
765; SEEM. FI. Vit. (1866) 114, 115; J. Bot. 4 (1866)
363; MerRR. FI. Manila (1912) 358, incl. var.
plumatum (Hort.) MERR. et var. victoriae (Hort.)

7. P. javanica

Me_rR.; Int. Rumph. (1917) 410; Sp. Blanc. (1918)
295; Harms, Bot. Jahrb. 56 (1921) 412; MERR. En.
Philip. 3 (1923) 233; HEYNE, Nutt. Pl. (1927) 1209;
OcHsE & BAKH. Veg. D.E.I. (1931) 64, f. 37;
CorNneER, Ways. Trees (1940). 150. — Nothopanax
obtusum (Bt.) Mia. FI. Ind. Bat. 1, 1 (1856) 766;
SEEM. FI. Vit. (1866) 114. — P. obtusa (BL.) HARMS
in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45, nom. illeg.,
non BLANCO, 1837, quae est Schefflera odorata
(BLANCO) Merr. & ROLFE; Koorp. Exk. Fl. Java 2
(1912) 716; Atlas 4 (1916) t. 679 S; BACK. & BAKH.
f. Fl. Java 2 (1965) 168.

Glabrous shrub or, small tree, up to 5 m, with
branches bearing spirally arranged leaves towards
their ends. Leaves tripinnate, variable in size to
c. 75 cm long; petiole up to c. 25 cm, with a sheath-
ing base to 5 cm long; pinnae to c. 25 cm long;
leaflets shortly petiolate, very variable in shape and
size, oblong or linear-lanceolate, or 1-12 by */,-
4 cm, more or less deeply serrate or lobed, or
irregularly pinnatisect, apex attenuate, acuminate
or rounded, base cuneate, truncate or subcordate,
midrib and lateral veins evident. Inflorescence a
diffuse panicle; primary axis up to c. 60 cm with

74 FLORA MALESIANA

[ser. I, vol. 9

secondary branches mostly in verticils at intervals
along its length; secondary branches up to 30 cm
bearing umbellules in an irregularly branched
system towards their extremities; umbeilules with
12-20 flowers on pedicels c. 3 mm long. Calyx a
minute rim. Petals 5, 2 mm long. Stamens 5. Ovary
turbinate, c. 1 mm high, 2—3(-4)-celled; styles at
first erect, later spreading. Fruit subglobose,
fleshy, c. 5 mm @ when dry.

Distr. Native country not properly known,
cultivated throughout the region, and in other parts
of the Indo-Pacific tropics.

Ecol. Grown at low and moderate altitudes
(c. 1000 m) as an ornamental or hedge shrub and
for culinary use.

Vern. (from various sources). Sumatra: orang
aring, Medan, kédongdong mekka, Palembang;
Malaya: daun girang, siku kluang, M; Java:
kédongdong alus, k. batur, k. laut, M, imba,
kédongdong laki, k. tjina, randa nunut, S, tjakar
kutjung, t. tjikri, J, kadongdung, k. laut, k. petédhan,
Md; Celebes: boka ula risé, kéndém rinték,
Minahasa, Alfur lang., bombu, Makassar; Philip-
pines: papud, Tag., Bik., bani, makan, Bik.;:
Moluccas: daun papeda papua, pagar pagar,
Ambon, guarbati, tampusong, Ternate.

The name kedongdong belongs properly to
species of the fruit tree genus Spondias, but is
sometimes also applied to trees of other families
with pinnate leaves; papua means ‘curly’.

Notes. The foliage of this plant is extremely
variable in size and form. Typical plants are figured
by Koorpers (1916, /.c.) and by OcHsE & BaAK-
HUIZEN VAN DEN BRINK (/.c.). These may be readily
identified by the intricately compound leaf.

The individual leaflets are characteristically
ovate-lanceolate and serrate to deeply pinnatifid.
However, broader leafiets with simpler outlines
are not uncommon. The more rotund, blunter
leaflets of P. obtusa are considered here to be an
extreme form of this species (the type is bipinnate).

fe
| z q
ae we
DOYS, Sig Geant, meet 6
Ss =<
SG BSE
Se oii gx
= Sa Se
ee s Na

<1 os —— oe

Fig. 29. Species density of Polyscias J. R. & G.

FORSTER in Malesia; above the hyphen the number

of endemic species, below it the non-endemics.
Only the native species.

Occasional specimens have leaves so much
reduced that they are simply pinnate or unifoliolate
when they approach forms of 6. P. cumingiana.

A number of names have been applied to horti-
cultural forms belonging to this section of the
genus (cf. BAILEY, Rhodora 18, 1916, 153), but the
interrelations of these will be understood only
after intensive biosystematic study of the many
cultivars.

2. Polyscias macgillivrayi (SEEM.) HARMs in E. &
P. Nat. Pfl. Fam. 3, 8 (1894) 45; Puitipson, Bull.
Br. Mus. Nat. Hist. Bot. 1 (1951) 14; Blumea 24
(1978) 169. — Nothopanax macgillivrayi SEEM. FI.
Vit. (1866) 114, nomen. — Panax macgillivraei
(SEEM.) BTH. Fl. Austr. 3 (1866) 382. — P. grandi-
folia VoLKENS, Bot. Jahrb. 31 (1901) 471;
KANEHIRA, En. Micron. Pl. (1935) 385. — Tieghe-
mopanax macgillivrayi VIGUTER, Bull. Soc. Bot.
Fr. 52 (1905) 313. — Polyscias sp. C. T. Wurrte,
J. Arn. Arb. 10 (1929) 255.

Glabrous shrub or small tree, up to 15 m, with
few branches bearing terminal clusters of large
leaves. Leaves imparipinnate, up to 1 m (or more)
long, uppermost leaves smaller; petiole c. 15 cm,
with a sheathing base extending for 6-10 cm along
the petiole; petiolules c. 10-15 mm; lamina oblong
often broader near the base (lower and the terminal
pinnules more ovate) c. 20-25 by 8-10 cm, slightly
succulent when fresh, margin entire, revolute or
occasionally minutely dentate, apex rounded and
shortly apiculate, base rounded, truncate or sub-
cordate, midrib and widely spaced laterals promi-
nent. Inflorescence a large panicle; primary axis
stout, rather short (c. 3-10 cm), often bearing
reduced leaves or cataphylls basally, and a few
lateral inflorescence branches, terminating in an
umbel of long diffusely branched rays; lateral
branches and rays c. 40-50 cm, with secondary
branches c. 6-12 cm borne singly or in subverticils
along their length; secondary branches with numer-
ous tertiary branches bearing lateral and terminal
umbellules (or the branching may be of a high
order); umbellules with c. 6-12 flowers on delicate
pedicels c. 3 mm long. Calyx a minute rim. Petals 5,
2'/, mm long. Stamens 5, anthers oblong 11/, mm
long, filaments delicate, 11/, mm long. Ovary
turbinate in bud, 1!/, mm long, rapidly becoming
rotund, compressed with prominent veins, 2-celled;
styles 2, divergent. Fruit fleshy, black, compressed,
c. 5 by 6 mm; styles persistent, recurved.

Distr. Micronesia, Solomon Is., Queensland; in
Malesia: New Guinea (Papua, Eastern and Milne
Bay Distr.; Territory of New Guinea, Morobe
Distr., Musi I.), also in the D’Entrecasteaux and
Trobriand Is., New Britain, and Louisiades.

Ecol. Strand vegetation and littoral rain-forest,
often behind mangrove.

Vern. Aikove, Mimific lang., ane, D’Entre-
casteaux Is., gambou, Musi I., Morobe Distr.,
nakaigwoo, Trobriand Is., raumonas, Onjob lang.

1979]

3. Polyscias yerticillata Stone, J. Arn. Arb. 47
(1966) 272, f. 1; PHILIPSON, Blumea 24 (1978) 169.

Glabrous, small tree to 7 m with few branches
bearing spirally arranged leaves towards their ends.
Leaves imparipinnate, up to 1 m long; petiole
c. 24-30 cm, terete with a sheathing base c. 6-7 cm
long; petiole 5-20 mm, leaflets oblong often
broader near the base, c. 16-27 by 5-13 cm, margin
entire, slightly revolute or sparsely denticulate,
apex acuminate, base subcordate or truncate, often
oblique, midrib and lateral veins evident. /nflores-
cence a large panicle; primary axis stout, rather
short, with broad cataphylls, bearing several long
radiating secondary branches in a sub-umbel;
secondary branches c. 50 cm, with numerous
tertiary branches borne in well defined verticils and
in a terminal umbel, bracts triangular c. 8 mm long,
caducous; tertiary branches c. 4-7 cm with small
bracts near the middle; umbellules with c. 10-15
flowers, on pedicels 1-4 mm long. Calyx a minute
rim. Petals 5, 2'/, mm long. Stamens 5, anthers
oblong, c. 1-1'/, mm long. Ovary turbinate,
c. 1 mm long, 2-celled; styles 2, at first erect, later
recurved. Fruit globose, fleshy, black, compressed,
c. 4 by 7 mm when dry; styles persistent.

Distr. Solomon Is.; in Malesia: New Guinea
(New Britain, New Ireland, New MHanover,
Admiralty Is., and Bagabag I.).

Ecol. Usually near the beach or in lowland
forest, to 140 m.

Uses. The young foliage is eaten fresh or boiled.

Vern. Babagula, valagur, Kuana dial., la bara
kiku, Nakanai, tauwol, Gasmata, pal/u, Vairamana,
vela vela, Pomio.

Note. Cut surfaces produce a sweet scented
sticky exudate. The petals are violet within, the
fruits purple-black.

4. Polyscias scutellaria (BuRM. f.) Foss. Un.
Hawaii Occ. Pap. 46 (1948) 9; Stone, Taxon 14
(1965) 284; PHILIPSON, Blumea 24 (1978) 169. —
Scutellaria prima RuMPH. Herb. Amb. 4: 75, t. 31.
— Scutellaria secunda latifolia RUMPH. I.c. 76. —
Crassula scutellaria Buro. f. Fl. Ind. (1768) 78. —
Aralia cochleata LAMK, Encycl. 1 (1783) 224. —
Panax scutellarioides REINW. ex BL. Bijdr. (1826)
880; SPAN. Linnaea 15 (1841) 208. — Panax
cochleatum (LAMK) DC. Prod. 4 (1830) 253;
Boer. Handl. 1 (1890) 647. — Panax conchifolium
Roxs. Fl. Ind. ed. Carey 2 (1832) 77. — Notho-
panax cochleatum (LAMK) Mia. PI. Jungh. 3 (1855)
425; Fl. Ind. Bat. 1, 1 (1856) 766; SEEM. J. Bot. 4
(1866) 296; FI. Vit. (1866) 116; Koorp. Minah.
(1898) 490; Exk. Fl. Java 2 (1912) 717; Atlas 4
(1916) f. 697. — Nothopanax tricochleatum Mia.
Fl. Ind. Bat. Suppl. (1860) 135, 340; Merr. Int.
Rumph. (1917) 409; En. Philip. 3 (1923) 234;
OcusE & BAKH. Veg. D.E.I. (1931) 69, f. 39A. —
Panax rumphii Hassk. Abh. Naturf. Ges. Halle 9
(1866) 220. — Nothopanax scutellarium (BuRM. f.)
Me_rr. Int. Rumph. (1917) 409; En. Philip. 3

ARALIACEAE—I (Philipson) 19

1923) 234; HEYNE, Nutt. Pl. (1927) 1209; OcHsE &
BAKH. Veg. D.E.I. (1931) 67, f. 39; CoRNER, Ways.
Trees (1940) 156; BAcK. & BAKH. f. FI. Java 2
(1965) 169. — P. tricochleata (MiQ.) Foss. Phyto-
logia 5 (1955) 290.

Glabrous shrub or small tree, up to 6 m, with
branches bearing spirally arranged leaves towards
their ends. Leaves simple or trifoliolate, variable in
size; petiole often c. 6 cm, but as long as 28 cm, with
a sheathing base 1-6 cm long; lamina rotund or
reniform, often c. 8 cm @ but as wide as 28 cm,
margin usually serrate or becoming sub-lobed
towards apex, in larger leaves the serrations often
inconspicuous, apex rounded, base + cuneate,
midrib and lateral veins evident. Inflorescence
a diffuse panicle; primary axis usually long (up to
1 m) with secondary branches (mostly in verticils, at
intervals along its length; secondary branches
15-30 cm, bearing umbellules in an irregularly
branched system towards their extremities;
umbellules with c. 8-16 flowers, on pedicels c. 3 mm
long. Calyx a minute rim. Petals 4-5, 2 mm long.
Stamens 4-5, anthers oblong, 1 mm long. Ovary
turbinate, c. 1 mm high, (2—)3—4-celled, styles at
first erect, later recurved. Fruit subglobose, fleshy,
c. 5 mm @ when dry.

Distr. Native country not properly known,
possibly East Malesia; cultivated throughout the
region, extending through the tropical Pacific.

Ecol. Grown at low and moderate altitudes
(800 m) as an ornamental or hedge shrub.

Uses. The foliage is aromatic and is used to
furnish perfume. Forms with variegated foliage are
in cultivation. Used medicinally as a diuretic,
against breast cancer, and to prevent baldness
(HEYNE, /.c.). Also for culinary purpose (OCHSE &
BAKH. /.c. 67).

Vern. Cf. HEYNE: memangkokan, (pohon)
mangkok, Mal. (after the dish(—= mangkok)-shaped
leaves which are in the Moluccas sometimes used
as dishes); Java: godong mangkokan, M, mama-
mékan, S, puring mangkok, Md; Lesser Sunda Is.:
lanido, ndalido, ndari, ramido, Roti; Celebes:
(daun) mangko, tuwo mangku, M, bobohang, boku
ula, kéndém wéwéné, woworan, Manado, angko
mangko, Bug. & Mak.; Philippines: salapiin,
Mindanao; Moluccas: daun koin, d. papéda, M,
Ambon, ai laun niwél, ai lohoi, Alf., Ambon, goma
ma tari, Gal., Halmaheira, sawoko, Loda, Halmah.,
rau paroro, Ternate.

Note. This species is usually readily distin-
guished by its simple, orbicular, saucer-shaped
leaves, but plants with some or all of their leaves
trifoliolate occur. STONE (Micronesica 2, 1965, 51)
advanced evidence for uniting these trifoliolate
plants with P. pinnata (= 6. P. cumingiana) while
retaining the unifoliolate plants as a distinct species.
Possibly all are forms of one polymorphous species
complex, but here it is considered convenient to
adhere to the view which unites all forms having

76 FLORA MALESIANA

[ser. I, vol. 9!

orbicular usually bowl-shaped leaves under the
concept P. scutellaria.

5. Polyscias guilfoylei (COGN. & MARCHE) L. H.
BAILEY, Rhodora 18 (1916) 153; STONE, Micro-
nesica 2 (1965) 57; PHILIPSON, Blumea 24 (1978)
169. — Aralia guilfoylei [BULL, Cat. (1873)] CoGn.
& MARCHE, PI. Ornam. 2 (1874) t. 58. — Notho-
panax guilfoylei (COGN. & MARCHE) MErRR. Philip.
J. Sc. 7 (1912) Bot. 242; Fl. Manila (1912) 357; En.
Philip. 3 (1923) 234.

Glabrous shrub to 3 m high, with few branches,
bearing spirally arranged leaves towards their ends.
Leaves imparipinnate, with 3-4 pairs of leaflets,
c. 60 cm long; petiole c. 18 cm, terete, with a
sheathing base 3-4 cm long; petiolules c. 11/,-
2'/, cm; leaflets rotund, oblong or rhomboidal,
c. 10-14 by 6-7 cm, rather thin and flaccid, some-
times rugose, margin sharply serrate to broadly
cuneate (often decurrent on the petiole), often
oblique, midrib and lateral veins evident. Inflores-
cence a diffuse panicle; primary axis short (c. 3-
4 cm) with one or few lateral flowering branches,
terminating in an umbel of long much-branched
rays; lateral branches and rays c. 40-50 cm, with
secondary branches (c. 5—8 cm) singly or mainly in
subverticils along their length and in a terminal
umbel; secondary branches with one or more pairs
of small bracts and ending in umbellules with a
few lateral tertiary branches also ending in umbel-
lules; umbellules with c. 8-12 flowers on pedicels
c. 8-10 mm long. Calyx a minute rim. Petals 5,
2'/, mm long. Stamens 5, anthers oblong, 1!/; mm
long; filaments 2 mm. Ovary turbinate, c. 1 mm
high, usually 3-celled, styles at first erect, but soon
elongating and recurved. Fruit fleshy, globose,
c. 4 by 5 mm.

Distr. Native country unknown, possibly from
East Malesia. Cultivated throughout the region,
but less commonly than other cultivated species of
this section. General throughout the tropical
Pacific.

Ecol. Usually grown as a hedge plant. Flowers
only when left untrimmed or when growing as an
escape from cultivation.

Note. Recognizable by the shape, texture and
serrations of the leaflets, which are usually varie-
gated with whitish or yellowish blotches near the
margins. The flowers are brown in bud but yellow-
green when open.

6. Polyscias cumingiana (PRESL) F.-VILL. Nov.
App. (1880) 102; PHILIPSON, Blumea 24 (1978) 169.
— Scutellaria secunda angustifolia RUMPH. Herb.
Amb. 4: 76, t. 32. — Panax pinnatum LAMK,
Encycl. 2 (1788) 715, non P. pinnata J. R. & G.
ForsTER, 1776; DC. Prod. 4 (1830) 254; SPAN.
Linnaea 15 (1841) 208; Mig. Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 15; BorerL. Handl. 1 (1890)
647; Koorp. Minah. (1898) 15. — Panax secundum
SCHULT. Syst. 6 (1820) 215, nom. superft. illeg. —

Panax bandanense Zipp. ex SPAN. Linnaea 15
(1841) 208, nom. inval. in synon. — Paratropia
cumingiana PRESL, Epim. (1851) 250. — Notho-
panax pinnatum (LAMK) Mia. Bonplandia 4 (1856)
139; Mia. Fl. Ind. Bat. 1, 1 (1856) 766; Merr. Int.
Rumph. (1917) 409; En. Born. (1921) 458; HEYNE,
Nutt. Pl. (1927) 1209; OCHsE & BAKH. Veg. D.E.I.
(1931) 67, f. 38; CORNER, Ways. Trees (1940) 156.
— Nothopanax cumingii (PRESL) SEEM. FI. Vit.
(1865) 114. — Aralia filicifolia C. Moore, Ill.
Hortic. 23 (1876) 72, t. 240. — Arthrophyllum
pinnatum (LAMK) CLARKE, FI. Br. Ind. 2 (1879) 734,
pro basionym. — Panax cumingiana (PRESL) ROLFE,
J. Linn. Soc. Bot. 21 (1884) 310; VipAL, Phan.
Cuming. (1885) 117. — ? Aralia naumannii E.
MARCHAL, Bot. Jahrb. 7 (1886) 469. — Panax
crispatum BULL, Cat. (1888) 9. — Panax ornatum
BULL, /.c. — P. cumingii (PRESL) HARMS in E. & P.
Nat. Pfl. Fam. 3, 8 (1894) 45. — P. rumphiana
Harms, /.c. 45; in K. Sch. & Laut. Fl. Schutzgeb.
(1900) 485; Baku. f. & OoststR. in Back. Bekn.
Fl. Java (em. ed.) 7 (1948) fam. 159, p. 13; BACK.
& BAKH. f. Fl. Java 2 (1965) 167. — Nothopanax
crispatum (BULL) MEeERR. Philip. J. Sc. 7 (1912)
Bot. 241. — Nothopanax ornatum (BULL) MERR.
lic. — P. filicifolia (C. Moore) BAILEY, Rhodora
18 (1916) 153. — P. sorongensis Grpps, Arfak (1917)
216. — Anomopanax cumingianus (PRESL) MERR.
Philip. J. Sc. 17 (1920) 300; En. Philip. 3 (1923) 236.

Glabrous shrub or small tree, to c. 4 m, un-
branched or with few branches bearing spirally
arranged leaves towards their ends. Leaves impari-
pinnate, up to 100 cm; petiole to 20 cm, with a
sheathing base c. 5-6 cm long; petiolules to 3 cm;
leaflets ovate-oblong or elliptic, 10-30 by 2-13 cm,
apex attenuated or acuminate, base rotund, trun-
cate or broadly cuneate, often oblique, margin
entire or minutely and distantly dentate (sometimes
pinnatilobed or almost pinnatipartite); midrib and
lateral veins evident. Inflorescence a large terminal
panicle, or with flowering branches also in the axils
of the upper leaves; primary axis up to 140 cm
with secondary branches mostly in verticils at
intervals along its length; secondary branches up
to 120 cm, bearing umbellules in an irregularly
branched system towards their extremities,
peduncles of the umbellules with one or more pairs
of small bracts; umbellules with c. 10-20 flowers;
pedicels 4-8 mm. Calyx a minute rim with
4-5(-6) teeth. Petals 4-5(—6), 21/.-31/. mm long.
Stamens 4-5(-6), 2 mm long. Ovary turbinate,
c. 2 mm long, 2-5-celled; styles at first erect, later
spreading. Fruit subglobose, fleshy, 3-S mm @
when dry.

Distr. Cultivated throughout the region, and
also apparently growing as part of the indigenous
vegetation.

Ecol. Rain-forest and secondary growth,
usually at low altitude but ascending to 1700 m,
native range not very clear.

Uses. A common ornamental shrub, especially

1979]

the cultivars with dissected foliage. Also for
culinary use.

Vern. (from various sources). Celebes: daun
grisik, d. mangko, d. papéda pandang, M, Manado;
Philippines: bani, Luzon, Albay; Moluccas:
kéndém, Ternate, gurabati, M; papua, Jappen I.;
New Britain: awalagu, Gazelle Pen.

OcHsE & BAKHUIZEN VAN DEN BRINK record
that the vernacular names applied to P. fruticosum
also include P. cumingianum.

Notes. A complex of forms requiring intensive
biosystematic study. I adhere to the view that the
Indo-Malayan material is specifically distinct from
the Polynesian Polyscias pinnata J. R. & G.
Forster; cf. HARMS in E. & P. Nat. Pfl. Fam.
3, 8 (1894) 45; Puitipson, Bull. Br. Mus. Nat.
Hist. Bot. 1 (1951) 9, but some cultivated forms
included here may have been derived from the
Pacific rather than the Indo-Malayan species.
Existing herbarium material, though voluminous,
is inadequate to determine whether the complex
includes species with more limited ranges.

As in 1. P. fruticosa, cultivated forms with dis-
sected foliage occur.

See also under 4. P. scutellaria for a discussion of
the relationship with that species.

I have not seen the type of Aralia naumannii
MARCHAL which, from the description, I assume to
be a synonym.

7. Polyscias javanica K. & V. Bijdr. 7 (1900) 13;
Koorp. Atlas 4 (1916) f. 679 A-R; BAKH. f. &
OoststrR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 159, p. 12; Back. & Baku. f. Fl. Java 2
(1965) 167; PHILIPson, Blumea 24 (1978) 169.

ARALIACEAE—I (Philipson) vi |

Glabrous shrub or sparsely branched small tree,
bearing spirally arranged leaves towards the ends of
the branches. Leaves imparipinnate, c. 70 cm long;
petiole c. 17 cm, with a wide membranous sheath-
ing base (c. 7 cm long); petiolules c. 1 cm; leaflets
ovate to elliptic-oblong, up to 22 by 8 cm, mem-
branous, apex attenuated, base rounded to broadly
cuneate, often oblique, margin entire or minutely
distantly dentate, slightly revolute, midrib and
lateral veins prominent. Inflorescence a panicle;
rachis 40 cm, with caducous bracts (reduced leaves
or leaf-sheaths), bearing few secondary branches
and terminating in a compound umbel; secondary
branches with one or more verticils of tertiary
branches; peduncles of the umbellules with one
obscure bract or none; umbellule of c. 10-20
flowers, pedicels c. 5 mm. Calyx an undulate rim,
indistinctly 5-dentate. Petals 5, 2'/,-3 mm long.
Stamens 5, with very short filaments. Ovary turbi-
nate, 1'/, mm high, 5-celled, styles at first erect,
later recurved. Fruit a globose or obovoid fleshy
drupe (prominently 5-ribbed when dry) c. 7 by
5 mm; persistent styles united below, strongly
reflexed above.

Distr. Malesia: East Java, Lesser Sunda Is.
(Lombok, Sumbawa, Sumba).

Ecol. In forest up to 1650 m.

Uses. The root is said to be made into a face
powder in Sumbawa.

Vern. Lesser Sunda Is.: pulosari, Sumbawa,
amdari, Sumba.

Note. The vegetative parts are rather similar
to those of 6. P. cumingiana, but the smaller and
simpler inflorescence is distinctive.

2. Section Gelibia

(HUTCH.) PHILIPSON, Blumea 24 (1978) 169. — Gelibia Hutcu. Gen. Fl. Pl. 2

(1967) 57.

Tree with bipinnate leaves. Flowers arranged racemosely; style arms free and

divergent after anthesis.

Distr. In New South Wales, Queensland, and Malesia (New Guinea) | sp.

8. Polyscias elegans (C. Moore & F.v.M.) HARMS
in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; J. Arn.
Arb. 20 (1939) 323; PHILIPSON, Blumea 24 (1978)
169. — Panax elegans C. Moore & F.v.M. Trans.
Phil. Inst. Vict. 2 (1858) 68; Bru. FI. Austr. 3
(1866) 383; C. Moore, Handb. FI. N.S.W. (1893)
215. — Nothopanax elegans (C. Moore & F.v.M.)
SEEM. FI. Vit. (1866) 114; J. Bot. 4 (1866) 294. —
Tieghemopanax elegans (C. Moore & F.v.M.)
VIGUIER, Buli. Soc. Bot. Fr. 52 (1905) 308. —
P. branderhorstii HARMS, Nova Guinea 8 (1910)
274; Bot. Jahrb. 56 (1920) 42; J. Arn. Arb. 20
(1939) 323. — Gelibia branderhorstii (HARMS)

Hutcu. Gen. FI. Pl. 2 (1967) 57. — Gelibia elegans
(C. Moore & F.v.M.) Hutcu. I/.c. 58

Tree to 20 m high, with a spreading crown of
thick branches bearing terminal clusters of large
leaves, young parts with fawn to grey scurfy
tomentum. Leaves bipinnate, becoming glabrous,
up to 110 by 50 cm, or larger; petiole c. 13 cm,
with a slightly clasping base; petiolules up to
c. 1/, cm; leaflets ovate or elliptic, c. 6 by 3 cm,
chartaceous to somewhat leathery, margin entire
and slightly revolute, apex with an obtuse apiculum,
base cuneate, midrib prominent, lateral veins
rather obscure. Inflorescence a large panicle,

78 FLORA MALESIANA

[ser. I, vol. 91

tomentum persistent especially on the pedicels;
primary axis stout, c. 30 cm, bearing secondary
axes along its length and in a terminal subumbel-
late cluster, bracts caducous; secondary axes
c. 20-30 cm, bearing tertiary axes (c. 10 cm) along
their length; flowers borne racemosely along the
tertiary axes on pedicels 1-2 mm long. Calyx a
minute rim. Petals 5, oblong, rather fleshy,
spreading at anthesis and soon falling. Stamens 5,
2 mm long on short filaments, soon falling. Ovary
scurfy, at first turbinate, globose at anthesis,
c. 4 mm high, 2-celled, disk rising to the 2 styles;
style arms at first short and pressed together,
elongating and diverging after the anthers have
fallen. Fruit leathery, globose, compressed, c. 5 by
5 mm and irregularly ribbed when dry.

Distr. New South Wales and Queensland; in

Malesia: New Guinea (West Irian, Southern Distr. ;
Papua, Western & Central Distr.).

Ecol. Gallery and second growth forest in
(seasonal) monsoon regions, at low altitudes, up
to 300 m.

Notes. The racemose flower arrangement has
sometimes been considered sufficiently distinctive
to justify the accommodation of this species in a
separate genus.

The bark is described as grey or patched grey-
brown; the wood as soft and pale; and the cut
surfaces having a clear sticky exudate which has a
pleasant fragrance. The inflorescence branches are
purplish, the petals maroon, the anthers cream to
yellow on reddish filaments, and the fruit purplish
and smooth.

3. Section Eupteron

(MiQ.) PHILIPSON, Blumea 24 (1978) 170. — Eupteron Mia. Pl. Jungh. 3 (1855) 420;

Fl. Ind. Bat. 1, 1 (1856) 762.

Trees or shrubs with imparipinnate or bipinnate leaves. Flowers in umbellules or
capitula; style arms divergent at least in fruit.

Distr. In Malesia 6 spp., but the section probably includes the majority of the genus in other parts of

the Old World tropics.

Note. With the exception of 14. P. philipsonii, the Malesian spp. of sect. Eupteron have umbellules or
capitula arranged racemosely along the main inflorescence branches. The type species of the section
(P. nodosa) has its flowers in capitula, but this is not considered a sufficient difference to require a separate

section.

KEY TO THE SPECIES

1. Flowers in capitula
1. Flowers in umbellules.

2. Umbeilules arranged racemosely along the principal rays.

3. Leaf margins crenate.
4. Leaflets oblong (Luzon) .

4. Leaflets lanceolate (New Guinea) . ait ;

3. Leaf margins entire (rarely with few minute dentations).

5. Leaflets ovate (Sabah, Palawan)
5. Leaflets elliptic (New Guinea)

2. Umbellules in a short corymbose compound umbel . . .

9. Polyscias nodosa (BL.) SEEM. J. Bot. 3 (1865)
181; F.-VitL. Nov. App. (1880) 102; VIDAL,
Phan. Cuming. (1885) 117; Rev. Pl. Vasc. Filip.
(1886) 145; BoerRL. Handl. 1 (1890) 647; HARMs in
E. & P. Nat. Pfl. Fam. 3, 8 (1894) 44; Koorp.
Minah. (1898) 491; K. & V. Bijdr. 7 (1900) 11;
Merk. Philip. J. Sc. 1 (1906) Suppl. 110; Koorp.
Exk. Fl. Java 2 (1912) 716; Atlas 4 (1916) f. 677 &
678; Merr. Int. Rumph. (1917) 408; Sp. Blanc.
(1918) 294; En. Philip. 3 (1923) 233; HEyNeg, Nutt.
Pl. (1927) 1208; Baku. f. & OoststRr. in Back.
Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 12;

. 12. P. borneensis
13. P. belensis

Back. & BAKH. f. Fl. Java 2 (1965) 167; PHILIPSON,
Blumea 24 (1978) 170. — Papaya silvestris RUMPH.
Herb. Amb. 1: 149, t. 53 f. 1. — Aralia nodosa
BL. Bijdr. (1826) 872. — Paratropia nodosa (BL.)
DC. Prod. 4 (1830) 265; PresL, Epim. (1851) 250.
— Aralia umbraculifera Roxs. (Hort. Beng. (1814)
22, nomen;] Fl. Ind. ed. Carey 2 (1832) 108. —
Aralia pendula BLANCO, FI. Filip. (1837) 223; ed. 2
(1845) 157; ed. 3, 1 (1877) 283. — Hedera nodosa
(BL.) Hassk. Tijd. Nat. Gesch. Phys. 10 (1843)
131. — Eupteron nodosa (Bu.) Mig. Pl. Jungh.
3 (1855) 420; Fl. Ind. Bat. 1, 1 (1856) 762; Ann.

179)

Mus. Bot. Lugd.-Bat. 1 (1864) 220; HutcH. Gen.
Fl. Pl. 2 (1967) 68. — ? P. acuminata VIDAL,
Sinopsis Atlas (1883) 28, t. 55 f. A, non SEEM. 1865;
F.-VILL. Nov. App. (1880) 102. — P. floribunda
Exner, Leafl. Philip. Bot. 10 (1939) 3819, nom.
inval. in synon. — Fig. 28.

Tree to 25 m, unbranched or with few thick
branches, bearing tufts of long leaves at their ends,
young parts scurfy-tomentose. Leaves imparipin-
nate, multijugate, becoming glabrous or minutely
villose on the nerves beneath, up to 2 m (or even
3 m); petiole c. 30 cm, to 2 cm @, with a short
sheathing base; leaflets sessile, ovate-oblong com-
monly 15 by 4 cm, or larger, chartaceous, margin
slightly crenate, apex ++ apiculate, base truncate.
Inflorescence a large panicle (sometimes with
additional flowering branches in the axils of the
upper leaves), tomentum + persistent; primary
axis stout, c. 1'/, m, bearing secondary axes along
its length, bracts triangular, c. 5 mm long; secon-
dary axes c. 20-40 cm; capitula borne racemosely
along the secondary branches on peduncles c. 6—
15 mm long. Flowers c. 8-12 in a capitulum. Calyx
a minute rim. Petals 5, triangular, 2 mm long.
Stamens 5, anthers broadly ovate on short fila-
ments. Ovary turbinate, c. 2'/, mm high, 5-celled;
disk flat, rising to the 5 central erect styles. Fruit
subglobose (5-ribbed when dry), styles spreading
and reflexed.

Distr. Solomon Is. (Bougainville); in Malesia:
Sunda Straits (Krakatau), Java, Lesser Sunda Is.
(Lombok), Celebes, throughout the Philippines
and Moluccas (Talaud, Ternate, Sulu Is., Ceram,
Ambon, Banda, Tenimber, Aru Is.), New Guinea
(eastwards to Milne Bay Distr.).

Recorded doubtfully, on sterile material, from
the Andaman Is. by Kurz (Veg. Andam. Is. 1870,
39) but unlikely as it is absent from Sumatra,
Malaya, and Borneo.

Ecol. Open thickets and rain-forest, mostly at
low altitudes and on small islands, but recorded
to 1000 m.

Uses. Used medicinally against purpuric fever
and to delay pregnancy (Mindanao). The leaves are
used to stupify fish and the wood makes durable
fence posts and handles for rice-knives.

Vern. (from various sources). Java: ki langit,
S, déleg, djaranan, manglé, puténgan, J, pénang-
pénangan, Md, kaju djaran, k. lanang, rangit,
tjaliru, tua kalap; Philippines: bias-bias, bingliu,
Tag., Bis., bungloi, Bis., bon liu, bungliu, goyung-
goyung, Tag., bungdieu, Ig., hagdan-anak, C.Bis.,
mano-mano, Yak., malapapdya, Tag., Bis., Pang.,
tukod-langit, Tag., Pamp.; Celebes: kambowa,
lalusuhan, pasusinggala, (pohon) mamalapa, tama-
lola, tundu; Sulu Is.: lua; Moluccas: kobo-kobo,
Morotai, batatopus, Ceram, papaya utan, pata
tulan, p. tulong, Ambon, matanglolan, Tenimber;
West New Guinea: amorigh, laulako, sonomdoro,
totja.

Note. A widespread, common and noticeable

ARALIACEAE—I (Philipson) 79

species. The flowers are described as yellow and as
having a fragrant scent. The wood is soft and white.

10. Polyscias florosa PHILIPSON, Blumea 24 (1978)
170.

Small tree, up to 10 m, with thick branches
bearing terminal clusters of large leaves, glabres-
cent. Leaves imparipinnate, multijugate, up to
2 m; petiole c. 25 by 11/, cm, with a short sheathing
base; leaflets sessile, oblong, c. 22 by 8 cm,
chartaceous, margin crenate, apex slightly to long
acuminate, base truncate or subcordate. Inflores-
cence a large glabrescent panicle; principal rays
over 1 m, up to 3 cm @, each bearing secondary
branches (20-40 cm) along its length; bracts
broadly ovate, 1 cm long, caducous; umbellules
borne racemosely along the secondary branches on
peduncles 1-2 cm long (in fruit to c. 4 cm).
Flowers c. 6-12 in an umbellule, pedicels 3-5 mm
with 2 minute bracts near the middle. Calyx an
undulate rim. Petals 5, oblong, 3 mm long.
Stamens 5. Ovary turbinate, c. 2 mm long, 5-celled,
disk flat, rising to the 5 central erect styles. Fruit
subglobose (5-ribbed when dry), 8 by 6 mm, styles
spreading and reflexed.

Distr. Malesia: Philippines (Luzon; Mt
Bulusan, Sorsogon Prov. and Mt Malinao, Albay
Prov.).

Ecol. Rarely collected in montane forest,
c. 560 m.

Note. The leaves are very similar to those of
P. nodosa, with which this species has often been
confused. However the pedicelled flowers are quite
distinctive.

11. Polyscias ledermannii HARMs, Bot. Jahrb. 56
(1921) 409; Puitipson, Blumea 24 (1978) 170. —
Panax murrayi (non F.v.M.) F.v.M. Descr. Pap.
Pl. 7 (1886) 29; BoerL. Handl. 1 (1890) 647. —
P. forbesii BAKER f. J. Bot. 56 (1923) Suppl. 22. —
P. clemensiana Harms, Bot. Jahrb. 69 (1938) 283.

Glabrous tree, to 28 m, with few thick branches
bearing terminal clusters of large leaves. Leaves
imparipinnate, up to 1'/, m; petiole to c. 16 cm,
base slightly dilated; petiolules c. 2-4 mm; leaflets
lanceolate or oblong-lanceolate, c. 10-20 by
2-4 cm, papyraceous, margin crenulate, apex
acuminate, base truncate or rounded, often oblique,
midrib prominent, lateral veins numerous, faint.
Inflorescence a panicle; primary axis stout, c. 15 cm,
bearing crowded secondary axes along its length
and in a subumbellate cluster at its apex, bracts
triangular, c. 3 mm long, caducous; secondary
axes 20—30(-40) cm; umbellules borne racemosely
along the secondary axes (often in subverticils) on
peduncles 2-5 cm long, usually bearing 2 minute
bracts. Flowers c. 8-12 in an umbellule, pedicels
c. 5mm. Calyx a minute rim. Petals 2'/, mm long.
Stamens 5. Ovary turbinate, 2 mm high, 3-4-
celled, disk rising to the central styles. Fruit

80 FLORA MALESIANA

[ser. I, vol. 92

globose, 4-5 mm_ high (when dry), 3—4-ribbed,
styles 3-4, divergent, persistent.

Distr. Malesia: New Guinea (Vogelkop to
Milne Bay).

Ecol. In rain-forest, gallery forest and secondary
growths, usually above 1500 m (to 3000 m), but
occasionally as low as 500 m.

Vern. Agugwa, Hagen, gapin, Morobe Distr.,
gowi, Efogi lang., panda panda, Mendi lang., puri,
Kepilan.

Notes. A common small tree. CARR estimated
the height of one specimen as 28 m, but most are
considerably smaller. The flowers are described as
green. The wood is soft and pale. The plant has a
scent resembling celery.

The species closely resembles P. murrayi (F.v.M.)
Harms from Queensland.

12. Polyscias borneensis PHILIPSON, J. Bot. 78
(1940) 118; Blumea 24 (1978) 170.

Glabrous shrub or small tree with few thick
branches bearing terminal clusters of leaves.
Leaves imparipinnate, c. 50 cm; petiole c. 15 cm,
base slightly dilated; petiolules c. 1 cm; leaflets
ovate, up to 10 by 4 cm, coriaceous, margin entire
or slightly undulate, apex attenuate, base rounded
or broadly cuneate, often asymmetrical, midrib
prominent, lateral veins rather obscure. Jnflores-
cence a large panicle; primary axis stout, up to
c. 40 cm, bearing secondary axes along its length
and in a subumbellate cluster at its apex, bracts
triangular c. 5 mm long; secondary axes c. 35 cm;
umbellules borne racemosely along the upper part
of the secondary branches on peduncles 2-5 cm
long. Flowers about 10 in an umbellule, pedicels
c. 10 mm long. Calyx a minute undulate rim.
Petals oblong 3 mm long. Stamens 5, oblong
c. 21/, mm long, filaments 2 mm. Ovary turbinate,
2 mm high, 5-celled, disk rising to the 5 central
erect styles. Fruit unknown.

Distr. Malesia: N. Borneo (Mt Kinabalu),
Philippines (Palawan, Mt Mantalingahan).

Ecol. Montane forest, c. 1500 m.

Note. Specimens collected on Palawan by
EDANO were distributed with the specific epithet
‘palawanensis’, but this name was never published.

13. Polyscias belensis PHILIPSON, Bull. Br. Mus.
Nat. Hist. Bot. 1 (1951) 13; Blumea 24 (1978) 170.

Glabrous tree, to 14 m. Branches bearing termi-
nal clusters of large leaves. Leaves imparipinnate,
up to 80 cm long; petiole c. 12-16 cm, base slightly
dilated; leaflets subsessile or with a petiolule to
1 cm; leaflets elliptic to narrowly elliptic, c. 12 by
31/,-6 cm, subcoriaceous, margin entire or sparsely
denticulate, slightly revolute, narrowed to an
obtuse apex, base cuneate, midrib prominent,
lateral veins c. 12. Inflorescence a panicle; primary
axis stout, c. 18-22 cm, bearing many secondary
branches along its length and in a subumbellate
cluster at its apex, bracts caducous; secondary

axes c. 22-35 cm, bearing umbellules racemosely
along their length, bracts triangular c. 3 mm long;
peduncles c. 1!/,-3 cm. Flowers c. 10 per umbellule,
pedicels 5-6 mm. Calyx a minute rim. Petals 4,
oblong. Stamens 4, anthers oblong, 2 mm long,
filaments very short. Ovary turbinate, 11/,-2 mm
high (in anthesis), 4~S-celled, disk flat with 4—5 free
styles, at first erect, later divergent. Fruit unknown.

Distr. Malesia: New Guinea (West Irian,
Bele R. near Lake Habbema; Papua, Morobe
Distr., Mt Kaindi and Aseki Patrol area).

Ecol. Infrequent tree in montane forest and
regrowth, 1450-2650 m.

Note. The inflorescence is similar to that of 11.
P. ledermannii, but the shape of the leaflets is
distinctive. In the type (from West Irian) the leaf
margins are entire, but in the three gatherings from
Morobe there are some dentations, and it is
possible these are not conspecific. The corolla is
maroon (Morobe). The plant has an odour like
celery.

14. Polyscias philipsonii BERNARDI, Ber. Schweiz.
Bot. Ges. 74 (1966) 364; PHiLiIpson, Blumea 24
(1978) 170. — P. fraxinifolia PHILIPSON, Bull. Br.
Mus. Nat. Hist. Bot. 1 (1951) 13, non Harms, 1894.

A sympodial, unbranched, glabrous shrub or
epiphyte to 2 m high, with leaves spirally arranged
towards the apex, new shoots arising below the
inflorescence. Leaves imparipinnate, or occa-
sionally 3 leaflets inserted together, or the leaves
bipinnate, up to 35 cm long; petiole to 10cm, witha
very short sheathing base; petiolules to 7 mm,
leaflets variable in size and shape, broadly or
narrowly elliptic, up to 9 by 4!/, cm, but usually
smaller, narrowed to the apex or apiculate, base
cuneate or rounded, margin minutely setose-
crenulate, midrib prominent, lateral veins few,
arched-ascending. Inflorescence a corymb;
peduncle short, 1-2 cm, sometimes bearing a
reduced leaf or cataphyll distally; primary rays
c. 5, subequal, c. 3-4'/, cm, bearing terminal com-
pound umbels (occasionally with a lateral umbel);
ultimate umbellules with 6-12 flowers on pedicels
c. 4mm long. Calyx a minute rim with indistinct
teeth. Petals 5, triangular, 2 mm long. Stamens 5,
anthers oblong, 1 mm long. Ovary turbinate,
c. 2 mm high, 5-celled, the fleshy disk rising to the
5 central erect styles. Fruit fleshy, globose, 5-ribbed
when dry, c. 4 by 6 mm, styles persistent, recurved.

Distr. Malesia: West New Guinea (Idenburg R.).

Ecol. In mossy forest at 1800 m, and on an
exposed slope at 2150 m.

Notes. The short corymbose inflorescence is
similar to that in sect. Palmervandenbroekia, but the
free recurved style arms exclude it from that sec-
tion.

The shrub is variable in aspect, and may grow
as an epiphyte or as an undershrub. The panicle
branches are described as purple and the flowers as
green.

1979]

ARALIACEAE—I (Philipson) 81

4. Section Kissodendron

(SEEM.) PHILIPSON, Blumea 24 (1978) 170. — Kissodendron Stem. J. Bot. 3 (1865)

201.

Trees or shrubs with pinnate or bipinnate leaves. Umbellules arranged in diffuse
panicles. Style arms fused, forming a beak-like projection on the fruit.

Distr. In Malesia (Moluccas and New Guinea) 5 spp., three of which extend to Queensland.

KEY TO THE SPECIES

1. Leaves bipinnate.
1. Leaves simply imparipinnate.
2. Leaflets in 3-4(-S) pairs .
2. Leaflets in more than 5 pairs.
3. Fruits rotund, c. 8 mm broad
3. Fruits ovoid, c. 4-5 mm broad.
4. Base of leaflets cuneate

4. Base of leaflets truncate, subcordate, or rounded .

15. Polyscias bipinnata (GrBBs) PHILIPSON, Blumea
24 (1978) 170. — Kissodendron, bipinnatum G1sBs,
Arfak (1917) 161; Harms, Bot. Jahrb. 56 (1921)
413.

A small tree; branches with spirally arranged
leaves towards their ends; young parts brown fur-
furaceous. Leaves bipinnate, 70 by 68 cm; petiole
26 cm, with a short sheathing base; rachis with a
pair of leaflets at each articulation; petiolules to
11/, cm; leaflets narrowly oblong or ovate, c. 6-12
by 2!/,-4(-5) cm, apex attenuated, base broadly
cuneate or rounded, margin entire, irregularly
undulate slightly revolute, midrib prominent.
Inflorescence a diffuse panicle (sometimes also with
flowering branches in the axils of the upper leaves);
peduncle c. 6 cm, terminating in few (3) primary
rays; primary rays 40-50 cm, bearing verticils of
pairs of secondary branches along their length and
ending in a compound umbel; umbellules of
c. 10-15 flowers on pedicels c. 10 mm long. Calyx
an undulate rim. Petals 5, 2'/, mm long. Stamens 5,
filaments 1 mm, anthers 1'/, mm long. Ovary
turbinate 2-3-celled, disk with a central stylo-
podium 1 mm long. Fruit ovoid, fleshy, 5 by 3 mm
(when dry) crowned by the calyx rim and the beak-
like stylopodium, stigmas minutely capitate.

Distr. Malesia: W. New Guinea (Vogelkop:
Angi Lakes).

Ecol. Montane forest, at c. 2000 m.

Vern. Louklouwko.

Note. The individual leaflets are similar to those
of 18. P. schultzei, but their bipinnate arrangement
is distinctive. The flowers are white.

16. Polyscias zippeliana (MiQ.) VALETON, Bull.
Dép. Agr. Ind. Néerl. 10 (1907) 42; PHILIPSON,
Blumea 24 (1978) 170. — Panax zippelianum Mia.
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 15; BoERL.

15. P. bipinnata
16. P. zippeliana
17. P. royenii

. . 18. P. schultzei
19. P. australiana

Handl. 1 (1890) 645; Harms in E. & P. Nat. Pfl.
Fam. 3, 8 (1894) 46. — Nothopanax zippelianum
(Miq.) SEEM. FI. Vit. (1866) 115. — Kissodendron
australianum [non (F.v.M.) SEEM.] BoERL. Hand.
1 (1890) 650. — P. caroli Harms, Bot. Jahrb. 56
(1920) 411.

Shrub, 1!/,(-5) m, with few branches, bearing
leaves spirally arranged near their ends, glabrous
except a slight rufous tomentum on young parts.
Leaves imparipinnate, with 3—4 pairs of leaflets, up
to c. 60 cm; petiole to 17 cm, with a very short
sheathing base; petiolules '/,-1'/, cm; leaflets
oblong, ovate or elliptic, 6-13 by 31/,-6 cm,
Papyraceous, apex attenuate or apiculate, base
cuneate, often oblique, margin entire, slightly
revolute, midrib and few lateral veins evident.
Inflorescence a panicle, often appearing lateral by
rapid growth of a bud at its base; branches rather
fine and wiry, rachis often angled at the nodes, up
to c. 60 cm; secondary branches borne singly at
wide intervals and in a few rayed terminal umbels,
c. 20 cm long, bearing umbellules on side branches
and terminally; umbellules with c. 10-15 flowers on
pedicels 1/,-1 cm long. Calyx a minute rim. Petals
5, broadly oblong, 2 mm long. Stamens 5, anthers
oblong, 1 mm long, filaments 1 mm. Ovary turbi-
nate, 2 mm high, 2-celled, the fleshy disk rising to
the central fused stylar column. Fruit ovoid,
crowned by the prominent stylopodium with a
capitate stigma, c. 7 by 4 mm when dry.

Distr. E. Malesia: SE. Moluccas (Aru Is.) and
New Guinea (S. Irian and southern part of Western
Distr. of Papua; also in the Sepik Distr.).

Ecol. In the Aru Is. and S. New Guinea in
obviously seasonal savannah country associated
with Acacia and Melaleuca at low altitudes, in the
Sepik Distr. in the mountains at c. 1000 m.

Vern. Mirmur, uwah, perdi-perdi, W. Irian.

82 FLORA MALESIANA [ser. I, voli9*

Fig. 30. Polyscias schultzei HARMS. a. Habit, x 2/;, b. flower and ditto in LS, x 10, c. petal, d. stamens,
e. fruit and ditto in CS, x 5 (a-d HoOGLAND & PULLEN 5841, e KALKMAN 4200). Drawn by HELENE
MULDER.

1979]

ARALIACEAE—I (Philipson) 83

Notes. The identity of P. zippeliana has been
discussed by BOERLAGE, VALETON, and Harms,
ll.cc. The short leaf-sheath and the connivent styles
place it within sect. Kissodendron.

A comparison of the type specimen with the
species usually known as P. caroli HARMS shows
them to be conspecific. The type of P. caroli is no
longer available but specimens collected by BRAss
can be considered authentic as they were identified
by HARMS. It should be noted, however, that the
type of P. caroli is from the Sepik Distr. (probably
in rain-forest) at 1000 m, while all other specimens
are from the south of the island and from seasonal
savannahs at much lower altitudes.

The single gathering from the Aru Is. approaches
19. P. australiana having larger leaves with more
pairs of leaflets than other gatherings of P. zippe-
liana.

17. Polyscias royenii PHILIPSON, Blumea 24 (1978)
170.

Small tree up to 5(-15) m, monocaulous sym-
podial, persistently rufous-furfuraceous, with
leaves spaced along the upper part of the stem.
Leaves imparipinnate, with c. 9 pairs of leaflets,
up to 100 cm long or more; petiole to 30 cm, witha
short sheathing base; petiolules of mid-leafiets
c. 8-20 mm; leaflets normally oblong-ovate, broad-
est near the base, up to 24 by 11 cm, coriaceous,
apex attenuated or apiculate, base truncate to
subcordate; often oblique, margin entire, irregu-
larly undulate, slightly revolute; midrib prominent,
reticulations depressed above (rugose) raised and
furfuraceous below. Inflorescence a_ panicle,
flowering branches also often present in the axils of
the uppermost leaves; rachis up to 60 cm with
verticils of secondary branches subtended by
caducous unifoliolate bracts and a terminal com-
pound umbel; lower secondary branches up to
50 cm, with verticillate tertiary branches and
terminal compound umbels; umbellules with
10-20 flowers on furfuraceous pedicels c. 8 mm
long. Calyx a furfuraceous rim, with 5 small teeth.
Petals 5, strap-shaped, c. 3 mm long, soon falling.
Stamens 5. Ovary subcylindric, furfuraceous,
c. 3 mm long, 2(-3)-celled, the fleshy disk forming
a projecting conical stylopodium 2 mm long;
stigmas 2, appressed at anthesis. Fruit rotund,
compressed, with prominent ribs when dry 7-9 by
8-10 mm, crowned by the prominent persistent
calyx and a short stout beak-like stylopodium;
stigmas short, divergent.

Distr. Malesia: New Guinea (Cyclops Mts to
the E. Highlands Distr.).

Ecol. Terrestrial or epiphytic in primary or
secondary montane forest, 1200-2400 m.

Vern. Irian: ato, Kapauku lang.; Papua New
Guinea: habia, S. Highlands, magabin, W. Sepik,
momin, Mendi lang.

Note. A species readily characterized by the
sympodial monocaul habit with thick leathery

leaflets, a large rigid furfuraceous inflorescence, and
large, compressed, black fruits with a glaucous
bloom.

18. Polyscias schultzei HARMs, Bot. Jahrb. 56
(1921) 410; PHiLipson, Blumea 24 (1978) 171. —
P. gjellerupii HARMS, Bot. Jahrb. 56 (1921) 410. —
Fig. 30.

Shrub or small tree, often unbranched, 4-20
(-26) m high, young parts densely brown furfura-
ceous, branches with spirally arranged leaves
towards their ends. Leaves imparipinnate, to
80 cm long, with c. 8-11 pairs of leaflets; petiole
up to 35 cm, with a short sheathing base; petiolules
c. 8 mm; leaflets ovate-lanceolate, oblong or
elliptic, up to 14(-17) by 4-5 cm, apex attenuated
or acuminate, base broadly cuneate, oblique,
margin entire, midrib prominent. Jnflorescence a
diffuse, repeatedly compound umbel, with reduced
leaves + persistent at the nodes; peduncle usually
rather short (c. 5 cm) ending in a group of several
primary rays (and sometimes with one or more
lateral rays); primary rays 50-60 cm, with verticils
and a terminal umbel of secondary rays which in
turn are branched; umbellules with c. 5—10 flowers
on fine pedicels c. 5-8 mm long (elongating in fruit
to 12-15 mm). Calyx a minute rim with 4-5 small
teeth, usually glabrous but occasionally furfura-
ceous. Petals 4-5, c. 3 mm long. Stamens 4-5,
2 mm long. Ovary + cylindric, glabrous or
occasionally furfuraceous, c. 2 mm long, 2-celled,
the fleshy disk forming a blunt stylopodium. Fruit
ovoid, fieshy, c. 5 mm long, crowned by the incon-
spicuous calyx and the projecting beak-like stylo-
podium (c. 2 mm long); stigmas slightly divergent
or capitate.

Distr. Queensland; in Malesia: Moluccas
(Morotai) and New Guinea (throughout most of
the island, from the Vogelkop Peninsula and
Japen I. to the Central and Morobe Distr.).

Ecol. Usually in lower and mid-montane rain-
forest (with Castanopsis, Nothofagus, Lithocarpus),
also in regrowth, usually above 1200 m, ascending
to 2400 m, but also descending to near sea-level.

Vern. New Guinea: atok, Wissel Lakes, funim,
Telefomin, houklouwké, Angi Lakes, paul, Wabag,
agugwa, Hagen.

Notes. A widespread and frequently collected
species recognized by the multijugate leaves with
rather small, thin, attenuate leaflets with cuneate
base. The flowers are white to yellowish and the
fruits black. The grey bark exudes a small amount
of gummy sap; the wood is white.

Harms recognized that P. gjellerupii was very
similar and the abundant material now available
indicates the variability of this common species.

19. Polyscias australiana (F.v.M.) PHILIPSON,
Blumea 24 (1978) 171. — Hedera australiana
F.v.M. Fragm. 4 (1864) 120; Bru. Fl. Austr. 3
(1866) 384. — Kissodendron australianum (F.v.M.)

84 FLORA MALESIANA

[ser. I, vol. 91

SEEM. J. Bot. 3 (1865) 201; ibid. 6 (1868) 129. —
Irvingia australiana (F.v.M.) F.v.M. Fragm. 5
(1865) 18.

var. disperma (F.v.M.) PHILIPSON, Blumea 24
(1978) 171. — Kissodendron australianum (F.v.M.)
SEEM. var. disperma F.v.M. Descr. Not. Pap. Pl. 5
(1877) 88; Harms, Bot. Jahrb. 56 (1921) 412.
Small tree, often unbranched, branches with
leaves arranged spirally towards their ends, young
parts brown furfuraceous. Leaves imparipinnate,
multijugate, to 1 m long; petiole c. 25 cm, with a
short sheathing base; petiolules to 11/, cm;
leaflets broadly ovate or oblong, broadest near the
base, up to 14 by 6 cm, + coriaceous, apex attenu-
ated to an obtuse apex, or with an obtuse apiculum,
base rounded, truncate, or subcordate, oblique,
margin entire, slightly revolute, midrib and lateral
veins prominent. Inflorescence a diffuse panicle
(sometimes also with flowering branches in the axis

of the upper leaves); peduncle short, terminating
in several primary rays, reduced leaves subtending
the principal branches; primary rays c. 50 cm,
bearing verticils of secondary branches along their
length and ending in a compound umbel; umbel-
lules of c. 10-15 flowers on pedicels c. 8-10 mm
long. Calyx an undulate rim. Petals 5, c. 11/, mm
long. Stamens 5, c. 1 mm long. Ovary turbinate,
2-celled, a fleshy disk rising to a low conical stylo-
podium. Fruit ovoid, fleshy, 6 by 4 mm (when dry),
crowned by the persistent calyx and beak-like
stylopodium (2 mm long).

Distr. Malesia: New Guinea (Vogelkop Penin-
sula and Southern Distr. in Papua).

Ecol. Primary rain-forest at low altitudes and
also in lower montane forest at 1750 m.

Note. Distinguished from 18. P. schultzei by the
leathery leaflets with a truncate base, and from 17.
P. royenii by the smaller fruits.

5. Section Palmervandenbroekia

(GIBBS) PHILIPSON, Blumea 24 (1978) 171. — Palmervandenbroekia Gtsps, Arfak

(1917) 162.

Trees or shrubs with imparipinnate leaves. Umbellules arranged in short corym-
bose compound umbels. Style arms fused, forming a beak-like projection on the

fruit.

Distr. Malesia: New Guinea (Vogelkop Peninsula and Cyclops Mts in W. Irian, Western Distr. in

Papua).

KEY TO THE SPECIES

1. Pedicels tomentose (at anthesis).

2. Inflorescence a weer regular compound umbel with rigid rufous branches (Vogelkop and Cyclops

Mts)

DE EE ee | ge ee ee 20. P. sleumeri

2. Inflorescence more open, less ee branched, with branches becoming + glabrous (Mt Bosavi,

Papua)
1. Pedicels glabrous (at anthesis).

21. P. jacobsii

3. Flower buds lanceolate, stylopodium at anthesis c. 2 mm long (Vogelkop, Arfak Mts)

22. P. palmervandenbroekii

3. Flower buds ovoid, stylopodium at anthesis c. 1 mm long (Vogelkop, Nettoti Range and Aifat R.)

20. Polyscias sieumeri PHILIPSON, Blumea 24
(1978) 171.

Monocaulous sympodial shrub, 1-1'/, m, with
leaves spaced along the upper part of the stem;
young parts red-furfuraceous. Leaves imparipin-
nate, with 3-4 pairs of leaflets (leaflets occasionally
in threes), c. 35 cm long; petiole to 9 cm, with
a very short sheathing base; petiolules 8-15 mm;
leaflets oblong, ovate or elliptic, up to 16 by 6 cm,
chartaceous, apex attenuate or apiculate, base
rounded or cuneate, margin entire, irregularly
undulate, slightly revolute, midrib prominent.
Inflorescence a compound umbel, red-furfuraceous

23. P. vogelkopensis

(stem growth renewed by a bud between the
inflorescence and the uppermost leaf); peduncle
3-6 cm, bearing scars of caducous reduced leaves,
occasionally with a lateral umbellule, and ending
in an umbel of c. 7 primary rays; primary rays
1—21/, cm, sometimes with a minute bract near the
middle; umbellule with 10-20 flowers on pedicels
2-3 mm long, densely furfuraceous. Calyx a rim
with 5 broad teeth. Petals 5, broadly oblong,
3-4 mm long. Stamens 5, anthers 1 mm long,
filaments 2 mm. Ovary turbinate, 2 mm high,
2-celled, the fleshy disk forming a projecting coni-
cal stylopodium; stigmas 2 appressed at anthesis.

1979]

Fruit rotund to ovoid compressed 5-6 by 5 mm
crowned by the prominent persistent calyx and the
beak-like stylopodium (2—2'/, mm long); stigmas
not prominent.

Distr. Malesia: New Guinea (Vogelkop Penin-
sula and Cyclops Mts).

Ecol. Undergrowth in primary forest, 450-
1000 m.

Note. The flowers are creamy white, the fruit
dark red.

21. Polyscias jacobsii PHILIPSON, Blumea 24 (1978)
Ale

A monocaulous sympodial shrub to 3 m high,
with leaves spaced along the upper part of the stem,
young parts furfuraceous. Leaves imparipinnate,
4-5 pairs of leaflets (leaves of juveniles smaller
with fewer leaflets), c. 45 cm long; petiole to 12 cm,
with a short sheathing base; petiolules 1—1!/, cm;
leaflets oblong or elliptic, c. 12 by 4 cm, charta-
ceous, apex attenuate or caudate, base truncate or
cuneate, margin entire, irregularly undulate,
slightly revolute, midrib prominent. Inflorescence
a compound umbel (stem growth renewed by a
bud between the inflorescence and the uppermost
leaf); peduncle 1-4 cm (sometimes bearing a uni-
foliolate leaf about the middle with an axillary
flowering branch), ending in an umbel of 2-4
primary rays; primary rays sometimes subtended
by 1-2 unifoliolate leaves, c. 4 cm long, ending in
2-3 tertiary rays which may branch again before
ending in umbellules; umbellules with 6-10 flowers
on furfuraceous pedicels c. 8 mm long. Flower
buds with an apical umbo. Calyx a rim with 5
acute teeth. Petals 5, attenuated. Stamens 3 mm
long, anthers 11/, mm long. Ovary subcylindric,
furfuraceous, 4 mm high, 2-celled, disk forming a
projecting conical stylopodium (1!/, mm long at
anthesis), stigmas 2. Fruit rotund, compressed,
8 by 9 mm, crowned by the small calyx and the
persistent stylopodium (2 mm long) with sub-
capitate stigmas.

Distr. Malesia: New Guinea (Papua: Mt
Bosavi). Only known from the type.

Ecol. On old well-drained volcanic soil in
primary mixed forest, mostly on ridges and upper
slopes, 1600-2100 m.

Note. An aromatic, single-stemmed shrub. The
flowers are white and the fruit dark purple.

22. Polyscias palmervandenbroekii BERNARDI,
Candollea 26 (1971) 16; PHiLirson, Blumea 24
(1978) 171. — Palmervandenbroekia papuana
Gipss, Arfak (1917) 162, f. 15, non P. papuana
SEEM. 1865.

Glabrous shrub, c. 1!/, m, sparsely branched,
with leaves spaced along the upper parts of the
stems. Leaves imparipinnate, with 3-5 pairs of
leaflets, up to 13 cm long; petiole to 4 cm, slightly
channelled above, with a very small sheathing

ARALIACEAE— I (Philipson) 85

base; petiolules 0-5 mm; leaflets lanceolate, obo-
vate to elliptic, 1'/,-4 by */,-2 cm, coriaceous or
chartaceous, apex attenuated (sometimes emargi-
nate) to apiculate, base cuneate, margin entire,
revolute, midrib prominent, lateral veins obscure.
Inflorescence a simple or compound umbel;
peduncle slender, c. 2 cm; primary rays (when
present) few, c. 1 cm; umbellules of c. 10 flowers
on glabrous pedicels 3-6 mm long. Flower buds
lanceolate, acute, 3-6 mm. Calyx a rim with 5
broad teeth. Petals 5, attenuate, 5-8 mm long.
Stamens 5, filaments c. 3 mm, anthers c. 1 mm.
Ovary turbinate, striate, 2 mm high, 2-celled, disk
projecting as a conical stylopodium (2 mm long).
Fruit ovoid, compressed, 6 by 5 mm, crowned by
the persistent calyx and the beak-like stylopodium;
stigmas 2, small, divergent.

Distr. Malesia; New Guinea (Vogelkop: Arfak
Mts).

Ecol. In summit heath vegetation (Tristania-
Dacrydium scrub) and Nothofagus forest, 1900-
2400 m.

Note. Coilections made since the original de-
scription show that this interesting plant is quite
variable as regards leaf-shape, texture and amount
of rolling of the margin, and also in the size of its
flowers. The calyx is purplish brown, the petals
reddish purple outside and white within. The ripe
fruit is purplish black. The thin papery bark is light

grey.

23. Polyscias vogelkopensis PHILIPSON, Blumea 24
(1978) 171.

An often monocaulous sympodial shrublet
usually under 1 m high, with leaves spaced along
the upper parts of the stem; young parts slightly
furfuraceous, but soon becoming glabrous.
Leaves imparipinnate, with 3-4 pairs of leaflets
(leaflets occasionally in threes) variable in size;
petiole 2-7 cm, slightly channelled above with a
short inconspicuous sheathing base; petiolules
2-8 mm; leaflets elliptic or ovate, 3-12 by 1'/,-4cm,
chartaceous, apex attenuated or apiculate, base
broadly to narrowly cuneate, margin entire,
irregularly undulate (occasionally with isolated
dentations) slightly revolute, midrib prominent.
Inflorescence a small compound umbel; peduncle
up to 8 cm, occasionally with a lateral or basal
umbellule, ending in c. 2-3 primary rays (some-
times subtended by a reduced leaf); primary rays
1-2 cm; umbellules with c. 10-15 flowers on
glabrous pedicels c. 3-6 mm long. Flower buds
ovoid, obtuse. Calyx a rim with 4-5 broad undula-
tions. Petals 4-5, broadly oblong, 4 mm long.
Stamens 4-5, anthers 1 mm, filaments 3 mm. Ovary
turbinate, 2 mm high, 2-celled, the fleshy disk
rising to a conical stylopodium (1 mm long);
stigmas 2, appressed at anthesis. Fruit ovoid,
compressed, 5 by 4 mm, crowned by the persistent
calyx and the prominent beak-like stylopodium;
stigmas inconspicuous.

86 FLORA MALESIANA

[ser. I, vol. 9!

Distr. Malesia: New Guinea (Vogelkop and
Wandammen Peninsula, Nettoti Range and Aifat
R.).

Ecol. Primary submontane forest (Nothofagus,
Castanopsis, and conifers), heath vegetation, or in
open places, 1200-2000 m.

Note. A delicate subshrub, often unbranched.
The material from Aifat R. has larger leaves than
that from the Nettoti Range. The flowers are cream
or light yellow, and the fruits orange with black
stylopodium.

Insufficiently known
Polyscias disperma BLANCO, Fl. Filip. (1837)

226; MeErRR. Sp. Blanc. (1918) 384; En. Philip. 3
(1923) 233. — Philippines.

MERRILL (1923, /.c.) stated that this name was
excluded by BLANCo from his 2nd edition, and that
it is unrecognizable from the very short and im-
perfect description. Possibly a Rubiacea.

Polyscias roemeriana Harms, Bot. Jahrb. 56
(1921) 411. — New Guinea.

J have seen no authentic specimens of this
species. Evidently it is related to P. palmervanden-
broekii by reason of its connivent styles, its short
inflorescence, and its small leaflets. However,
bipinnate leaves are not known in that species.

Excluded

Polyscias joskei L. S. Gisss, J. Linn. Soc. Bot. 39
(1909) 149. — This species was by error ascribed by
Index Kewensis to the Philippines; it is from Fiji.

11. MACROPANAX

Mia. FI. Ind. Bat. 1, 1 (1856) 764; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 13; BTH. in
B. & H. Gen. Pl. 1 (1865) 945; BoerL. Handl. | (1890) 643; Harms in E. & P. Nat.
Pfl. Fam. 3, 8 (1894) 47; Hutcn. Gen. FI. Pl. 2 (1967) 80. — Hederopsis CLARKE,
Fl. Br. Ind. 2 (1879) 739; HARMs in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 56; KING,
J. As. Soc. Beng. 67, 1i (1898) 62; Hutcu. Gen. FI. Pl. 2 (1967) 78; Stone, Gard.
Bull. Sing. 30 (1977) 141. — Fig. 31.

Small, unarmed trees. Leaves digitately compound or unifoliolate; petiole
terete, with a sheathing base; stipules inconspicuous; leaflets entire or denticulate.
Inflorescence a terminal panicle of umbellules. Pedicels articulated below the ovary.
Flowers bisexual or on lower branches male. Calyx 5—6-dentate. Petals 5-6,
valvate. Stamens 5-6, extrorse. Ovary inferior, 2—6-celled. Disk fleshy with the
styles united below. Fruit ellipsoid or obovate, surmounted by an awl-shaped or
conical stylopodium; exocarp fleshy, endocarp crustaceous. Endosperm ruminate.

Distr. 4 spp. from India, Burma and southern China to West Malesia: 3 spp., Sumatra, Malaya and
ope Lowland and montane rain-forest.

Notes. Hederopsis is united with Macropanax because the species are very similar in appearance and
in their technical characters. The sole difference is the number of cells in the ovary, 2 (or sometimes 3) in
Macropanax and 5-6 (sometimes 4) in Hederopsis. This distinction does not seem adequate to split a few
very similar species into distinct genera.

This genus is distinguished from Pseudopanax (China, New Caledonia, Tasmania, New Zealand, Chile)
by the ruminate endosperm. Cf. PHILIPSON, New Zeal. J. Bot. 3 (1965) 333.

KEY TO THE SPECIES
1. Ovary 2-celled.
2. Ovary broadly campanulate, not ribbed; ‘epicalyx’ below ovary distinct. Leaf margin usually dis-
tinctly serrate. Inflorescence branches usually bearing lateral umbellules. . . . . 1. M. dispermus
2. Ovary narrowly turbinate, strongly ribbed; ‘epicalyx’ below the ovary absent. Leaf margin entire or
minutely dentate. Inflorescence branches usually without lateral umbellules 2. M. concinnus
imOvarvis-.o0-celledwamir Artarms: <<cal cals, a bostotee.«teieaiey alte «me 3. M. maingayi

1. Macropanax dispermus (BL.) O. K. Rev. Gen.
Pl. 1 (1891) 271; Koorp. Exk. Fl. Java 2 (1912)
716, 717; Atlas 4 (1916) f. 681 & 682 F-P; FI.

Tjib. 2 (1923) 227; Baku. f. & Ooststr. in Back.
Bekn. FI. Java (em. ed.) 7 (1948) fam. 159, p. 14;
BAck. & BAKH. f. Fl. Java 2 (1965) 168. — Aralia

1979]

ARALIACEAE— I (Philipson) 87

disperma BL. Bijdr. (1826) 872. — Hedera disperma
(BL.) DC. Prod. 4 (1830) 265. — Aralia calyculata
Z. & M. in Mor. Syst. Verz. (1846) 41. — M. flori-
bunda Mia. FI. Ind. Bat. 1, 1 (1856) 764. —
M. oreophilus Mia. I.c. 764; Suppl. (1860) 135; Ann.
Mus. Bot. Lugd.-Bat. 1 (1863) 13; Kurz, For. FI.
Burma 1 (1877) 541; CLARKE, FI. Br. Ind. 2 (1879)
738; BoeRL. Handl. 1 (1890) 644; K. & V. Bijdr. 7
(1900) 16.

Small tree, up to 18 m. Leaves dispersed along
the shoots; petiole to 25 cm, striate, with a small
basal sheath, the connate stipules forming a small
ligule within the petiole; leaflets 5—7, petiolules up
to c. 5 cm (the lateral shorter); blade elliptic or
oblanceolate, usually c. 10-23 by 2'/,9 cm,
gradually tapered to an acute apex, base cuneate,
rounded or oblique, margin coarsely dentate or
sometimes only finely serrate, glabrous when
mature. Jnflorescence a panicle of umbellules, often
stellate-furfuraceous, with a stout rachis to 40 cm
and lateral (secondary) branches which terminate
in umbellules, and which commonly bear tertiary
branches. Pedicels c. 5-7 mm, with a distinct
involucre (‘epicalyx’) around the base of the flower.
Calyx rim undulate or indistinctly dentate,
puberulous. Petals 2-3 mm long. Ovary subglobose,
c. 2'/, mm high. Disk fleshy, surmounted by an
awl-shaped stylar column, which bifurcates at
apex. Fruit ovate, 1 by 1/, cm, crowned by the
persistent stylar column.

Distr. India, Burma, and southern China; in
Malesia: Sumatra, Malay Peninsula, and through-
out Java.

Ecol. Mountain rain-forests, 1000—2300 m, in
Java especially in Central and East Java.

Vern. Java: panggang puju, p. serem, p. Siju,
ramo-gentjel, tjerem, S, pampung, sahang, tanganan,
J, konjingal, Md.

2. Macropanax concinnus Mig. Ann. Mus. Bot.
Lugd.-Bat. 1 (1864) 220; Boeri. Handl. 1 (1890)
643; K. & V. Bijdr. 7 (1900) 18; Koorp. Exk. FI.
Java 2 (1912) 717; Atlas 4 (1916) f. 682 Q-W. —
M. undulatus (WALL. ex G. Don) SEEM. J. Bot. 2
(1864) 294, pro specim. malac.; BAKH. f. & OOSTSTR.
in Back. Bekn. FI. Java (em. ed.) 7 (1948) fam. 159,
p. 14; Back. & Baku. f. Fl. Java 2 (1965) 168.
Small tree, up to 15 m. Leaves dispersed along
the shoots; petiole slender, to 20 cm, with a small
basal sheath, the connate stipules forming a small
ligule within the petiole; leaflets 5 or 3, petiolules
slender, up to c. 5'/, cm (the lateral shorter); blade
elliptic, usually c. 7-15 by 2!/,-5 cm, gradually
tapered to a caudate-falcate acuminate apex, base
narrowly cuneate, or rarely obtuse, often oblique,
Margin entire or sometimes a few very fine serra-
tions, glabrous, membranous. Inflorescence a
panicle of umbellules, minutely stellate-pubescent,
with a stout rachis to 30 cm, and lateral (secondary)
branches which terminate in umbellules; pedicels
c. 6-7 mm, only slightly swollen at the articulation.

Calyx rim undulate or indistinctly dentate. Petals
c. 2 mm long. Ovary narrowly turbinate with
distinct longitudinal ribs, c. 2 mm high. Disk fleshy,
surmounted by an awl-shaped stylar column. Fruit
ovate, 8 by 4 mm, crowned by the persistent stylar
column.

Distr. Malesia: widespread in Java, but local
and uncommon.

Ecol. Damp mountain forests, descending to
lower altitudes (400 m) than the preceding species.

Vern. Djampang tjerem, panggang puju, S,
tanganan, J.
Notes. This species has sometimes been

identified with M. undulatus (WALL. ex G. Don)
Seem. from the Himalayan region, but that species
differs in the more verticillate inflorescence
branches, the large involucre around the umbellule,
the hermaphrodite flowers more confined to the
terminal umbellules, and the broader less promi-
nently ribbed fruit. A considerable geographical
gap separates the two species.

A specimen collected by GrirritH labelled
‘Malacca’ is perhaps M. undulatus, but the
locality may be incorrect.

3. Macropanax maingayi (CLARKE) PHILIPSON,
comb. nov. — Hederopsis maingayi CLARKE, FI.
Br. Ind. 2 (1879) 739; RipL. Fl. Mal. Pen. 1 (1922)
888; Stone, Gard. Bull. Sing. 30 (1977) 287, f. 7. —
Arthrophyllum trifoliatum Ripi. J. Fed. Mal. St.
Mus. 7 (1916) 42. — Hederopsis major RIDL. Kew
Bull. (1929) 124. — Fig. 31.

Small tree, up to 15 m. Leaves dispersed along
the shoots; petiole to 20 cm, striate, with a small
basal sheath; leaflets 5, or (below the inflorescence)
3 or 1, petiolule of the central leaflet to 6 cm, of the
lateral leaflets much shorter (c. 1 cm); blade
elliptic or ovate, up to 20 by 8 cm, base rounded
or broadly cuneate, apex acuminate, margin
denticulate or nearly entire, membranous or
chartaceous. Inflorescence a terminal panicle of
umbellules, with a stout rachis to 20 cm, termi-
nating in an umbellule and bearing lateral secon-
dary branches (c. 12 cm) which also terminate in
umbellules and bear racemosely arranged tertiary
branches (c. 4 cm) which terminate in umbellules;
umbellules terminating the primary and secondary
branches with hermaphrodite flowers, those
terminating the tertiary branches smaller and
apparently mostly functionally male. Flowers of
small umbellules usually 5-merous, of larger
umbellules usually 6-7-merous, pedicels c. 1 cm at
anthesis, slightly elongating in fruit. Calyx rim
obscurely dentate. Petals of terminal flowers
c. 3 mm long, fleshy. Ovary subglobose, c. 2 mm
high; disk fleshy surmounted by the style arms
which become divergent in their upper part at
anthesis. Fruit enlarging to a drupe 2 by 1 cm with
a persistent calyx rim and a prominent conical
stylopodium tapering into the stylar column bear-
ing the recurved distal parts of the styles.

88 FLORA MALESIANA [ser. I, vol. 9!

Fig. 31. Macropanax maingayi (CLARKE) PHILIPSON. a. Leaf and upper branches of inflorescence, x 2/<,
b. flower bud and CS of ovary, x 2, c. flower, x 2, d. fruit, slightly enlarged (Phytochem. Surv. Mal.
2519). Drawn by W. R. PHILIPSON.

1979] ARALIACEAE—I (Philipson) 89

Note. The trunk may have small buttresses, the
bark is rugose with large lenticels, and produces a
watery exudate when cut. The flowers are greenish
yellow.

Distr. Tonkin; in Malesia: Malay Peninsula
(from Kedah southwards) and Central Sumatra
(East Coast Res. and Mt Kerintji).

Ecol. Uncommon, in forest, up to 1050 m.

12. ANAKASIA

W. R. PHILIPSON, Blumea 21 (1973) 87, fig. on p. 88. — Fig. 32.

Shrub with large, simple, exstipulate leaves. Inflorescence axillary; rachis bearing
racemosely arranged umbellules. Pedicels very short with an articulation below the
flower. Calyx a short rim with minute lobes. Petals 5 or 6, valvate, triangular.
Stamens 5 or 6, filaments thick, anthers large, dorsifixed. Ovary inferior, broadly
obconic, (4-)5—6-celled. Disk fleshy with (4-)5-6 stylar arms. Fruit broadly obo-
void, strongly ribbed when dry; exocarp fleshy. Pyrenes compressed, crustaceous;
endosperm smooth.

Distr. Malesia: West New Guinea. Monotypic.

Note. The large, simple, oblanceolate leaves clustered at the ends of the branches recall Meryta, but
the flowers do not share the highly distinctive features of that genus. The floral and fruit characters are

not unlike those of Polyscias (e.g. there is an articulation below the flower, the style arms are free, and the
endosperm is smooth), but the general facies is unlike that genus, and this, together with the distinctive

inflorescence and leaf, make the plant quite distinct from any species of Polyscias.

1. Anakasia simplicifolia W. R. PHILIPSON, Blumea
21 (1973) 87, fig. on p. 88. — Fig. 32.

Glabrous shrub, 5 m, branches marked with
prominent lenticels. Leaves crowded at the ends of
the branches, sessile or with a very short petiole,
lanceolate, up to 135 by 18 cm, narrowed gradually
to the base, apex acuminate, margin entire or
undulate, midrib prominent, lateral veins arching
upwards, reticulation rather indistinct. /nflores-
cence rachis simple or forked, to 70 cm, at maturity
5 mm @, bearing small lanceolate bracts. Peduncles
arising from the axils of all but the lowest bracts,
3-4 mm long, elongating to c. 15 mm in fruit.
Umbellules with c. 11 minute bracts. Flower buds
when dry c. 3 mm long. Calyx rim with 5-6 minute
teeth. Petals slightly fleshy, 2 mm long. Stamens
5-6, filaments broad; anthers 4-celled, 3/, mm long.
Ovary glabrous. Disk with a central boss formed
by closely appressed subulate arms which soon

recurve and spread beyond the calyx. Fruit with
(4-)5-6 prominent ridges when dry, c. 2 by 1!/, cm,
with the persistent style arms in a _ terminal
depression.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula and near Babo).

Ecol. Primary forest near sea-level.

Note. BEcCCARI described (in sched.) the in-
florescence branches as erect in flower and
reflexed in fruit. He noted that the outer flowers of
the umbellules are probably male and those at the
centre female, though he could not be certain
of this. His observations cannot be confirmed from
the material available. Evidently all flowers are
structurally hermaphrodite, but all developing
fruits seen are attached to central pedicels. The
corolla is green and soon falls, the disk yellow, the
anthers cream, and the mature fruit blue (azure-
violet) and aromatic.

13. BRASSAIOPSIS

DECNE & PLANCH. Rev. Hort. IV, 3 (1854) 106; Bru. in B. & H. Gen. Pl. 1 (1865)
945; CLARKE, FI. Br. Ind. 2 (1879) 735; BoeRL. Handl. 1 (1890) 643; HARMs in
E. & P. Nat. Pfl. Fam. 3, 8 (1894) 42; Kina, J. As. Soc. Beng. 67, ii (1898) 61;
NoGoc-SANH Bul, Adansonia 6 (1966) 437, pl. 1 (map); Hurcu. Gen. FI. Pl. 2
(1967) 79; Stone, Gard. Bull. Sing. 30 (1977) 280. — Araliopsis KURZ, Rep.
Andam. (1870) 39, nom. inval., in synon., non ENGLER, 1896. — Wardenia KING,
J. As. Soc. Beng. 67, ii (1898) 60; HARMs in E. & P. Nat. Pfl. Fam. Nachtr. 2

90 [ser. I, vol. 9!

4 b

Fig. 32. Anakasia simplicifolia PHILIPSON. a. Leaf, x '/,, b. inflorescence, x 1/3, c. single umbellule with
an immature fruit, d. flower bud, x 12, e. immature fruit, x 2 (BECCARI PP 282).

1979] ARALIACEAE—I (Philipson) 91

(1900) 51; Rupr. Fl. Mal. Pen. 1 (1922) 887; Hutcn. Gen. FI. Pl. 2 (1967) 381;
cf. FRODIN, Misc. Rec. Found. Fl. Males. 3 (1973) 8. — Euaraliopsis HUTCH. Gen.
Fl. Pl. 2 (1967) 80, 624. — Pseudobrassaiopsis BANERJEE, J. Bomb. Nat. Hist. Soc.
71975) 71. — Fig. 33/34.

Shrubs or trees, usually prickly and tomentose. Leaves digitately compound,
palmately lobed or simple, with a sheathing base and a usually bicuspid ligule;
petiole terete. Inflorescence usually a terminal panicle of umbellules, often with
persistent small bracts; pedicel not articulated below the ovary. Calyx rim 5-den-
tate. Petals 5, valvate. Stamens 5, extrorse. Ovary inferior, 2-celled; disk shallow;
styles united into a usually long column. Fruit globose, exocarp fleshy, endocarp
chartaceous; seed 2, not compressed, endosperm smooth.

Distr. More than 20 spp., extending from India, Tibet and southern China to West Malesia: Sumatra,
Malay Peninsula and W. Java. In Malesia 6 spp., 1 endemic in Sumatra, 3 endemic in the Malay Penin-
sula, 2 in Indo-Malaya.

Ecol. Understorey of damp evergreen forest, mostly in mountainous districts, up to 2400 m.

Notes. A genus characterized by the 2-celled ovary, columnar style, and non-articulated pedicel, com-
bined with leaves which may be either entire, palmately lobed, or digitately compound. Possibly close to
Trevesia, which is similar vegetatively, but which has more massive inflorescences and ovaries with more
numerous cells.

In dividing the species into two genera merely on the basis of leaf shape, HUTCHINSON /.c. fragmented
what appears to be a coherent assemblage.

KEY TO THE SPECIES
1. Inflorescence shorter than the petioles (or at most about equal to them). Leaves palmately lobed, but

Vaniaplenmtnenvaniety™ = sian. © seen 1. B. sumatrana
1. Inflorescence much longer than the petioles.

2. Leaves simple, entire, minutely and remotely dentate.

3. Leaf base subcordate; blade to 35 cm or more long; petioles c.15cm,5 mm2@ .
3. Leaf base cuneate or rounded; blade to 25 cm long; petioles c. 7 cm, 2 mm @
2. Leaves palmately lobed or digitately compound (occasionally some simple).

4. Leaves palmately lobed
4. Leaves digitately compound.

. 2. B. simplex
3. B. minor

. 4. B. polyacantha

5. Branches of the inflorescence bearing several umbellules racemosely (as well as one at end);

umbelilules with c. 25-35 flowers

es. Se 5. B. glomerulata

5. Branches of the inflorescence bearing a terminal umbellule only (or occasionally with separate

flowers below it); umbellule with c. 10-13 flowers

1. Brassaiopsis sumatrana RIDL. J. Fed. Mal. St.
Mus. 8, 4 (1917) 43. — Kalopanax sumatranum
Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 17,
heterotypic; BoERL. Handl. 1 (1890) 647; HARMs
in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 51. —
? Kalopanax resectum Mia. Ann. Mus. Bot. Lugd.-
Bat. 1 (1863) 18, see note below. — Euaraliopsis
sumatrana (RIDL.) HuTcH. Gen. FI. Pl. 2 (1967) 624.

var. sumatrana.

Shrub or small tree, to 15 m, with stout prickly
branches, rufous-tomentose in its younger parts.
Leaves palmately lobed, clustered at the ends of the
shoots; petiole c. 15-32 cm, striate, sometimes
with prickles, with an enlarged basal sheath
and a small bicuspid ligule; blade rounded, cor-
date, c. 14-22 by 20-28 cm, palmately 5—7-lobed,

6. B. elegans

lobes elliptic acuminate, midrib prominent and
lateral veins widely spaced and arched-ascending,
margin irregularly spinulose-serrate. Inflorescence
a short, rufous-tomentose panicle, arising below
the leaves (actually terminal but overtopped by
sympodial innovations); rachis c. 4—6 cm, lateral
branches ascending, c. 2-4 cm, subtended by small
linear bracts and sometimes branching to the third
degree; umbellules with c. 8-20 flowers; pedicels
c. 1 cm long, subtended by numerous minute
bracts. Calyx a minute rim with 5 indistinct teeth.
Petals 5, triangular, 2'/, mm long. Stamens 2 mm
long, anthers oblong, 1 mm long. Ovary obconic,
c. 2 mm high; disk fleshy, cushion-shaped, sur-
mounted by a slender awl-shaped stylar column.
Fruit subglobose, c. 8 mm high, with a persistent
calyx-rim and stylar column.

92 FLORA MALESIANA [ser. I, vol. 91

Fig. 33. Brassaiopsis elegans RiDL. a. Habit, x 2/;, b. fruit and ditto in CS, 4 (KLoss 1912). Drawn by
P. PRENDERGAST.

1979]

Distr. Malesia: West .Sumatra (Mts Kerintji,
Merapi, Singalang).

Ecol. Montane, evergreen forest, 1500-2400 m.

Vern. Semontong, Karo.

Notes. The foliage is similar to that of 4.
B. polyacantha and of Trevesia sundaica, but the
small inflorescence is distinctive. The prickles persist
on the older branches and the trunk. The petals
and filaments are white, the anthers light purple,
and the disk yellow.

Kalopanax resectum Mia. l.c., described on a
KoRTHALS collection from Sumatra, possibly
belongs here, but the inflorescence may be larger
and most leaves are decidedly digitately compound.
Possibly it represents a hybrid between B. suma-
trana and 5. B. glomerulata.

var. variaefolia PHILIPSON, var. nov. — B. ovalifolia
RIpL. J. Fed. Mal. St. Mus. 8, 4 (1917) 43.

Frutex var. sumatrana humilior et tenerior, foliis
secus ramis dispersis, inermis. Folia minora (10-
20 cm longa) forma variantia etiam in eodem ramo,
vel simplicia et ovata, vel profunde bi- vel trilobata,
vel digitatim composita. — Typus: W. MEWER 7200
(L), Mt Sago near Pajakumbuh, W. Sumatra,
1000-1200 m, 8—vim—1957.

A smaller, more delicate and unarmed shrub
than var. sumatrana, with the leaves scattered along
the branches. Leaves smaller (10-20 cm long) and
variable in shape, even on the same branch, being
either simple and ovate, or deeply bi- or trilobed,
or digitately compound.

Distr. Malesia: Central West Sumatra.

Ecol. Montane, evergreen forest, 1000-1200 m.

Note. It is possible that RIDLEY was correct in
regarding this as a distinct species, but the size and
shape of the inflorescence links it with B. sumatrana,
and the variable form of the leaf suggests that it
may be no more than an extreme form of B. suma-
trana. Nevertheless, typical forms of that species,
with their stout prickly stems and rather coarse
palmately lobed leaves, are very distinct. On the
evidence available it is not possible to decide
whether this variety consists of shade and/or
juvenile forms of B. sumatrana, or whether it should
constitute a distinct species.

2. Brassaiopsis simplex (KING) STONE, Gard. Bull.
Sing. 30 (1976) 282, f. 3. — Wardenia simplex Ki1nG,
J. As. Soc. Beng. 67, ii (1898) 60; HARMS in E. &
P. Nat. Pfl. Fam. Nachtr. 2 (1900) 51; Ript. FI.
Mal. Pen. 1 (1922) 887; HutcH. Gen. FI. Pl. 2
(1967) 81; PxHitipson, J. Linn. Soc. Bot. 63,
Suppl. (1970) 90.

Shrub, to 5 m high, branches prickly, rufous-
tomentose in its younger parts. Leaves large,
simple, coriaceous, clustered towards the tips of
the branches; petiole c. 14-20 cm, stout, striate,
with a basal sheath and a short, bicuspid ligule;
blade ovate, c. 35-40 by 12-18 cm, apex shortly
acuminate, base slightly cordate, rufous-stellate

ARALIACEAE—I (Philipson) 93

hairs persistent beneath, midrib prominent,
lateral veins very widely spaced, arched ascending,
margin minutely dentate. Inflorescence a panicle of
umbellules; rachis c. 10-20 cm; lateral branches
reflexed, 5-12 cm, subtended by small lanceolate
bracts and often bearing 1—2 bracts from which
tertiary branches arise; umbellules with c. 10-20
flowers; pedicels 1'/,—-2 cm, slender, subtended by
numerous minute bracts. Calyx a narrow rim with
5 indistinct teeth. Petals 5, falling as a calyptra.
Stamens 2 mm long, anthers oblong. Ovary ob-
conic, c. 2 mm high, disk cushion-like, surmounted
by an awl-shaped stylar column 1'/, mm long.
Fruit globose, fleshy, c. 5 mm @, with a persistent
calyx rim and a stylar column c. 2 mm long.

Distr. Malesia: Malay Peninsula (Perak,
Selangor).

Ecol. In open bamboo forest, 100—800 m.

Note. Originally described as a distinct genus
because of its supposedly 1-celled ovary. RIDLEY
(1922) did not include this feature in his descrip-
tion. He distinguished Wardenia from Brassaiopsis
because of its simple leaves, but several species of
Brassaiopsis from further north also have undivided
leaves. Both Harms (1900) and HuTCHINSON
(1967) accepted the original statement that the
ovary is l-celled at the time of flowering, but I
found the ovary to be 2-celled even in the young
state (1970, /.c.).

3. Brassaiopsis minor STONE, Gard. Bull. Sing. 30
(1976) 282, f. 5.

Small shrub, c. 60 cm, with very few small
prickles, rufous tomentose on its young parts.
Leaves simple, spaced towards the ends of the
branches, subcoriaceous; petiole 5-8 cm, c. 2 mm
@, with a basal sheath and a small ligule; blade
elliptic or ovate 20-26 by 10-12 cm, apex shortly
acuminate, base rounded to cuneate, rufous stellate
hairs persistent beneath, midrib prominent,
lateral veins widely spaced, arched ascending,
margin minutely and remotely dentate. /nflores-
cence: peduncle short, bearing lanceolate bracts;
umbellules on short lateral branches, with 15-20
flowers on slender pedicels c. 11/, cm long, sub-
tended by minute puberulent bracts. Calyx with
5 indistinct teeth. Ovary rotund, surmounted by a
slender stylar column 1 mm long.

Distr. Malesia: Malay Peninsula (Trengganu).

Ecol. Humid situations in lowland to montane
forest, to 1100 m.

4. Brassaiopsis polyacantha (WALL.) BANERJEE,
Ind. For. 93 (1967) 341; Stone, Gard. Bull. Sing.
30 (1977) 282. — Hedera polyacantha WALL. Pl. As.
Rar. 2 (1831) 82, t. 190. — Panax palmatum Roxs.
[Hort. Beng. (1814) 21, nomen;] Fl. Ind. ed.
Carey 2 (1832) 74. — B. palmata (ROxB.) KuRZ, J.
As. Soc. Beng. 39, ii (1870) 77; CLARKE, FI. Br.
Ind. 2 (1879) 735; Kinc, J. As. Soc. Beng. 67, ii
(1898) 61; Ripv. FI. Mal. Pen. 1 (1922) 887. —

94 FLORA MALESIANA

[ser. I, vol. 91

oo

Fig. 34. Brassaiopsis polyacantha (WALL.) BANERJEE. In flower at Kuala Lumpur (Photogr. PHILIPSON,
1975).

Euaraliopsis palmata (ROxB.) HutcH. Gen. FI.
Pl. 2 (1967) 80. — Pseudobrassaiopsis polyacantha
(WALL.) BANERJEE, J. Bomb. Nat. Hist. Soc. 72
(1975) 72. — Fig. 34.

A small sparsely branched tree with buttresses,
to c. 13 m; branches prickly, rufous-tomentose on
the younger parts and persisting on the leaves and
inflorescence. Leaves palmately lobed, clustered at
the ends of the shoots; petiole to c. 70 cm, longi-
tudinally grooved, with an enlarged basal sheath
and a pair of small stipular processes; blade
rounded, cordate, c. 18-30 by 20-40 cm, deeply
palmately 5—9-lobed, lobes oblong-elliptic, acumi-
nate, midrib prominent and main lateral veins
widely spaced and arched-ascending, margin
inconspicuously serrate. Inflorescence a _ large
panicle often overtopped by sympodial innova-
tions; rachis c. 40-60 cm, lateral branches c. 8-
15 cm, subtended by very small caducous bracts
and often bearing a pair of minute bracts near the
middle, from which short tertiary branches may
arise, branches terminating in spherical umbellules;
umbellules with c. 15-25 flowers, subtended by
numerous minute bracts; pedicels 3/,-11/, cm.

Calyx of 5 small teeth, rusty tomentose. Petals 5,
triangular, 2'/, mm long. Stamens 2 mm long,
anthers oblong, 1 mm. Ovary obconic, c. 2 mm
high; disk fleshy, cushion-shaped, surmounted by
an awl-shaped stylar column 1!/, mm long. Fruit
globose, 6-8 mm @, with a persistent calyx rim and
stylar column.

Distr. Widespread in SE. Asia (India, Nepal,
Burma, Andamans); in Malesia: Malay Peninsula
(from Kedah to Selangor and Pahang).

Ecol. Montane rain-forest, to c. 1800 m.

Note. The inflorescence is similar to that of 5.
B. glomerulata but the leaves of these two species
cannot be confused.

5. Brassaiopsis glomerulata (BL.) REGEL, Gartenfl.
12 (1863) 275, t. 411; Koorp. Exk. Fl. Java 2
(1912) 715; Atlas 4 (1916) f. 674; FI. Tjib. 2 (1923)
226; BACK. & BAKH. f. Fl. Java 2 (1965) 167;
NoGoc-SANH Bul, Adansonia 6 (1966) 437; STONE,
Gard. Bull. Sing. 30 (1977) 282. — Aralia glomeru-
lata BL. Bijdr. (1826) 872. — Hedera glomerulata
(BL.) DC. Prod. 4 (1830) 265; Hook. Bot. Mag. 80
(1854) t. 4804. — B. speciosa DECNE & PLANCH.

1979]

Rev. Hort. 4, 3 (1854) 106; BoerL. Hand. 1 (1890)
643; HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894)
43, f. 1 B; K. & V. Bijdr. 7 (1900) 8; Baku. f. &
OoststTrR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 159, p. 11. — Macropanax glomerulatum
(BL.) Mia. FI. Ind. Bat. 1, 1 (1856) 764. — Macro-
panax cyrtostylum Miq. Ann. Mus. Bot. Lugd.-
Bat. 1 (1863) 13. — B. cyrtostyla (MIQ.) SEEM. J.
Bot. 2 (1864) 293.

Tree up to 10 m, with stout often sinuous
branches prickly in their upper parts, the young
parts of the shoots and inflorescences rufous-
tomentose. Leaves digitately compound, clustered
at the ends of the shoots; petiole c. 25-60 cm, with
an expanded basal sheath with two aculeate
stipules; leaflets 5—9, petiolules c. 8-10 cm (the
laterals shorter), blade elliptic-oblong, c. 15-25 by
6-10 cm, apex acuminate, base broadly cuneate to
rounded, midrib prominent, lateral veins widely
spaced, arched-ascending, margin serrulate. /n-
florescence a large panicle; rachis 30-60 cm, lateral
branches c. 20-30 cm, with umbellules arranged
racemosely along them on peduncles c. 2-5 cm
long, bracts numerous, lanceolate, c. !/, cm long,
mostly persistent; umbellules with c. 25-35 flowers,
pedicels 1—-1!/, cm, subtended by numerous minute
ferruginous bracts. Calyx of 5 small teeth. Petals
5, triangular, 3-4 mm long. Filaments 2—2!/, mm,
anthers oblong, 1 mm. Ovary obconic at anthesis,
c. 2 mm high; disk fleshy, cushion-shaped, sur-
mounted by an awl-shaped stylar column. Fruit
globose, 6-7 mm high, with a persistent calyx rim
and a stylar column usually 2 mm long but occa-
sionally much shorter (even in the same inflores-
cence).

Distr. India to SW. China; in Malesia: Malay
Peninsula (Pahang, Negri Sembilan), Sumatra,
W. Java.

ARALIACEAE—I (Philipson) 95

Ecol. Humid, evergreen, montane rain-forest,
1200-2200 m.

Vern. Sumatra: kayu aro, M (Kerintji); Java:
panggang, p. pujut (or puju), p. ranti (or ranto), S.

Note. Juvenile plants bear leaves which are
simple, irregularly lobed, or with 2, 3 or 5 leaflets.
The flowers are pale yellowish cream. The cut
stems exude a yellowish sap.

6. Brassaiopsis elegans RIDL. J. Linn. Soc. Bot. 41
(1913) 291; Fl. Mal. Pen. 1 (1922) 888; Strong,
Gard. Bull. Sing. 30 (1977) 282, f. 4. — Fig. 33.

Shrub, stems with few to many prickles, young
parts rufous-tomentose, becoming glabrous. Leaves
digitately compound, or occasionally simple;
petiole c. 14 cm, striate, with a small basal sheath
and a short, bicuspid ligule; leaflets 3—5 (or 1),
petiolules c. 3 cm (laterals shorter); blade elliptic,
c. 12-16 by 4-5 cm, apex long acuminate, base
cuneate, midrib prominent, lateral veins widely
spaced, arched-ascending, margin minutely denti-
culate. Inflorescence a lax gracefully pendent
raceme of umbellules; rachis c. 23 cm, lateral
branches c. 5 cm, subtended by small lanceolate
bracts and bearing 1-2 minute bracts near the
middle, terminating in spherical umbellules and
occasionally with separate flowers arising below
the apex; umbellules with c. 10-13 flowers, sub-
tended by numerous minute bracts; pedicels at
anthesis c. 1/, cm elongating to c. 11/,-11/, cm in
fruit. Calyx of 5 minute teeth. Petals 5, triangular,
21/, mm long. Stamens 2 mm long, anthers oblong.
Ovary obconic, c. 2 mm high; disk fleshy, sur-
mounted by an awl-shaped stylar column. Fruit
globose, 6 mm high, with a persistent calyx rim and
stylar column.

Distr. Malesia: Malay Peninsula (Selangor,
Pahang), very local.

Ecol. Montane, evergreen rain-forest, 800 m.

14. TREVESIA

VISIANI, Giorn. Tosc. Sc. Med. | (1840) 72; Mem. Accad. Torino I, 4 (1842) 262;
Mia. FI. Ind. Bat. 1, 1 (1856) 747; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 10; BTH. in
B. & H. Gen. Pl. 1 (1865) 942, p.p.; BoerRL. Ann. Jard. Bot. Btzg 6 (1887) 107;
Handl. 1 (1890) 639; HArMs in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; KING, J. As.
Soc. Beng. 67, 11 (1898) 57; Koorpb. Atlas 4 (1916) f. 664-666; Hutcu. Gen. FI. Pl. 2
(1967) 71. — Petasula NORONHA, Verh. Bat. Gen. 5 (1790) ed. 1, art. 4: 3, p.p.,
cf. STEEN. Regn. Veget. 71 (1970) 376. — Fig. 35, 36.

Shrubs or trees, unarmed or prickly, tomentose at least on young parts; buds
enclosed in cataphylls. Leaves large, palmately lobed or digitately compound,
stipules forming a bicuspid ligule within the terete petiole. Flowers in umbellules
which are arranged racemosely along a rachis or in panicles; pedicels not articu-
lated below the ovary. Calyx a small rim. Petals 7-12, valvate, often falling at

96 FLORA MALESIANA [ser. I, vol. 9!

Fig. 35. Trevesia burckii BOERL. a. Habit, */;, b. flower, x 2, c. fruit and ditto in CS, x 1'/, (@ RAHMAT
st Toroes 5048, b YapP 99, c RAHMAT sI ToroEs 5049). Drawn by P. PRENDERGAST.

1979] ARALIACEAE—I (Philipson) 97

anthesis as a cali ies Stamens as many as the petals; Bothers large ovat dorsi-
fixed, introrse, filament strap-like. Ovary inferior, broadly obconic or subglobose,
7-12-celled; disk rising gradually to a central boss formed by the united styles.
Fruit globose or ovoid, crowned by the disk and prominent stylar column; pyrenes
coriaceous, compressed, endosperm smooth.

Distr. 6 spp. ranging from eastern India and SW. China to Malesia; 5 of the species occur in W.

Malesia: Sumatra, Malay Peninsula, Java, Borneo, Lesser Sunda Is. (Lombok).

Formerly erroneously credited to the Philippines, cf. MERRILL, En. Philip. 3 (1923) 237.

Ecol. Primary rain-forest and second growth, especially in humid, shady localities.

Note. The genus comprises three well known species, each occupying distinct areas, except that the
ranges of two overlap in Sumatra. In addition there are three very local and imperfectly known species.

KEY TO THE SPECIES

1. Leaves digitately compound.

2. Petiolules connected by a web of tissue for most of their length

2. Petioles free. Continental SE. Asia. .
1. Leaves palmately lobed.

3. Inflorescence branches and petioles densely covered with long red setae

: Sk peck
: T. palmata (DC.) Vis.

2. T. rufo-setosa

3. Inflorescence branches not as above (but sometimes setulose, furfuraceous, pubescent or glabrous).

4. Flowers sessile or subsessile . .
4. Flowers pedicelled.

5. Ovary c. 14-celled, pedicels stout,c.4mm@ ..
5. Ovary c. 8-10-celled, pedicels more slender, 1-2 mm @

1. Trevesia burckii BoERL. Ann. Jard. Bot. Btzg 6
(1887) 110, pl. 12(-14); Handl. 1 (1890) 649; Merr.
En. Born. (1921) 456; MASAMUNE, En. Phan. Born.
(1942) 566. — T. palmata (DC.) Vis. var. cheirantha
CLARKE, FI. Br. Ind. 2 (1879) 732; KiNG, J. As.
Soc. Beng. 67, ii (1898) 58. — T. cheirantha
(CLARKE) O. K. Rev. Gen. Pl. 1 (1891) 272; RIDL.
Fl. Mal. Pen. 1 (1922) 882. — Fig. 35.

Shrub or small, sparsely branched tree up to
10 m, branches stout with small prickles, young
parts rufous-furfuraceous, cataphylls_ prickly.
Leaves crowded at the ends of the shoots, digitately
compound; leaflets usually 7-9; petioles c. 60 cm,
striate, sometimes with small prickles or bristles,
dilated into a clasping base, ligule with two lanceo-
late lobes; petiolules united for all or most of
their length by a foliaceous web; leaflets oblong-
lanceolate or elliptic, up to 30 by 10 cm, apex
acuminate, base cuneate or rounded, margin
finely serrate especially in the upper part, principal
veins arched-ascending, prominent. /nflorescence a
large terminal panicle, often overtopped by lateral
shoots; rachis to 60 cm, slightly prickly or not,
bearing branches along its length and usually end-
ing an umbel of branches, bracts caducous or
persistent; secondary branches (peduncles) c. 10—
18 cm, sometimes with 1 or few bracts along their
length and terminating in spherical umbellules of
40-50 flowers; pedicels c. 20-35 mm, slender.
Calyx an irregular obsolete rim. Corolla hemi-
spherical, c. 6 mm high, falling as a calyptra,
petals c. 7-10. Stamens c. 7-10, filament flattened
c. 21/, mm, anthers broadly triangular, c. 3 mm

3. T. beccarii

4. T. sundaica
. 5. T. arborea

long. Ovary at anthesis broadly obconic, usually
rufous-furfuraceous, c. 7—10-celled; disk broadly
conical surmounted by the connate styles, stigmas
slightly swollen. Fruit subglobose 1*/, by 1*/, cm,
surmounted by the stylopodium and connate
styles.

Distr. Malesia: Sumatra, Malay Peninsula
(from Kedah southwards), Borneo (Sarawak).

Ecol. Primary rain-forest, mostly below 500 m,
rarely up to 1000 m.

Vern. Sumatra: tapa arimau, tada lada,
Malaya: kakabu, tapak itek, t. rimau, M.

Note. The stamens are described as cream on an
orange disk. Seedling plants have leaves which are
palmately lobed or entire.

M;

2. Trevesia rufo-setosa RIDL. J. Str. Br. R. As. Soc.
n. 86 (1922) 294; Fl. Mal. Pen. 1 (1922) 883.

A shrub with stout prickly stems, young parts,
petioles and inflorescence densely covered with
red-brown flat setae. Leaves palmately lobed,
clustered at the ends of the shoots; petioles
c. 60 cm, dilated into a clasping base with a bicus-
pid ligule; blade c. 30 by 30 cm, with 7 lobes;
lobes c. 20 by 9 cm, elliptic, shortly cuspidate, mid-
veins and widely spaced arched-ascending laterals
prominent, margin with many upwardly directed
spinulous teeth. Inflorescence terminal; rachis
flexuous, c. 20 cm, bearing lateral umbellules,
bracts linear c. 12 mm; pedicels short. Flowers
heterosexual. Ca/yx rim obscure. Petals 5, triangu-
lar, 6 mm long, setose on the outside, spreading.
Stamens 5. Ovary in the fertile flowers obconic,

FLORA MALESIANA [ser. I, vol. 9!

Fig. 36. Trevesia sundaica Mia. in mixed evergreen mountain forest in E. Java at Sarangan, c. 1200 m
altitude, with Quercus (Photogr. JEswieT, 1925).

1979]

setose, 5-celled; disk with cylindrical projecting
stylar column. Fruit unknown.

Distr. Malesia: Malay Peninsula (Selangor:
Semangkok Pass).

Ecol. Montane forest, at c. 800 m.

3. Trevesia beccarii BoeRL. Ann. Jard. Bot. Btzg 6
(1887) 110, pl. 11.

Shrub, c. 3 m, with stout prickled stems. Leaves
palmately lobed, clustered at the ends of the
branches; petiole c. 50 cm, hirsute with short
patent bristly hairs when young, some persisting,
dilated into a clasping spinulose base with a bicus-
pid ligule; blade c. 30 by 40 cm, with 7-9 lobes,
base cordate, lobes c. 15 by 7 cm, broadly elliptic-
oblong, apex rather blunt, rusty stellate-pubescent
when young, sometimes persisting, mid-veins and
rather numerous arched-ascending lateral veins
prominent, margin irregularly serrate. Inflorescence
when young hirsute like the petioles, sometimes
persistent; rachis c. 90 cm, bearing many lateral
branches (peduncles) along its length, bracts
lanceolate, caducous; peduncles c. 5-15 cm with
some caducous, linear bracts along their length and
around the base of the umbellules (capitula).
Capitula of c. 10-20 sessile (or subsessile) flowers.
Calyx rim short, undulate. Corolla conical in bud,
falling as a calyptra. Stamens 10-12, anthers
oblong, filaments stout. Ovary obconic, angular
from contact with neighbouring flowers, 10—12-
celled; disk broadly conical; surmounted by the
massive connate styles, stigmas slightly swollen.
Fruit obconic angular from mutual pressure, c. 10
by 7 mm.

Distr. Malesia: West Central Sumatra.

Ecol. Forest, ascending to 1200 m.

Vern. Likabau, M.

4. Trevesia sundaica Mia. Pl. Jungh. 3 (1855) 420;
Fl. Ind. Bat. 1, 1 (1856) 747; DE Vriese, PI. Ind. Or.
(1857) 81; Mie. Ann. Mus. Bot. Lugd.-Bat. 1
(1863) 11; Boer. Ann. Jard. Bot. Btzg 6 (1887)
111, pl. 12 (16); Handl. 1 (1890) 649; K. & V.
Bijdr. 7 (1900) 4, incl. var. glomerata K. & V. l.c.
5; Koorp. Exk. Fl. Java 2 (1912) 710; Atlas 4
(1916) f. 664-666; FI. Tjib. 2 (1923) 222; Baku. f.
& Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7
(1948) fam. 159, p. 4; Back. & BAKH. f. Fl. Java 2
(1965) 163. — Aralia palmata REINW. ex BL. Cat.
Btzg (1823) 43, nomen; ex DE VRIESE, Pl. Ind. Or.
(1857) 82, nomen in synon.; BoOERL. Ann. Jard. Bot.
Btzg 6 (1887) 111, in synon., non LAMK, 1783, nec
Lour. 1790. — Sciadophyllum palmatum BL. Bijdr.
(1826) 875, non T. palmata (Roxs.) Vis. 1842. —
Aralia reinwardtiana STEUD. Nomenct. 1 (1840)
118, nomen. — Gastonia sundaica (MiIQ.) BAILL.
Hist. Pl. 7 (1880) 161, f. 202. — Actinophyllum
palmatum BL. ex BoerRL. Ann. Jard. Bot. Btzg 6
(1887) 111, in synon. — Fig. 36.

Shrub or small, sparsely branched tree, up to

ARALIACEAE—I (Philipson) 99

8 m; branches stout, with small prickles, young
parts stellate-tomentose, becoming glabrous, buds
with cataphylls. Leaves palmately lobed, clustered
at the ends of the branches; petiole striate, c. 50—
60 cm, unarmed or with a few prickles near the
base, dilated with a clasping, heavily lenticellate
base, ligule bicuspid; blade c. 60 by 60 cm, with
7-11 lobes, base cordate, lobes c. 30 by 40 cm,
elliptic-oblong, apex acute to acuminate, separated
by wide sinuses, mid-veins and widely spaced
arched-ascending lateral veins prominent, margin
serrate, occasionally irregularly incised. Inflores-
cence arising among or below the leaves, a raceme
of umbellules, at first rusty pubescent, glabrescent;
rachis up to 60 cm, often much shorter, bearing
branches along its length and ending in an umbel,
bracts lanceolate, caducous; secondary branches
(peduncles) c. 8-16 cm, bearing small bracts along
their length and around the umbellules; umbellules
of c. 20-35 flowers, pedicels c. 2-5 cm, lateral
umbellules sometimes bearing small sterile (male)
flowers. Calyx rim very short, undulate. Corolla
hemispherical in bud, c. 6 mm high, falling as a
calyptra, petals 8-12. Stamens 8-12, filaments
stout, anthers ovate, c. 3 mm long. Ovary at
anthesis broadly obconic, rufous-furfuraceous,
c. 8-10-celled; disk broadly conical passing into the
stylar column. Fruit semiglobose, 1*/,-1'/, cm 2,
surrounded by the stylopodium and connate

styles.

Distr. Malesia: Sumatra, Java, Lesser Sunda Is.
(Lombok).

Ecol. Evergreen rain-forest, especially in

ravines, ascending to 1500 m.

Vern. Sumatra: ahab, tapa arimau, M; Java:
borang, djemporang, dorang, gabus, gorang,
lontanglanting, panggang, p. lembur, p. puju, p.
tjutjuk, papanggangan, S.

5. Trevesia arborea MeERR. Contr. Arn. Arb. 8
(1934) 116.

Tree up to 15 m, young parts shortly reddish
stellate-tomentose, prickly. Leaves palmately lobed,
clustered at the ends of the branches; petioles
17-50 cm, shortly rusty tomentose, glabrescent,
unarmed, dilated into a sheathing base, ligule bi-
cuspid; blade rotund, 25-40 cm, base cordate,
glabrous above, shortly red-pubescent below,
coriaceous, usually 9-lobed, sinuses narrow, lobes
oblong-elliptic or oblong-oblanceolate, acuminate,
margin distantly serrate distally. Inflorescence a
raceme of umbellules, at first reddish pubescent,
glabrescent, sometimes with a few spines below;
rachis stout (c. 1 cm @), c. 35 cm, bearing branches
along its length, and ending in an umbel, bracts
oblong-ovate, acuminate 1-2 cm long; primary
branches (peduncles) few, spreading, stout, without
spines, to c. 15 cm; umbellules of c. 8-13 flowers,
pedicels 11/,-2!/, cm, stout (3-4 mm @), reddish
tomentose, basal bracts triangular, c. 5 mm long.
Flowers hermaphrodite. Calyx an irregular short

100 FLORA MALESIANA [ser. I, vol. 91

Fig. 37. Dendropanax borneensis (PHILIPSON) MERR. a. Habit, x '/,, b. flower, x 5, c. fruit and ditto in CS,
x 5, d. bifid leaf with area enlarged to x 100 to show glands (a—b NOOTEBOOM 2258, c CLEMENS 51039,
d CLEMENS 28927). Drawn by P. PRENDERGAST.

1979] ARALIACEAE—I (Philipson) 101

hemispherical, shortly furfuraceous, 14-celled.
Fruit unknown.
Distr. Malesia: N. Sumatra (Atjeh).

Ecol. Primary, evergreen forest, 1100 m.

rim. Corolla hemispherical in bud, c. 6 mm high,
12 mm diam., densely rusty pubescent outside,
falling as a calyptra. Stamens c. 13, filaments stout,
4 mm, anthers ovate, c. 4 mm long. Ovary broadly

15. DENDROPANAX

DECNE & PLANCH. Rev. Hort. IV, 3 (1854) 107; Bru. in B. & H. Gen. Pl. 1 (1865)
943; CLARKE, FI. Br. Ind. 2 (1879) 733; MeRR. Brittonia 4 (1941) 129; Hut-Lin Lt,
Sargentia 2 (1942) 38; PHILIPsoN, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 18;
Hutcu. Gen. Fl. Pl. 2 (1967) 71; Srone, Gard. Bull. Sing. 30 (1977) 148. —
Gilibertia Ruiz & Pav. Prod. FI. Peruv. (1794) 50, non J. F. GMEL. 1791; BTH. in
B. & H. Gen. Pl. | (1865) 944; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 40 &
Nachtr. 2 (1900) 254; PHILtIpson, J. Bot. 78 (1940) 116. — Textoria Miq. Ann. Mus.
Bot. Lugd.-Bat. 1 (1863) 12. — Fig. 37.

Unarmed, usually glabrous small trees, or erect or subscandent shrubs. Leaves
simple, entire, or sometimes palmately 3—5-lobed, often with pellucid glands, with-
out articulation between blade and petiole; stipular sheath inconspicuous. Umbels
solitary or compound, or on racemosely arranged branches or paniculate; pedicels
not articulated below the ovary. Flowers sexually dimorphic. Calyx on entire rim
or 5-dentate. Petals 5, valvate. Stamens 5-8, extrorse. Ovary inferior, 4-8-celled;
styles united throughout all or part of their length, or free. Disk fleshy, conical.
Fruit globose or ellipsoid, usually strongly ribbed when dry; exocarp fleshy;
endocarp cartilaginous. Endosperm smooth or rarely weakly ruminate.

Distr. About 30 spp. in tropical America and eastern Asia south to Malesia (3 spp.): N. Sumatra,

Malay Peninsula, Borneo. Fig. 38.

Ecol. Rain-forest in lowland and montane regions, 300-3100 m.
Note. Distinguished from Hedera by the normally erect habit and the smooth (or only weakly

ruminate) endosperm.

KEY TO THE SPECIES

1. Leaves without pellucid gland-dots.
2. Inflorescence a compound umbel
Zainnorescencea simple umbel.. 4) 2) ei.

1. Leaves with pellucid gland-dots

1. Dendropanax lancifolius (RIDL.) RmDL. Fl. Mal.
Pen. 1 (1922) 884; Stone, Gard. Bull. Sing. 30
(1977) 148, f. 6 C. — Arthrophyllum lancifolium
RIDL. J. Str. Br. R. As. Soc. n. 75 (1917) 38.

Tree to c. 14 m. Leaves dispersed along the
shoots; petiole c. 5 cm, slightly dilated at the base
and with an obscure ligular rim; blade to c. 15 by
33/, cm, chartaceous, lanceolate, base narrowly
cuneate, apex gradually tapered to an acute point,
margin entire but undulate, midrib well defined,
secondary and tertiary veins delicate. Umbel
terminal, compound; peduncle c. 2 cm; secondary
rays few, 1-2 cm; pedicels 5—10, c. 1-2 cm. Calyx a
rim with indistinct teeth. Petals triangular, falling
as a cap or separating. Ovary 4~5-celled, styles
forming an erect column.

1. D. lancifolius
2. D. maingayi
3. D. borneensis

Distr. Malesia: Malay Peninsula (Perak) and
N. Sumatra (Toba Lands).

Ecol. Montane rain-forest, c. 300-1500 m.

Vern. Sumatra: modang simarla siak, Batak
lang.

2. Dendropanax maingayi KING, J. As. Soc. Beng.
67, ii (1898) 48; RipL. Fl. Mal. Pen. 1 (1922) 884;
Stone, Gard. Bull. Sing. 30 (1977) 48, f. 6 B. —
D. parviflorus [non (CHAMP.) BTH.] CLARKE,
Fl. Br. Ind. 2 (1879) 733. — Gilibertia maingayi
PHILIPSON, J. Bot. 78 (1940) 117.

Slender shrub to c. 1!/, m, with few branches.
Buds enclosed in small cataphylls. Leaves dis-
persed along the shoots; petiole !/,-5 cm, chan-
nelled above, slightly dilated at the base and with

102

FLORA MALESIANA

[ser. I, vol. 91

Fig. 38. Range of Dendropanax DECNE & PLANCH.

a small to obscure ligule within the petiole; blade
5-15 by 2-53/, cm, thinly coriaceous to membran-
ous, ovate, elliptic, oblong or lanceolate, base
rounded to narrowly cuneate, apex acute, margin
entire, often undulate, midrib prominent, some-
times with three distinct principal veins. Umbel
terminal, simple, peduncle c. “1-2 cm; pedicels
1-2 cm, slightly elongating in fruit, bracts cadu-
cous. Flowers sexually dimorphic, males pre-
dominate in lateral umbellules. Calyx margin
minute. Petals triangular, c. 2'/, mm long. Stamens
5. Ovary 5—6-celled, reduced in male flowers, disk
fleshy, conical; styles 5—6, at first erect, but the
free upper half spreading in fruit. Fruit globose or
ellipsoid, succulent, 5—6-ribbed when dry, c. 10
by 8 mm, crowned by the persistent spreading
styles.

Distr. Malesia: Malay Peninsula (from Kedah
southwards).

Ecol. Montane rain-forest, 1000-2000 m.

Note. Very like the next species in the general
appearance of leaf and inflorescence. However, all
specimens from the Malay Peninsula lack pellucid
glands in the lamina of the leaf, whereas these are
present in all specimens from Borneo.

3. Dendropanax borneensis (PHILIPSON) MERR.
Brittonia 4 (1941) 132. — Gilibertia borneensis
PHILIPSON, J. Bot. 78 (1940) 116, fig. — Fig. 37.
Slender shrub or small tree, with few or no
branches, occasionally prostrate or subscandent,
sometimes reaching 5-6 m. Buds enclosed in few
broadly ovate cataphylls. Leaves dispersed along

the shoots; petiole 4-10 cm, channelled above,
slightly dilated at base and with a small to obscure
ligule within the petiole; blade 7-15 by 3-7 cm,
coriaceous, with pellucid glands, usually ovate,
elliptic or oblong, occasionally irregularly lobed,
base rounded to broadiy cuneate, apex acute,
margin entire, often undulate, midrib and lateral
veins prominent, frequently with 3 distinct princi-
pal veins. Umbel terminal, simple, or occasionally
compound, single or two borne side by side, with
c. 20 flowers; peduncle and secondary rays (if
present) usually rather short (1'/,-3'/, cm), but
occasionally much longer (to 13 cm); pedicels
c. 11/,-11/, cm (occasionally to 3 cm), elongating
slightly in fruit, bracts ovate caducous. Flowers
sexually dimorphic, males predominate in lateral
umbellules. Calyx margin minute. Petals triangular,
c. 21/, mm long. Stamens 5. Ovary 4~6-celled,
reduced in male flowers; disk fleshy, conical with
4-6 styles, at first erect, but the free upper half
spreading in fruit. Fruit globose or ellipsoid, succu-
lent, 4~6-ribbed when dry, 5-10 by 6-12 mm,
crowned by the persistent spreading styles.

Distr. Malesia: N. Sumatra (Mts Goh Lembuh
and Pinto) and northern Borneo (Sabah: Mt
Kinabalu; Brunei: Pagon Ridge; Sarawak: Mt
Murut and Kalabit Highlands).

Ecol. Local in montane, mossy forest, and alpine
thickets, 1700-3100 m.

Vern. Borneo: merit, Kalabit Highlands.

Notes. Flower greenish or cream, fruit black.

This species exhibits considerable variation in
size and shape of leaf, inflorescence and fruit, and

1979]

ARALIACEAE—I (Philipson)

103

also in venation pattern, but these variations are
not correlated with geographical distribution.
Specimens from the two Sumatran localities have

in Borneo. The presence of pellucid glands between
the reticulations of the veins of the lamina dis-
tinguishes this species from the other two species

small fruits and leaves, but similar specimens occur in the area.

16. ACANTHOPANAX

(DECNE & PLANCH.) H. Witte, Ann. Hort. Bot. 4 (1861) 89; Mia. Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 10; Bru. in B. & H. Gen. Pl. 1 (1865) 938; Seem. J. Bot. 5
(1867) 238; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 49; Merr. Philip. J.
Sc. 1 (1906) Suppl. 217; Hutcu. Gen. FI. Pl. 2 (1967) 69. — Panax subg. Acantho-
panax DECNE & PLANCH. Rev. Hort. IV, 3 (1854) 105. — Fig. 39.

Small trees or scandent shrubs, prickly or unarmed. Leaves digitately compound,
stipules absent, glabrous or with bristles. /nflorescence terminal, with umbels
either solitary or more usually umbellately or racemosely arranged; pedicels not
articulated below the flower. Flowers hermaphrodite or sexually dimorphic. Calyx
margin minutely dentate. Petals 4 or 5, valvate. Stamens as many as the petals,
anthers dorsifixed, introrse. Ovary 2—4(—5)-celled; disk fleshy; styles 2, connate to
about the middle. Fruit a subglobose drupe, crowned by the persistent bifid style,
2-4(-S)-seeded ; exocarp fleshy, endocarp chartaceous or cartilaginous. Endosperm
smooth or undulate.

Distr. About 30 spp. in eastern Asia and the Himalayan region, south to Malesia (2 spp.): Malay

Peninsula, N. Sumatra (Gajo Lands), Philippines (N. Luzon).

Ecol. Usually in montane forest or in secondary growth.

Note. A sterile specimen from Gunong Iran, Cameron Highlands (SyMINGTON CF 36300) in the
Kepong Herbarium, probably represents an undescribed species.

KEY TO THE SPECIES

1. Twigs with prickles. Leaf margins without bristles
1. Twigs without prickles. Leaf margins with bristles

1. Acanthopanax trifoliatus (L.) MerRR. Philip. J.
Sc. 1 (1906) Suppl. 217. — Zanthoxylum trifoliatum
LINNE, Sp. Pl. (1753) 270. — Panax aculeatus Att.
Hort. Kew. ed. 1, 3 (1789) 448. — A. aculeatus
H. Witte, Ann. Hort. Bot. 4 (1861) 89, nom. illeg.;
SEEM. J. Bot. 5 (1867) 238; Harms in E. & P. Nat.
Pfl. Fam. 3, 8 (1894) 50.

Scandent shrub, c. 2-8 m, with broadly based
recurved prickles sparsely disposed on the branches,
usually below the nodes; buds with small brown
cataphylls. Leaves disposed along the branchlets;
petiole up to c. 5 cm, flattened above, with a
slightly dilated base, glabrous, usually with a few
prickles; leaflets (1—-)3(—5); petiolules up to 8 mm,
articulated with the petiole, channelled above;
blade up to 31/, by 2!/, cm, broadly ovate to subro-
tund, apex acute, base cuneate, margins serrate,
chartaceous, glabrous, lateral veins conspicuous on
both surfaces, pinnate, reticulation inconspicuous.
Inflorescence terminating the main or lateral
branches, a sessile compound umbel; primary

1. A. trifoliatus
2. A. malayanus

rays c. 4-5, or rarely solitary, slender or the central
ray stronger (the laterals sometimes apparently
male), with minute furfuraceous bracts at their
bases, occasionally with a few prickles, glabrous,
c. 3-6 cm long; secondary rays (pedicels) numerous,
slender, c. 1-13/, cm. Calyx a rim with 5 minute
teeth. Petals 5, ligulate, c. 2 mm long. Stamens 5,
filaments c.2 mm. Ovary turbinate, c. 11/, mm high,
2-celled; styles 2, connate to about the middle.
Fruit a spheroidal drupe, c. 5 mm @ when dry,
crowned by the persistent bifid style; endocarp
chartaceous. Endosperm surface slightly undulate.

Distr. From the Himalayas through S. China
to Japan and Formosa; in Malesia: Philippines
(N. Luzon: Benguet; Bontoc).

Ecol. In montane forest and thickets, 1100-
1400 m.

2. Acanthopanax malayanus M. R. HENDERSON,
Gard. Bull. S. S. 7 (1933) 105, pl. 22. — Fig. 39.
Unarmed tree to 17 m, trunk to 1 m @; branch-

104 FLORA MALESIANA [ser. I, vol. 91

Fig. 39. Acanthopanax malayanus HENDERSON. a. Habit, x 1/2, b. flower, x 7, c. fruit, x 4 (a, b DE WILDE
c.s. 16725, c DE WILDE c.s. 15925). Drawn by W. R. PHILIPSON.

1979]

ARALIACEAE—I (Philipson)

105

lets rather stout (c. 5 mm @); bark greyish brown,
glabrous, smooth; buds with small brown cata-
phylls. Leaves crowded at the ends of the branch-
lets; petiole terete, striate, glabrous, with a slightly
dilated base, up to 12'/, cm; leaflets 3—5; petiolules
up to 7 mm, articulated with the petiole, channelled
above; blade up to 17 by 5'/, cm, ovate-lanceolate
to narrowly elliptic, apex acute, base cuneate or
one side truncate, chartaceous, the principal veins
pinnately arranged and conspicuous, the minor
veins forming a dense reticulation, margin with
numerous spinulose teeth. /nflorescence a terminal,
sessile, compound umbel, often on short lateral
shoots, apparently dioecious; primary rays c. 3-7,
slender, terete, glabrous, striate, c. 4-8 cm;
secondary rays (pedicels) of male flowers numer-
ous, slender, 1'/,-2 cm, of female flowers stouter.
Calyx a rim with 4-5 minute teeth. Petals 4—5,
ligulate, c. 2'!/, by 1'/, mm long. Stamens 4-5,
filaments c. 23/, mm long. Ovary turbinate, 3—4-
celled; styles 2, at first connate, but becoming free
to about the middle. Fruit an oblate spheroidal

drupe, c. 10 by 8 mm when dry, with a small stylo-
podium bearing a persistent bifid style; endocarp
cartilagineous. Endosperm surface strongly undu-
late.

Distr. Malesia: Malay Peninsula (Pahang;
Cameron Highlands), N. Sumatra (Gajo Lands).

Ecol. Montane rain-forest and mossy forest,
1400-2600 m.

Vern. Berlaki, Sakai lang.

Notes. The interpretation of this species as
having sexually dimorphic inflorescences cannot be
proved with the available material. This appears to
consist of twigs bearing either inflorescences of
male flowers, or infructescences. Appearances
suggest that the putative male flowers have rudi-
mentary ovaries. There is no evidence whether the
fruiting flowers had produced pollen.

The tree is said to be conspicuous when flowering
by reason of its delicate feathery foliage, which is
pale green with a reddish tinge, the petioles and
inflorescence also being reddish.

Excluded

Meryta colorata F. M. BAtLey, Queensl. Agric. J. 3 (1898) 283; HARMs, Bot. Jahrb. 56 (1920) 384.
The type specimen of this species cannot be located, but the original description does not seem to relate
to a member of the Araliaceae, and certainly not to a Meryta.

Panax ? anisum DC. Prod. 4 (1830) 254. — Anisum moluccanum RumpH. Herb. Amb. 2: 131, t. 42. —
Nothopanax ? anisum Mia. FI. Ind. Bat. 1, 1 (1856) 766; SEEM. FI. Vit. (1866) 114. — Polyscias anisum

Harns in E. & P. Nat. Pfi. Fam. 3, 8 (1894) 45.

According to MERRILL (Int. Rumph. 1917, 289) these names are all based on the Rumphian description
and plate and must refer to some species of Fagara, possibly F. avicennae LAMK = Zanthoxylum avicennae

(LaMk) DC. (Rutaceae).

VaR ays,

fj ana t 3) i

ek? 3 @ipe) “4
z nd Used ali

o heaee @ atin ik rwterpoen F
(iva Aw

1.” tage

ee

be tay

oo i faliwey Oa

&
iL ‘ my |

a :
ag :

CYPERACEAE—IDI (J. H. Kernt & H. P. Nooteboom, Leyden)

28. CAREX

LINNE, Gen. Pl. ed. 5 (1754) 420; Sp. Pl. (1753) 972; ENDL. Gen. (1836) 110;
KUNTH, En. 2 (1837) 368; SteuD. Syn. 2 (1855) 182; Mia. Fl. Ind. Bat. 3 (1856)
346; Boorr, Illustr. Genus Carex I-IV (1858-67); Boeck. Linnaea 39 (1875) 14;
ibid. 40 (1876) 327; ibid. 41 (1877) 145; B. & H. Gen. Pl. 3 (1883) 1073; Pax in
E. & P. Nat. Pfil. Fam. 2, 2 (1887) 122; CLARKE, FI. Br. Ind. 6 (1894) 699; J. Linn.
Soc. Bot. 37 (1904) 1; Philip. J. Sc. 2 (1907) Bot. 107; KUK. Pfl. R. Heft 38 (1909)
67; Philip. J. Sc. 6 (1911) Bot. 57; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11
(1936) 232; NELMES, Kew Bull. (1946) 5; Reinwardtia 1 (1951) 221; ibid. 2 (1954)
373; RAYM. Mém. Jard. Bot. Montréal n. 53 (1959) 17; KoyAma, J. Fac. Sc. Un.
Tokyo III, 8 (1962) 149; KERN in Back. & Bakh. f. FI. Java 3 (1968) 487. —
Fig. 118-135.

Perennial herbs with tufted or creeping rhizome, monoecious, rarely dioecious.
Stems arising centrally or laterally, erect or obliquely erect, mostly triquetrous or
trigonous, rarely subterete, solid or sometimes hollow, often clothed at the base by
persistent leaf-sheaths or their fibrous remains. Leaves tristichous, usually narrowly
linear, sheathing at the base, with a ligule at the junction of blade and sheath,
rarely lanceolate or elliptic with a more or less distinct petiole and eligulate, mostly
basal and subbasal, 0-several higher on the stem, the lower ones often reduced to
bladeless sheaths; sheaths of the stem-leaves and bracts closed. /nflorescence
paniculiform, racemiform or spiciform, more rarely reduced to a single spikelet.
Spikelets 1-very numerous, terete, sessile or peduncled, few- to many-flowered,
wholly male, wholly female, or bisexual (androgynous when male flowers above,
gynaecandrous when female flowers above). Bracts foliaceous or glume-like, often
sheathing, sometimes wanting. Base of the branches of the inflorescence usually
with a utriculiform or ocreiform bracteole (cladoprophyllum) surrounding it.
Flowers unisexual, naked, solitary in the axils of the spirally arranged glumes; male
flowers consisting of 3 free or rarely more or less connate stamens; anthers linear;
female flowers consisting of a single pistil enclosed in a bottle-shaped prophyll
(utricle, perigynium). Style either continuous with the ovary and persistent, or
articulated with it and deciduous, straight or flexuous, often incrassate at the base;
stigmas 2 or 3, protruding through the small terminal orifice of the utricle. Vestigial
rachilla (see Uncinia) rarely present. Utricles membranous, chartaceous, or
coriaceous, bicarinate, sometimes winged, sessile or stipitate, beakless to strongly
beaked, nerveless, nerved, or ribbed, glabrous, or pubescent or hispid, papillose or
puncticulate or smooth, sometimes spongy at the base; beak truncate, obliquely
cleft, bidentate, or bifurcate at the top. Nut trigonous (when stigmas 3), or lenticu-
lar (plano-convex or biconvex; when stigmas 2), enclosed within the utricle.

Distr. A large genus with 600 to 1000 spp., the majority of them outside the tropics. However,
the most primitive section, Vigneastra with a compound, paniculate inflorescence and andro-
gynous spikelets, occurs mainly in the tropics of the Old World, from sea-level up to 3000 m. This
section is represented in Malesia with 11 spp., and is there by far the largest section.

Subg. Carex is, besides with the 11 spp. of sect. Vigneastra, represented in Malesia by 25
sections and 49 spp., subg. Vignea with 6 sections and 6 spp.

(1) Continued from volume 7, part 3 (1974) p. 753.
(107)

108 FLORA MALESIANA [ser. I, vol. 9!

Of the 66 spp. in all, 12 spp. are endemic and mostly confined to one island. Of the other 54
spp. the majority occur also in the northern hemisphere, either widely or only in SE. and/or
E. Asia. Of these, 25 spp. extend without noticeable disjunctions into Malesia, either to its western
part or further eastward. Another 4 of the E. Asian spp. occur in Formosa and in Malesia only
in the Philippines (mostly only in Luzon), viz 12. C. satzumensis, 30. C. rhynchachaenium, 32.
C. dolichostachya, and 33. C. formosensis.

Among the Asian spp. 4 show a distinct disjunction between SE. Asia and Java, viz 11. C. vesi-
culosa, 14. C. helferi, 63. C. nubigena, and 58. C. longipes (which is also once found in Celebes
and once in New Guinea); all are mountain species.

A similar disjunction is found in the lowland species 10. C. stramentitia and 38. C. tricephala;
this disjunction is understandable as both are bound to a distinct seasonal climate which shows a
similar pattern (see VAN STEENIS, Reinwardtia 5, 1961, 420-429).

Still larger disjunctions are shown by 5 E. Asian spp. which are in Malesia almost only found
in New Guinea, Formosa being mostly the nearest station, viz 46. C. brachyathera, 47. C. finitima
(also in N. Sumatra), 48. C. michauxiana, 56. C. bilateralis, and 61. C. duriuscula; all are mountain
plants.

Not a few Carex spp. occur in both the northern and southern hemisphere (Australia, often
Tasmania and New Zealand) and are also found in the intervening tropical zone in Malesia.
There are 11 of such species in all. Among these the following 7 are found in more than one island,
viz 6. C. indica, 31. C. breviculmis, 39. C. maculata, 40. C. capillacea, 57. C. brunnea, 5. C. hors-
fieldii, and 64. C. echinata, although the latter two are very rare. Four others are also transtropical
but are in Malesia only found in New Guinea, and thus show almost the same disjunction
between Formosa and New Guinea as mentioned above; these are 43. C. brownii, 49. C. pseudo-
cyperus, 55. C. gaudichaudiana, and 66. C. curta. They occur in Malesia only in bogs and marshes
on (high) mountains.

The southern hemisphere has contributed very little to the Malesian Carices, as there is only
one southern species from Australia, New Zealand, and New Caledonia, viz 62. C. appressa,
which is found in Malesia, and then again only in New Guinea at very high altitude.

Summarizing, one cannot withdraw oneself from concluding that only the presence of moun-
tain bogs, marshes and grasslands gives the opportunity for the maintenance of Carex in the
Malesian tropics. This would explain why the big mountainous island of New Guinea — where
this habitat is most abundantly represented, much more than in any other island — is so rich in
species. It is then also understandable that this is the place where relict areas could maintain
their last stand in the tropical mountains of Malesia.

Ecol. Most species prefer moist or wet localities, some are found in rain-forest, others in open
habitats like in mountain marshes, grassland, heaths, etc. Only 22 spp. grow below 1000 m, 8 of
them belonging to sect. Vigneastra, 13 are found below 500 m, 4 of them of sect. Vigneastra;
14 spp. occur exclusively above 2000 m. 57. C. brunnea, which is found from 80 m up to 3400 m,
has the largest altitudinal range.

The majority of the Carex spp. is growing on more or less acid soils. Three species of the
lowland tropics occupy a different habitat, and are bound to a seasonal climate, viz 10. C. stra-
mentitia, 38. C. tricephala, and 37. C. speciosa. The first two occur only in Java or Madura I.,
amongst others with the teak-forest; the third of these drought species occurs also in some local
dry spots in E. Sumatra and SE. Borneo, and is obviously adapted to a slightly less severe dry
season.

Of the three mentioned species, C. tricephala and C. speciosa are confined to limestone, like
one other species, 35. C. malaccensis, an endemic of N. Malaya (the Langkawi Is.).

One species, 59. C. teinogyna, seems in Malesia to be confined to banks of swift streams and
occupies a rheophytic habitat.

Notes. The distinction of the sections is not always unanimously agreed on, and several
species were put into different sections by different authors. Some sections are closely allied,
the only difference being sometimes 2 versus 3 carpels. As this has great influence on the shape of
the nut, it often is considered a very important character. In my opinion the difference is due to a
minor genetic difference, and is taxonomically not very important. Besides, in some collections
2-carpelled and 3-carpelled fruits are found in the same inflorescence, e.g. in species of sect.

1979] Cy PERACEAE—II (Kern & Nooteboom) 109

Graciles, which has normally 2 carpels, and of sect. Oligostachyae (Decorae) which has normally
3 carpels (Noort.).

Acknowledgement. The manuscript of Carex was for the greater part elaborated by Dr.
J. H. KERN before he died in 1974. The obstacle for finishing it in time was his worry about the
status cq. evaluation of a number of names in sect. Vigneastra, which also prohibited the finishing
of the practical key to the species. Dr. H. P. NooTEBOom has done this evaluation and he made
the final draft of the key, for which he is responsible. He is also responsible for the treatment
offered here for several species, viz 3. C. cruciata, 18. C. perakensis, 19. C. turrita, 20. C. verticil-
lata, and 31. C. breviculmis, and for some critical notes which have been marked ‘‘Noor.”
between brackets. The division in subgenera and sections is largely from Dr. KERN. The revision
of Uncinia is entirely by Dr. NOOTEBOoo. (Ed.)

Explanation of some terms. The term ‘stems’ refers to the flower-bearing stems, except
when stated otherwise, whereas ‘inflorescence’ means the entire flower-bearing stem including all
the spikelets. The terms used for the inflorescence (‘panicle’, ‘raceme’) indicate the disposition
of the spikelets, not of the individual flowers. ‘Stems central’ means that the leaves are arranged
around the flower-bearing stem (usually towards the base). ‘Stems lateral’ means that the leaves
are placed on a vegetative shoot, and that the flower-bearing stems arise separately from the
rhizome and bear few short-bladed leaves. The flowering stems can also be situated in the axils
of the leaves. This situation is sometimes difficult to distinguish from ‘stems central’, because the
leaves are often crowded in a basal rosette.

The bracts are inserted on the stem and they are sheathing or not; they bear in their axils a
spikelet or a partial inflorescence, the peduncles of which are often basally surrounded by a
cladoprophyllum; this is a bracteole which is utriculiform or ocreiform. If the cladoprophyllum
is ocreiform, it usually is hidden entirely in the sheath of the bract. Each flower is solitary in the
axil of a glume; several flowers together form a spikelet which can be J, 9, or bisexual; in the
latter case the 3 flowers are usually placed only at the base or at the apex of a spikelet. Gynaecan-
drous spikelets have 2 flowers above, 3 below, androgynous spikelets have 3 flowers above and
2 below.

The figures 118-123, depicting utricles and nuts of nearly all the species, and some glumes
(fig. 132), are given to facilitate use of the keys.

KEY TO SUBGENERA AND SECTIONS
as represented in Malesia

1. Stigmas 3 and nut trigonous, or, when stigmas 2 and nut lenticular (plano-convex or biconvex) some or
all of the spikelets peduncled or terminal spikelet g. Spikelets either dissimilar in appearance (the
terminal one ¢ and some or all the lateral ones 9), or similar in appearance (all bisexual). Base of the
peduncles of the lateral spikelets or of the branches of the inflorescence surrounded by a utriculiform
or ocreiform bracteole (cladoprophyllum); inflorescence sometimes reduced to a single terminal spikelet

I. SuUBG. CAREX
2. Stigmas 3; nut trigonous.
3. Inflorescence consisting of more than one spikelet.
4. Lateral spikelets arising from a gaping, utriculiform, fertile (i.e. nut-bearing) cladoprophyll.
Spikelets androgynous, up to 15 mm long. Bracts not sheathing. Stems 3-20 cm tall. Sp. 12
2. Sect. Japonicae
4. Lateral spikelets arising from a sterile cladoprophyll, or sessile without cladoprophyll.

5. Stems scapiform, surrounded at the base by spadiceous sheaths in place of leaves. Leaves wide,
conduplicate-petioled below. Secondary panicles contracted, very dense. Utricles with very long
linear beak and very oblique mouth. Sp.14...... 2.2... . . . 4. Sect. Mapaniifoliae

5. Otherwise.

6. Spikelets peduncled, in fascicles in the axils of at least some of the sheathing bracts, all androgy-
nous or some wholly ¢ or 9. Utricles long-beaked. Spp.17-20 . . . . . 6. Sect. Oligostachyae
6. The peduncled or sessile spikelets or secondary panicles single or binate in the axils of the bracts.
7. Spikelets similar in appearance, androgynous (upper flowers 3, lower ones 9).
8. Lower bracts not or only shortly (some millimeters) sheathing.
9. Apex of nut truncate, 1/,-*/, mm @, hollowed out. Style centred in the apical hollow of the
nut. Utricles lageniform or rhomboid-lageniform. Spp. 27-30. . . 10. Sect. Lageniformes

110 FLORA MALESIANA [ser. I, vol. 91

9. Apex of nut not truncate. Style base pyramidally thickened, often broader than apex of nut.
‘Spps 35-300 See oan os ae a ee ee er. Ce 2 sSecratiemes
8. Lower bracts long sheathing.

10. Flowering stems arising from the axils of the leaves which are crowded on a short stem and
much longer than the flowering stems. Style flexuous. Nut with irregularly excavated sides,
seated ona spongy, disk-like stipe. Sp.26 .. . . . . 9. Sect. Cryptostachyae

10. Flowering stems arising either from the centre of the leaves, or from lateral shoots bearing a
few short-bladed leaves.

11. Spikelets small, up to 15(—20) mm long.
12. Stems central. Spikelets numerous, ovoid or oblong-ovoid, arranged in often diffuse,

compound panicles. Bracts long. Spp. 1-11 .. . .. . . 41. Sect. Vigneastra
12. Stems lateral. Spikelets few, globose-ovoid, in a simple panicle. Bracts shorter than 8 cm,
the sheaths not included:Sp:'21-....:. 2). &. bine =) s cn. ty) TenSeeSeeeeionste

11. Spikelets longer.
13. Utricles densely many-nerved. Spikelets 1—8(-14).
14. Utricles ellipsoid, short-beaked, more or less ciliate on the margins, spongy-thickened
at the base. Spikelets 1-3(-4). Spp. 35-38... ..... . . 12. Sect. Radicales
14. Utricles narrowly ellipsoid, long-beaked, more or less hispidulous on the faces, not
spongy-thickened at the base. Spikelets 2-8(-14). Spp. 17-20 . 6. Sect. Oligostachyae
13. Utricles distinctly but not densely nerved. Leaves with reddish sheaths splitting into
fibres. Spikelets 6-numerous. Spp. 15-16... . ... . 5. Sect. Polystachyae
7. Spikelets dissimilar in appearance, unisexual (or rarely gynaecandrous), the terminal one or
some upper ones 4, the remainder 2 (occasionally the 2 spikelets may have a few 3 flowers, and
the ¢ spikelets a few 2 flowers). Or most of the spikelets 9, shortly cylindrical, the 3 spikelets
lateral, 1-5 just below some of the terminal 9 spikelets.

15. Leaves not crowded in a basal rosette but inserted all over the stem, the upper ones merging
into leafy bracts, the lower ones gradually decreasing in length and merging into bladeless
sheaths.

16. Inflorescence a compound panicle consisting of 5-8 secondary panicles each containing
numerous @ spikelets and 1-5 3 spikelets. Spikelets small, 4-8 mm long. Utricles glabrous,
with short, !/, mm long beak. Leaves sparsely hairy beneath. Sp. 13

3. Sect. Hypolytroides

16. Inflorescence simple, racemose, consisting of 4-9 spikelets; terminal spikelet J (sometimes a
second, smaller one added), remaining spikelets 9, 11/.-4 cm long. Utricles densely hispid,
with distinct, c. 1 mm long beak. Leaves glabrous. Sp. 50... . . . 21. Sect. Occlusae

15. Leaves crowded at the base of the stem, often also some higher on the stem.

17. Nut excavated in the middle, the apex abruptly narrowed into a conspicuous, hastiform,
i.e. cylindric, !/,—'/, mm long beak expanding into the annulate apex. Spp. 22-25

8. Sect. Rhomboidales
17. Nut without hastiform beak.
18. Apex of nut truncate, 3/,-1 mm wide, hollowed out. Style centred in the apical hollow of
the nut. Utricles lageniform or rhomboid-lageniform. Spp. 27-30 10. Sect. Lageniformes
18. Apex of nut not truncate, at most '/, mm wide.
19. Nut mitrate, i.e. contracted at the apex and then expanded into a discoid annulus.
Wtricles ovoidiorellipsoid.,Spp. 31-34 . .: :.-. .. . +222 11. Sect. Mitratae
19. Nut not mitrate.
20. Utricles densely papillose, erostrate or but shortly beaked. Sp. 39
13. Sect. Trachychlaenae
20. Utricles not papillose, distinctly beaked.
21. 2 Spikelets broadly ovoid or subglobose, with few narrowly lanceolate, c. 1 cm or more
long utricles. Bracts sheathing. Rhizome short. Sp.48 . . . . 19. Sect. Folliculatae
21. 2 Spikelets not broadly ovoid-subglobose. Utricles smaller.
22. Spikelets linear-cylindric, loosely flowered. Utricles nerveless (except for 2 submar-
ginal nerves) or obscurely nerved, with long beak and oblique mouth. Bracts sheathing.
23. Utricles hispidulous, brownish. Sp.46 ...... . .. 17. Sect. Ferrugineae
23. Utricles glabrous, light green. Sp.47 .... . ... . . 18. Sect. Sylvaticae
22. Spikelets oblong or cylindric. Utricies many-nerved.
24. Spikelets short-cylindric, few-flowered. Utricles fusiform-ellipsoid or rhomboid, at
least towards the base many-nerved. Flowering stems lateral or central. Spp. 22-25
8. Sect. Rhomboidales

24. Spikelets cylindric, densely many-flowered.

1979] CyPERACEAE—II (Kern & Nooteboom) 111

25. Utricles firm, of thick texture, short-beaked. Rhizome emitting strong stolons.
Leaves conspicuously septate-nodulose. une erect. Bracts not or scarcely
sheathing. Sp. 60 . . . . . 26. Sect. Paludosae
25. Utricles membranous, with long beak (except in C. brownii ssp. brownii).
26. Spikelets erect. Utricles patulous, subinflated, many-nerved, with shortly bidentate
beak. Style straight. Rhizome often creeping. Spp. 42-45 . 16. Sect. Anomalae
26. Lower spikelets cernuous. Utricles widely patent to reflexed, closely many-ribbed,
with bifurcate beak. Style flexuous. Rhizome cespitose. Sp. 49
20. Sect. Pseudocypereae
3. Inflorescence consisting of a single terminal spikelet.
27. Utricles with whitish pubescent margins or hispidulous, densely and strongly many-nerved,
A—1Oenim longs: Spps5—36il 2) we doe ee eles ib, elZo Sect Radicales
27. Utricles glabrous, 1'/,-S mm long.
28. Utricles rather large (4'/,-5 mm long), with slender, linear, bidentate beak. Sp. 41
15. Sect. Rhizopodae
28. Utricles small (1!/,4 mm long), with short, subentire or slightly emarginate beak. Sp. 40
14. Sect. Capitellatae
2. Stigmas 2; nut lenticular (plano-convex or biconvex).
29. Bracts sheathing. Spikelets often in fascicles. Utricles often hispidulous, membranous, nerved,
usually long-beaked. Stigmas often very long. Spp. 56-59 ...... . . 25. Sect. Graciles
29. Bracts not sheathing (or very shortly).
30. Lateral spikelets sessile or the lowest shortly peduncled, erect. Terminal spikelet ¢. Glumes
PHULICOUS: SPSS Estee >. Sg (Meee Ss ee Sn MISS te 2A Seck Carex:
30. Spikelets peduncled, cernuous. ‘Glumes often awned.
31. Spikelets 3-8, single at the nodes, cylindric, short peduncled. Terminal spikelet 3, or (often with
exception df basal spikelets) spikelets gynaecandrous. Spp. 52-54. . . . 23. Sect. Praelongae
31. Spikelets 6—50, single or binate at the nodes, the lower ones long-peduncled, all androgynous or
some 3, linear-cylindric. Sp.51.... . .... . 22. Sect. Longispicae
1. Stigmas 2 and nut lenticular. Spikelets similar in appearance, bisexual, sessile. Cladoprophylla usually
absent . =. . (foie ote = see SupG<VIGNEA
32. Spikelets androgynous (upper flowers 3 lower ones 9).
33. Rhizome long-creeping. Spikelets 4-7. Utricles coriaceous, ppg sharp-edged but not
winged=S7-.61.o0 420. 4 ped 2) ees Sect. Divisae
33. Rhizome short, forming dense tufts. Spikelets numerous or ‘very numerous.
34. Utricles coriaceous, with rounded margins. Bracts inconspicuous. Sp. 62 28. Sect. Paniculatae
34. Utricles membranous, with acute or winged margins. Lower bracts foliaceous. Sp. 63
29. Sect. Multifiorae
32. Spikelets gynaecandrous (upper flowers, 2, lower ones 3). Utricles membranous.
35. Utricles with rounded margins, densely whitish-puncticulate, beakless or very shortly beaked.
Spikelets ebracteate (bracts glumiform). Sp.66 ........ .. . .32. Sect. Heleonastes
35. Utricles with acute or winged margins, not puncticulate.
36. Utricles obliquely erect, also when mature. Spikelets more or less distinctly bracteate. Sp. 65.
31. Sect. Elongatae
36. Utricles widely spreading or reflexed when mature. Spikelets not or inconspicuously bracteate
(eencts clumiform)., Sp.64. . 2 2 < 2dse he Ns. Se, ee eer Sicliatae

KEY TO THE SPECIES

Only complete specimens with nearly or fully mature utricles are identifiable. As the distribution of
sexes is variable, especially in the species with a terminal male spikelet, it is advisable to examine several
specimens if possible.

In the measurements of the length of the utricles the beak is included. Unless stated otherwise ‘glumes’
refers to the glumes subtending the female flowers.

1. Inflorescence consisting of a single (androgynous) spikelet terminating the stem. Stigmas 3; nut
trigonous.

2. Utricles with whitish pubescent margins, or hispidulous, or winged and scabrid margined, 4-8 mm
long. Glumes (many-)nerved. Style pyramidally thickened towards the base, persistent on the nut.
Leaves (2—)3-10 mm wide.

3. Spikelets 2-8 cm long. Utricles with whitish pubescent margins ........ 37. C. speciosa
3. Spikelets 6-12 mm. Utricles hispidulous or glabrous with scabrid margins.

£12 FLORA MALESIANA [ser. I, vol. 91

4. Utricles hispidulous in the upper 2/,, 4-6 mm, with conic-cylindric, 1-2!/, mm long beak.
38. C. tricephala
4. Utricles glabrous with scabrid, distinctly winged Sagan. 6-8 mm, gradually narrowed into the
3mm long beak .. . . . 36. C. ramosii
2. Utricles glabrous, membranous and otherwise differed 2 5 mm n long. Glumes gi nerved in the centre,
otherwise nerveless. Style not or but slightly thickened towards the base. Leaves !/,-3 mm wide.
5. Utricles (11/,-)21/,-3'/,(-4) mm long, rather gradually tapering into a very short, subentire or

slightly emarginate beak. Leaves '!/,-2 mm wide. Glumes 1!/,-3:mm long . . . 40. C. capillacea
5. Utricles 4'/,-5 mm long, abruptly narrowed into a slender, linear, bidentate, c. 1!/, mm long beak.
Leaves 2-3 mm wide. Glumes 3-4 mm long ........... .. .. 41. C. eremostachya

1. Inflorescence consisting of 2 or more spikelets.
6. Terminal and most of the other spikelets 2, shortly cylindrical, the ¢ spikelets lateral, 1-5, just below
some of the terminal 2 spikelets. Inflorescence a compound panicle. Leaves all over the stem
13. C. hypolytroides
6. Terminal spikelet bisexual or wholly 2.
7. Terminal spikelet wholly 3 (exceptionally in some specimens with a few © flowers), or when spikelets
in fascicles, 1-2 spikelets of the upper fascicle wholly 3.
8. Stigmas 3.

9. Leaves inserted all over the stem, the upper ones merging into leafy bracts, the lower ones gradu-
ally decreasing in length and merging into bladeless sheaths. Inflorescence simple, racemose,
consisting of 4-9 spikelets. Spikelets oe cm long. Utricles densely hispid, with distinct, c. 1 mm
long beak. Leaves glabrous. . . ~ Sees 29502 @imankernare

9. Leaves basal and often also some higher o: on the stem.

10. Nut excavated in the middle, the apex abruptly narrowed into a conspicuous, cylindric,
'/,-1'/, mm long beak expanding into the annular apex. . ... . . . 22. C. anomocarya
10. Nut different.
11. Apex of nut truncate, 3/,-1 mm wide, hollowed out. Style centred in the apical hollow of the
nut. Utricles lageniform or rhomboid-lageniform.
12. Utricles 5-7 mm long. Apex of nut contracted into a cylindric, truncate, ?/,-1 mm long and
wide neck.
13. Leaves 5-10 mm wide. Beak of utricles 1'/,-2 mm long. Stems arising from basal leaf axils
28. C. gracilispica
13. Leaves 1-3 mm wide. Beak of utricles 3/,-1 mm long. Stems central 30. C. rhynchachaenium
12. Utricles 3!/,-4'/, mm long. Apex of nut hardly or not contracted into a cylindric neck
27. C. breviscapa
11. Apex of nut not truncate, at most '/, mm wide.
14. Utricles densely papillose, subabruptly contracted into a very short, up to '/, mm long beak
39. C. maculata
14. Utricles not papillose, nut either mitrate or distinctly beaked.
15. Nut mitrate, 7.e. contracted at the apex and then expanded into a discoid annulus.
16. Glumes of ¢ spikelets cup-shaped, the margins more than halfway connate in front.
Stamens monadelphous (not always so in the lowermost flowers). Utricles many-nerved
34. C. tristachya
16. Glumes of ¢ spikelets with free margins. Stamens free.
17. Nut rhomboid with deeply concave faces. . . ...... ... . 33. C. formosensis
17. Nut (long-)ellipsoid to ovoid or cbovoid.
18. Leaves 3-10 mm wide. Stems arising from basal leaf-axils. Utricles strongly many-nerved
32. C. dolichostachya
18. Leaves 1-4(-6) mm wide. Stems central. Utricles nerveless to multinerved.
31. C. breviculmis
15. Nut not mitrate.
19. Spikelets in fascicles of 3-20 from the axils of at least one of the bracts.
20. Utricles narrowly ellipsoid, 4-6'/, mm long, gradually tapering below into a stipe-like,

3/,-1 mm long base, above into a 1-3 mm long beak . . ... . . 20. C. verticillata
20. Utricles ovoid or ellipsoid, 2!/,-3'/, mm long, scarcely stipitate; beak 3/,-1 mm long.
17. C. celebica
19. Spikelets solitary (exceptionally binate) from the axils of the bracts, or crowded at the apex
of the stem.

21. Utricles (8—)12-13 mm, many-nerved. 2 Spikelets broadly ovoid to subglobose, 15-25 mm
long and wide, with few, divergent, narrowly lanceolate utricles. Bracts long-sheathing
48. C. michauxiana

1979) Cy PERACEAE—II (Kern & Nooteboom) 113

21. Utricles at most 7'/, mm long. Other characters not so combined.
22. Broader leaves 1—2 cm wide. Spikelets 5!/,-16 cm long. Glumes vinaceous to dark red
with a green, central stripe. Stoloniferous . . . . . . 45. C. olivacea
22. Broader leaves at most 10 mm wide. Spikelets usually much shorter
23. Stems lateral. Lateral spikelets 6-12 by 4-7 mm, or androgynous. Utricles at least
below many-nerved, 5-7 mm long. Glumes 2!/,-4 mm. Nut 3-4 mm.
24. Leaves 3-7 mm wide. Glumes ae mm, with a 1'/,-6 mm long awn. Lateral

spikeletssGy 3 ses eld Be eae, Atenas
24. Leaves pe 3 mm wide. Glumes Cc 4 mm Tong, muticous or mucronulate. Lateral
spikelets androgynous . . . A spear Bi7 eek Oral (1 1

23. Stems central. Lateral poiteleion more fhe) 10 mm fone! ee or realy androgynous, when

shorter than 12 mm nut 2!/,-3 mm.

25. Utricles 5—7'/, mm long; nut at least 2'/, mm long.

26. Glumes 2—2!/, mm, with an antrorsely scabrous, 2-4 mm long awn. Utricles with
deeply furcate 1'/,-2 mm long beak, strongly and densely many-nerved, when ripe
widely spreading or reflexed. Lower bracts much overtopping the inflorescence,
scarcely sheathing. . . . .... . . 49. C. pseudocyperus

26. Glumes 3-6 mm long, whether c or not awned.

27. Utricles nerveless or obscurely few-nerved, except for the 2 marginal veins, 5—71/, mm
long. 2 Spikelets linear-cylindric, lax-flowered, 2-9 cm by 3-7 mm. Glumes 3?/,—
6 mm, oblong to ovate, acute to very obtuse, translucent, with broad white margins
and greenish midrib, the latter not reaching the apex, but sometimes excurrent into
ashortiawniup to 1@2)imm long = > 3098 see ee 47 Go initia
27. Utricles (strongly) many-nerved.

28. Utricles 5'/,—7!/, mm, fusiform. Glumes oblong-ovate, acute, very thin, dirty white
with 3-nerved greenish central stripe, 3-5!/, mm long, muticous or mucronulate,
rarely witha shortawn . . . sce 41235°C. jackiana

28. Utricles 5—S!/, mm. Glumes ovate, deeply emarginate at the top, pale with purplish
margins, c. 3 mm long, the strong midrib excurrent into a firm, smooth or scabrid
awn as longasthe glume. . . . .. . . . 60. C. sp. (§ Paludosae)

25. Utricles 3—-5(—7) mm long; nut at most 21), mm iene but mostly shorter; if the utricles
are longer than 5 mm, the nut is at most 1?/,; mm long.

29. Utricles sparsely to rather densely hispidulous, nerveless except for 2 submarginal
nerves, 3—4!/, mm long, with stout, 1-1!/, mm long beak. 2 Spikelets linear-cylindric,
1!/,-6 cm by 2!/,-3!/, mm. Glumes 3—4!/, mm, with an hispidulous awn up to | mm
long .. .. ss. . . . 46. C. brachyathera

29. Utricles glabrous, many- -nerved | or ‘-tibbed.

30. Utricles 4-5(-7) mm long, strongly many-nerved. Glumes 2-2!/, mm, with a 2-4 mm
long awn. 2 Spikelets 2!/,-5 cm by 7-15 mm. Leaves 6-10 mm wide.
49. C. pseudocyperus
30. Utricles 3-4 mm (up to 6 mm in C. brownii ssp. transversa), strongly many-ribbed
or plurinerved. Glumes 11/,-2!/, mm, excurrent into a 1/,-3'!/, mm long awn.
2 Spikelets 1!/,-12 cm by 4-6 mm. Leaves 2-10 mm wide.

31. Utricles strongly many-ribbed. 9 Spikelets short-cylindric to cylindric, 11/,-3 cm
by 5-6 mm. Glumes 1-2 mm with a !/,-3'/, mm long awn. Nut 2!/,-2'/, mm. Leaves
3-4mmwide .. . . . 43. C. brownii

31. Utricles plurinerved. 9 Spikelets (narrowly) cylindric, ( Eyes psy 12 cm by 4-6 mm.
Nut 1!/,-2!/, mm. Leaves 2-10 mm wide.

32. Plant densely cespitose, without stolons. Lower bracts long sheathing. Utricles

olive-brown to fuscous. 2 Spikelets 2!/,-12 cm long .. 44. C. oedorrhampha
32. Plant stoloniferous. Bracts not sheathing. Utricies pale greenish to stramineous.
9 Spikelets (11/.-)2-S5 cm ............. . .42. C. alopecuroides

8. Stigmas 2.
33. Utricles densely covered with raised glandular papillae, slenderly nerved or nerveless. At least the
lower spikelets subcernuous to pendulous. Lowest bract much overtopping the inflorescence.
34. Glumes acute to obtusish, muticous or only the lower ones excurrent in a short, up to 1 mm long

awn. Utricles slenderly nerved, 3-4 mm long . . ... . . 53. C. pruinosa
34. Glumes truncate or bilobed-emarginate, distinctly awned (awn 55 mm long). Utricles mostly
nerveless, 21/,-3!/, mm. . . ee CO era eS2i. Ce phacota

33. Utricles not papillate, whether minutely punctulate o or not.

114 FLORA MALESIANA [ser. I, vol. 9#

35. Stems 3-40(-75) cm by 1-1'/, mm. Utricles distinctly 8—14-nerved, 2-31/, mm. Spikelets erect
or suberect, terminal 3, remainder 2 or with a few 3 flowers at the top, 1-6 cm. Leaves
11/4 mmwide..... .... . . 55. C. gaudichaudiana

35. Stems 30-110 cm by up to A mm elo: Spikelets poddines all lateral ones androgynous or
1(-2) at the base of the uppermost much smaller and wholly 3, (3—)6-13 cm long. Leaves
3-12) mimbwide-wane lee) - 1 Sy Seat, oe, Were ce ken @ woractleania

7. Terminal spikelet always Gisextall

36. Terminal spikelet (often all the spikelets) gynaecandrous (i.e. 2 flowers above, 3 ones below);
3 flowers often so few that the spikelets have an entirely 2 appearance.
37. Stigmas 2. Nut lenticular (biconvex or planoconvex).

38. Spikelets all sessile, 1/.-11/. cm long.

39. Utricles distinctly winged on the margins. Lower bracts foliaceous, long, far we the up

to 15 cm long inflorescence . . Dae . C. remota

39. Utricles wingless. Bracts glumiform, the lowest exceptionally subulate to ccheee some-
times exceeding the 1!/,-3'/,(-6) cm long inflorescence.

40. Utricles (3-)4~-5(—5!/,) mm long, when mature widely patent to reflexed, distinctly beaked,

not whitish-punctulate . . . . . 64. C. echinata

40. Utricles 2-2!/, mm long, suberect also when mature, ‘seareely ‘beaked, densely whitish-
punctulateze. oe: ales: 9 OU ieee OOo Ce CORES

38. At least the lowest poikeley pedunelede cernuous, 3 6(-8 ) cm foney «aod pf oe SAGs tenes

37. Stigmas 3. Nut trigonous.

41. Utricles hispidulous or sparsely pubescent. Nut mitrate (i.e. contracted at the apex and then

expanded into a discoid annulus).
42. Utricles many-nerved. Glumes of 3 flowers infundibuliform ... . . . 34. C. tristachya
42. Utricles obscurely nerved or nerveless. Glumes of 3 flowers with free margins
31. C. breviculmis
41. Utricles glabrous. Nut not mitrate.
43. Lower bracts sheathing. Plants cespitose.
44. Utricles broadly ellipsoid to subglobose, 3-4 mm long or ovoid-ellipsoid, 6 mm long
43. C. brownii
44. Utricles fusiform or fusiform- ce 5-7'/, mm long.
45. Utricles strongly many-nerved . . . .. 23. C. jackiana
45. Utricles nerveless except 2 marginal 1 nerves or “obscurely few- nerved . . . 47. C. finitima
43. Bracts not sheathing. Plants stoloniferous . . . . . . . 42. C. alopecuroides
36. Terminal spikelet and all or most of the other spikelets androgynous (i.e. 3 flowers above, 2 ones
below).
46. Stigmas 3.

47. Leaves pseudopetiolate, 1'/,-3!/, cm wide, the base conduplicate. Stems central, erect, scapiform,
the base surrounded by some brown, infundibuliform, up to 10 cm long leaf sheaths. Utricles
very long beaked, the beak curved, bulbous at the base (to hold the thickened conical style-base),
nearly linear towards the very oblique mouth, the whole beak 3-3'/, mm long, the utricle
including the beak 6-7 mm long. Inflorescence paniculate with head-like very dense partial
inflorescences . . . / ss « » 140 Cl helfers

47. Leaves not pseudopetiolate (except i in C. corecray Other characters not so combined.

48. Spikelets peduncled, in fascicles from at least one of the sheathing bracts, sometimes com-

pound.
49. Utricles densely many-nerved, 5—91/, by 1-21/; mm. 5 3-8 mm thick. Nut 3-4 mm
long. . . . . 18. C. perakensis
49. Utricles menvcless or r slenderly fogs neared! ce G by 3/ aay mm. Spikelets 2-31/, mm thick. Nut
FSi 5 italy § 5 « ooo sy) pee « 19S CS torrta

48. Inflorescence racemose or SHEXE at eeikelers aeierdlet « or sesgiie single or binate at the
nodes or inflorescence a panicle and the secondary panicles single or binate at the nodes.
Bracts sheathing or not.

50. Flowering stems arising from the axils of the basal leaves, which are crowded on a short stem
and much longer than the flowering stems. Inflorescence racemiform or paniculiform;
spikelets 6-30, 8-30 mm long. Bracts usually shorter than the spikelets, with funnel-shaped
sheaths and short blade. Nut irregularly rhomboid-ellipsoid or rhomboid-obovoid with
faces excavated at base and apex; style flexuous . . .. . . 26. C. eryptostachys

50. Flowering stems not arising from the axils of the basal leaves.

1979} Cy PERACEAE—II (Kern & Nooteboom) 115

51. Flowering stems developing from lateral shoots. Inflorescence a narrow, simple panicle,
consisting of 6-12 head-like or racemose partial inflorescences each containing 1-8 spikelets
4-10 mm long. Utricles strongly many-nerved, 2'/,4 mm long. . . . 21. C. oligostachya

51. Flowering stems central.

52. Nut with a stout, cylindric, '/.-*/, mm wide neck. Style centred in the apical hollow of the

nut.
53. Inflorescence spiciform, with 2-10 sessile 2 garni Utricles rhomboid-lageniform,
4'/,-5mmlong ... . . . . 29, C. palawanensis
53. Inflorescence a slender compound panicle with numerous spikelets. Utricles ellipsoid-
rhomboid,c.4 mmlong . . . . 1. C. cirrhulosa

§2. Nut without stout cylindric neck. Style not centred i in an ‘apical hollow.
54. Lower bracts not or only very shortly sheathing.

55. Inflorescence with 12 to numerous spikelets, spiciform, the lateral spikelets arising from
a utriculiform, gibbous prophyll as ia a 2 flower or a nut. Stems 3—20(-—30) cm
talbe e 2 . . . 12. C. satzumensis

55. Inflorescence with 1-4 spikelets, or 1-4 heads, each consisting of 1-4 crowded sessile
spikelets. The lateral spikelets not arising from a fertile prophyll.

56. Utricles densely hispidulous in the upper 2/;. Glumes densely setulose. Lateral spikelets
of the inflorescence sessile, ovoid or subglobose (sometimes absent) 38. C. tricephala

56. Utricles at most scabrous at the margins towards the apex, the rest glabrous.
57. Utricles 6-8 mm long, with a c. 3 mm be: beak. Glumes 3-4'/, mm long, with a

1-17/,; mm longawn . . . . 36. C. ramosii
57. Utricles 4-6!/, mm long, with a a 12 mm 1 long beak. Glumes ‘21,3 mm long, with a
/=imor lonmiawn: <2) 3s oe ee ee ee Rees 8352 'Comalaccensis

54. Lower bracts long-sheathing.
58. The longer spikelets at least 20 mm.
59. Spikelets many. Utricles 3-4'/, mm. Nut 2-3 mm.

60. Beak of utricles straight. Utricles not inflated, ellipsoid or ellipsoid-obovoid, trigo-
nous, pale to castaneous, slenderly nerved, sparsely to subdensely hispidulous, at least
towards the apex. Leaves 2-10 mm broad ... a. =,” 16. myosurus

60. Beak of utricles curved. Utricles inflated, obscurely trigonous, ovoid to subglobose,
strongly nerved, glabrous except the hispidulous margins towards the apex, at first
yellowish green, ultimately red and more or less succulent. Leaves 5-18 mm broad

15. C. baccans
59. Spikelets 1-many. Utricles 4-9!/, mm (if utricles less than 5 mm, spikelets few). Nut
3-4 mm.

61. Spikelets 2-many, branched (or when not branched the beak of the utricle 2-3 mm).

Utricles 5—9!/, mm long, the beak 1'/,-3 mm. Style-base slightly thickened
18. C. perakensis
61. Spikelets not branched, 1—3(-4), 5—20 cm distant. Utricles 4-7 mm, the beak shorter.

Style-base pyramidically thickened. . . . . : a. & <2 26375 Gy speciosa
58. Spikelets never becoming longer than 20 mm.
62. Leaves inserted all over the stem.. . . ...... . 13. C. hypolytroides

62. Leaves (sub)basal, often 1-3 higher on the a.
63. Utricles densely pale to golden hispidulous, 5—-6(—8) mm long. Spikelets at least 8 mm
NON Oe re ee er suc vial gare Zoe cstNies, Cree e = ocn€Gn MEPAKEMSIS
63. Utricles different, 21/,-6 mm long.
64. Nut narrowly discoid-annulate at the apex. Utricles whitish setulose, not inflated.
Secondary panicles mostly spiciform ........ . . . 8. C. nodiflora
64. Nut not narrowly discoid-annulate at the apex, or utricles glabrous, inflated.

65. Leaves often pseudopetiolate, the broader leaves more than 15 mm wide. Spikelets
+ patent, rather few-flowered, 5-10 mm long. Glumes ovate or oblong-ovate,
obtuse or slightly emarginate, 2-3 mm long, the awn 1/,-2 mm. Utricles distinctly
trigonous, ellipsoid, with prominent angles and flattish faces, membranous, c. 5-
nerved on each face, 4-5(-6, extra-Mal.) by 11/,-13/, mm, subabruptly beaked, the
beak 1!/,-2(-3, extra-Mal.) mm long, straight, bidentulate on the ventral side with
very ODHquc mouth eae ey ee ee ee ne ee 2. C. commixta

65. Leaves not pseudopetiolate, rarely more than 15 mm wide.

66. Utricles glabrous, whether inflated or not, shiny and conspicuously 24-nerved on
each face and then glumes muticous or mucronulate, or dull, 3-5-spongy-ribbed or
strongly 5—10-nerved on each face, with oblique, often scabrid beak.

116 FLORA MALESIANA [ser. I, vol. 9!

67. Utricles shiny, triquetrous, not inflated, conspicuously few-(c. 2-4-)nerved on each
face, olivaceous to reddish-castaneous, 2!/,—5 by !/,-1 mm, with slender, subulate
or scarcely tapering beak which is about as long as the body or slightly longer
(1-2!/, mm long). Glumes 1!/,—2!/,(-4!/,) mm, about as long as the body of the
utricle, muticous or rarely mucronulate .. . eine’ (4C Anica

67. Utricles rather dull, inflated or subinflated, or at east not triquetrous, strongly
spongy 3—5-ribbed on each face or strongly 5—10-nerved. (if different proceed
under 66 second lead.)

68. Utricles spongy 3-—5-ribbed on each face, stramineous or brown, 2-4 by
1(-1'/,) mm, abruptly beaked, the beak aged canan with oblique mouth

often becoming bifid, 3/,-11/, mmlong. . . . be, Sei yomentsa
68. Utricles strongly many-nerved (5—7 or c. 10 nerves on each face), 3'/,-5 by
1—2}/; mm.

69. Utricles distinctly inflated, obscurely trigonous, c. 10 nerves on each face. The
rather compact ¢ part of the spikelets as long as to usually much longer than
the 2 part, rarely shorter. Glumes aimee mm, with an awn to 3!/, mm. Nut
often rostrate .... . s+ fap, Os Gaindica

69. Utricles rhomboid, fconoust 5a 7 nerves on pact fice Nut erostrate.

70. The (often plumose) 3 part of the spikelets shorter than to about as long as

the 2 part. Glumes 1!/,-2 mm, with an awn of 1-3 mm. . 5. C. horsfieldii
70. The 3 part longer than the 2 part of the spikelets. Glumes 2—23/, mm, with an
awn of 3/,-2mm.. . ... . . 10. C. stramentitia

66. Utricles glabrous, or hairy, noe cadaeed seen to conspicuously nerved or
spongy-ribbed, the beak bidentate or bidentulate.

71. Glumes distinctly awned, broadly ovate. At least in the lowest glumes of a large
part of the spikelets the awn longer than the glume. Utricles with (3—)4—6 strong
nerves on each face, glabrous and smooth (glossy), or hispidulous on the margins
in the upper part, light green to stramineous, (2!/,-)3-4 by 1-11/, mm, the beak
compressed, shorter to about as long as the body of the utricle, 1-11/, mm long.
(If the utricles are hispidulous, see 3. C. cruciata.) . . . 7. C. lamprochlamys

71. Glumes whether awned or not, awn never longer than glume.

72. Glumes muticous or mucronulate, rarely awned. Utricles obscurely nerved, only
2 nerves more prominent, glabrous at the base, otherwise densely scabrid-
pubescent, blackish fuscous or greenish, 23/,—4"/, by c. 1 mm; the beak straight,
compressed, deeply bidentate, but often originally oblique, blackish fuscous,
the mouth with whitish hyaline margins . . . . . .. 9. C. sarawaketensis

72. Glumes often awned. Beak of utricles not blackish fuscous, or utricles not
scabrid-pubescent except at the base, or characters otherwise combined.

73. Glumes ovate or broadly ovate, acutish or obtuse. Utricles obtusely trigonous,
strongly spongy 3—5-ribbed on each face, glabrous, except sometimes on the
margins ... 2s 4 Sh Cxemciata

73. Glumes ovate or nerowia orate aeate! ltrcles distinctly trigonous or
triquetrous, more or less distinctly several-nerved to obscurely nerved, glabrous
or hairy.

74. The 3 and @ part of the spikelets of same length. Glumes 1'/,-21/, mm
3. C. cruciata
74. In the longer spikelets the g ie much ES than the few-flowered 9 part.
Glumes 21/.-41/, mm .... . + « + « » M0 (Cavesicalosa

46. Stigmas 2.
75. Rhizome long-creeping, slender. Leaves canaliculate or convolute, c. 1 mm wide. Spikelets
approximate, forming an oblong, 1—1'/, cm long head. Utricles nerveless, glabrous
61. C. duriuscula
75. Rhizome short, plant forming dense tufts. Leaves usually wider than 1 mm.

76. Spikelets very numerous, forming a slender, 5—25 cm long, oblong-cylindric, contracted, spike-
like 1-2 cm broad panicle. Leaves with very scabrous margins, 3-10 mm wide. Utricles coria-
ceous . . sos) o 62 G@aporessa

76. Spikelets 4-10(-25), forming a “head- like or ‘spike-like 1 -3(-5) cm long inflorescence, or
spikelets up to 50, in a lax raceme or in (2-)4-8 fascicles of 2-7 spikelets each and inflorescence
up to 50 cm. Utricles membranous.

1979] Cy PERACEAE—II (Kern & Nooteboom)

117

77. Spikelets all sessile. Cladoprophylls (see below) absent. Utricles ovate or ovate-lanceolate,
not cordate at the base, winged on the margin, 3'/,-4'/, mm long. Glumes mucronate
63. C. nubigena
77. At least the lower spikelets distinctly peduncled. Peduncles of the lateral spikelets at the base
surrounded by an ocreiform or utriculiform bracteole (cladoprophyllum) which is often
hidden in the sheath of the bract subtending the spikelet.

78. Bracts not sheathing. Spikelets linear-cylindric, very densely flowered, (2—)6-13 cm long.
79. Utricles densely beset with ferrugineous papillae. . . a . 52. C. phacota
79. Utricles epapillate . . . 51. C. graeffeana

78. Bracts sheathing. Spikelets loosely or somewhat densely flowered, up to 4'/, cm long.

80. Inflorescence very lax, only 1 spikelet at each node. Glumes with a stoutish, up to 5 mm
long awn. Utricles glabrous except for the sparsely hispid beak (rarely the margin setulose)
58. C. longipes
80. Spikelets often in fascicles at the nodes. Glumes muticous or minutely apiculate (rarely an
awn up to 2 mm present). Utricles setulose at least on the margins.
81. Lower bract setaceous, suddenly widening into a spathaceous, reddish brown base
clasping the base of the spikelets. Glumes 4-6 mm 56. C. bilateralis
81. Lower bracts foliaceous.
82. Glumes distinctly shorter than to about as long as the utricle, 2-4 mm, rarely some of
them to 5 mm. Stigmas shorter than to about as long as the utricles, up to c. 5 mm
57. C. brunnea
82. Glumes about as long as the utricles, 31/,-5 mm. Stigmas very long (7-12 mm), always
much longer than the utricles . . 59. C. teinogyna

I. Subgenus Carex

Carex subg. Indocarex BAILL. Hist. Pl. 12 (1893) 345. — Primocarex KUK. Verh.
Bot. Ver. Brandenb. 47 (1905) 204. — Carex subg. Primocarex KUK. Pfl. R. Heft 38
(1909) 68.

Type species: Carex acuta L.

1. Section Vigneastra

TUCKERM. En. Meth. (1843) 10. — Sect. Vigneastra [grex| Indicae [TUCKERM. I.c.,
nomen]; ex BAILEY, Proc. Am. Ac. 22 (1886) 98. — Sect. Indicae CLARKE, FI. Br.
Ind. 6 (1894) 713; Kuk. Pfl. R. Heft 38 (1909) 260; RAym. Mém. Jard. Bot.
Montréal n. 53 (1959) 20, 38; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 491, in
nota. — Sect. Polystacheae CLARKE [ser.] Stramentitiae, Cruciatae & Filicinae
CLARKE, J. Linn. Soc. Bot. 37 (1904) 4. — Sect. Stramentitiae (CLARKE) NELMES,
Reinwardtia | (1951) 250, p.p. — Sect. Cruciatae (CLARKE) NELMES, /.c. 275. —
Sect. Filicinae (CLARKE) NELMES, /.c. 286; RAYM. Mém. Jard. Bot. Montréal n. 53
(1959) 20, 48. — Sect. Indicae subsect. Indicae KOYAMA, J. Fac. Sc. Un. Tokyo III,
8 (1962) 151.
Type species: Carex indica L. (lectotype).

1. Carex cirrhulosa Nees in Hook. J. Bot. Kew
Misc. 6 (1854) 29; NeLMES, Reinwardtia 1 (1951)
262. — ? C. densiflora PREsL, Rel. Haenk. 1 (1828)
204; F.-ViLL. Nov. App. (1882) 310. — C. fuire-
noides (non GAUDICH.?) F.-VILL. Nov. App. (1882)
310; CLarKE, J. Linn. Soc. Bot. 37 (1904) 11,
p.p.; Philip. J. Sc. 2 (1907) Bot. 107, p.p. —
— C. fibrata Boott ex VIDAL, Phan. Cuming.
(1885) 156; Rev. Pl. Vasc. Filip. (1886) 286, nomen.

— C. fuirenoides GaupicH. var. cirrhulosa KUK.
Pfil. R. Heft 38 (1909) 287; Philip. J. Sc. 6 (1911)
Bot. 61; Merr. En. Philip. 1 (1923) 138, p.p. —
Fig. 118.

Inflorescence a slender, compound panicle,
continuous above, interrupted below, c. 18 cm long;
secondary panicles 6, single at the nodes, erect,
oblong, rather dense, 11/,-31/, cm long, their lower
branches again branched into several sessile, patent,

118

FLORA MALESIANA

[ser. I, vol. 9!

crowded spikelets on trigonous, smooth or sparsely
scabrid peduncles; rachis sparsely scabrid above.
Lower bracts foliaceous, much exceeding the in-
florescence, stiff, flat or with revolute margins,
long-attenuate, long-sheathing, 5-8 mm wide,
upper ones much reduced, shortly sheathing.
Spikelets numerous, androgynous, oblong, ovoid,
or subglobose, dense, 4-5 mm long, their 3 part
about as long as the 9, their bracteoles glumiform,
with hispid, often curved awns, c. 10 mm long.
Glumes ovate or suborbicular, translucent, erose-
ciliolate at the apex, otherwise glabrous, 3/,-1 mm
long, the midnerve excurrent in a stout, flat,
sparsely hispid, straight or slightly curved,
11/,-21/, mm long awn. Urtricles trigonous, ellipsoid-
rhomboid, subcoriaceous, patent, strongly many-
nerved, glabrous, smooth or very sparsely scabrid
at the apex, usually slightly curved, somewhat
spongy-thickened at the base, subabruptly
narrowed into the beak, pale stramineous to
brown, 4 by 1-1!/, mm; beak scarcely tapering,
compressed, sparsely scabrid, straight or slightly
curved, bidenticulate, with straight mouth, 11/, mm
long. Nut triquetrous, with prominent angles and
concave faces, ellipsoid-rhomboid, stipitate, the
apex narrowed into a short, thick neck expanding
into a discoid ring c. '/, mm diam., 2!/, by 1-
11/, mm. Style-base broadly pyramidal, persistent
on the nut, centred in the apical hollow of the nut.
Stigmas 3.

Distr. Malesia: Philippines (Cebu).

Note. Only known from the type collection
(CUMING 1764, collected in 1841), which is re-
presented in BM and K by very defect specimens
(rhizome, leaves and parts of the inflorescence
missing), so that it is difficult to ascertain its status.
CLARKE considered it synonymous with C. fuire-
noides GAuDICH. from the Marianas, which is
C. indica L. NELMES supposed affinity with
C. cryptostachys BRONGN. and the species of sect.
Lageniformes (OHWi) NELMES, wrongly as I think.

2. Carex commixta STEUD. [in Zoll. Syst. Verz. 1
(1854) 60, nomen] Syn. 2 (1855) 207; Mia. FI. Ind.
Bat. 3 (1856) 349; Kern, Blumea 15 (1967) 427,
f. 1; in Back. & Bakh. f. Fl. Java 3 (1968) 491. —
C. horsfieldii (non Bootr) Mia. Fl. Ind. Bat. 3
(1856) 349, p.p: (quoad specim. Jungh.); Kix. Pfi.
R. Heft 38 (1909) 273; Back. Bekn. Fl. Java
(em. ed.) 10 (1949) fam. 246, p. 67. — C. spatiosa
Bootrt, Ill. 2 (1860) 86, t. 246; Boeck. Linnaea 40
(1876) 349; CLarKE, J. Linn. Soc. Bot. 37 (1904)
12, incl. var. bogorensis CLARKE; KUK. Pfl. R. Heft
38 (1909) 265; Camus, FI. Gén. I.-C. 7 (1912) 188;
NELMES, Kew Bull. (1946) 21, 23; Mém. Mus. Hist.
Nat. Paris n.s. B4 (1955) 111; RAym. Mém. Jard.
Bot. Montréal n. 53 (1959) 40. — C. blepharolepis
NELMES, Kew Bull. (1946) 18, 23; Reinwardtia 1
(1951) 265. — C. smitinandii RAYM. Dansk Bot.
Ark. 23 (1965) 255, f. 1. — Fig. 118.

Rhizome shortly creeping, stout, woody. Stems

loosely tufted, triquetrous, smooth, 40-100 cm by
2-3 mm, the base clothed with membranous, pale
or fuscous bladeless sheaths or their fibrous
remains. Leaves herbaceous, subbasal and a few
higher on the stem, equalling to much longer than
the stems, linear-lanceolate, long-attenuate, taper-
ing below (often into a pseudo-petiole), flat, light
green, scabrid on the margins, 1'/,-3 cm wide.
Inflorescence a pale, compound, erect, much inter-
rupted, 15-40 cm long panicle; secondary panicles
2-6, single at the nodes, erect, pyramidal, loose,
3-7'/, by 1'/,-5 cm, upper continuous, lower
(when more than 2) distant, on exserted, hispidu-
lous peduncles; rachis densely whitish hispid.
Lower bracts foliaceous, exceeding the inflores-
cence, long-sheathing, upper reduced. Spikelets
androgynous, subsessile, patulous, rather few-
flowered, 5-10 mm long, 3 and ? parts about equal
in length. Glumes ovate or oblong-ovate, thinly
membranous, obtuse or slightly emarginate,
slenderly nerved, ciliolate at least at the top, other-
wise glabrous or minutely adpressed-hispidulous,
pale stramineous to light brown, 2-3 mm long, the
midnerve excurrent in an antrorsely scabrid,
1/,-2 mm long awn. Utrricles distinctly trigonous,
ellipsoid with prominent angles and flattish faces,
membranous, not inflated, patulous, many-nerved
(nerves c. 5 on each face), glabrous or very sparsely
hispidulous, rounded at the base, straight or
slightly recurved, subabruptly beaked, 4—5(-6) by
1!/,-13/, mm; beak sparsely scabrid on the margins,
or smooth, 1!/,-2(—3) mm long, bidenticulate on the
ventral side, with very oblique mouth. Nut trique-
trous, ellipsoid-rhomboid, not cuneate at the base,
erostrate, dark brown with yellowish angles, 21/,—
23/, by 1'/,-14/,; mm. Style-base pyramidally
thickened, subpersistent on the nut. Stigmas 3.

Distr. Burma, N. Thailand, Tonkin, Annam,
Hainan; in Malesia: Sumatra (Bencoolen, Lam-
pongs), West and Central Java.

Ecol. Moist places in primary and, less fre-
quently, secondary forests, 800-1500 m, along
water-courses sometimes descending to 300 m.

Vern. Java: ilat harashas, S.

Note. The type of C. spatiosa is from Annam
(GAUDICHAUD 67). According to CLARKE, l.c.,
the Malesian plants should differ by the ovoid
utricles with a scabrid beak '/,-*/, as long as the
body, and according to Nemes (1946, /.c.) by the
shorter, glabrous glumes and the shorter utricles.
On the whole the Indochinese specimens have
longer utricles (up to 6 mm) because of the
slenderer beak, but some of them are indistinguish-
able from the Malesian plants, which were
described as C. spatiosa var. bogorensis CLARKE and
C. blepharolepis NELMES.

3. Carex cruciata WAHLENB. Vet. Akad. Handl.
Stockh. 24 (1803) 149; Crarke, FI. Br. Ind. 6
(1894) 715; J. Linn. Soc. Bot. 37 (1904) 9; RIDL.
Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 118; KUK.

1979] Cy PERACEAE—II (Kern & Nooteboom) 119

E VIJSMA ° 78

Fig. 118. Utricles and nuts of Carex. Species numbered as in the text (J CumINnG 1764, 2 Jacoss 8046,

3a CHEW, CORNER & STAINTON 1706, 3b (left) VERHEIWJEN 2585, 3b (right) VAN OosTsTROOM 13154,

4 HarTLeY 11122, 5 NGF 21261, 6 Ripiey 15721, 7 BRAss 24684, 8 B. F. HERNAY 652, 9 PULLEN 6106,
10 BAKHUIZEN VAN DEN BRINK 6565, 7 VAN STEENIS 4140). All x 10.

120

FLORA MALESIANA

[ser. I, vol. 9!

Pfil. R. Heft 38 (1909) 265; Camus, Fl. Gén. I.-C. 7
(1912) 189, f. 27, 5-9; RipL. Fl. Mal. Pen. 5 (1925)
185; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 11
(1936) 460; KUx. Bull. Jard. Bot. Btzg III, 16
(1940) 315; Netmes, Reinwardtia 1 (1951) 277;
Mem. Mus. Hist. Nat. Paris n.s. B4 (1955) 129;
RaymM. Mém. Jard. Bot. Montréal n. 53 (1959) 46;
KoyaMaA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 153;
RAyYM. Dansk Bot. Ark. 23 (1965) 254; KERN in
Back. & Bakh. f. Fl. Java 3 (1968) 492. — Fig. 118,
124a-d.

Further synonyms under the varieties.

Rhizome short or shortly creeping, stout, woody.
Stems tufted, stout, trigonous, smooth, up to
200 cm by 2-5 mm, surrounded below the leaves by
bladeless, dark reddish to blackish sheaths and
their fibrous remains. Leaves (sub)coriaceous,
mostly basal but 1-3 higher on the stem, much
exceeding to shorter than the stems, linear, long-
attenuate, flat or with revolute margins, scabrid on
the margins and the nerves, (3—)5-14 mm wide.
Inflorescence a slender, usually much elongated,
interrupted, decompound, ferrugineous or strami-
neous panicle up to 70 cm long; secondary panicles
3-12, at 3-8 nodes, all or the middle ones binate at
the nodes or all single, erect or suberect, oblong-
linear or (broadly) oblong-lanceolate or ovoid-
subpyramidal, loose to dense, 5-10 cm long, the
upper approximate, the lower distant on unequal
scabrid or smooth peduncles; rachis scabrid-hispid
or hispidulous on the angles. Lower bracts folia-
ceous, equalling or exceeding the inflorescence,
long-sheathing, upper much reduced; bracteoles
excurrent in a slender, ciliolate-scabrid, often
recurved awn. Spikelets (very) numerous, andro-
gynous, obliquely patent or divaricate, 4-8
(-10) mm long, ovoid to oblong, the ¢ part as
long as or somewhat longer than the @ part.
Glumes broadly ovate, ovate or ovate-lanceolate,
acute to very obtuse, membranous, (sub)translu-
cent, glabrous or sparsely hispidulous, with reddish
brown to dark brown streaks, 1'/,-3 mm long,
several-nerved, the midnerve of the lower ones
acute or excurrent in a smooth or hispidulous, up
to 1 mm long awn. Utricles distinctly or obtusely
trigonous, not inflated or subinfiated, (broadly)
ellipsoid to ovoid, membranous or subcoriaceous,
patulous or patent, straight or slightly recurved,
often finally recurved, more or less distinctly
several-(spongy-)nerved, glabrous or _ sparsely
hispidulous, (sub)abruptly beaked, stramineous
or brown with reddish brown streaks and spots,
2-4 by */,-1(-1/,) mm; beak slender, smooth or
scabrid on the margins, (3/,—)1—-1!/, mm long, with
oblique but soon bidenticulate to bifid mouth. Nut
distinctly trigonous, with prominent angles and
flat or concave sides, (ovoid-)ellipsoid, not or
scarcely beaked, scarcely stipitate, stramineous to
brown or blackish, 11/,-21/, by 3/,-1 mm. Style-
base not or slightly thickened. Stigmas 3.

Distr. Widely spread, from India through SE.

Asia to S. China, Formosa and the Ryu Kyu Is.;
throughout Malesia.

Note. Several collections from Sumatra: VAN
BORSSUM WAALKES 2753, BUNNEMEIJER 2530 (type
of C. buennemeijeri NELMES), 3644, 3880, 4126,
4651, are intermediate between var. cruciata and
var. rafflesiana.

KEY TO THE VARIETIES

1. Leaves usually much exceeding the stem.
Inflorescence stramineous. Utricles strongly
spongy 3—5-ribbed on each face

a. var. cruciata

1. Leaves usually as long as the stems or shorter.
Inflorescence ferrugineous. Utricles more or
less distinctly 3-7-nerved on each face

b. var. rafflesiana

a. var. cruciata. — C. cruciata WAHLENB. Vet.
Akad. Handl. Stockh. 24 (1803) 149; Ripv. FI.
Mal. Pen. 5 (1925) 185, incl. var. condensata RIDL.
— C. bengalensis Roxs. FI. Ind. ed. Carey 3 (1832)
572; Boorrt, Ill. 2 (1860) 85, t. 240-243; BoEck.
Linnaea 40 (1876) 346, excl. varieties. — C. con-
densata NEES in Wight, Contr. (1834) 123; CLARKE,
Fl. Br. Ind. 6 (1894) 716; Boorr, Ill. 2 (1860) 86,
t. 247-248. — C. vacua BooTt ex Boeck. Linnaea
40 (1876) 343, p.p. (excl. pl. jav.). — C. valida NEES
in Wight, Contr. (1834) 123, p.p.; KUNTH, En. 2
(1837) 513, p.p. — C. continua CLARKE, FI. Br.
Ind. 6 (1894) 717. — ? C. repanda CLARKE var.
implumis CLARKE, J. Linn. Soc. Bot. 37 (1904) 9. —
C. galactolepis Ne-mMes, Kew Bull. (1946) 20;
Reinwardtia 1 (1951) 279. — C. spongocrepis
NeELMes, Mém. Mus. Hist. Nat. Paris n.s. B4 (1955)
128. — Fig. 118, 124a-b.

Rhizome shortly creeping. Stems 40-150 cm.
Leaves coriaceous, usually much exceeding the
stems. Inflorescence stramineous, continuous
above, usually interrupted below, 15-60 cm long;
secondary panicles 3-11, at 3-8 nodes, usually all
single at the nodes, sometimes binate at the middle
nodes, erect, broadly lanceolate or ovoid-sub-
pyramidal, rather dense to dense, on stiff, smooth
or scabrid peduncles long-exserted from the
sheaths, up to 10 cm long; rachis hispidulous on
the angles. Lower bracts slightly to much exceeding
the inflorescence. Spikelets very numerous, divari-
cate, rather dense, 5-8(—10) mm long, the 3 part as
long as or somewhat longer than the @ part.
Glumes ovate or broadly ovate, acutish to very
obtuse, thinly membranous, translucent, several-
nerved, glabrous, with ferrugineous to dark brown
streaks, 13/,-3 mm long. Urtricles obtusely trigo-
nous, subinflated, ovoid or broadly ellipsoid,
membranous, patent, finally recurved, strongly
spongy-ribbed (nerves 3-5 on each face), glabrous,
rarely sparsely setulose at the apex, often with
spongy-thickened base, abruptly beaked, strami-
neous or brown, purplish spotted, 21/,-3(-4) by

1979] Cy PERACEAE—II (Kern & Nooteboom) 121

Ei VIISMA ’78

2)

ern ae

s in the text (12 SANTos 5776, 13 BUNNEMEUJER

9820, 14 BACKER 22996, 15 ELBERT 62, 16 VAN OosTSTROOM 13293, 17 NGF 39555, 18a VAN STEENIS 8277,
18b BROOKE 8561, 18c VAN STEENIS 8461, 19a ANU 15532, 19b SANTOS 5785, 20 MEIJER 6127). All x 10.

Fig. 119. Utricles and nuts of Carex. Species numbered a

122

FLORA MALESIANA

[ser. I, vol. 9!

1(-1'/,) mm; beak usually straight, smooth or
sparsely scabrid on the margins, with oblique
mouth often becoming bifid, (7/,—-)1—1'/, mm long.
Nut with flat or slightly concave sides, ovoid-
ellipsoid, olive-brown to blackish, 2—2!/, by °/10-
1 mm. Style-base slightly thickened.

Distr. Widely spread, from India through SE.
Asia to S. China, Formosa, the Ryu Kyu Is., and
Queensland; in Malesia: Malay Peninsula,
Sumatra, N. Borneo.

Ecol. In savannahs, open places in mountain
forests, forest-borders, along river-banks, 1100-
1500 m.

Vern. Rija-rija.

Note. Very variable; sometimes hard or im-
possible to distinguish from var. rafflesiana,
especially in Sumatra. The type specimens of C.
galactolepis (KiNG’s coll. 106) and C. repanda
var. implumis (WRAY 1982) are too poor to be
certain of their affinity.

b. var. rafflesiana (BooTT) Noot., stat. nov. —
C. rafflesiana Boott, Trans. Linn. Soc. 20 (1846)
132; Ill. 1 (1858) 12, t. 33; CLARKE, J. Linn. Soc.
Bot. 37 (1904) 10, incl. var. tenuior CLARKE;
KUK. Pfl. R. Heft 38 (1909) 282, incl. var. macro-
thyrsa KUK. et var. scaberrima KUxK.; Philip. J. Sc.
6 (1911) Bot. 59, incl. var. scaberrima KUK. et var.
continua KUK.; MERR. En. Philip. 1 (1923) 140, incl.
var. scaberrima KUK. et var. continua KUK.; KUK.
in Hochr. Candollea 6 (1936) 430, incl. var. minor
KUx.; Nevmes, Reinwardtia 1 (1951) 290, incl.
var. macrothyrsa KUxk. et var. virgata NELMES;
ibid. 2 (1954) 376; KoyAMA, J. Fac. Sc. Un. Tokyo
III, 8 (1962) 153, incl. var.; KERN in Back. & Bakh.
f. Fl. Java 3 (1968) 492. — C. macrothyrsa Mia.
FI. Ind. Bat. 3 (1856) 351. — C. virgata Mia. l.c.,
non SOL. ex BootTt, 1853. — C. pentacarpa BOECK.
Flora 58 (1875) 265. — C. vacua Bootr ex BOECK.
Linnaea 40 (1876) 343, p.p. (pl. jav.). — C. benga-
lensis var. virgata BOECK. et var. scaberrima
Boeck. /.c. 347. — C. continua [non CLARKE, FI.
Br. Ind. 6 (1894) 717] CrLarkeE, J. Linn. Soc. Bot.
37 (1904) 11; Philip. J. Sc. 2 (1907) Bot. 107; KUK.
Pfi. R. Heft 38 (1909) 282, p.p.; NELMES, Rein-
wardtia 1 (1951) 299; ibid. 2 (1954) 376. — C. gem-
bolensis CLARKE var. timorensis CLARKE, J. Linn.
Soc. Bot. 37 (1904) 10. — C. scaberrima CLARKE,
I.c. 10; Philip. J. Sc. 2 (1907) Bot. 107; Kew Bull.
add. ser. 8 (1908) 72. — C. clarkeana KUx. Bull.
Herb. Boiss. II, 4 (1904) 52, ex descr.; Pfl. R. Heft
38 (1909) 282; RipL. Fl. Mal. Pen. 5 (1925) 183;
NELMES, Reinwardtia 1 (1951) 288. — C. pycno-
thyrsos Kx. Philip. J. Sc. 6 (1911) Bot. 60; MERR.
En. Philip. 1 (1923) 140; Nemes, Reinwardtia 1
(1951) 282; ibid. 2 (1954) 374, descr. — C. semi-
glomerata Kx. Bull. Jard. Bot. Btzg III, 16 (1940)
315; in Fedde, Rep. 53 (1944) 106; NELMEs,
Reinwardtia 1 (1951) 285; ibid. 2 (1954) 376. —
C. sarawaketensis KUx. var. brevirostris KUK. Bot.
Jahrb. 70 (1940) 464. — C. timorensis (CLARKE)

NeLMes, Kew Bull. (1946) 24; Reinwardtia 1

(1951) 242, 287. — C. spongoneura NELMES, Kew

Bull. (1946) 18; Reinwardtia 1 (1951) 281. —
C. xestogyne NELMES, Kew Bull. (1946) 16;
Reinwardtia 1 (1951) 311. — C. buennemeijeri
NeELMeEs, Kew Bull. (1950) 191; Reinwardtia 1
(1951) 283; ibid. 2 (1954) 376. — C. oblonga
NeLMes, Kew Bull. (1950) 192; Reinwardtia 1
(1951) 297. — C. ceramica NeLmeEs, Kew Bull.
(1950) 193; Reinwardtia 1 (1951) 319; ibid. 2 (1954)
376. — Fig. 118, 124c-d.

Rhizome short. Stems up to 200 cm. Leaves
subcoriaceous, mostly as long as or shorter than
the stems. Inflorescence ferrugineous, interrupted,
up to 70 cm long; secondary panicles 8-12, at
4-6 nodes, all or the middle ones binate at the
nodes, or single in depauperate specimens, erect or
suberect, oblong-linear or oblong-lanceolate,
loose or rather dense, 5-10 cm long, the lower on
unequally exserted scabrid peduncles; rachis
scabrid-hispid on the angles. Lower bracts equal-
ling or exceeding the inflorescence. Spikelets
numerous, obliquely patent, 4-7 mm long, the 3
and @ part about equal in length. Glumes ovate or
ovate-lanceolate, acute, membranous, subtrans-
lucent, glabrous or sparsely hispidulous, with
ferrugineous streaks, 1!/,-2'/, mm long. Utricles
distinctly trigonous, not inflated, ellipsoid or
ovoid-ellipsoid, membranous or subcoriaceous,
patulous, straight or slightly recurved, more or less
distinctly 3—7-nerved, glabrous or sparsely his-
pidulous, subabruptly beaked, stramineous with
reddish brown streaks and spots, 2-4 by */,-1 mm;
beak slender, scabrid on the margins, 1—-1'/, mm
long, with oblique but soon bidenticulate mouth.
Nut with concave sides, ellipsoid, stramineous to
brown, 1'/,-2 by 3/,-1 mm. Style-base not or
scarcely thickened.

Distr. Thailand, Formosa (Kotosho Is.),
Queensland; throughout Malesia.

Ecol. Primary forests, forest edges, grassy
slopes, sometimes in dry sunny places, 500-2400 m.

Vern. Java: ilateun, S; Philippines: chidak, Ig.,
ikidsan, Klg., talayid, tamalang, Bag.

4. Carex filicina NEES in Wight, Contr. (1834) 123;
KuntH, En. 2 (1837) 510; Boortr, Ill. 3 (1862) 105,
t. 311-318; Boeck. Linnaea 40 (1876) 352; O. K.
Rey. Gen. Pl. 2 (1891) 748, incl. var. laevis O. K.
et var. ciliata O. K.; CLARKE, Fl. Br. Ind. 6 (1894)
717; J. Linn. Soc. Bot. 37 (1904) 11; KUx. Pfi.
R. Heft 38 (1909) 274, incl. var. saturata (CLARKE)
Kwux.; Camus, Fl. Gén. I.-C. 7 (1912) 191; MERR.
En. Philip. 1 (1923) 137; NeLMeEs, Reinwardtia 1
(1951) 304, incl. var. angustifolia NELMES et var.
zipelii NELMES; ibid. 2 (1954) 376; AKIYAMA, Car.
Far East Reg. Asia (1955) 138, t. 124; RayM.
Mém. Jard. Bot. Montréal n. 53 (1959) 48;
Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 152.
— C. nilagirica SteuD. Syn. 2 (1855) 207.) —
C. neoguineensis CLARKE, J. Linn. Soc. Bot. 37

1979] CYPERACEAE—II (Kern & Nooteboom) 123

E VIJSMA ’7&

Fig. 120. Utricles and nuts of Carex. Species numbered as in the text (2/ VANOVERBERGH 323, 22 DE
WILDE c.s. 13535, 23 VAN STEENIS 4579, 24 KAUDERN 333, 25 BS 45002, 26 MEWER 688, 27 LORZING 6678,
29 ELMER 13146, 30 MERRILL 512). All x 10.

124

FLORA MALESIANA

{ser. I, vol. 9!

(1904) 12; Ktx. Pfl. R. Heft 38 (1909) 280; S. T.
BLAKE, J. Arn. Arb. 28 (1947) 105; NELMEs,
Reinwardtia 1 (1951) 308; KERN in Back. & Bakh.
f. Fl. Java 3 (1968) 492. — C. saturata CLARKE,
J. Linn. Soc. Bot. 37 (1904) 12; Netmes, Rein-
wardtia 1 (1951) 302. — C. filicina NEEs f. saturata
Kwx. Philip. J. Sc. 6 (1911) Bot. 59. — C. sclerioides
Ripi. Trans. Linn. Soc. Bot. II, 9 (1916) 247;
NeELMES, Reinwardtia 1 (1951) 307. — C. ceylanica
BOECK. var. saturata KUx. in Hochr. Candollea
6 (1936) 431. — C. sarawaketensis KUK. var.
glabrinux Kx. Bot. Jahrb. 70 (1940) 464. — Fig.
118.

Rhizome short, stout, woody. Stems loosely
tufted, slender to stout, trigonous, smooth, up to
150 cm by 6 mm near the base, surrounded below
the leaves by bladeless, reddish or blackish sheaths.
Leaves herbaceous to subcoriaceous, mostly basal
but some spaced on the stem, as long as or shorter
than the stems, linear, long-attenuate, flat or with
revolute margins, scabrid on the margins, (2-)7-
20 mm wide. /nflorescence a slender, interrupted or
continuous, decompound, fuscous panicle up to
60 cm long; partial panicles 5-13, at 4-8 nodes,
single and (middle ones) binate, oblong-lanceolate
or pyramidal, erect, rather loose to very dense, up
to 12 by 2-5 cm, the upper approximate, the lower
usually distant on smooth or scabrid, unequally
exserted peduncles; rachis smooth towards the
base, hispidulous above. Lower bracts foliaceous,
shorter than to slightly exceeding the inflorescence,
long-sheathing, the upper much _ reduced;
bracteoles excurrent into a hispidulous awn.
Spikelets very numerous, androgynous, oblong,
rather loose, 4-15(-20) mm long, the ¢ part
shorter than to about as long as the @ part.
Glumes ovate-lanceolate, acuminate, muticous,
rarely minutely mucronulate, membranous, sub-
translucent, glabrous or the lower ones hispidulous,
finely nerved, ferrugineous, often darker streaked,
11/,-21/,(—4"/,) mm long (about as long as the body
of the utricle). Utricles triquetrous, not inflated,
ellipsoid, membranous, patent, eventually
arcuately recurved, conspicuously 2—4-nerved on
each face, glabrous, smooth, often shiny, sub-
abruptly beaked, olivaceous to reddish-castaneous,
(2/,-)3-5 by +/,-1 mm; beak slender, subulate
(scarcely tapering), smooth, or very sparsely scab-
rid on the margins, (1—)11/,-2(-2!/,) mm long
(about as long as the body or slightly longer);
mouth very oblique (the base 1/,—/, mm from the
apex). Nut trigonous, ellipsoid, beaked, closely
filling the utricle, 11/,-17/, by ‘/.-1 mm. Style-
base slightly thickened. Stigmas 3.

Distr. Ceylon and India to S. China, Formosa,
Tonkin, Laos and Annam; probably throughout
Malesia, in the Lesser Sunda Is. only known from
Lombok and in the Malay Peninsula from Pahang.

Ecol. Wet openings in primary forests, in sub-
alpine shrub vegetation, on grassy slopes, along
river-banks, 1350-3750 m.

Vern. Philippines: silak, taan, Ig., taldyig,
Bag.

Notes. In the circumscription here accepted
very variable in all its parts. Numerous segregates
have been described, but typical C. filicina from
India and the aberrant forms from New Guinea are
connected by all sorts of intermediates. Typical
C. neoguineensis is characterized by its dense, often
darker inflorescence, and the utricles more scabrid
on the beak (usually smooth or almost so in typical
C. filicina).

The high variability of C. filicina also in India is
amply discussed and excellently figured by Boortr,
Ke:

The few specimens on which C. sarawaketensis
var. glabrinux KUk. was based are more or less
transitional to 9. C. sarawaketensis.

5. Carex horsfieldii Boott, Proc. Linn. Soc. Lond.
1 (1845) 257; Ill. 1 (1858) 11, t. 32; Mia. FI. Ind.
Bat. 3 (1856) 349, p.p. (quoad specim. Horsf.);
CLARKE, J. Linn. Soc. Bot. 37 (1904) 11; NELMEs,
Reinwardtia 1 (1951) 269 (incl. var. major NELMES?);
ibid. 2 (1954) 374; RAym. Mém. Jard. Bot. Mon-
tréal n. 53 (1959) 44; KERN in Back. & Bakh. f.
Fl. Java 3 (1968) 492. — C. fleckeri NELMES, Kew
Bull. (1939) 313, fide NeLmMes 1951. — Fig. 118,
124e-¢.

Rhizome very short, stout, woody. Stems loosely
tufted, obtusely trigonous, smooth, 50-100 cm by
2-4 mm, the base clothed with fuscous, bladeless
sheaths and comose by their fibrous, fuscous to
blackish remains. Leaves herbaceous, subbasal
and a few higher on the stem, exceeding the stems,
linear, long-attenuate, flat or with revolute mar-
gins, greyish-glaucescent, scabrid on the margins in
the upper part, 5-15 mm wide. /nflorescence a pale,
decompound, erect, much interrupted, 20-40 cm
long panicle; secondary panicles 5-8, single at the
nodes, erect, oblong-ovoid, loose, distant, up to
10 cm long; peduncles smooth below, scabrid
above, the lower much exserted from the sheaths,
upper shortly; rachis hispid on the angles. Spikelets
androgynous, widely patent, finally almost squar-
rose, sessile, 5-15 mm long, the ¢ part shorter than
to about as long as the 2 part. Glumes ovate, obtuse
(the lower ones truncate to emarginate), membra-
nous, glabrous or sparsely hispidulous, not ciliate,
slenderly nerved, whitish, 11/,-2 by 1—11/, mm, the
midnerve excurrent in an antrorsely scabrid,
curved, 1-3 mm long awn. Utricles distinctly trigo-
nous, rhomboid, with prominent angles and flat
faces, membranous, not inflated, strongly many-
nerved (nerves c. 5-7 on each face), glabrous,
patent, arcuately recurved, gradually tapering to
the base, suddenly narrowed above into the beak,
olive-brown, 3!/,—4'/, by 1—1'/, mm; beak strongly
recurved, smooth or very sparsely scabrid, 1'/.-
2 mm long; mouth dorsally very oblique, not
bidentate. Nut trigonous, rhomboid-ellipsoid,
erostrate, dark brown with prominent pale angles,

1979} Cy PERACEAE—II (Kern & Nooteboom) 125

E.VIISMA ?78

Fig. 121. Utricles and nuts of Carex. Species numbered as in the text (3/a BRASS 4697, 31b EyMA 864,

31c BRASS 9032, 32 SANTOS 5394, 33 EDANO 17857, 34 ROBBINS 5.1., 35 ROBINSON 6186, 36 RAmMos 20990,

37 LORZING 16274, 38 BACKER 20462, 39 DE WILDE c.s. 13325, 40 KALKMAN 5213, 4] Brass 10255, 42
JERMY 4274). All x 10.

126

FLORA MALESIANA

[ser. I, vol. 9!

2-21/, by 1-1!/; mm. Style-base pyramidally thick-
ened at the base. Stigmas 3.

Distr. Burma, Thailand, Annam (var. annamica
RaymM.), Queensland (C. fleckeri NELMEs); in Mal-
esia: West to East Java, Moluccas, New Guinea.

Ecol. Primary and secondary forests, 100-
1100 m.

Vern. Java: irissan, J; New Guinea: homuma,
Garaina.

Notes. C. horsfieldii var. major NELMES, from
Tanimbar Is., P. Jamdena (‘“‘utricles 5.2-5.8 mm
long, straight or slightly recurved, beak 2.5—
2.8 mm long’’), was based on a young, very poor
collection.

KUKENTHAL and others misapplied the name
C. horsfieldii to the Javan plants of C. commixta
STEUD., while C. horsfieldii itself has often been
misidentified as C. indica L., from which species it
chiefly differs in the shorter ¢ part of the spikelets,
and the strongly recurved, distinctly trigonous, not
inflated, less densely nerved utricles.

6. Carex indica LINNE, Mant. 2 (1771) 574; Mia.
Fl. Ind. Bat. 3 (1856) 350; Boorr, Ill. 2 (1860) 87,
t. 250-254; Boeck. Linnaea 40 (1876) 347; CLARKE,
Fl. Br. Ind. 6 (1894) 714, incl. var. ? laetebrunnea
CLARKE et var. milnei Boott ex CLARKE; J. Linn.
Soc. Bot. 37 (1904) 8; RipL. Mat. Fl. Mal. Pen.
(Monoc.) 3 (1907) 117; Kx. Pfl. R. Heft 38 (1909)
262, f. 40, incl. var. fissilis (BOOTT) KUK., saltem
quoad specim. males.; Camus, Fl. Gén. I.-C. 7
(1912) 187; Merr. En. Philip. 1 (1923) 138; Rw.
Fl. Mal. Pen. 5 (1925) 185; NeLMes, Reinwardtia 1
(1951) 271; Raym. Mém. Jard. Bot. Montréal
n. 53 (1959) 42; Dansk Bot. Ark. 23 (1965) 253;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. —
C. fuirenoides GAUDICH. Freyc. Voy. Bot. (1826)
412. — C. moritzii StEuD. [in Zoll. Syst. Verz. 1
(1854) 60, nomen] Syn. 2 (1855) 207; Mia. FI Ind.
Bat. 3 (1856) 350; Boreck. Linnaea 40 (1876) 350.
— C. dietrichiae Boeck. Flora 58 (1875) 122;
CLARKE, J. Linn. Soc. Bot. 37 (1904) 8; S. T.
BLAKE, J. Arn. Arb. 28 (1947) 102; NeELMEs,
Reinwardtia 1 (1951) 273; Kew Bull. 2 (1955) 303.
— C. fuirenoides (non GAUDICH.?) CLARKE, J.
Linn. Soc. Bot. 37 (1904) 11; Philip. J. Sc. 2 (1907)
Bot. 107, p.p.; KUx. Pfl. R. Heft 38 (1909) 287,
p.p.; MEeERR. En. Philip. 1 (1923) 138, p.p. —
Fig. 118, 124j-k, 125.

Rhizome shortly creeping, woody. Stems loosely
tufted, triquetrous, smooth, (15-)60-100 cm by
1-3 mm, surrounded below the leaves by bladeless,
brownish to blackish sheaths and their fibrous
remains. Leaves subcoriaceous, mostly basal but
some higher on the stem, as long as or longer than
the stems, linear, long-attenuate, flat or with
slightly revolute margins, scabrid on the margins
and the nerves above, 5-15 mm wide; sheaths
blackish-nerved. Jnflorescence a decompound,
erect, interrupted panicle occupying the upper half
of the stem; secondary panicles 3-8, single at the

nodes, erect, oblong-pyramidal, rather loose to
dense, the upper approximate, the lower distant on
long-exserted peduncles, up to 10 cm long; rachis
sparsely hispidulous on the angles. Lower bracts
foliaceous, exceeding the inflorescence, long-
sheathing, their sheaths glabrous, or hispidulous
at the mouth; upper bracts much reduced; brac-
teoles with a filiform, usually long and recurved
awn. Spikelets divaricate, androgynous, rather
dense, (S—)10—20 mm jong, the ¢ part as long as to
(usually) much longer than the 2 part. Glumes
thinly membranous, ovate or ovate-lanceolate,
acute or obtuse, sometimes slightly emarginate,
nerved, glabrous, stramineous to brownish,
2-3'/, mm long, the midnerve excurrent into an
antrorsely scabrid, recurved, up to 3'/, mm long
awn. Utricles inflated, obsoletely trigonous,
broadly ellipsoid to subglobose, patent, straight or
almost so, subcoriaceous, strongly many-nerved
(nerves c. 10 on each face), glabrous, scarcely
stipitate, abruptly beaked, olive-brown, 31/,-5 by
17/,-2'/, mm; beak straight or slightly bent,
1'/,-2 mm long; mouth oblique, not bidentate.
Nut triquetrous, with very prominent angles and
concave faces, broadly ellipsoid-obovoid to sub-
globose, pyriform, or rhomboid, scarcely stipitate,
sometimes slightly discoid-annulate at the apex,
often with a curved or straight, dark brown to
blackish rostrum, 2-3 by 1!/,-2 mm. Style-base
pyramidally thickened, subpersistent on the nut.
Stigmas 3.

Distr. Widely distributed, from Ceylon and
India through Burma, Thailand, Indo-China and
S. China to Queensland, New Caledonia, Caro-
lines, Solomon and Fiji Is.; in Malesia: Malay
Peninsula, W. Java (only once collected near
Tjikoya by ZOLLINGER), Borneo, SE. & N. Celebes
(2 collections), Philippines (Luzon, Samar,
Palawan), New Guinea.

Ecol. Moist places in forests, along streams, at
low and medium altitudes, up to 1000 m.

Vern. Bundung, sesayak, M.

Notes. Very variable, especially in the length of
the awns on bracteoles and glumes. When KOYAMA,
Micronesica 1 (1964) 108-109, says that in C. indica
the glumes gradually taper to the cuspidate apex
without any conspicuous awn, in contradistinction
to the Micronesian C. fuirenoides GAUDICH. with
truncate or shallowly emarginate glumes with a
long, scabrous awn, he cannot have had true
C. indica before him.

Segregation of C. dietrichiae (= C. indica var.
laetebrunnea) on account of the darker glumes
appears to be impossible. See RAYMOND 1959, /.c.,
who is in all probability right in supposing that in
general NELMEs referred young plants to C. indica,
and those with mature fruits, in which the glumes
have often become darker, to C. dietrichiae.

7. Carex lamprochlamys S. T. BLAKE, J. Arn. Arb.
28 (1947) 104, f. 2A; NELMEs, Kew Bull. (1949)

1979 ] Cy PERACEAE—II (Kern & Nooteboom) 127

E VIJSMA ’78

Fig. 122. Utricles and nuts of Carex. Species numbered as in the text (43a WoMERSLEY 5122, 43b ANU
614, 44 Brass 4867, 45 MEIJER 6656, 46 BRASS 9803, 47 LAE 61655, 48 SCHODDE 1992, 49 Eyma 4709,
50 KOSTERMANS 14005, 5J SINCLAIR 9797, 52 JeRMY 4634). All x 10.

128

FLORA MALESIANA

[ser. I, vol. 9!

379, incl. var. diplocolea NELMES. — C. rafflesiana
var. continua [non (CLARKE) KUK.] KUx. Bull.
Jard. Bot. Btzg III, 16 (1940) 315. — C. continua
(non CLARKE) S. T. BLAKE, J. Arn. Arb. 28 (1947)
104, p.p. — C. papuana NELMES, Kew Bull. (1949)
379; Reinwardtia 1 (1951) 314. — C. tyttholepis
NELMES, Kew Bull. (1949) 388; Reinwardtia 1
(1951) 320. — Fig. 118, 124h-i.

Rhizome short. Stems tufted, rather stout,
triquetrous, smooth, 50-100 cm by 3 mm at the
base. Leaves subcoriaceous, mostly basal but some-
times one halfway up the stem, shorter than the
stems, linear, long-attenuate, with strongly revo-
lute margins, vesiculose-asperous by whitish
vesicles especially on the upper surface, (3-)7-
10 mm wide; lower sheaths reddish brown, densely
but minutely hispidulous, eventually fraying into
fibres. Inflorescence a slender, interrupted or sub-
continuous, decompound panicle up to 50 cm
long; partial panicles 4-12, at up to 8 nodes,
mostly single, sometimes binate, oblong-lanceolate,
erect, rather dense, 3-9 cm long, upper ones
approximate, lower ones distant on slender, rigid,
scabrid peduncles more or less exserted from the
sheaths; rachis hispid. Lower bracts foliaceous, as
long as or longer than the inflorescence, long-
sheathing, upper much reduced; bracteoles glumi-
form, with hispidulous awns. Spikelets numerous,
androgynous, oblong or ovoid, 5-12 mm long, the
3 part shorter than to about as long as the @ part.
Glumes broadly ovate, often broader than long,
obtuse, translucent, glabrous or the lower ones
hispidulous, pale fulvous to dark reddish, finely
several-nerved, (3/,-)1-17/, mm, the midnerve
excurrent into a strong, antrorsely scabrid, up to
2 mm long awn. Utricles trigonous, not inflated,
ellipsoid or slightly obovoid, membranous,
patent, straight or more or less recurved, with
(3—)4—6 strong nerves on each face, glabrous and
smooth, or hispidulous on the margins in the upper
part, subabruptly beaked, light green to strami-
neous, (2!/,-)3-4 mm by 1-1!/, mm; beak slender,
compressed, somewhat tapering, straight or slightly
curved, scabrid on the margins, shorter than to
about as long as the body of the utricle, (1-)1!/, mm
long; mouth bidentate, not or scarcely oblique.
Nut ellipsoid or slightly obovoid, triquetrous with
shallowly concave sides, brown, 11/,-2 by 9/19-
1 mm. Style-base slightly thickened. Stigmas 3.

Distr. Malesia: throughout New Guinea, also
known from some adjacent islands.

Ecol. Primary forests (often Fagaceae forests),
secondary growths, rarely in wet grassland, 850-
2700 m.

Vern. Anifi, Wapi lang.

Note. NELMes distinguished C. papuana from
C. lamprochlamys by the narrower leaves and the
darker glumes and utricles of the former. To me
those differences are insufficient for specific
separation. The type collection of C. tyttholepis 1
cannot distinguish from C. lamprochlamys.

8. Carex nodiflora Boeck. Bot. Jahrb. 5 (1884)
516; KUx. Pfl. R. Heft 38 (1909) 288; Philip. J.
Sc. 6 (1911) Bot. 61; Merr. En. Philip. 1 (1923)
140; NetMes, Reinwardtia 1 (1951) 260; ibid. 2
(1954) 374. — C. cumingii [‘Boott’ in VIDAL,
Phan. Cuming. (1885) 156; Rev. Pl. Vasc. Filip.
(1886) 286, nomen]; ex CLARKE, J. Linn. Soc.
Bot. 37 (1904) 11; Philip. J. Sc. 2 (1907) Bot. 107. —
C. vulcanica ELmer, Leafl. Philip. Bot. 10 (1938)
3526. — Fig. 118.

Rhizome shortly creeping, woody, covered with
the fibrous remains of old scales. Stems tufted,
triquetrous, smooth, surrounded below the leaves
by a few bladeless, brown sheaths or their fibrous
remains, 50-70 cm by 1'/,-2 mm. Leaves basal,
sometimes 1-2 higher on the stem, much exceeding
the stems, rigid, flat, long-attenuate, scabrid on the
margins in the upper part, 7-16 mm wide; cauline
leaves long-sheathing. Inflorescence a_ slender,
compound panicle, continuous above, interrupted
below, 15-35 cm long; secondary panicles 4-6,
single at the nodes, erect, spiciform or the middle
ones again branched, oblong, dense, 2-6 by 1-2 cm;
lower peduncles more or less exserted from the
sheaths, smooth, or scabrid above; rachis hispidu-
lous especially above. Lower bracts foliaceous,

overtopping the inflorescence, long-sheathing,
upper much _ reduced. Spikelets numerous,
androgynous, ovoid, oblong, or subglobose,

5-10 mm long, the 2 part rather longer than the ¢
part. Glumes ovate-lanceolate, acutish or obtuse,
membranous, nerved, shortly setulose above or
glabrescent, sparsely ciliolate, greenish white or
light brown with whitish hyaline margins, 2 mm
long, the midnerve excurrent in a smooth or his-
pidulous, '/, mm long awnlet. Utricles much
overtopping the glumes, trigonous, ellipsoid-
rhomboid, with prominent angles and flattish faces,
membranous, suberect, many-nerved, straight,
whitish setulose (except at the base), scarcely
stipitate, rather abruptly beaked, finally dark
brown, 31/,-4'/, by 11/,-2 mm; beak oblong-
conical, bidenticulate, with slightly oblique mouth,
1 mm long. Nut triquetrous, ellipsoid-rhomboid,
sessile, narrowly discoid-annulate at the apex,
ferrugineous to blackish, 2!/,—2!/, by 11/,-13/, mm.
Style-base pyramidally thickened, persistent on the
nut. Stigmas 3.

Distr. Malesia: Philippines (Luzon; according
to MERRILL, /.c., also in Alabat and Mindanao:
Agusan).

Ecol. Primary forests at low and medium alti-
tudes.

Note. According to NELMES (1951: 262) in this
species it is the style-base which is discoid-annulate,
not the apex of the nut. I do not see any difference
with the nut and style-base of the allied species.

9. Carex sarawaketensis Kix. Bot. Jahrb. 69
(1938) 262; ibid. 70 (1940) 464, incl. var. minor;
NELMES, Reinwardtia 1 (1951) 316. — C. melano-

1979] Cy PERACEAE—II (Kern & Nooteboom) 129

E VISSMA ’78

Fig. 123. Utricles and nuts of Carex. Species numbered as in the text (53 VAN STEENIS 4624, 54 DE WILDE

c.s. 13329, 55 Brass 9284, 56 Brass 9515, 57 EyMA 3862, 58 herb. VAN SOEST 88, 59 VAN STEENIS 9804,

62 HOooGLAND & SCHODDE 7470, 63 JESWIET 39, 64 DE WILDE c.s. 13323, 65a KOORDERS 43403, 66 BRASS
9539). All x 10.

130

FLORA MALESIANA

[ser. I, vol. 9!

phora S. T. BLAKE, J. Arn. Arb. 28 (1947) 106, f.
2B. — Fig. 118.

Rhizome emitting slender stolons covered with
brown or blackish scales. Stems erect or somewhat
curved, solitary or tufted, slender, obtusely trigo-
nous, smooth or scaberulous at the top, 3-40
(—60) cm by '/,-1 mm, the base surrounded by the
fibrous, reddish-black remains of old leaf-sheaths.
Leaves basal, rarely 1 higher on the stem, rigid,
shorter than the stems, linear, long-attenuate, flat
or with revolute margins, conspicuously keeled,
scabrid on the margins and keel in the upper part,
2-4 mm wide; lower sheaths fuscous to blackish-
purple. Inflorescence a decompound, narrow
panicle interrupted at the base, 10-20(-30) cm
long; partial panrcles 3-14, at 2-8 nodes, single and
binate, oblong, dense, 1-5 cm long, upper ones
approximate, lower on slender, erect, scabrid, up
to 8 cm long peduncles exserted from the sheaths.
Lower bracts foliaceous, not or scarcely over-
topping the inflorescence, long-sheathing, upper

ones much reduced; bracteoles glume-like,
hispidulous, awned; rachis hispid. Spikelets
androgynous, suberect or appressed, oblong,

4-10 by 2-2'/, mm, the g part inconspicuous,
mostly shorter than the 2 part. Glumes ovate,
membranous, acute, muticous or minutely mucro-
nulate, rarely awned, slenderly nerved, glabrous or
the lower ones minutely hispidulous, blackish-
fuscous with white-hyaline margins, 2-3 mm long.
Utricles trigonous, not inflated, ellipsoid, narrowed
at both ends, suberect, membranous, straight,
obscurely nerved (only 2 nerves more prominent),
glabrous at the base, otherwise densely scabrid-
pubescent, gradually narrowed into the beak,
blackish-fuscous, 23/,-41/, by 4/,.-11/; mm; beak
straight, compressed, deeply bidentate (teeth
1/,*/; mm) but often originally oblique, scabrid on
the margins, 1—-1°/, mm long, the mouth with whit-
ish-hyaline margins. Nut triquetrous with shallowly
concave sides, ellipsoid or broadly ellipsoid,
stipitate, conspicuously apiculate, stramineous-
brown, 1'/,-13/; by 4/;-1 mm. Style scabrid,
not or scarcely incrassate at the base. Stigmas 3.

Distr. Malesia: New Guinea (W. New Guinea:
Carstensz Mts, Mt Wilhelmina, Hellwig Mts; NE.
New Guinea: Mt Sarawaket, Lake Naho).

Ecol. Wet places in alpine grassland, steep
mountain slopes, open places in mountain forests,
moist grassy cliffs, 2700-3950 m.

10. Carex stramentitia Bootr ex Boeck. Linnaea
40 (1876) 351; CLARKE, Fl. Br. Ind. 6 (1894) 717;
J. Linn. Soc. Bot. 37 (1904) 9; Kix. Pfl. R. Heft
38 (1909) 264; Camus, FI. Gén. I.-C. 7 (1912) 188;
NELMES, Kew Bull. (1950) 191; Reinwardtia 1
(1951) 267; Mém. Mus. Hist. Nat. Paris n.s. B4
(1955) 120; Raym. Mém. Jard. Bot. Montréal
n. 53 (1959) 45, 99, f. 28 (map); Dansk Bot. Ark. 23
(1965) 254; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 492. — Fig. 118, 124m-n.

Rhizome shortly creeping, stout, woody. Stems
loosely tufted, trigonous, scaberulous below the
nodes to almost smooth, 30-130 cm by 1!/,-3 mm,
the base clothed with fuscous, bladeless sheaths
and comose by their fibrous remains. Leaves
subcoriaceous, subbasal and 1-2 higher on the
stem, exceeding the stems, linear, long-attenuate,
flat or with revolute margins, scabrid on the mar-
gins in the upper part, pale green, 5—10(-15) mm
wide. Inflorescence a pale, compound, erect, much
interrupted, narrow, 10-40 cm long panicle;
secondary panicles 2-4, single at the nodes, erect,
lanceolate or oblong, very dense, distant, up to
10 cm long; peduncles smooth or scaberulous, the
lower ones much exserted from the sheaths, the
upper ones scarcely so; rachis hispid. Lower bracts
foliaceous, exceeding the inflorescence, long-
sheathing, hispidulous in front or at the mouth,
upper reduced. Spikelets androgynous, sessile,
suberect, 5-15 mm long, the 2 part few-flowered,
rather shorter than the g part. Glumes oblong or
oblong-ovate, thinly membranous, obtuse or
slightly emarginate, slenderly nerved, glabrous or
sparsely hispidulous, not ciliate, pale stramineous
to whitish, 2—23/, mm long, the midnerve excurrent
into an antrorsely scabrid, */,-2 mm long awn.
Utricles distinctly trigonous, rhomboid-ellipsoid,
with prominent angles and flattish faces, mem-
branous, not inflated, patulous, many-nerved
(nerves 5-7 on each face), glabrous, straight to
slightly recurved, curved-tapering below into a
cuneate basal part, suddenly narrowed above into
the beak, greenish to light brown, 4-5 by c. 11/, mm;
beak straight or slightly curved, often somewhat
inflated at the base, glabrous or very sparsely
scabrid, 11/2 mm long; mouth dorsally very
oblique, not bidentate. Nut trigonous, rhomboid-
ellipsoid, erostrate, curved-tapering below into a
stout cuneate part, with prominent angles and
concave faces, brown, 2'/,-3 by 11/,-14/; mm.
Style-base pyramidally thickened, persistent on the
nut. Stigmas 3.

Distr. India, Lower Burma, S. China (Kwei-
chow), Thailand, Laos, Tonkin, Annam; in
Malesia only known from a single collection from
W. Java (Krawang: Pléréd near Purwakarta); see
map in RAYMOND, 1959, /.c.

Ecol. In Krawang abundant in jungle at low
altitude under seasonal climatic conditions.

Vern. Lilisungan, S.

Note. In spite of its different facies because of
the stiff, linear leaves, C. stramentitia is very near
C. commixta, which it closely resembles in the size
and shape of the utricles. According to NELMES the
leaves of C. stramentitia can reach a width of
20 mm.

11. Carex vesiculosa Bootrt, Ill. 3 (1862) 107, t. 323;
Boeck. Linnaea 40 (1876) 345; CLARKE, FI. Br.
Ind. 6 (1894) 717, incl. var. paniculata CLARKE;
Kwx. Pfi. R. Heft 38 (1909) 283, f. 43, inc! var.

1979] Cy PERACEAE—II (Kern & Nooteboom) 13]

Fig. 124. Inflorescence (nat. size), spikelets and (occasionally) venation (x 2). — 3a. Carex cruciata
WAHLENB. var. cruciata, a—b. — 3b. var. rafflesiana (Boott) Noot., c-d. — 5. C. horsfieldii Boott, e-g. —
7. C. lamprochlamys S. T. BLake, h-i. — 6. C. indica L., j-k. — 13. C. hypolytroides Riv. 1. — 10.
C. stramentitia Boott, m-n. — 11. C. vesiculosa Boot, 0-p. (a-b CHEW, CORNER & STAINTON 1706,
¢ VAN OosTsTROOM 13154, d VERHEUEN 2585, e-g NGF 21261, A-i Brass 24684, j-k RipLey 15721, /
BUNNEMEVJER 9820, m—n BAKHUIZEN VAN DEN BRINK 6565, 0-p VAN STEENIS 4140).

132

FLORA MALESIANA

[ser. I, vol. 9!

congesta KUK.; BACK. Bekn. FI. Java (em. ed.) 10
(1949) fam. 246, p. 68; KERN in Back. & Bakh. /.
Fl. Java 3 (1968) 492. — C. impunctata Bootr,
Ill. 3 (1862) 107, t. 326 (pl. depauperata); BoEcK.
Linnaea 40 (1876) 342. — C. gembolensis CLARKE,
J. Linn. Soc. Bot. 37 (1904) 10; Netmes, Rein-
wardtia 1 (1951) 294, incl. var. crebra NELMES;
ibid. 2 (1954) 376. — C. rhizomatosa var. impunctata
KUK. Pfl. R. Heft 38 (1909) 291. — C. vesiculosa
var. latifolia KUxK. in Hochr. Candollea 6 (1931)
341. — Fig. 118, 1240-p.

Rhizome shortly creeping, woody, covered with
the fibrous remains of decayed sheaths. Stems
loosely tufted, stiff, trigonous, smooth, 30-150 cm
by 2-3 mm, surrounded below the leaves by blade-
less, reddish or fuscous sheaths and their fibrous,
often reticulate remains. Leaves coriaceous, basal
and 1-2 higher on the stem, usually much shorter
than the stems, narrow, long-attenuate, keeled,
with strongly revolute margins when dry, scabrous
on the margins, often vesiculose-asperous above
especially towards the apex, 2-8 mm wide. /nflores-
cence a decompound, interrupted, ferrugineous,
15—40 cm long panicle; secondary panicles 5—10, at
4-6 nodes, lowest 1-2 often single, remainder
usually unequally binate, erect, often with nodding
top, oblong-pyramidal, dense or very dense,
rarely loose, up to 10 by 5 cm, upper ones approxi-
mate, lower distant on exserted, smooth or scabrid,
up to 15 cm long peduncles; rachis hispidulous on
the angles. Lower bracts foliaceous, narrow,
shorter than the inflorescence, long-sheathing, the
upper ones much reduced; bracteoles pilose,
excurrent in a curved, hispidulous awn. Spikelets
numerous, androgynous, obliquely patent, 5—

15 mm long, the g part in the longer spikelets much
longer than the few-flowered 92 part. Glumes
lanceolate-ovate, acutish, sometimes lightly emar-
ginate, membranous, translucent, glabrous or
somewhat hispidulous, ferrugineous or castaneous,
2'/,-41/, mm long, the midnerve in the upper
glumes excurrent in a hispid, up to 13/, mm long
awn. Utricles triquetrous, narrowly ellipsoid, sub-
coriaceous, not inflated, obliquely erect, straight or
but slightly recurved, slenderly or obscurely nerved,
rather densely setulose in the upper 3/,, scarcely
stipitate, rather abruptly beaked, reddish with
castaneous flecks, 3-5 by 1—-13/, mm; beak slender,
scabrid on the margins, mouth not oblique, 11/,-
2 mm long. Nut triquetrous, ellipsoid or slightly
obovoid, with concave faces, shortly stipitate,
abruptly beaked, 1!/,-2'/, by 1-11/, mm. Style-
base scarcely thickened. Stigmas 3.

Distr. Nepal to Thailand; in Malesia: Java (in
W. Java on Mt Papandajan, not rare in the moun-
tains of Central and E. Java), Lesser Sunda Is.
(Bali, Lombok, Flores, Timor).

Ecol. In dry grass-lands, open stony places,
mountain scrub, Casuarina forests, 1200-3500 m.

Notes. In old specimens the leaves are often
less asperous because the vesiculose (bulbous-
based) hairs have broken off. The former presence
of vesicles is indicated by small, circular scars.

If C. gembolensis is specifically distinct from
C. vesiculosa, its correct name is C. impunctata
Boott, based on ZOLLINGER 2563 from Java. This
collection NELMEs rightly included in C. gembolen-
SES

Var. congesta KUK. (= C. gembolensis var.
crebra NELMES) has no taxonomic value.

2. Section Japonicae

KUK. Pfl. R. Heft 38 (1909) 252. — Sect. Paciricae OHw1, Mem. Coll. Sc. Kyoto
Imp. Un. B11 (1936) 458; NeLmes, Reinwardtia 1 (1951) 329; Raym. Mém. Jard.
Bot. Montréal n. 53 (1959) 20, 52. — Sect. Indicae subsect. Japonicae (KUK.)
Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 152.

Type species: Carex nikoensis FRANCH. & SAVAT.

12. Carex satzumensis FRANCH. & SAVAT. En. PI.
Jap. 2 [(1877) 132, nomen] (1878) 558; FRANCH.
Nouv. Arch. Mus. Hist. Nat. Paris III, 8 (1896)
259 (t. 4, f. 1 ut C. nikoensis); AKIYAMA, J. Fac.
Sc. Hokk. Imp. Un. V, 2 (1932) 84, f. 36; Car. Far
East. Reg. Asia (1955) 135, t. 121; On#wi, Mem.
Coll. Sc. Kyoto Imp. Un. B11 (1936) 458; NELMes,
Reinwardtia 1 (1951) 330; Koyama, Nat. Canad.
82 (1955) 197; Contr. Inst. Bot. Un. Montréal
n. 70 (1957) 10; YosHIKkAwa, Ic. Jap. Carex 2
(1958) 236, t. 118; RAym. Mém. Jard. Bot.
Montréal n. 53 (1959) 52. — C. nikoensis FRANCH.
& SAVAT. En. Pl. Jap. 2 [(1877) 132, nomen] (1878)

558; KUx. Pfl. R. Heft 38 (1909) 252, f. 383A-E
(nikkoensis’); Philip. J. Sc. 6 (1911) Bot. 59;
Merr. En. Philip. 1 (1923) 140. — C. contracta
Boeck. Cyp. Nov. 2 (1890) 34. — Fig. 119.
Rhizome long-creeping, clothed with large,
brown scales. Stems erect, triquetrous, ribbed,
smooth, or scabrid just below the inflorescence,
3-20(-30) cm by 1-1!/, mm, surrounded at the
base by the fibrous remains of old leaf-sheaths.
Leaves subbasal, rigid, flattish to conduplicate,
longer than the stems, often recurved, scabrid on
the margins and on the upper surface towards the
long-attenuate apex, 2-5 mm wide. Inflorescence

133

Fig. 125. Carex indica L. a. Habit, x *+/3, b-c. glumes, d. fruit in utricle, e. fruit, all x 7 (SCHODDE 2972).

simple, spiciform, sometimes slightly branched at
the base, cylindrical-conical, acute, 2-8 cm by
1—2'/, cm at the base; rachis smooth. Lower bracts
subulate to subfoliaceous, shorter than the in-
florescence, not sheathing, upper filiform.
Spikelets 12-numerous, androgynous, _ sessile,
patent, ovate to oblong, densely flowered, lower
5-14 mm, upper 3-7 mm long, all the lateral ones

arising from a + utriculiform, gibbous prophyll
containing a 2 flower. Glumes lanceolate-ovate,
acute to obtuse, glabrous, pale, slenderly nerved,
2-3 mm long, the midrib sometimes slightly excur-
rent. Utricles oblong-ellipsoid, obtusely trigonous,
hardly inflated, glabrous, slenderly plurinerved,
greenish, 2!/,—3!/, by 3/,-1 mm, rather gradually
narrowed into a long, conical, bilobed beak with

134

FLORA MALESIANA

[ser. I, vol. 9!

oblique mouth. Nut oblong-obovoid, triquetrous,
shortly beaked, dark brown, 1'/,-1'/, mm long.
Style pyramidally thickened at the base. Stigmas 3.

Distr. Japan, Formosa, Tonkin; in Malesia:
Philippines (Luzon: Benguet).

Ecol. Along trails in the mossy forest, at
c. 2300 m, also along roadsides under pine-trees
at 400 m.

Notes. Readily recognizable by its spike-like
inflorescence with non-sheathing bracts.

Carex nikoensis FRANCH. & SAVAT., with slightly
larger spikelets in denser spikes and somewhat

longer beak of the utricle than in typical C. satzu-
mensis, was already reduced to C. satzumensis by
FRANCHET (1896).

The specific epithet is frequently spelt ‘satsumen-
sis’, but the spelling ‘satzumensis’, employed when
the species was validly published, must be retained.

In an abnormal inflorescence of SANTOS 5776
from Luzon I found in all branches between the
fertile cladoprophyll and the normal utricles some
deeply split utricles containing one or some ¢
flowers besides the 2 one, a situation normal in
Schoenoxiphium and Kobresia.

3. Section Hypolytroides

NELMES, Kew Bull. (1951) 121; Reinwardtia 1 (1951) 246; Raym. Mém. Jard. Bot.

Montréal n. 53 (1959) 20, 21.

Type species: Carex hypolytroides RIDL.

13. Carex hypolytroides RipL_. J. Fed. Mal. St.
Mus. 8 (1917) 124; NeLmes, Reinwardtia 1 (1951)
246; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 96;
RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 21,
f. 3, 102, f. 31 (map). — C. hypolytroides Gross &
Mattr. Notizbl. Berl.-Dahl. 14 (1938) 190. —
C. hypolytropsis Gross in Fedde, Rep. 50 (1941)
213. — Fig. 119, 1241, 126.

Rhizome woody, emitting strong stolons
covered with lanceolate, fuscous sheaths. Stems
rigid, erect, rather acutely trigonous, smooth,
100-250 cm by 3-5 mm, surrounded at the base by
a few reddish brown, bladeless sheaths. Leaves at
regular intervals throughout the stem, shorter than
the stem, stiff, flattish, or with revolute margins
when dry, smooth, or scabrid on the margins,
sparsely to rather densely covered with long, pale
hairs beneath, 4-8 mm wide; sheaths smooth, hairy
above, membranous in front, with concave mouth;
ligule elongate, hairy, ferrugineous. Inflorescence a
compound, interrupted panicle, 25-40 cm long;
secondary panicles 5-8, erect, broadly pyramidal,
single at the nodes, or lowest sometimes binate,
rather loose, upper approximate, lower distant, on
exserted, hispidulous peduncles with patent
branches. Lower bracts foliaceous, slightly
exceeding the inflorescence, long-sheathing, upper
reduced. Spikelets usually unisexual; 2 ones
numerous, shortly cylindrical, very dense, 5-8 by
31/,-4'/, mm, sometimes with a few 3 flowers at the
top; d ones inconspicuous, few, lateral, 1-5 just
below some of the terminal 9 spikelets, sessile,
ellipsoid, 4 by 1-2 mm. Cladoprophylls of branches
and peduncles utriculiform, hairy, those of the
3 spikelets sometimes enclosing a 2 flower, those of
the partial panicles ocreiform. Glumes thinly
membranous, ovate-lanceolate, acute, hairy, slen-
derly nerved, with narrow whitish-hyaline margins

and shortly excurrent midnerve, 2—23/, mm long,
in ripe spikelets almost completely hidden among
the utricles. Utricles obtusely trigonous, obovoid,
not or but slightly inflated, horizontally patent,
straight, glabrous or very sparsely hispidulous,
slenderly nerved (2 nerves more prominent),
fuscous, densely reddish glandular-spotted, 11/,—
1*/, by c. 1 mm, subabruptly narrowed into a very
short, bidenticulate beak. Nut triquetrous, obovoid,
filling the utricle, sessile, brown, whitish papillose,
1?/<-1'/; by °/;>-1 mm. Style-base thickened.
Stigmas 3.

Distr. Tonkin, Annam; in Malesia: Central
Sumatra (Mt Kerintji), Borneo (Mt Kinabalu:
Lumu-lumu, Mesilau Cave). See the distribution
map by RAYMOND, /.c.

Ecol. In wet spots in forests, on Mt Kerintji
between 2200 and 2750 m, on Mt Kinabalu at
1800 m.

Notes. The infructescence so strongly recalls a
Hypolytrum that RIDLEY and Gross independently
of each other chose the epithet hypolytroides for it.

The ¢ spikelets are inserted laterally, as they are
borne from a utriculiform prophyll, and there-
fore cannot have degraded from an originally
terminal position as was supposed by NELMES,
Reinwardtia 1 (1951) 248.

Carex hypolytroides is closely related to C.
moupinensis FRANCH. Nouv. Arch. Mus. Hist. Nat.
Paris II, 10 (1888) 102; ibid. III, 8 (1896) 257, t. 7
f. 2; CLARKE, J. Linn. Soc. Bot. 36 (1904) 209;
Kok. Pfl. R. Heft 38 (1909) 289; Raym. Mém.
Jard. Bot. Montréal n. 53 (1959) 21, f. 2. —
Homalostachys sinensis BOECK. Cyp. Nov. 1 (1888)
38, non C. chinensis Retz. — Scleria sinensis
H. PreirF. in Fedde, Rep. 26 (1929) 263, only
known from China (Hupeh, Szechuan, Yunnan).

1979] CYPERACEAE—II (Kern & Nooteboom) 135

Fig. 126. Carex hypolytroides Rwv. a. Habit, x */3, b. terminal ¢ and lateral ¢ spikelet, x 3, c-d. fruit in
utricle, e. fruit, both x 13 (Jacoss 4332).

The latter species is much lower, with narrower the slightly larger (2 by 1!/; mm), much inflated
leaves, and usually all the spikelets of the terminal _utricles truncate or depressed at the top, and the
partial panicle 3; it is especially characterised by nut not filling the utricle.

136

FLORA MALESIANA

[ser. I, vol. 9!

4. Section Mapaniifoliae

NELMES & AIRY SHAW in Hook. Ic. Pl. 35 (1943) t. 3434; NeELMEs, Reinwardtia 1
(1951) 248; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 27; 93, f. 23 (map). —
Sect. Indicae subsect. Scaposae KUx. Pfl. R. Heft 38 (1909) 285, p.p.

Type species: Carex helferi BOECK.

14. Carex helferi Boeck. Linnaea 40 (1876) 365;
CLARKE, FI. Br. Ind. 6 (1894) 714; KUx. Pfl. R.
Heft 38 (1909) 286; Netmes in Hook. Ic. Pl. (1947)
t. 3468; Kew Bull. (1950) 189; Reinwardtia 1
(1951) 248; Mém. Mus. Hist. Nat. Paris n.s. B4
(1955) 99; Raym. Mém. Jard. Bot. Montréal n. 53
(1959) 30; Dansk Bot. Ark. 23 (1965) 252; KERN
in Back. & Bakh. f. Fl. Java 3 (1968) 491. —
C. mapaniifolia Rip. J. Fed. Mal. St. Mus. 10
(1920) 124; Ne_mes, Mém. Mus. Hist. Nat. Paris
n.s. B4 (1955) 98; RayM. Mém. Jard. Bot. Montréal
n. 53 (1959) 30. — Fig. 119.

Rhizome short, stout, woody, clothed with some
brown bladeless sheaths or their fibrous remains.
Stems central, erect, scapiform, trigonous or sub-
terete, smooth, or scabrid on the angles, (S—)20—
40 cm by 1-2 mm, the base enveloped by some
brown, infundibuliform, up to 10 cm long sheaths.
Leaves overtopping the stems, broadly linear,
conduplicate below (narrowed into a _ pseudo-
petiole), otherwise flat, long-acuminate, minutely
scabrid on the margins in the upper part, with the
midnerve and 2 mid-lateral nerves prominent,
11/,-31/, cm wide; ligule elongate-triangular,
brown-bordered, 1!/,-2 cm long. Inflorescence
paniculate, consisting of (2—)3-7 partial inflores-
cences, up to 25 cm long; partial inflorescences
erect, oblong-ovoid, very dense, head-like, upper
approximate, lower distant, single at the nodes,
with hispid rachis, 2—31/, by 1—21/, cm, on exserted,
smooth or finely scabrid peduncles. Bracts sheath-
like, ampliate, infundibuliform, short-bladed,
light brown or spadiceous. Spikelets densely
crowded, androgynous, ovoid, patent, 6-10 by
5-8 mm, the ¢ part usually much longer than the
few-flowered 2 part. Glumes thinly membranous,
oblong-ovate, obtuse to truncate-bilobed, many-

nerved, glabrous or hispidulous, pale brown with
broad, whitish hyaline margins, 2-4 mm long, the
midnerve excurrent in a 1-4 mm long, scabrous,
often recurved awn. Urtricles trigonous, ellipsoid,
with shallowly concave faces, patulous, glabrous
at the base, hispidulous above, shortly stipitate,
strongly 6—-8-nerved on each face, stramineous to
brown, 6-7 by 1'/,-14/; mm (the long beak
included); beak hispidulous, curved, bulbous at the
base (to hold thickened style-base), nearly linear
above, with very oblique mouth (its base 11/,—
13/, mm from the apex), 3—-3'/, mm long. Nut tri-
gonous, ellipsoid or slightly obovoid, scarcely
stipitate, shortly beaked and suddenly expanded
into the conical style-base, dark brown with pale
angles, 21/,-23/, by 11/,-13/, mm. Style-base
pyramidally thickened, subpersistent on the nut.
Stigmas 3.

Distr. Lower Burma, SE. & Peninsular
Thailand; in Malesia: W. Java (Priangan).
Wrongly recorded for Sumatra and Borneo by
NeEwMES (1950, p. 100).

Ecol. In forests, 1100 m.

Notes. This is the only Malesian member
of the wide-leaved sect. Mapaniifoliae, which
section appears to be almost confined to the Indo-
Chinese Peninsula, where it is represented by several
species.

The few Javan specimens hitherto collected are
vegetatively much less developed than those from
the continent, but agree with them in floral and
fruit characters.

By its broad leaves, scapiform flowering stems,
and shape of the inflorescence Carex helferi
deceivingly resembles Hypolytrum humile (STEUD.)
Boeck. It can be distinguished by the long-beaked
utricles, the 3 stigmas, and the trigonous nuts.

5. Section Polystachyae

[TUCKERM. En. Meth. (1843) 10, nomen (‘Polystachae’)|; ex KUK. Bot. Jahrb. 27
(1899) 517, quoad basion.; Pfl. R. Heft 38 (1909) 257; NELMEs, Reinwardtia 1 (1951)
322; RAYM. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 49. — Sect. Acrarrhenae
Fries Sippe Longebracteatae PAx in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 124. — Sect.
Polystacheae CLARKE [ser.] Longispicae CLARKE, J. Linn. Soc. Bot. 37 (1904) 4,
p.p. — Sect. Extensae Fries subsect. Baccantes KOYAMA, J. Fac. Sc. Un. Tokyo HI,

8 (1962) 151.

Type species: Carex myosurus NEES.

1979]

Cy PERACEAE—II (Kern & Nooteboom)

iS7

15. Carex baccans NEEs in Wight, Contr. Bot. Ind.
(1834) 122; KuntH, En. 2 (1837) 513; Boortr, Ill. 2
(1860) 83, t. 234-236, 238, 239; Boeck. Linnaea 40
(1876) 339; O. K. Rev. Gen. Pl. 2 (1891) 747, incl.
var. nigra O. K.; CLARKE, Bot. Mag. III, 49 (1893)
t. 7288; Fl. Br. Ind. 6 (1894) 722; J. Linn. Soc. Bot.
37 (1904) 14, incl. var. siccifructus CLARKE;
Philip. J. Sc. 2 (1907) Bot. 108; KUx. Pfl. R. Heft
38 (1909) 258, f. 39E-H; Philip. J. Sc. 6 (1911)
Bot. 59; Koorp. Exk. Fl. Java 1 (1911) 211;
Camus, FI. Gén. I.-C. 7 (1912) 185; Merr. En.
Philip. 1 (1923) 136; Rm. Fl. Mal. Pen. 5 (1925)
184; Onwi1, Mem. Coll. Sc. Kyoto Imp. Un. B11
(1936) 462; Back. Bekn. Fl. Java (em. ed.) 10
(1949) fam. 246, p. 68; NeLMes, Kew Bull. (1950)
194; Reinwardtia 1 (1951) 322; Raym. Mém. Jard.
Bot. Montréal n. 53 (1959) 50; Koyama, J. Fac.
Sc. Tokyo Un. III, 8 (1962) 216; Dansk Bot. Ark.
23 (1965) 257; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 493; STEEN. Mt. Fl. Java (1972) pl. 15-1. —
C. curvirostris KUNZE, Suppl. Riedgr. (1840-50)
79, t. 20; Mia. FI. Ind. Bat. 3 (1856) 350; CLARKE,
J. Linn. Soc. Bot. 37 (1904) 14; Streup. [in Zoll.
Syst. Verz. 1 (1854) 60, nomen] Syn. 2 (1855) 207
(‘recurvirostra’). — C. javanica BOECK. Cyp. Not. 1

(1888) 43. — C. walkeri (non ARN. ex Boott) KUx.
Pfl. R. Heft 38 (1909) 546; cf. BAcK. Brittonia 3
(1938) 76. — Fig. 119, 127, 131i.

Rhizome short, stout, woody. Stems usually
robust, loosely tufted, erect, triquetrous or
trigonous, smooth, 60-150 cm by up to 5 mm, the
base surrounded by reddish, bladeless sheaths
splitting in front into reticulate fibres. Leaves all
over the stem, often overtopping the inflorescence,
long-attenuate, coriaceous, flat or with revolute
margins, asperous on the upper surface in the apical
part, with scabrous margins, 5-18 mm wide;
sheaths long, often splitting up in front. /nflores-
cence paniculate, compound or decompound,
oblong, occupying 1/,—'/, of the stem, erect or
somewhat nodding at the top; partial panicles 5-8,
single at the nodes, much branched, upper approxi-
mate, lower distant on long-exserted, smooth or
scabrid peduncles. Lower bracts usually much
overtopping the inflorescence, foliaceous, long-
sheathing, upper much reduced. Spikelets very
numerous, androgynous, suberect to patent,
2'/,-8 cm long; 2 part cylindrical, densely flowered,
usually longer than the slenderer ¢ part. Glumes
ovate or oblong-ovate, acute to subobtuse, glab-

Fig. 127. Carex baccans NEES in pioneer tufts with sand-binding capacity in the sand plain and dunes of
volcanic ash of the Tengger Sandsea, East Java, c. 1900 m altitude (CLAsoN).

138

FLORA MALESIANA

[ser. I, vol. 9!

rous or hispidulous, strongly nerved, purplish or
castaneous with whitish-hyaline margins, muticous
or the midrib excurrent in a scabrid mucro,
21/,-31/, mm long. Utricles inflated, obscurely
trigonous, obovoid to subglobose, patent, sub-
coriaceous, strongly nerved, glabrous except for
the hispidulous margins at the apex, shining, at
first yellowish green, ultimately red and more or
less succulent, 31/,-4'/, mm long, abruptly beaked;
beak recurved, bidentate. Nut triquetrous, ellip-
soid, with flattish or shallowly concave faces, dark
brown, 23/,-3 mm long. Style-base not thickened.
Stigmas 3.

Distr. Ceylon, India (Sikkim and Khasia), S.
China and Formosa, and through Thailand and
Indo-China to Malesia; in Malesia: Sumatra,
Malay Peninsula (Pahang), Java, Lesser Sunda Is.
(Bali, Lombok), Philippines (Luzon), New Guinea.

Ecol. In damp thickets, open places in forests, in
the mossy forest on the higher mountains, some-
times in mountain savannahs, or as pioneer in the
Sandsea and on fresh landslides, on volcanoes,
talus; (600—)1000-—3300 m.

Vern. Ladingan, ria-ria-batu, sukét kérisan, J,
ilat beureum, ilateun téki, S; Philip.: gihidsan,
Bon., mankat, silak, Ig.; New Guinea: djigudjigufa,
Dunantina, djugudjuguha, Asoro: Kefamo, pul,
Chimbu: Masue, koimin, Wahgi: Minj.

Note. This beautiful species seems to be some-
times cultivated as an ornamental in temperate
regions; see NICHOLSON, III. Dict. Gard. 1 (1885-
89) 267, f. 367).

16. Carex myosurus NEES in Wight, Contr. Bot.
Ind. (1834) 122; KuNTH, En. 2 (1837) 507; STEUD.
Syn. 2 (1855) 207; Boorrt, Ill. 2 (1860) 82, t. 229,
230, 232; BoEcK. Linnaea 40 (1876) 334; CLARKE,
Fl. Br. Ind. 6 (1894) 723, incl. var. eminens (NEES)
CLARKE; KUk. Pfl. R. Heft 38 (1909) 258; MeErrR.
En. Philip. 1 (1923) 139; NELMEs, Kew Bull. (1950)
195, incl. var. celebica NELMES; Reinwardtia 1
(1951) 325; ibid. 2 (1954) 377; Raym. Mém. Jard.
Bot. Montréal n. 53 (1959) 51; KERN in Back. &
Bakh. f. Fl. Java 3 (1968) 493; STEEN. Mt. Fl. Java
(1972) pl. 14-2. — C. eminens NEES in Wight,
Contr. Bot. Ind. (1834) 122. — C. longibracteata
STEUD. [in Zoll. Syst. Verz. 1 (1854) 60, nomen]
Syn. 2 (1855) 205 (‘longebracteata’), non SCHLEICH.
Cat. Pl. Helv. ed. 4 (1821) 11, nomen; Mia. FI. Ind.
Bat. 3 (1856) 348, incl. var. major MiQ.; BOECK.
Linnaea 39 (1875) 108; CLARKE, J. Linn. Soc. Bot.
37 (1904) 15; Kx. Pfl. R. Heft 38 (1909) 260, incl.
f. distans Kix. et f. angustifolia KUK.; Bull. Jard.
Bot. Btzg III, 16 (1940) 314, incl. var. gigantea
KUK.; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949)
fam. 246, p. 70; NELMES, Reinwardtia 1 (1951) 327;
ibid. 2 (1954) 377. — C. floribunda Boeck. Linnaea
40 (1876) 335. — C. kuntzeana BoEcK. Cyp. Nov.
1 (1888) 51. — C. composita (non BooTT) CLARKE,
J. Linn. Soc. Bot. 37 (1904) 14. — Fig. 119.
Rhizome short, stout, woody. Stems slender,

loosely tufted, erect, trigonous, smooth, (20—)50—
150 cm by up to 4 mm at the base, clothed at the
base with fuscous to purplish sheaths or their
fibrous, reticulate remains. Leaves mainly subbasal,
a few widely spaced on the stem, equalling or
exceeding the inflorescence, long-attenuate, coria-
ceous, flat or with revolute margins, asperous on
the upper surface in the upper part, with scabrous
margins, (2-)5-10 mm wide; sheaths long, often
reddish, frayed in front into reticulate fibres.
Inflorescence from almost simple to decompound,
narrow, (10—)25-100 cm long, erect or somewhat
nodding at the top; secondary panicles 5-9, single
at the nodes, upper approximate, lower distant on
exserted, more or less scabrid peduncles, sometimes
all reduced to simple spikelets. Lower bracts
foliaceous, long-sheathing, exceeding the stem,
upper much reduced. Spikelets androgynous, sub-
erect to patulous, the longer ones on a plant 3 to
10 cm long, 3 and @ parts often about equal in
length, but sometimes the terminal spikelet almost
wholly 3 and some of the lateral almost wholly
2. Glumes oblong or oblong-ovate, acute to obtuse,
glabrous, or sparsely hispidulous, slenderly nerved,
pale to castaneous, with whitish-hyaline margins,
2'/,-4 mm long, the midrib usually excurrent in a
hispidulous awn up to 2 (rarely 3) mm long.
Utricles trigonous, ellipsoid or ellipsoid-obovoid,
patulous, membranous, slenderly nerved, sparsely
to subdensely hispidulous at least towards the apex,
straight or slightly bent, pale to castaneous,
3-4'/, mm long, subgradually to subabruptly
narrowed into a hispidulous-margined, bidentate,
3/,-1 mm long beak. Nut triquetrous, ellipsoid,
oblong-ellipsoid, or slightly obovoid, with flattish
to slightly concave sides, dark brown, abruptly
beaked, 2-3 mm long; stipe and beak straight to
bent at the base. Sty/e-base not thickened. Stigmas
3:

Distr. India, Burma, Indo-China; in Malesia:
Sumatra (Atjeh, W. Coast Res.), Java, Lesser
Sunda Is. (Lombok: Mt Rindjani), Philippines
(Luzon: Bontoc), SW. Celebes (Mt Bonthain).

Ecol. In open or lightly shaded places, on
forested slopes, in mountain scrub, heath forma-
tions, a pioneer on sterile crater soil, sometimes
dominant; 1700-3300 m.

Notes. Carex myosurus in the wide circumscrip-
tion here accepted is extremely polymorphous.
CLARKE, FI. Br. Ind. 6 (1894) 723, distinguished
between typical C. myosurus (from Nilghiri Mts
and Coromandelia) and C. myosurus var. eminens
(NEES) CLARKE (occurring throughout the Hima-
layas), the latter mainly characterized by the
shorter and usually broader utricles with often
rather deeply bifid beak, although in some Sikkim
and Bhotan plants (distinguished by BOECKELER as
C. floribunda) the beak is not more notched than in
the Nilghiri plants. Carex spiculata Boott, with
narrower leaves, denser spikelets with obliquely
ascending utricles and more rigid panicles he

1979]

Cy PERACEAE—II (Kern & Nooteboom)

139

considered specifically distinct. Obviously he had
not seen Malesian specimens of C. /ongibracteata
STEuD. (cf. J. Linn. Soc. Bot. 37, 1904, 15).

KUKENTHAL, Pfl. R. Heft 38 (1909) 259, distin-
guished var. eminens by the shorter and broader
utricles and the usually fuscous glumes, and var.
floribunda (BoECK.) KUK. by its profusely branched
inflorescences; C. spiculata was reduced to sub-
specific rank. On the other hand, C. /ongibracteata
was upheld as a species. As in numerous Indian
myosurus specimens the inflorescence is paniculate
or even spicate (see KUKENTHAL’s description!) it is
clear that C. longibracteata cannot be separated
from C. myosurus on account of the characters used
in KUKENTHAL’s key (“‘inflorescentia subdepanicu-
lata, spiculae numerosae” in C. myosurus, versus
“‘inflorescentia paniculata rarius spicata, spiculae
haud numerosae”’ in C. longibracteata).

No more can I understand in what way NELMES

distinguished between C. myosurus and C. longi-
bracteata, for the former is said to have leaves
5-10 mm wide and secondary panicles composed of
3 to rather numerous spikelets, whereas the latter
should differ in having leaves 2-6 mm wide and
secondary panicles composed of 1-7 spikelets.
Besides, in the Lombok specimens referred to C.
myosurus, the leaves are only 2-3 mm wide.

Carex longibracteata is very similar to C. myo-
surus var. eminens; usually the inflorescence is less
compound (but profusely branched inflorescences
occur, see KERN 8376!) and the utricles are still
somewhat broader. In the Celebes and Lombok
specimens referred by NELMES to C. myosurus the
utricles are considerably narrower.

The only Javan collection NeLMes referred to
C. myosurus (VAN STEENIS 12267) was gathered
together with VAN STEENIS 12269, which was named
C. longibracteata for reasons I do not understand.

6. Section Oligostachyae

CLARKE, J. Linn. Soc. Bot. 37 (1904) 4. — Sect. Polystachyae CLARKE [ser.] Longis-
picae CLARKE, /.c., p.p. — Sect. Frigidae Fries subsect. Decorae KUK. Pfi. R. Heft
38 (1909) 541. — Sect. Decorae (KUK.) OHWI, Mém. Coll. Sc. Kyoto Imp. Un.
B11 (1936) 338; NeL-mes, Reinwardtia 1 (1951) 332; Raym. Mém. Jard. Bot.
Montréal n. 53 (1959) 54, 71; KoyAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 154. —
Sect. Borneenses NELMES, Kew Bull. (1951) 121; Reinwardtia 1 (1951) 347; Ray.
Mém. Jard. Bot. Montréal n. 53 (1959) 54, 82.
Type species: Carex borneensis CLARKE (lectotype).

17. Carex celebica KUK. Bot. Jahrb. 70 (Jan. 1940)
465; Bull. Jard. Bot. Btzg III, 16 (Feb. 1940) 318;
NELMES, Kew Bull. (1949) 389; Reinwardtia 1
(1951) 345. — C. constricta S. T. BLAKE, J. Arn.
Arb. 28 (1947) 112, f. 4A. — Fig. 119.

Rhizome creeping, emitting slender stolons.
Stems solitary or somewhat tufted, erect or curved,
triquetrous, smooth, often hidden in the leaf-
sheaths, 2-25 cm by c. 1'/, mm, surrounded below
the leaves by purplish cataphylls or their fibrous
remains. Leaves subbasal, often recurved, longer
than the stem, flattish or canaliculate, rigid, scabrid
in the upper part, gradually attenuate into a firm
point, 1!/,-5 mm wide; sheath purplish. Jnflores-
cence consisting of 5-15 spikelets, 2-10 cm long.
Lower bracts foliaceous, much exceeding the
inflorescence, shortly sheathing, upper much
reduced. Spikelets suberect, 1/.-3 cm long, upper
approximate on hardly or not exserted,; smooth
peduncles; lower sometimes distant on longer
peduncles, terminal dg, linear, 1-2 mm thick,
lateral 2 or witha few ¢ flowers at the top, cylindric,
3-4 mm thick, lowest solitary, upper in fascicles of
2-3 (at least one of the fascicles of 3). Glumes
ovate or oblong-ovate, obtuse to emarginate, more

rarely acute, reddish brown with whitish-hyaline
margins, slenderly few-nerved, 2-3 mm long, the
midrib excurrent in a mucro or in an up to 3 mm
long awn. Utricles triquetrous, ovoid or ellipsoid,
with flat faces, nerveless (except for two marginal
nerves), glabrous, smooth, patulous, slightly
recurved, finally dark brown, 2!/3—3?/, by ?/;,.-1 mm,
subabruptly narrowed into a smooth or sparsely
hispidulous-margined 3/,-1 mm long beak; mouth
subentire or bidenticulate. Nut ellipsoid or ovoid,
triquetrous with flat faces, densely puncticulate,
dilate-annulate at the apex, 1!/,-1/, by */;-1 mm.
Style-base thickened. Stigmas 3.

Distr. Malesia: Sumatra (G. Leuser), SW.
Celebes (Mt Pokapindjang), New Guinea (W.:
Carstensz Mts; NE.: Mt Hagen, Mt Michael,
Mt Piora, Mt Wilhelm, Mt Sarawaket, Sattelberg;
Papua: S. Highlands, Mt Ambua, Mt Giluwe,
Mt Victoria, Owen Stanley Range.

Ecol. Open places, mountain heaths, alpine
grasslands and swamps; 2400-4040 m.

Vern. New Guinea: tudik, Mendi language.

Note. The original descriptions of both Carex
celebica and C. constricta were based on dwarfy
specimens a few cm tall. Collections recently made

140

FLORA MALESIANA

[ser. I, vol. 9!

in New Guinea have shown that the species can
reach a height of 25 cm and the leaves a width of
5 mm. Stout specimens are very similar to 20.
C. verticillata, a close ally of C. celebica, but easily
distinguished by the long stipitate 4'/,-6'/, mm
long utricles in the latter.

18. Carex perakensis C. B. CLARKE, FI. Br. Ind. 6
(1894) 720; J. Linn. Soc. Bot. 37 (1904) 9; RipDL.
Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 116; FI.
Mal. Pen. 5 (1925) 184; Uitr. Rec. Trav. Bot.
Néerl. 32 (1935) 201; NeLMeEs, Kew Bull. (1950)
189; Reinwardtia 1 (1951) 253; Mém. Mus. Hist.
Nat. Paris n.s. 4 (1955) 114; RAym. Mém. Jard.
Bot. Montréal n. 53 (1959) 74, f. 18; Koyama, J.
Fac. Sc. Un. Tokyo III, 8 (1962) 156; Raym.
Dansk Bot. Ark. 23 (1965) 259. — ? C. arridens
CLARKE, FI. Br. Ind. 6 (1894) 726; Kuk. Pfl. R.
Heft 38 (1909) 548; RipL. Mat. Fl. Mal. Pen.
(Monoc.) 3 (1907) 117; Fl. Mal. Pen. 5 (1925) 184;
NELMES, Reinwardtia 1 (1951) 333; Mém. Mus.
Hist. Nat. Paris n.s. B4 (1955) 139; RAvym. Mém.
Jard. Bot. Montréal n. 53 (1959) 74. — C. tonkinen-
sis FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris
III, 8 (1896) 251; Kix. Pfl. R. Heft 38 (1909) 292;
NELMES, Kew Bull. (1950) 190; Reinwardtia 1
(1951) 254; Ray. Nat. Canad. 82 (1955) 165, f. 5.
— C. wightiana NEES var. perakensis KUK. Pfi. R.
Heft 38 (1909) 288. — C. leucostachys RipL. Kew
Bull. (1928) 77; NeELMEs, Reinwardtia 1 (1951) 251;
Mém. Mus. Hist. Nat. Paris n.s. 4 (1955) 113. —
C. nodiflora (non BorEcKk.) Ktx. Bull. Jard. Bot.
Btzg III, 16 (1940) 316. — C. pseudorivulorum KuxK.
l.c. 319, quoad specim. cit., non C. rivulorum RIL.
(basionym). — C. setulifolia NELMEs, Mém. Mus.
Hist. Nat. Paris n.s. B4 (1955) 116. — Fig. 119,
131b-d.

Further synonyms under the varieties.

Rhizome short or shortly creeping (thick)
woody. Stems tufted, erect, trigonous or triquet-
rous, smooth or sparsely scabrid on the angles,
especially above, 15-175 cm by !/,-4 mm below,
surrounded below the leaves by brown to purplish
brown bladeless sheaths or their fibrous remains.
Leaves basal and subbasal, rarely spaced through-
out the stem, shorter than to exceeding the stem,
flattish, rather stiff, scabrid or smooth on the
margins, asperous towards the long acuminate
apex, (2-)3-16 mm wide; sheaths often dark
nerved, hispidulous or puberulous at the mouth.
Inflorescence usually consisting of 2-7 fascicles, or
spikelets single, binate or ternate at each node; the
fascicles formed by single spikelets (2-14 spikelets
in the whole inflorescence), or by up to 20 panicles
(in the whole inflorescence) ; the panicles consist of a
very lax raceme of up to 10 secondary spikelets,
each secondary spikelet subtended by a whether or
not sheathing glumiform, often long awned
bractlet, and surrounded at the base by an ocrei-
form cladoprophyll. The lower fascicles or spike-
lets distant, on long exserted smooth peduncles,

the higher approximate. Lower bracts foliaceous,
long sheathing, the sheaths often ampliate; higher
bracts much reduced, when glumiform the inflores-
cence is terminated by a seemingly more compound
panicle. Spikelets androgynous, cylindric, but the
3 part tapering, up to 10 in each panicle, the 9 part
mostly lax flowered, from much shorter to longer
than the 3 part, 3-8 mm thick. Glumes oblong to
ovate or lanceolate acute to obtuse or acuminate,
glabrous or hispidulous, nerved, light or red brown,
or whitish to fulvous, usually with whitish hyaline
margins, 2'/,-7 mm long, the midrib just below the
apex apiculate or excurrent into an up to 5 mm
long awn. Urtricles trigonous, oblong to narrowly
ellipsoid, or obovoid, 5—9!/, by 1—21/; mm, sub-
erect to patulous, straight or obliquely bent at the
apex, many-nerved, hispidulous, shortly stipitate
or sessile, subabruptly beaked or gradually
narrowed, beak bidentate, often with oblique
mouth, 1'/,-3 mm long. Nut trigonous with flat to
slightly concave faces, ellipsoid to oblong ellipsoid
or obovoid, shortly stipitate and beaked, 3-4 mm
long. Style-base slightly thickened. Stigmas 3.

Distr. S. China (Yunnan and Kwangsi), For-
mosa, Tonkin, Annam, Laos, Lower Burma,
Thailand; in Malesia: Sumatra (Atjeh, East Coast,
Palembang), Malay Peninsula (Perak, Larut,
Pahang, Selangor), Borneo, SW. & SE. Celebes.

Ecol. See under the varieties.

Notes. The species is very variable, three varie-
ties being distinguished. Carex borneensis seems at
first sight to be a different species, although closely
allied to C. vansteenisii s.s., because in most of the
material of C. borneensis the spikelets are not com-
pound. In the Malay Peninsula, however, this
difference fades away, the spikelets of C. borneensis
becoming also compound. Carex vansteenisii, a
very rare plant, possesses exactly the same inflores-
cence as C. perakensis, the utricles are narrower,
and their length overlaps with the range of those of
C. perakensis (7-9 mm in C. vansteenisii, S5—
6(—8) mm in C. perakensis).

NELMES distinguished also between C. borneensis
and C. kinabaluensis, although there are no
differences at all between the two. The Celebes
collection (C. eymae NELMES) is young and differs
hardly from the Bornean specimens of C. borneen-
sis.

Carex kinabaluensis was considered an ally of 57.
C. brunnea, as STAPF made a mistake in describing
it as having 2 stigmas (in fact the number of stigmas
is the only difference between the two sections!).

In the specimens of C. borneensis from the Malay
Peninsula, as mentioned in the first note, some of
the spikelets are branched (into 2-4 secondary
spikelets). On account of this NELMES maintained
C. breviglumis RIDL. as a species distinct from the
Bornean plants of C. borneensis, in which the
spikelets are usually unbranched. However, this
difference does not always hold.

NELMES considered C. tonkinensis and C. leuco-

1979]

stachys specifically distinct from C. perakensis s.s.,
but I could not find differences, and his key
characters are unfit for discrimination.

Carex leucostachys has been recorded from
Pahang (P. Tioman), S. Sumatra (G. Pesagi), and
Tonkin. In the type collection the inflorescence is a
dense, head-like panicle, and the utricles are about
8 mm. The inflorescence is still too young for good
description, but it shows resemblance with C. van-
steenisii s.s. in the utricles, and several other
collections show transitions to C. perakensis s.s.

CLARKE based C. arridens on two collections
(Lower Burma, KuRzZ, n.v., and Perak, KING’s coll.
2801). The Perak specimen, the only one ever
collected in Malesia (in 1882) is very young with
quite undeveloped flowers. It has the dark-nerved
basal sheaths and the pale indumentum of the
young utricles of C. perakensis s.s. The glumes are
darker than is usual. In 1951 NeELMes placed it in
subg. Carex, but C. perakensis, C. tonkinensis and
C. leucostachys in subg. Indocarex. The two sub-
genera he distinguished by their cladoprophylls
(utriculiform in subg. Indocarex, ocreiform in
subg. Carex). He described the cladoprophylls of
C. perakensis as “‘utriculi-glumiform”, those of
C. tonkinensis as ‘“‘more or less ocreiform” of
C. leucostachys as “‘subocreiform below, glumiform
above” and of C. arridens as “subutriculiform
below, glumiform above”. In 1955 he removed
C. tonkinensis to subg. Carex, C. leucostachys and
C. perakensis he left in subg. Indocarex.

KEY TO THE VARIETIES

1. Spikelets single or binate at the nodes, not com”
pound. Utricles 5—91/, mm, gradually tapering
into the 2-3 mm long beak. Leaves 2-8 mm wide

b. var. borneensis

1. Spikelets compound, or when simple in fascicles
at the nodes.

2. Utricles densely pale to golden hispidulous,
subabruptly beaked, sessile, ellipsoid or
ellipsoid-obovoid, 5—6(-8) mm long. Glumes
translucent, whitish, or fulvous with broad
whitish margins, 31/,-51/, mm long. Inflores-
cence consisting of up to 20 panicles which are
single, binate or ternate at the nodes. Each
panicle consists of a raceme of up to 10 shortly
peduncled 8-40 mm long secondary spikelets
which is sometimes branched. The ¢ part of
the spikelets from much shorter to longer than
the 2 part. Number of 92 flowers usually less
than lO. tear. TREES, . a. var. perakensis

2. Utricles brown or olivaceous, gradually
beaked, shortly stipitate, narrowly ellipsoid or
fusiform, 5—9!/, mm. Glumes light brown or
reddish brown, the margin often whitish
hyaline, 2'/,-7 mm long. Inflorescence con-
sisting of 2-14 spikelets branched into 2-4
secondary spikelets, single or binate at the
nodes, or 4-7 fascicles of spikelets or panicles.

Cy PERACEAE—II (Kern & Nooteboom)

141

3. Inflorescence consisting of 2-14 spikelets,
which are often branched into 2-4 secondary
spikelets, the ¢ part shorter to much longer
than the 2 part. Glumes 2'/,-4 mm long, light
brown with whitish hyaline margins

b. var. borneensis

3. Inflorescence consisting of 4~7 fascicles of
spikelets or panicles. The panicles consisting
of a very lax raceme of up to 7 secondary
spikelets. Spikelets (or secondary spikelets)
up to 10 in each fascicle, lax flowered, the 2
part much longer than the ¢ part. Glumes
brown or reddish brown, 5—7 mm long

c. var. vansteenisii

a. var. perakensis. — All synonyms under the
species. — Fig. 119, 131c-d.

Stems tufted, erect, trigonous or triquetrous,
smooth or sparsely scabrid on the angles above,
60-175 cm by 2-4 mm. Leaves 4-12(-16) mm wide.
Spikelets 1-2(-4) cm long, the 3 part much shorter
than the 5-7 mm thick 2 part. Awn of glumes up to
11/, mm long. Beak of the 1'/,-21/; mm wide
utricles 11/,-2(-3) mm long, often obliquely bent.

Distr. China (Yunnan, Kwangsi), Formosa,
Tonkin, Annam, Thailand, Lower Burma; in
Malesia: Sumatra (Atjeh, E. Coast, Palembang),
Malay Peninsula (Perak: Larut; Pahang, Selangor),
Borneo (Sarawak: Mt Dulit; Sabah: Mt Kinabalu),
SW. Celebes (Mt Poka Pindjang).

Ecol. In swampy localities and primary forest,
750-1700 m, on Mt Kinabalu up to 2700 m.

b. var. borneensis (CLARKE) Noot., comb. nov. —
C. borneensis CLARKE, J. Linn. Soc. Bot. 37 (1904)
14; Kew Bull. add. ser. 8 (1908) 76; KUx. Bull.
Jard. Bot. Btzg III, 16 (1940) 320, incl. var.
clemensii (KUK.) KUtx. f. angustifrons KUK.;
NELMES, Reinwardtia 1 (1951) 351. — C. fusiformis
(non NEES) STAPF, Trans. Linn. Soc. II, 4 (1894)
246. — C. fusiformis NEES var. borneensis (CLARKE)
Kwx. Pfl. R. Heft. 38 (1909) 598. — C. kinabaluen-
sis STAPF, J. Linn. Soc. Bot. 42 (1914) 183;
NeLmes, Kew Bull. (1950) 200; Reinwardtia 1
(1951) 349; ibid. 2 (1954) 377; RayM. Mém. Jard.
Bot. Montréal n. 53 (1959) 82. — C. rivulorum
Rip. J. Fed. Mal. St. Mus. 6 (1915) 195, non
Dunn, 1908. — C. breviglumis Riwv. Fl. Mal. Pen.
5 (1925) 183; NELMES, Reinwardtia 1 (1951) 348. —
C. clemensii KUxK. in Fedde, Rep. 29 (1931) 202. —
C. pseudorivulorum KUx. Bull. Jard. Bot. Btzg III,
16 (1940) 319, quoad basionvm. — C. eymae
NeLmes, Kew Bull. (1950) 199; Reinwardtia 1
(1951) 352. — Fig. 119, 131b.

Stems erect, triquetrous, smooth or slightly
scabrid, (15—)40-140 cm. Leaves shorter than to as
long as the stem, (2-)3-5(-8) mm wide. Inflores-
cence narrow, up to 30 cm long, consisting of 2 to
14 spikelets. Spikelets up to 5 cm long, single or
binate at the middle nodes, erect or suberect, simple
or branched into 2-4 secondary spikelets, the ¢

142

FLORA MALESIANA

[ser. I, vol. 94

part shorter to much longer than the lax-flowered
3-5 mm thick 2 part. Glumes light brown with
whitish hyaline margins, 2'/,-4 mm long, apiculate
or midrib excurrent into an up to 5 mm long awn.
Utricles olivaceous, 5—91/, by 1-2 mm, gradually
tapering into a hispidulous margined, 2-3 mm
long beak.

Distr. Annam (Prov. Kontum: Ngoe Pang); in
Malesia: Malay Peninsula (Pahang: G. Tahan,
G. Jasar, G. Berumban), Borneo (Sabah: Mt
Kinabalu, Mt Tambuyokan, G. Alab; Sarawak:
Mt Poé, Mt Dulit; SE. Borneo: W. Kutei), SE.
Celebes (Enrekang). See the map by RAYMOND,
Kes ps03:

Ecol. Wet places in forests and in shady places
along streams; 800-3300 m.

Note. Specimens with very regularly puckered-
bullate leaves were twice collected on Mt Kinabalu.
The strange abnormality of the leaves may be due
to some disturbance during the growth period. The
same phenomenon has been observed in some
Aponogeton, Cryptocoryne, and Halophila species.

c. var. vansteenisii (KUK.) Noot., comb. nov. —
C. vansteenisii KUK. Bull. Jard. Bot. Btzg III, 16
(1940) 320, incl. var. brevispiculosa KUK. I.c. 321;
NELMES, Reinwardtia 1 (1951) 444; Mém. Mus.
Hist. Nat. Paris n.s. B4 (1955) 141; Raym. Mém.
Jard. Bot. Montréal n. 53 (1959) 73. — Fig. 119.

Stems rather stout, erect, triquetrous, smooth,
c. 150 cm by 3-4 mm. Leaves shorter than the stem,
stiff, 3-15 mm wide, sheaths shining membranous
in front. Inflorescence consisting of 4-7 fascicles
of panicles consisting of a very lax raceme of up to
7 secondary spikelets. Spikelets lax-flowered,
2-6 cm long, 4-8 mm thick, the 2 part much longer
than the ¢ part, from c. 4- to c. 10-flowered. Glumes
(reddish) brown, 5-7 mm long, awn 0-1 mm long.
Utricles exceeding the glumes, suberect to patulous,
gradually narrowed into the 2-3 mm long beak,
7-9 by 1-1!/, mm.

Distr. Tonkin, Laos; in Malesia: N. Sumatra
(Atjeh: Losir massif).

Ecol. Mountain heaths, 2100-2500 m. In the
Losir area this variety is found at higher altitude
than var. perakensis.

19. Carex turrita C. B. CLARKE, J. Linn. Soc. Bot.
37 (1904) 13; Philip. J. Sc. 2 (1907) Bot. 108; Kew
Bull. add. ser. 8 (1908) 74; Kix. Philip. J. Sc. 5
(1911) Bot. 63; Merr. En. Philip. 1 (1923) 142;
NELMES, Reinwardtia 1 (1951) 335. — Fig. 119,
131e-f.

See for synonyms under the varieties.

Rhizome short or shortly creeping, woody.
Stems erect, trigonous, smooth, up to more than
1 m by 4/,-3 mm below. Leaves basal and up to
3 higher on the stem, shorter than the stem, stiff,
flat or with slightly revolute margins, with scabrid
margins and asperous upper surface towards the
long attenuate apex, 2-7 mm wide; sheaths usually

reddish or purplish, glabrous, with concave to
convex mouth, the basal ones fraying into fibres.
Inflorescence an often interrupted slender panicle to
50 cm long, consisting of 3-8 fascicles, lower fas-
cicles distant, upper approximate. Lower bracts
foliaceous, shorter to longer than their fascicles
but much exceeded by the stem, long sheathing,
upper much reduced; sheaths similar to those of
the stem. Lower spikelets often single or binate,
upper 2-6 together, often branched into 2-6
secondary spikelets, all androgynous, the 3 part
from much shorter to longer than the 9 part,
2-8 cm by 2-3'/, mm, much exserted from their
sheaths. Glumes oblong-ovate, obtuse to emargi-
nate, glabrous or sparsely hispidulous towards the
apex, dark red with wide white hyaline margins,
nerved, 2—3'/, mm long, the midrib excurrent into
an up to 2 mm long hispidulous awn. Utricles
trigonous, oblong to ellipsoid, slenderly nerved to
nerveless except 2 submarginal nerves, dark red,
reddish or brown, straight or slightly curved,
glabrous or hispidulous, at least above, the margins
sparsely hispidulous from the middle or above the
middle upwards, 3-6 by */,-1 mm, gradually taper-
ing into a hispidulous margined, 1-2 mm long
bidentate beak. Nut triquetrous, ellipsoid, with
flattish or concave faces, brown or fulvous, stipi-
tate, 11/,-3!/, mm long. Style-base slightly
thickened. Stigmas 3.
Distr. Malesia: Philippines and New Guinea.

a. var. turrita. — C. turrita C. B. CLARKE. —
C. walkeri ARN. ex Bootr var. turrita (CLARKE)
Kix. Pfi. R. Heft 38 (1909) 546. — C. atrosanguinea
NeELMeES, Kew Bull. (1950) 197; Reinwardtia 1
(1951) 339. — Fig. 119, 131e-f.

Glumes 3-3!/, mm long, with an awn of ?/,-
2 mm. Utricles slenderly nerved, 5-6 mm long,
with a beak of 2 mm. Nut 3-3!/, mm long.

Distr. Malesia: Philippines (Luzon: Abra,
Bontoc, Ifugao, Benguet), New Guinea (W. New
Guinea: Mt Treub; NE. New Guinea: Morobe
Distr.; Papua: W. Highlands, Kubor Range,
E. Highlands, Central Div.).

Ecol. Open places in mossy forest, montane
rain-forest, 1500-3040 m.

Vern. Philippines: tangtafigo, Bon.

Notes. NELMES distinguished the New Guinean
C. atrosanguinea from the Philippine C. turrita by
the simple spikelets of the former. Additional
collections from New Guinea have shown that
also here specimens with branched spikelets occur;
it must, however, be admitted that in general
the spikelets in the Philippines are more com-
pound.

I have not seen C. gibbsiae RENDLE, J. Linn. Soc.
Bot. 39 (1909) 180, from Fiji, which is, according to
NELMES, closely related to his C. atrosanguinea, but
looking ‘“‘very distinct because of its different
colouring”. I suspect it to be also conspecific with
C. turrita.

1979]

CyYPERACEAE—II (Kern & Nooteboom)

143

b. var. merrillii (KUK.) Noort., comb. nov. —
C. merrillii KOK. in Fedde, Rep. 8 (1910) 7; Philip.
J. Sc. 6 (1911) Bot. 63; Merr. En. Philip. 1 (1923)
139; NeLMes, Reinwardtia 1 (1951) 336; ibid. 2
(1954) 377. — C. pullei NELMES, Kew Bull. (1950)
198; Reinwardtia 1 (1951) 338. — Fig. 119.

Glumes 2-2'/, mm long, awn up to '/; mm.
Utricles nerveless to slenderly few-nerved,
3-33/, mm long, with a beak of 1-1'/, mm. Nut
11/,-13/, mm long.

Distr. Malesia: Philippines (Luzon, Ifugao,
Benguet), New Guinea (W. New Guinea: Mt
Treub; NE. New Guinea: Morobe Distr.).

Ecol. Mossy forest, open places along trails,
also on steep slopes; 1800-2490 m.

Note. KERN (in manuscript) reduced C. pullei to
C. turrita with a question-mark. He noted: “It
seems to differ mainly by the smaller utricles only
3-4 mm long, and may represent a depauperated
state’. NELMES described the utricles as being
“almost nerveless to rather slenderly 1—-3-nerved”’.
As these were the only differences between C. tur-
rita and C. merrillii, | have reduced the latter to a
variety of the former.

20. Carex verticillata ZoLL. & Mor. in Mor. Syst.
Verz. (1846) 98; STEUD. Syn. 2 (1855) 222; Mia.
Fl. Ind. Bat. 3 (1857) 353; Boeck. Linnaea 41
(1877) 256; NeLMes, Kew Bull. (1950) 195, incl.
var. havilandii (CLARKE) NELMES ef var. lutescens
NELMES; Reinwardtia 1 (1951) 340; Raym. Mém.
Jard. Bot. Montréal n. 53 (1959) 74; KERN in Back.
& Bakh. f. Fl. Java 3 (1968) 488. — C. hypsophila
Mia. Fl. Ind. Bat. 3 (1857) 354; Boeck. Linnaea
41 (1877) 257; CLARKE, J. Linn. Soc. Bot. 37 (1904)
13; KUxK. Pfl. R. Heft 38 (1909) 546, f. 89, incl. var.
havilandii (CLARKE) KUK. et var. verticillata (ZOLL.
& Mor.) KuK.; in Hochr. Candollea 6 (1936) 432.
— C. walkeri (non Boott) Boeck. Linnaea 40
(1876) 332, p.p.; BACK. Bekn. Fl. Java (em. ed.) 10
(1949) fam. 246, p. 65. — C. tartarea RIL. J. Bot.
23 (1885) 35. — C. havilandii CLARKE, J. Linn.
Soc. Bot. 37 (1904) 13; Kew Bulli. add. ser. 8 (1908)
75. — C. sumatrensis CLARKE, J. Linn. Soc. Bot. 37
(1904) 13; Kew Bull. add. ser. 8 (1908) 75. —
C. turrita (non CLARKE) KUxK. Bull. Jard. Bot.
Btzg III, 16 (1940) 319. — C. decora BootrT var.
losirensis KUK. l.c. — C. phacelostachys NELMES,
Kew Bull. (1950) 195; Reinwardtia 1 (1951)
344, incl. var. losirensis (KUK.) NELMES; RAYM.
Mém. Jard. Bot. Montréal n. 53 (1959) 72. — Fig.
119.

Rhizome creeping, woody. Stems solitary or
somewhat tufted, erect, triquetrous, smooth,
25-100 cm by 1-2 mm, surrounded below the leaves
by dark reddish, entire or fibrous remains of old
leaf-sheaths. Leaves basal and 1-2 on the stem
proper, shorter than to equalling the stem, flattish
with often strongly revolute margins, stiff, coria-
ceous, with scabrid margins, asperous above
especially towards the long-attenuated apex,

3-8 mm wide; sheaths of the lower leaves reddish
to blackish red, upper ones pale. Inflorescence a
lax to dense panicle, 15-50 cm long, consisting of
4~7 fascicles of up to 20 spikelets, lower fascicles
rather distant, upper approximate. Bracts of the
lower fascicles foliaceous, equalling or exceeding
their fascicles but usually much exceeded by the
whole inflorescence, long-sheathing, upper much
reduced; sheaths ampliate, glabrous, the mouth
often prolonged into a short tongue. Spikelets
erect or somewhat nodding, slenderly cylindric,
lax- to rather dense-flowered, up to 6 cm long,
simple or longest ones sometimes branched into
1-3 short secondary spikelets, on slender, smooth
peduncles more or less exserted from the sheaths;
1-2 spikelets of uppermost fascicle (sometimes of
all the fascicles) g, remaining ones wholly 2 or
with a few ¢ flowers at the top; g spikelets 1-2 mm
thick, 2 3-5 mm. Glumes oblong-lanceolate, ob-
tuse, glabrous, fulvous to dark reddish with wide
whitish-hyaline margins, 3-43/, mm long, the
midrib often excurrent in a smooth or hispidulous
awn up to 1 mm long. Utricles trigonous, narrowly
ellipsoid, tapering at both ends, glabrous but the
margins often hispidulous, nerveless or faintly
nerved (except for the marginal or submarginal
nerves), suberect to patulous, often somewhat
recurved, long-stipitate, stipe (*/,-)3/,-1 mm,
subabruptly beaked, blackish red to golden,
4-6'/, by */,-1 mm; beak almost linear, sparsely
hispidulous-margined, bidenticulate, (1-)2-3 mm
long. Nut triquetrous with flattish faces, ellipsoid,
brown, 1°/,-2 mm long. Style not or scarcely
thickened at the base. Stigmas 3.

Distr. Laos, Tonkin; in Malesia: Sumatra,
Malay Peninsula (Perak: G. Kerbau), Java,
N. Borneo (Mt Kinabalu).

Ecol. Damp open places in forests, open places
in the subalpine region, near craters; 2000-3750 m.

Notes. Variable as to the colour of glumes
and utricles. In most of the Mt Kinabalu speci-
mens the glumes and utricles are very dark; they
were distinguished as var. havilandii (CLARKE)
NELMES.

A collection from E. Java (G. Semeru), in which
the glumes and utricles are pale golden to fulvous,
was distinguished as var. /utescens NELMES.

Carex phacelostachys NELMES only could be
distinguished by the utricles which are faintly
nerved against nerveless in C. verticillata. The
collection VAN STEENIS 9624 from Atjeh has been
identified as C. verticillata by both NeLMes and
Kern; after careful examination of the utricles I
could not find any difference with C. phacelostachys.
I examined many utricles of C. verticillata with
translucent light. Most of them are nerveless
indeed, but others are faintly nerved, often only at
the base, at least in plants from Sumatra and
Borneo. — (Noot.)

The species might also be conspecific with
C. walkeri ARNOTT ex Bootr. — (Noot.)

144

FLORA MALESIANA

[ser. I, vol. 91

7. Section Surculosae

Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 21. — Sect. Scabrellae KUx. Pfl.
R. Heft 38 (1909) 286, p.p.; NELMES, Reinwardtia | (1951) 243.
Type species: Carex oligostachya NEES (lectotype).

21. Carex oligostachya Nees in Hook. J. Bot. Kew
Misc. 6 (1854) 29; NeLmes, Reinwardtia 1 (1951)
244; ibid. 2 (1954) 373; Kew Bull. (1955) 301;
Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 95;
RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 22.
— C. cumingiana StEuD. Syn. 2 (1855) 206; Mia.
FI. Ind. Bat. 3 (1856) 349; Boorr, Ill. 3 (1862) 107,
t. 324, 325; Boeck. Linnaea 40 (1876) 367;
F.-VILL. Nov. App. (1882) 310. — C. rhizomatosa
STeupD. [in Zoll. Syst. Verz. 1 (1854) 60, nomen]
Syn. 2 (1855) 206; Mia. Fl. Ind. Bat. 3 (1856) 348;
CLARKE, FI. Br. Ind. 6 (1894) 721; J. Linn. Soc.
Bot. 37 (1904) 12; Philip. J. Sc. 2 (1907) Bot. 108;
Kuk. Pfl. R. Heft 38 (1909) 289, f. 44, excl. var.
impunctata (BooTT) KUK.; Philip. J. Sc. 6 (1911)
Bot. 61; Camus, Fl. Gén. I.-C. 7 (1922) 193;
Merr. En. Philip. 1 (1923) 141; KUxK. Bull. Jard.
Bot. Btzg III, 16 (1940) 316, incl. var. aristulata
Kwvx.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 107;
NELMES, Kew Bull. (1949) 378, 387. — C. bukaénsis
PALLA in Rechinger, Bot.-Zool. Ergebn. Samoa &
Solomon Ins. (1913) 58. — C. breviceps KUK. Bot.
Jahrb. 69 (1938) 263. — Fig. 120.

Rhizome shortly creeping, woody, like the stem-
base clothed with brown or blackish, fibrous
remains of old leaf-sheaths. Flowering stems lateral,
approximate, subscapiform (bearing a few short-
bladed leaves), triquetrous, smooth, 20-90 cm by
3/,—-2 mm. Leaves of the sterile shoots shorter than
the flowering stems, stiffish, flat, rather abruptly
acuminate, scabrid in the upper part, glaucous,
2-5 mm wide. Inflorescence a narrow panicle
consisting of 6-12 head-like or racemose partial
inflorescences, 10-50 cm long; partial inflores-
cences erect, subglobose to oblong-ovoid, with
1-8 spikelets, the lower ones single at the nodes,
distant, on long-exserted, setaceous peduncles, the
upper ones more approximate, unequally binate

(rarely ternate); axis and top of peduncles scabrid.
Bracts foliaceous with blades shorter than 8 cm,
their sheaths ampliate, membranous near the
mouth, the upper ones much reduced, infundibuli-
form. Spikelets androgynous, ovoid to subcylindric,
sessile, 4-10 mm long, the g¢ and 2 parts usually
about equal in length. G/umes ovate or lanceolate,
acutish to slightly notched, usually sparsely hispi-
dulous towards the apex, 5—10-nerved, brown, often
with reddish spots, whitish hyaline-margined,
2-3 mm long, the midnerve usually excurrent in a
short, scabrid awn up to 1 mm long. Utricles
obtusely trigonous, ellipsoid, subinflated, patent,
straight, glabrous or sparsely hispidulous in the
upper !/,, strongly many-nerved, rather abruptly
beaked, fuscous, 2!/,-4 by 1-2 mm; beak 3/,-1 mm
long, bidenticulate. Nut trigonous, ellipsoid or obo-
void, minutely stipitate, 2—21/, by 1-1!/, mm.
Style-base not thickened. Stigmas 3.

Distr. Assam, Upper Burma, S. China, Tonkin,
through Malesia to the Solomon Is.; in Malesia:
Sumatra (Atjeh, Tapanuli), Lesser Sunda Is. (Sum-
ba, Sumbawa, Flores), Philippines (Luzon, Negros,
Mindanao), SE. Celebes (Buton I.), New Guinea.
According to MERRILL, /.c., also in the Moluccas.

Ecol. In Jmperata-fields, on open grassy slopes,
often in places where the grass is burned annually;
from low altitude up to 1400 m.

Vern. New Guinea: simboro, Orokaiva lang.

Notes. The utricles vary from wholly glabrous
(C. breviceps Kix.) to sparsely hispidulous above.

I have not seen C. breviceps var. recurvirostris
Kx. Bot. Jahrb. 70 (1940) 464, based on CLEMENS
8032 from NE. New Guinea, Morobe, Sattelberg.
Stems 90 cm tall. Leaves 5-10 mm wide. Panicle
35 cm long. Spikelets longer, broadly oblong.
Utricles green, subabruptly excurrent in a long,
recurved beak.

8. Section Rhomboidales

KUK. Pfl. R. Heft 38 (1909) 622; NELMES, Reinwardtia 1 (1951) 383; RayM. Mém.

Jard. Bot. Montréal n. 53 (1959) 54, 66.

Type species: Carex chinensis RETZ. (lectotype).

22. Carex anomocarya NELMES, Kew Bull. (1950)
202; Reinwardtia 1 (1951) 383; Mém. Mus. Hist.
Nat. Paris n.s. B4 (1955) 163; RayM. Mém. Jard.
Bot. Montréal n. 53 (1959) 66, f. 16, map p. 102,
f. 32; Dansk Bot. Ark. 23 (1965) 259; KERN in
Back. & Bakh. f. Fl. Java 3 (1968) 489. — C. har-
landii (non Boott) Merr. & CHUN, Sunyatsenia 2

(1935) 208. — C. harlandii BootT var. angustior
Kuk. ex Back.; Gross, Notizbl. Berl.-Dahl. 14
(1938) 193; Back. Bekn. Fl. Java (em. ed.) 10
(1949) fam. 246, p. 65. — C. harlandii Boorr
f. longibracteata Gross, Notizbl. Berl.-Dahl. 14
(1938) 193. — C. manca Bootrt var. contigua GROSS,
l.c. — Fig. 120, 128.

1979]

145

\\ |
ite
1h}
FT
iE Bl}

TITAS

MMMM LL

Fig. 128. Carex anomocarya NELMES. @. Habit, x 2/3, b. glume, c. fruit in utricle, d. fruit, all x 6 (RAHMAT
st BoEEA 10601).

146

FLORA MALESIANA

[ser. I, vol. 9!

Rhizome short, not creeping. Stems central,
erect, triquetrous, smooth, 30-70 cm by 1'/,—
2 mm. Leaves basal (rarely one in the middle of the
stem), flat, weak, scabrid on the margins, 6-15
(-20) mm wide. Inflorescence with 3-5 spikelets,
erect; terminal spikelet g, cylindric, 21/,-5 cm
by c. 2 mm, peduncled, lateral ones 2, oblong-
cylindric, approximate with one another and with
the 3 spikelet, but lowest often distant, subdense-
flowered, on shortly exserted peduncles, 4-8 cm by
6-9 mm; peduncles stoutish, smooth. Bracts
foliaceous, lower usually much exceeding the
inflorescence, shortly sheathing; sheaths pale,
membranous in front or only at the mouth, sub-
ampliate. Glumes of the 2 spikelets oblong, sub-
truncate to slightly emarginate, dirty white with
hyaline margins and 3-nerved centre, ciliolate at
the apex, awned, 3-4'/, mm long, those of the 3
spikelets narrower, also long-awned; awns scabrid,
up to 1 cm long. Urtricles ellipsoid to rhomboid-
lageniform, obtusely trigonous, surface uneven
with concavities and convexities corresponding to
the surface of the nut, densely many-nerved,
coriaceous, glabrous, straight, suberect, shortly
stipitate, stramineous, 7—8(-10) by 2-23/, mm,
rather abruptly narrowed into a 2'/,-4 mm long,
conical, bidentate beak; teeth 1-1'/, mm long,
diverging, sparsely hispidulous at the mouth. Nut
unevenly ellipsoid or ovoid, distorted-trigonous,
excavate on the angles at the centre, with concave
faces below, stipitate, abruptly beaked, 4—5!/, mm
long; beak cylindric, '/,-11/, mm long, expanding
into the annulate apex. Style-base pyramidally
thickened, persistent. Stigmas 3.

Distr. N. Burma, NE. Thailand, Tonkin,
Annam, Hainan; in Malesia: N. Sumatra (E.
Coast: Dolok Singgalang and G. Batu Lopang;
Toba), W. Java (Priangan: Tjadas Malang S. of
Tjiandjur; Mt Bésér near Tjidadap).

Ecol. In damp shady forest, in W. Java 1000-
1200 m, in N. Sumatra at 1400-1700 m.

Note. Very near to C. harlandii Bootr, Ill. 2
(1860) 87, t. 255; Kix. Pfl. R. Heft 38 (1909) 630,
f. 107, and possibly better treated as a race of that
species. In C. harlandii the leaves are broader (up to
3 cm wide), the bracts much shorter, often not or
hardly overtopping spikelet, the glumes of the
3 spikelet rounded at the apex and muticous, those
of the 2 spikelets but shortly awned, the teeth of
the utricles shorter (c. 1/, mm long), and the beak
of the nut slenderer, c. 11/, mm long. It is known
from South and Central China.

23. Carex jackiana Boott, Proc. Linn. Soc. 1
(1845) 260; Trans. Linn. Soc. 20 (1846) 132; Ill. 1
(1858) 9, t. 25; SteuD. Syn. 2 (1855) 226; Mio. FI.
Ind. Bat. 3 (1856) 353; Boeck. Linnaea 41 (1877)
277; CLARKE, FI. Br. Ind. 6 (1894) 735, excl. var. 8;
J. Linn. Soc. 37 (1904) 15; Kix. Pfl. R. Heft 38
(1909) 638; in Hochr. Candollea 6 (1936) 432,
excl. var. tumens KUK.; BACK. Bekn. FI. Java

(em. ed.) 10 (1949) fam. 246, p. 66; NELMEs,
Reinwardtia 1 (1951) 384; Koyama, Bot. Mag.
Tokyo 70 (1957) 352, f. 10 & 12 A-A’; ibid. 72
(1959) 303; J. Fac. Sc. Un. Tokyo III, 8 (1962) 230;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 490;
STEEN. Mt. Fl. Java (1972) pl. 14-3.

ssp. jackiana. — Fig. 120.

Rhizome short. Stems central, tufted, erect,
triquetrous (angles prominent to narrowly winged),
smooth, 15-100 cm by 1-2 mm, surrounded at the
base by a few brownish sheaths or their fibrous
remains. Leaves subbasal, sometimes one higher on
the stem, shorter than to as long as the stem, flat,
rather weak, scabrid at the top, 3-10 mm wide;
sheaths pale to whitish. Inflorescence with 3-7
spikelets, erect; terminal spikelet 3 (sometimes
gynaecandrous), slenderly cylindric (or clavate),
1-3 cm by 2-4 mm, lateral ones 2 (rarely andro-
gynous), oblong-cylindric, sublax-flowered, 1-3 cm
by 5-8 mm, upper erect, crowded with the ¢
spikelet, sessile or very shortly peduncled, lower
distant on included to long-exserted, slender,
smooth peduncles, often with 1-3 shorter spikelets
branching from their peduncles. Bracts foliaceous,
exceeding the inflorescence, lower long-sheathing,
upper much shorter, scarcely to shortly sheathing.
Glumes oblong-ovate, acute, very thin, dirty white
with 3-nerved, greenish, central stripe, muticous
or mucronulate, rarely with a short awn, 3—5!/, mm
long. Utricles fusiform-ellipsoid, trigonous, sub-
coriaceous, densely and strongly many-nerved,
glabrous, straight, suberect, shortly stipitate,
olivaceous, 51/,-7!/, by 11/,-2 mm, gradually
narrowed into the conical, straight, bidentate
beak; teeth '/,*/, mm long, hardly diverging,
smooth. Nut oblong-obovoid to suborbicular,
triquetrous with prominent angles, with faces
shallowly concave below, very shortly stipitate,
abruptly shortly beaked, 2'/,-3 mm long. Style-
base not thickened. Stigmas 3.

Distr. Ceylon, India (Assam, Khasia), Yunnan;
according to NELMEs (1951) 386 also in Australia;
in Malesia: Sumatra (W. Coast: Mt Kerintji),
West and Central Java. Distribution maps in Bot.
Mag. Tokyo 72 (1959) 303; Phytologia 17 (1968)
408. Ssp. parciflora (BooTT) KUK., often considered
specifically distinct, differs from ssp. jackiana by its
smaller utricles and shorter glumes; in S. Sachalin,
Japes, S. Kuriles, Kyushu, and S. Korea.

Ecol. In marshes, swamps, and grassy plains;
1350-2550 m.

24. Carex lateralis KUK. Pfl. R. Heft 38 (1909)
639; Bull. Jard. Bot. Btzg III, 16 (1940) 322; Back.
Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 66;
NELMES, Kew Bull. (1950) 204; Reinwardtia 1
(1951) 388; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 488. — C. jackiana BootT var. minor
CLARKE, FI. Br. Ind. 6 (1894) 735, p.p. typ. —
C. elmeri KUxK. in Fedde, Rep. 8 (1910) 326;

1979]

Cy PERACEAE—II (Kern & Nooteboom)

147

Extmer, Leafl. Philip. Bot. 3 (1910) 853; Kuk.
Philip. J. Sc. 6 (1911) Bot. 64; Merr. En. Philip. 1
(1923) 137; KUK. Bull. Jard. Bot. Btzg III, 16
(1940) 322. — Fig. 120.

Rhizome short, tufted. Flowering stems lateral,
obliquely erect or somewhat cernuous, very slender
to subfiliform, weak, triquetrous, smooth, (S5—)20—
50 cm by '/,—1 mm, surrounded at the base by a few
brown sheaths or their fibrous remains, and
bearing a few small, bract-like or subfoliaceous
leaves. Leaves of the sterile shoots much over-
topping the stems, flat, revolute on the margins,
stiff, with very rough margins, asperous towards
the apex, 3-7 mm wide. Spikelets 3-4, crowded at
the apex of the stem, subsessile on shortly exserted
or wholly included peduncles, sometimes one
1-3 cm lower down; terminal spikelet 3, cylindric,
5-10 by '/,-1 mm, usually very inconspicuous when
the lateral spikelets have developed utricles, few-
flowered, lateral ones 9, oblong, few- (up to
5-)flowered, erect, 7-12 by 5S—7 mm. Bracts folia-
ceous, much exceeding the inflorescence, shortly
sheathing. Glumes oblong or _ oblong-ovate,
subacute to rounded, thin, whitish with 3-nerved
greenish central stripe, 2!/,—3'/, mm long, excurrent
in a flat, scabrous awn 1'/,-6 mm long. Utricles
ovoid-ellipsoid, trigonous, membranous, suberect,
many-nerved below but few nerves extending more
than halfway towards the apex, sparsely pubescent
to glabrous, stipitate, shiny, pale green, 5-7 by
2-2!/, mm, subabruptly contracted into a long,
conical, somewhat recurved, bidentate beak. Nut
ovoid-ellipsoid, triquetrous, with faces concave in
the lowest 1/,, brown to fuscous, stoutly stipitate,
scarcely or not beaked at the rounded apex, 3-4 mm
long. Style thickened at the base. Stigmas 3.

Distr. Ceylon, India; in Malesia: S. Sumatra
(Palembang: Air Njuruk), W. Java (Priangan:
Mt Papandajan), Central Celebes, Philippines
(Luzon, Mindoro, Negros), Lesser Sunda Is.:
Flores (Mt Ranaka).

Ecol. In forests, on road-sides; 1400-2200 m.

Note. The type of this species is THWAITES
CP 3198 p.p., not CLARKE 11061 as cited by
NELMES. CLARKE’S description of C. jackiana var.
minor, based on this THWAITES collection and on
CLARKE 11061 (!) refers to the species described
above. Carex jackiana var. minor is therefore
synonymous with C. /ateralis and cannot be main-
tained as a variety of C. jackiana alongside of C.
lateralis, as was done by KUKENTHAL. BOECKELER’S
description of C. jackiana (Linnaea 41, 1871, 277),
cited by CLARKE in the synonymy of his var.
minor, obviously refers to typical 23. C. jackiana.

VELDKAMP 7130 has the entirely hairy utricles of
C. lateralis, but lacks the awn on the glume; in that
respect it fits in 25. C. loheri. The leaves are also

intermediate between C. lJoheri and C. lateralis.
When more material becomes available, it is not
impossible that the two species will have to be
united. — (Noot.)

See also note under 25. C. loheri.

25. Carex loheri CLARKE, J. Linn. Soc. Bot. 37
(1904) 14; Philip. J. Sc. 2 (1907) Bot. 108; Kix.
Pfl. R. Heft 38 (1909) 487; Philip. J. Sc. 6 (1911)
Bot. 64, incl. f. grandimascula KUK.; MERR. En.
Philip. 1 (1923) 139; NeELMEs, Reinwardtia 1 (1951)
386. — Fig. 120.

Rhizome short, tufted. Flowering stems lateral,
suberect or cernuous, filiform, weak, triquetrous,
smooth, 10-40 cm by !/, mm, surrounded at the
base by a few brown scales or their fibrous remains.
Leaves crowded at the base, shorter than to exceed-
ing the stems, flat with revolute margins, stiff,
greyish or glaucous-green, with scabrid margins,
asperous towards the apex, 1!/,-3 mm wide. Spike-
lets 3-6, terminal g, cylindric, 7-15 by 1-2 mm,
lateral androgynous, 6-10 by 4-6 mm, with ¢ and
2 parts about equal in length and few-flowered,
upper one often approximate with the ¢ spikelet,
lower on long, subbasal ones pendulous on very
long, setaceous, smooth peduncles. Bracts of the
lower spikelets subbasal leaves, of the other lateral
spikelets subfoliaceous, rather long-sheathing.
Glumes oblong-ovate, acute, thin, many-nerved,
muticous or mucronulate, whitish, c. 4 mm long.
Utricles ovoid-ellipsoid, trigonous, membranous,
suberect, many-nerved, shining, glabrous below,
sparsely to densely pubescent above, shortly
stipitate, shining, pale green, 6-7 by 13/,-2 mm,
subabruptly contracted into a long, conical, some-
what recurved, bidentate beak. Nut ovoid or ellip-
soid, triquetrous with concave faces, brown to
fuscous, shortly stipitate, hardly beaked, 31/,4 mm
long. Style thickened at the base. Stigmas 3.

Distr. Malesia: Philippines (Luzon: Lepanto,
Bontoc, Benguet, Rizal, Zambales, Laguna,
Tayabas).

Ecol. Mossy forest, 1300-2400 m.

Vern. Silak, Ig.

Notes. Very similar to 24. C. lateralis, to which
it is certainly closely related, though KUKENTHAL
placed it in a different section.

Carex tatsutakensis HAYATA, Ic. Pl. Form. 6
(1916) 133, f. 45; OHw1, Mem. Coll. Sc. Kyoto Imp.
Un. B11 (1936) 396; Koyama, Nat. Canad. 82
(1955) 204; Contr. Inst. Bot. Un. Montreal n. 70
(1957) 19, not rare in Formosa and, according to
Koyama, also in Tonkin (not mentioned for this
country by RAYMOND!) is very near to C. loheri.

Also C. sublateralis KoyAMA, Jap. J. Bot. 15
(1956) 180, f. 9, from E. China (Kiangsu) belongs
to this group of very closely related species.

148 FLORA MALESIANA [ser. I, vol. 9!

9. Section Cryptostachyae

(OHw1) NELMES, Reinwardtia | (1951) 363; RAYM. Mém. Jard. Bot. Montréal n. 53
(1959) 52, 61. — Sect. Praecoces CHRIST subsect. Cryptostachydeae FRANCH. ex

OuwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 340.
Type species: Carex cryptostachys BRONGN.

26. Carex cryptostachys BRONGN. in Duperrey,
Voy. Coq. Bot. (1828) 152, t. 25; Mig. FI. Ind.
Bat. 3 (1856) 352; Boortrt, Ill. 3 (1860) 103, t. 310;
BENTH. FI. Hongk. (1861) 403; Boeck. Linnaea 40
(1876) 327; CLarKe, FI. Br. Ind. 6 (1894) 714
(‘cyrtostachys’); J. Linn. Soc. Bot. 36 (1903) 281;
ibid. 37 (1904) 8; RipL. Mat. Fl. Mal. Pen. (Monoc.)
3 (1907) 116; Kix. Pfl. R. Heft 38 (1909) 471;
Camus, FI. Gén. I.-C. 7 (1922) 195, f. 27, 10-13;
Merr. En. Philip. 1 (1923) 137; Rip. Fl. Mal.
Pen. 5 (1925) 181, f. 221; KUK. in Hochr. Candol-
lea 6 (1936) 432; OHwi, Mem. Coll. Sc. Kyoto
Imp. Un. B11 (1936) 342; Etmer, Leafl. Philip.
Bot. 10 (1938) 3526; Back. Bekn. Fl. Java (em.
ed.) 10 (1949) fam. 246, p. 66; NELMES, Reinwardtia
1 (1951) 363; AKIYAMA, Car. Far East. Reg. Asia
(1955) 176, t. 177; Raym. Nat. Canad. 82 (1955)
151, f. 2; Mém. Jard. Bot. Montréal n. 53 (1959)
61, map p. 102, f. 34; Dansk Bot. Ark. 23 (1965)
258; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 491.
—Fig. 120.

Rhizome elongate, horizontal or ascending,
woody, clothed with fibrous remains of sheaths.
Flowering stems arising from the axils of the leaves,
and often almost hidden among them, single or
binate, scapiform, suberect, flexuous, obtusely
trigonous, smooth, 10—50 cm tall, with sheaths and
peduncles almost from the base. Leaves crowded on
a short stem, much longer than the flowering stems,
firm, flat, long-acuminate, scabrid on the margins,
grey- or glaucous-green, 3-18 mm wide, surrounded
at the base with fibrous remains of older leaves.
Inflorescence racemiform or paniculiform, with
6-30 spikelets; spikelets androgynous, oblong-
lanceolate, lax-flowered, 8-30 by 3-5 mm; ¢ part

much shorter than the 2 one. Bracts usually shorter
than their spikelets, with funnel-shaped sheaths
and short blades; peduncles sparsely scaberulous,
lower exserted, upper included. G/umes ovate,
amplexicaul, many-nerved, obtuse, ciliolate, often
minutely appressed-hairy, whitish or stramineous,
apiculate or mucronulate, 21/,-23/, mm _ long.
Utricles oblong-rhomboid to obovoid-fusiform,
obscurely trigonous, suberect, densely many-
nerved, subcoriaceous, sparsely puberulous,
ciliolate-scabrid on the margins, stramineous-
green, stoutly stipitate, abruptly shortly beaked,
with oblique, bidentate mouth, 31/,-5!/, by 1!/,-
2 mm. Nut irregularly rhomboid-ellipsoid or
oblong-obovoid, trigonous, with faces excavated at
base and apex, broadly stipitate, seated on a
spongy, disk-like body, c. 2 mm long; beak short,
strongly deflexed; style bent upwards, thickened
above, papillose. Stigmas 3. Vestigial rachilla
sometimes present (according to KUKENTHAL).

Distr. Formosa, Hainan, S. China (Kwang-
tung), Tonkin, Annam, Thailand, Queensland; in
Malesia: Sumatra (also Banka), Malay Peninsula,
W. Java, E. Borneo, Philippines (Luzon, Minda-
nao), Aru Is., New Guinea; the specimen in L from
the Lesser Sunda Is. (leg. PLOEM) is probably mis-
labelled.

Ecol. In primary forests, on banks in woods, in
open jungle, at low and medium altitude; in Malaya
between 350 and 1200 m, in Banka at 40 m, in the
Aru Is. at a few m above sea-level, in W. Java
between 500 and 1250 m, in the Philippines
ascending to 1000 m.

Vern. Rumput ringgin, Mal. Pen.; ilat kampadn,
Ss

10. Section Lageniformes

(OHWI1) NELMES, Reinwardtia | (1951) 366; Raym. Mém. Jard. Bot. Montréal
n. 53 (1959) 54, 63. — Sect. Praecoces CHRIST subsect. Lageniformes OHWI1, Mem.
Coll. Sc. Kyoto Imp. Un. B11 (1936) 340.

Type species: Carex formosensis LEV. & VAN. (lectotype).

27. Carex breviscapa C. B. CLARKE, FI. Br. Ind. 6
(1894) 736; Kutx. Pfl. R. Heft 38 (1909) 474;
Emer, Leafi. Philip. Bot. 10 (1938) 3525; MErRR.
En. Philip. 1 (1923) 137; KUx. Bot. Jahrb. 69
(1938) 265; Back. Bekn. FI. Java (em. ed.) 10
(1949) fam. 246, p. 64; NELMES, Kew Bull. (1949)

390; Reinwardtia 1 (1951) 369; ibid. 2 (1954) 380;
Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 63,
f. 14, map p. 102, f. 33; Dansk Bot. Ark. 23 (1965)
258; KoyAMA, Phytologia 17 (1968) 405, t. 14;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 489. —
C. jackiana Bootr var. breviculmis THw. En. Pl.

1979]

Cy PERACEAE—II (Kern & Nooteboom)

149

Zeyl. (1864) 356. — C. curtisii RipL. Mat. Fl. Mal.
Pen. (Monoc.) 3 (1907) 117; Fl. Mal. Pen. 5 (1925)
183. — C. lutchuensis OHW1, Mem. Coll. Sc. Kyoto
Imp. Un. B5 (1930) 270; ibid. B11 (1936) 343, f. 5,
t. 9 f. 8; AKryAMA, Car. Far East. Reg. Asia
(1955) 177, t. 178. — Fig. 120.

Rhizome short, cespitose. Stems central, some-
times more stems together, more or less hidden
amongst the leaves, suberect, triquetrous, smooth,
5—20(—30) cm by c. 1 mm. Leaves basal, very much
longer than the stems, flat, long-acuminate, scabrid
on the margins and upper surface, 3-6(-8) mm
wide; lower leaves reduced to bladeless brown
sheaths or their fibrous remains. Spikelets usually
5-7, single at the nodes, erect or suberect, cylindric,
lax-flowered, lower ones somewhat distant; termi-
nal spikelet g, 1-2 cm long, 1 mm thick, often
overtopped by some of the 2 spikelets; lateral
spikelets 2 or with some ¢ flowers at the top,
1-3 cm by 3-4 mm; peduncles scaberulous, lower
ones exserted. Lower bracts foliaceous, much
exceeding the inflorescence, shortly sheathing,
upper much reduced. Glumes broadly ovate to
oblong-ovate, usually rounded at the erose apex,
much shorter than the utricles, slenderly nerved,
with not or slightly (up to 1 mm) excurrent mid-
nerve and hyaline margins, ciliolate, 2-3 mm long.
Utricles rhomboid-lageniform, trigonous, broadest
about the middle, subcoriaceous, straight, suberect,
strongly multinerved, glabrous or sparsely puberu-
lous above, stramineous or greenish, stipitate,
31/,—4'/, by 11/,-11/, mm; beak hispid, bidenticulate,
1/,-1 mm long. Nut narrowly rhomboid, triquetrous
with concave faces, stipitate, truncate and hollowed
out at the apex, dark brown with pale stipe, apex,
and angles, 2'/,-3 by 1'/,-1'/, mm. Style-base
slightly thickened. Stigmas 3.

Distr. Ceylon, NE. Thailand, Annam, Ryu Kyu
Is., Formosa, N. Queensland; in Malesia: Sumatra
(E. Coast Res., Lampongs), Malay Peninsula
(Penang, Pahang, Negri Sembilan), W. Java,
Borneo (Sarawak), Philippines (Luzon: Sorsogon;
Palawan), W. & NE. New Guinea.

Ecol. In secondary forest, on forested ridges,
from low altitude up to 1250 m.

Vern. Jlat daun eurih, S.

Note. In some of the ¢ flowers I observed
stamens with connate filaments. — (Noort.)

28. Carex gracilispica HAYATA, Ic. Pl. Form. 10
(1921) 62, f. 39; Onwi, Mem. Coll. Sc. Kyoto
Imp. Un. B11 (1936) 345; Akiyama, Car. Far East.
Reg. Asia (1955) 178, t. 179; RAymM. Mém. Jard.
Bot. Montréal n. 53 (1959) 65. — C. ligata (non
Bootr) Riwi. Fl. Mal. Pen. 5 (1925) 181. — ?
C. tristachya THUNB. var. pseudopocilliformis
Gross, Notizbl. Berl.-Dahl. 14 (1938) 191. —
C. malayana NeELMeES, Kew Bull. (1950) 209;
Reinwardtia 1 (1951) 366.

Rhizome short, cespitose. Flowering stems
arising from basal leaf-axils, erect, compressed-

trigonous, smooth or scaberulous just below the
inflorescence, 10—30 cm by */,—-1 mm. Leaves basal,
much longer than the stems, plicate to flat, long-
attenuated, with scabrid margins, 5-10 mm wide,
surrounded at the base by brown, fibrous remains
of old leaf-sheaths. Spikelets 4-6, approximate or
lowest somewhat distinct, erect or suberect,
cylindric, 1—3!/, cm long; terminal spikelet 3, 1 mm
thick, lateral ones single at the nodes, wholly 2 or
with some ¢ flowers at the top, lax-flowered,
2-3 mm thick, their peduncles shortly exserted,
smooth or scaberulous. Bracts foliaceous, lower as
long as or exceeding the inflorescence, shortly
sheathing, upper much reduced. Glumes oblong-
ovate, acute to obtuse, slenderly nerved, with not
or slightly excurrent midnerve, whitish hyaline
margins, and erose-ciliate apex, 3-4 mm long.
Utricles \ageniform, obscurely trigonous, sub-
coriaceous, broadest 2-21/, mm from the base,
straight, suberect, strongly multinerved, sparsely
puberulous, stramineous or greenish, shortly
stipitate, 5-7 by 11/,-14/; mm; beak 11/,-2 mm
long, bidenticulate. Nut oblong, triquetrous, with
faces concave below, stoutly stipitate, slightly
constricted at the apex and re-expanded into a
cylindric, truncate neck 7/,-1 mm long and wide,
the whole nut 3-4 mm long. Style-base thickened,
centred in the apical hollow of the nut. Stigmas 3.

Distr. Formosa, Indo-China (Annam,
?Tonkin); in Malesia: Malay Peninsula (Pahang:
Mt Tahan).

Ecol. On Mt Tahan in damp woods by streams,
at c. 1700 m.

29. Carex palawanensis KUK. in Elmer, Leafl.
Philip. Bot. 4 (1911) 1169; Merr. En. Philip. 1
(1923) 140; Netmes, Reinwardtia 1 (1951) 263;
ibid. 2 (1954) 374. — Fig. 120.

Rhizome probably shortly creeping. Stems
triquetrous, sparsely scabrid just below the
inflorescence, otherwise smooth, 25-50 cm by
1/,-1 mm, surrounded below the leaves by the
fibrous remains of old leaf-sheaths. Leaves basal
and one half-way up the stem, exceeding the stem,
long-attenuate, rigid, with revolute margins,
scabrid in the upper part, grey-green with a metallic
hue, 3-7 mm wide; cauline leaf long-sheathing.
Inflorescence spiciform (or almost so when its
lowest node bears 2 spikelets), 2-4 cm long. Lowest
bract foliaceous, patent, much overtopping the
inflorescence, shortly sheathing, upper one(s) much
reduced. Spikelets 2-10, androgynous, subglobose,
dense, sessile or on very short included peduncles,
5-8 mm long, the $ part about as long as the 9, but
inconspicuous when the utricles are fully developed.
Glumes ovate or ovate-lanceolate, acutish or ob-
tuse, nerved, minutely ciliolate, otherwise glabrous,
brownish with whitish hyaline margins, 11/,-2 mm
long, the midnerve excurrent in a hispidulous,
3/,-1 mm long awn. Utricles much overtopping the
glumes, trigonous, rhomboid-lageniform, with

150

FLORA MALESIANA

[ser. I, vol. 91

prominent angles and somewhat concave faces,
subcoriaceous, patulous, strongly many-nerved,
straight, glabrous, smooth except for a few setae in
the upper part, not stipitate, curved-tapering to the
base, rather gradually narrowed into the beak,
greenish brown, 4!/,—5 by 11/,-13/, mm; beak com-
pressed, bidentate, 1!/,-2 mm long, with slightly
oblique mouth. Nut trigonous, broadly rhomboid,
with thickened angles and concave faces, curved-
tapering downwards to a short cylindric stipe and
upwards to a stout, cylindric, 1/,/, mm long and
1/,-3/, mm wide neck, truncate at the apex, c. 21/,
by 11/, mm. Style-base slightly thickened, centred in
the hollowed apex of the nut. Stigmas 3.

Distr. Malesia: N. Borneo (Sabah: Lahad
Datu, Mt Silam; Sandakan) and SW. Philippines
(Palawan: Mt Pulgar).

Ecol. Common in wet, sandy, gravelly soilamong
shrubs bordering streams in the hills at 150-600 m.

Note. Only known from the type collection,
ELMER 13146 from Palawan and SAN 37905 and
43845 from Sabah.

30. Carex rhynchachaenium CLARKE in Merr.
Publ. Gov. Lab. Philip. n. 35 (1906) 5; Philip. J.
Sc. 2 (1907) Bot. 108; Kew Bull. add. ser. 8 (1908)
79; KUK. Pfl. R. Heft 38 (1909) 480; Philip. J. Sc. 6
(1911) Bot. 62; Merr. En. Philip. 1 (1923) 142;
NELMES, Reinwardtia 1 (1951) 368; RAyM. Mém.
Jard. Bot. Montréal n. 53 (1959) 65. — C. hatusi-
mana OHwI, Jap. J. Bot. 7 (1934) 196; Mem. Coll.
Sc. Kyoto Imp. Un. B11 (1936) 344, f. 6, t. 9 f. 7;
Koyama, Contr. Inst. Bot. Un. Montréal n. 70
(1957) 21, t.3; AKryAMA, Car. Far East. Reg. Asia
(1955) 178; Raym. Mém. Jard. Bot. Montréal
n. 53 (1959) 65, f. 15, map p. 102, f. 29. — Fig. 120.

Rhizome short, cespitose. Stems central, more or
less hidden amongst the leaves, erect, triquetrous,

scaberulous above, 5—10(—30) cm by !/,—'/, mm,
surrounded below the leaves by brown, fibrous
remains of old leaf-sheaths. Leaves basal, much
longer than the stems, plicate to flat, long-attenuate,
scabrid on the margins and upper surface, 1-3 mm
wide; sheaths pubescent. Spikelets 3-6, upper
approximate, lower distant, erect or suberect,
cylindric, lax-flowered; terminal spikelet ,
1(—2) cm long, 1 mm thick, lateral ones 2 or with
some ¢ flowers at the top, !/,-2 cm long, 3-4 mm
thick; peduncles smooth or scaberulous, lower
exserted. Lower bracts foliaceous, much exceeding
the inflorescence, upper much reduced. Glumes
elliptic-ovate to oblong-lanceolate, acute or
with rounded apex, slenderly nerved, with not
or scarcely excurrent midnerve and hyaline
margins, erose-ciliate towards the apex, 2-3 mm
long. Urtricles lageniform, trigonous, broadest
c. 2 mm from the base, subcoriaceous, straight,
suberect, strongly multinerved, more or less
puberulous above, stramineous or _ greenish,
shortly stipitate, 5'/,-6'/, by 11/,-1'/, mm; beak
3/,-1 mm long, bidenticulate. Nut rhomboid, tri-
quetrous, with faces concave below, stoutly
stipitate, the apex subabruptly contracted into a
cylindric, truncate neck c. 1 mm long and 3/, mm
broad, the whole nut c. 4 mm long. Style-base
scarcely thickened, centred in the apical hollow
of the nut. Stigmas 3.

Distr. Formosa, Tonkin, Annam; in Malesia:
Philippines (Luzon: Kalinga, Pampanga, Bataan,
Benguet, Laguna; Mindanao: Bukidnon, Zam-
boanga).

Ecol. On forested ridges in and near the mossy
forest, 800-2100 m.

Note. I have not seen CLEMENS 34431 from
Borneo (Mt Kinabalu?) which may belong here
(cf. NELMES, /.c. 369).

11. Section Mitratae

KUK. Pfl. R. Heft 38 (1909) 458; NeLMgs, Reinwardtia | (1951) 371; Raym. Mém.
Jard. Bot. Montréal n. 53 (1959) 52, 60. — Sect. Praecoces CHRIST, Bull. Soc. Bot.
Belg. 24 (1885) 14, nomen; MACKENZ. N. Am. FI. 18 (1935) 183; OHw1, Mem. Coll.
Sc. Kyoto Imp. Un. B11 (1936) 339. — Type species: Carex mitrata FRANCH.
Note. In this section many species have been described which differ only in slight characters and in my

opinion several must be combined or deserve at most varietal rank. With the existing keys of KUKENTHAL,
OxwIl, and Koyama identification appears often impossible. — (Noot.).

31. Carex breviculmis R. Br. Prod. (1810) 242;
Boortr, Ill. 4 (1867) 181; Boeck. Linnaea 41 (1877)
209; BENTH. FI. Austr. 7 (1878) 445; CLARKE,
Fl. Br. Ind. 6 (1894) 746; Kix. Pfl. R. Heft 38
(1909) 469; RipL_. Trans. Linn. Soc. Bot. II, 9
(1916) 247; S. T. BLaxe, J. Arn. Arb. 28 (1947)
112; NELMEs, Kew Bull. (1949) 383; Reinwardtia 1
(1951) 373; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 489. — Fig. 121.
For synonyms see under the varieties.

Rhizome short (obliquely descending, woody).
Stems tufted, slender, erect or oblique, (obtusely)
trigonous, smooth, or scaberulous above, 1-40 cm
by 1/,-1 mm, clothed at the base by old leaf sheaths
or their fibrous remains. Leaves basal and subbasal,
from much shorter to much longer than the stems,
often thickish, rigid, flat or with recurved or
revolute margins, gradually attenuate to the
triquetrous tip, smooth or mostly scabrid on
margins and keel, 1-4(-6) mm wide. IJnflorescence

1979]

simple, erect, with 3-7 approximate spikelets
(sometimes lowest spikelet remote on a peduncle
from the sheath of a basal leaf; in small plants
sometimes only one ¢ and one spikelet, the other
2 spikelets single on a long peduncle arising from
the centre of the basal leaves together with the
main inflorescence); terminal spikelet 3, sometimes
gynaecandrous (in var. perciliata), linear, 5—12(—20)
by 1-2 mm, lateral ones 9, sessile or (very) shortly
peduncled, subglobose to shortly cylindrical,
5-15 (-30) by 3-4 mm, peduncles smooth or scab-
rous; lower bracts usually overtopping the in-
florescence, foliaceous or setaceous, shortly sheath-
ing, upper much reduced. Glumes ovate or oblong-
ovate, acute to obtuse, whitish or brownish,
excurrent into a scabrid awn in var. breviculmis,
muticous or with a short mucro in the other
varieties, whitish or brownish and then with white
hyaline margins, 3-nerved in the centre, 2—3'/, mm
long. Utricles obtusely or obsoletely trigonous,
lanceolate to ellipsoid or obovoid, membranous,
except the 2 marginal nerves nerveless or obscurely
nerved to more distinctly nerved, subabruptly to
gradually beaked, from densely hispidulous to
only hispidulous on the margins or glabrous, not
or scarcely stipitate, light brown to stramineous or
fulvous, 2!/,-4 by c. 1 mm. Nut triquetrous, with
faces concave below, ellipsoid or ovoid to obovoid,
stramineous to fuscous, 1!/,-21/, by c. 1 mm,
suddenly contracted into a very short neck and
then suddenly expanded into an annulate apex.
Style-base thickened, persistent on the annulus
(but when the nuts are falling the style-base
probably has disappeared). Stigmas 3.

Distr. Widely distributed from the Himalayas
to China, Japan and Formosa through Malesia to
Australia and New Zealand.

Note. The length of the stem varies consider-
ably, even on the same plant. The same holds for
the nerves on the utricles and for their hairiness,
but these characters are rather constant in a single
specimen. For these reasons I consider C. brevicul-
mis R. Br., C. perciliata NELMES, and C. montivaga
S. T. BLAKE to be conspecific, although it is possible
to discriminate between them on varietal rank.

KEY TO THE VARIETIES

1. Glumes whitish, 2-3 mm long, excurrent into a
scabrid awn. Utricles greenish becoming light
brown, subabruptly beaked, several-nerved,
usually more or less pubescent or hispidulous,
21 (5-3/5, mmi long». 2. = a. var. breviculmis

1. Glumes brownish with white hyaline margin,
acute or obtuse, muticous or with a small mucro.
Utricles stramineous to fulvous, gradually
beaked, nerveless or faintly nerved, hispidu-
lous to glabrous, 2!/,-4 mm.

2. Utricles usually hispidulous, at least above on
the margins, 2'/,-3 mm . ._ b. var. perciliata

Cy PERACEAE—II (Kern & Nooteboom)

2. Utricles glabrous, rarely obscurely hispidulous
on the margins above, 3'/,-4 mm long
c. var. montivaga

a. var. breviculmis. — C. leuchochlora BUNGE, En.
Pl. Chin. Bor. (1833) 68; KoyAMA, Act. Phytotax.
Geobot. 16 (1955) 9; YosHiKAwa, Ic. Jap. Car. 1
(1957) 104, t. 52; RAyM. Mém. Jard. Bot. Montréal
n. 53 (1959) 60. — C. royleana NEES in Wight,
Contr. (1834) 127; Boorrt, Ill. 1 (1858) 6, t. 19;
Camus, FI. Gén. I.-C. 7 (1922) 195. — C. eggytera
STEuD. Syn. 2 (1855) 220. — C. breviculmis ssp.
royleana KUK. Pfl. R. Heft 38 (1909) 469; Philip.
J. Sc. 6 (1911) Bot. 62, incl. var. kingiana KUK.;
Merr. En. Philip. 1 (1923) 137. — C. jackiana
Boott var. tumens KUxK. in Hochr. Candollea 6
(1936) 432. — C. rugata (non OHWwt) NELMES,
Reinwardtia 1 (1951) 378. — C. conorrhyncha
NeELMES, Kew Bull. (1956) 182. — Fig. 121.

Distr. As for the species; in Malesia: E. Java
(Mt Tengger and Mt Jang), SW. Celebes (Mt Bon-
thain), N. Borneo (Mt Kinabalu), Philippines
(Luzon), and New Guinea.

Ecol. Open grassy slopes, among shrubs, 1750—
3900 m.

b. var. perciliata KUK. Pfl. R. Heft 38 (1909) 469. —
C. breviculmis: CLARKE, J. Linn. Soc. Bot. 37
(1904) 16. — C. bulbostylis var. ciliato-marginata
Kx. et var. hispidula KUx. Bot. Jahrb. 70 (1940)
464; Bull. Jard. Bot. Btzg III, 16 (1940) 318. —
C. tricholoma S. T. BLake, J. Arn. Arb. 28 (1947)
1105 f3AS— 1G) brevis S. de BEAKE SG) lee B=
NELMES, Kew Bull. (1950) 202; Reinwardtia 1
(1951) 375. — C. perciliata (KUK.) NELMES, Kew
Bull. (1946) 26; ibid. (1949) 383, 391; Reinwardtia
1 (1951) 374. — Fig. 121.

Distr. Malesia: N. Borneo (Kinabalu), SW.
Celebes (Latimodjong Range: Mt Rante Mario),
New Guinea (W. New Guinea: Star Mts; Papua
and Territory of New Guinea).

Ecol. Rock crevices, open bare ground, wet
grassland, forest glades, 2400-4200 m.

c. var. montivaga (S. T. BLAKE) Noot., comb. nov.
— C. montivaga S. T. BLakE, J. Arn. Arb. 28 (1947)
109; NeLmes, Kew Bull. (1949) 383; Reinwardtia 1
(1951) 377. — C. bulbostylis Kix. Bot. Jahrb. 69
(1938) 264; Bull. Jard. Bot. Btzg III, 16 (1940)
317, excl. var. et specim. Born., non MACKENZ.
1915. — Fig. 121.

Distr. Malesia: New Guinea (Lake Habbema,
Mt Wilhelmina, Star Mts, Mt Sarawaket).

Ecol. Bogs, alpine grassland, wet grassy slopes,
along water, also in mossy forest, 3200-3450 m.

32. Carex dolichostachya HayaTA, Ic. Pl. Form. 10
(1921) 61, f. 38; Ow1, Mem. Coll. Sc. Kyoto Imp.
Un. B11 (1936) 375; KoyAMA, Bull. Arts & Sc. Div.
Ryukyu Un. (Math. & Nat. Sc.) n. 3 (1959) 72;
AxtyaMA, Car. Far East. Reg. Asia (1955) 204,

152

FLORA MALESIANA

[ser. I, vol. 9!

t. 208, f. 2. — C. ligata BooTT var. nexa KUK.
Philip. J. Sc. 6 (1911) Bot. 63; MerrR. En. Philip. 1
(1923) 39, quoad specim. cit., non C. nexa BooTT.
— C. multifolia OHw1, Mem. Coll. Sc. Kyoto Imp.
Un. B5 (1930) 264; ibid. B11 (1936) 373; NELMEs,
Reinwardtia 1 (1951) 371; AktyAMa, Car. Far East.
Reg. Asia (1955) 202, t. 206; YosHIKAWA, Ic. Jap.
Carex 1 (1957) 122, t. 61. — C. foliosissima (non
F. SCHMIDT) FRANCH. Carex As. Or. (1898) n. 232;
KUK. Pfl. R. Heft 38 (1909) 478. — Fig. 121.

Rhizome short, cespitose, or shortly stoloni-
ferous. Stems arising from basal leaf-axils, trigo-
nous, smooth or sparsely scaberulous above,
(15—)30-60 cm by 1/,-1 mm, bearing a few short-
bladed, bract-like leaves. Basal /eaves about as
long as the stems, flat, with scabrid margins and
under surface above, 3-10 mm wide; sheaths
reddish brown to spadiceous, older fibrous.
Spikelets 3-7, distant, erect or suberect; terminal
spikelet g, linear, (1!/,-)3-6 cm by 1!/,-2 mm,
lateral 2 or with a few ¢ flowers at the top, lax-
flowered, (1!/,-)3-5 cm by 2!/,-3!/, mm, on
exserted, smooth or slightly scaberulous peduncles.
Bracts vaginiform with a short blade or reduced to
long, subampliate, bladeless sheaths. Glumes
oblong or oblong-obovate, truncate or rounded at
the erose-ciliate apex, translucent, whitish to
brownish, 21/,-3 mm long, the 3-nerved centre
excurrent in a short, hispidulous awn c. !/, mm
long. Utricles fusiform, trigonous with flat faces,
straight, membranous, suberect, strongly many-
nerved, hispidulous especially above, greenish
to light brown, 3-4 by c. 1 mm; beak conical,
1 mm long, bidenticulate. Nut ellipsoid to oblong-
ellipsoid, triquetrous with shallowly concave faces,
stoutly stipitate, contracted at the apex and re-
expanded into a discoid, '/, mm broad annulus,
c. 2 by 1 mm. Style-base pyramidally thickened.
Stigmas 3.

Distr. Japan, Ryu Kyu Is., Formosa; in
Malesia: Philippines (Luzon: Kalinga, Bontoc,
Rizal).

Ecol. Clearings and along edges of primary
forest, 1200-1600 m.

Notes. The distinction between C. multifolia
OHWI, common in the montane regions of Japan,
and C. dolichostachya HAYATA, occurring from the
Ryu Kyu Is. and Formosa to Luzon, is mainly
made on account of the different colour of the basal
leaf-sheaths. I have followed KoyAMA, who united
the two as neither this character nor the other slight
differences are constant.

HatTusimMa, Mem. Fac. Agric. Kagoshima Un. 5,
3 (1966) 59, referred specimens from the Batan Is.
(N. Luzon), mentioned above as C. ligata var. nexa
KUK. (non C. nexa BootTt), to C. sociata BoottT in
A. Gray, Bot. Jap. (1859) 420; Ill. 4 (1867) 200;
Ouwil, Cyp. Japon. 2 (1943) 376. If this identity is
true, C. sociata Boott would be the correct name
for this species.

A specimen from Sarawak (Mt Murud, 2400 m,

NooTesoom 2030) undoubtedly belongs to sect.
Mitratae, possibly in the vicinity of C. dolichos-
tachya. The utricles, however, are longer together
with the nuts, and the leaves are smaller. —
(Noot.)

33. Carex formosensis LEv. & VAN. Mém. Soc.
Nat. Sc. Nat. Math. Cherb. 35 (1906) 216, et in
Fedde, Rep. 5 (1908) 31; OHw1, Mem. Coll. Sc.
Kyoto Imp. Un. B11 (1936) 345; NELMegs, Rein-
wardtia 2 (1954) 379; AkryAMA, Car. Far East. Reg.
Asia (1955) 178, t. 180. — C. ligata Bootr var.
formosensis (LEV. & VAN.) Ktx. Pfl. R. Heft 38
(1909) 474; Merr. En. Philip. 1 (1923) 139. —
Fig. 121.

Rhizome short, cespitose. Stems central, erect,
trigonous, smooth, 10-25(—5S0) cm by !/,-1 mm.
Leaves basal, shorter to longer than the stems, flat,
long-attenuate, with scabrid margins, 2-4(-6) mm
wide, surrounded at the base by brown, fibrous
remains of old leaf-sheaths. Spikelets 3-7, sub-
approximate or lower more distant, erect or sub-
erect, cylindric; terminal spikelet usually 4,
1-2 cm long, 2-3 mm thick, lateral ones single at
the nodes, usually gynaecandrous, with very few 3
flowers at the base, subdensely flowered, 31/,-5 mm
thick, their peduncles smooth, upper shortly,
lower rather long-exserted. Lower bracts folia-
ceous, shorter to longer than the inflorescence,
lower long-sheathing, upper much reduced. Glumes
oblong, truncate to bilobed-emarginate, 11/,-2 mm
long, nerveless except for a greenish, strongly
3-nerved central stripe excurrent in a hispid, up to
1'/, mm long awn. Utricles rhomboid, slightly
lageniform or fusiform, trigonous, subcoriaceous,
broadest below the middle, straight, suberect,
multinerved, very sparsely puberulous, strami-
neous or greenish, shortly stipitate, c. 31/. mm by
1 mm; beak c. 1 mm long, bidentate. Nut rhom-
boid, trigonous, with deeply concave faces, shortly
stipitate, slightly constricted at the apex and slightly
re-expanding into a rather discoid-annulate
1/,1/, mm wide apex, the whole nut c. 2(—2!/,) mm
long. Style-base slightly thickened. Stigmas 3.

Distr. Korea, Japan (Kyushu, Hondo), For-
mosa; in Malesia: Philippines (Luzon: Benguet
Subprov., Ilocos Norte).

Ecol. Forested ridges, along streams and trails,
on cliffs and steep banks in and near the mossy
forest, 1000-2400 m.

Vern. Egegedan, Bon., silak, Ig.

34. Carex tristachya THUNB. FI. Jap. (1784) 38;
Scukunr, Riedgr. 2 (1806) 48, t. Ww f. 109;
Boorr, Ill. 4 (1867) 131, t. 424; KUx. Pfl. R. Heft
38 (1909) 471; Onw1, Mem. Coll. Sc. Kyoto Imp.
Un. B11 (1936) 363; AkryAMA, Car. Far East. Reg.
Asia (1955) 195, t. 198; YosHrkAwa, Ic. Jap.
Carex 1 (1957) 118, t. 59.
The few Malesian collections belong to:

1979]

var. pocilliformis (Bootr) KUxK. Pfl. R. Heft 38
(1909) 473, t. 75, f. A-F; Philip. J. Sc. 6 (1911) Bot.
62; Merr. En. Philip. 1 (1923) 142; Onwi, Mem.
Coll. Sc. Kyoto Imp. Un. B11 (1936) 364; Yosut-
KAWA, Ic. Jap. Carex 1 (1957) 120, t. 60. —
C. pocilliformis Bootr, Ill. 4 (1867) 175, t. 593;
Newmes, Kew Bull. (1949) 391; Reinwardtia 1
(1951) 381; AkryAMA, Car. Far East. Reg. Asia
(1955) 196, t. 199. — Fig. 121.

Rhizome short. Stems densely tufted, very slen-
der, erect, trigonous, smooth, 10-40 cm by
1/,-1 mm, clothed at the base with dark brown,
more or less fibrous remains of old leaf-sheaths.
Leaves basal and subbasal, as long as or shorter
than the stems, rather rigid, flat, gradually attenu-
ate, 11/4 mm wide. Inflorescence with 3-7
spikelets, erect, fastigiate or lower spikelets more
distant on exserted peduncles; terminal spikelet
6 or gynaecandrous, cylindrical, '/,-3 cm by
1/,-1 mm, lateral ones 9, slenderly cylindrical,
rather loosely flowered, 1-3 cm by 2-2!/, mm;
peduncles smooth. Lower bracts foliaceous,
shorter than to slightly exceeding the inflorescence,
upper reduced. G/umes of the 3 spikelet cup-shaped
(the margins more than halfway connate in front)
but not so in gynaecandrous spikelets, muticous,
those of the 2 spikelets ovate or oblong-ovate,
rounded at the ciliolate apex, glabrous, muticous
or mucronulate, light brown with whitish hyaline
margins and 3-nerved central stripe, 2 mm long.
Stamens monadelphous (filaments connate almost
throughout their length), not always so in lower
flowers. Utricles trigonous, ellipsoid, membranous,
many-nerved, sparsely pubescent, suberect, re-
curved at the top, shortly stipitate, gradually
tapering above, green to brownish, 2!/,-3 by
3/,-1 mm; beak subconical, glabrous or his-
pidulous-margined, bidenticulate, !/, mm _ long.
Nut oblong-ovoid or oblong-ellipsoid, triquetrous
with faces concave below, stipitate, c. 2 mm long,
stramineous to dark brown, rounded above and
abruptly contracted into a short neck which is
suddenly expanded into the annulate apex. Style-
base pyramidally thickened, persistent. Stigmas 3.

Distr. Var. tristachya occurs in Japan, Korea,

Cy PERACEAE—II (Kern & Nooteboom)

[53

Central and Eastern China, var. pocilliformis is also
in Japan and Korea, but extends more southwards,
through the Ryu Kyu Is. and Formosa to Malesia:
Philippines (Luzon: Benguet: Mt Pulog), Borneo
(Mt Kinabalu), and New Guinea (NE.: Mt
Sarawaket).

Ecol. Open meadows and grassy slopes, 2200-
2700 m.

Notes. Carex tristachya is remarkable for its
monadelphous anthers, a feature very rare in the
genus (according to BARRos also found in C. acaulis
D’URv. of the Falkland Is. [cf. BRONGN. in
Duperrey, t. 28, f. A; RoEPER, Zur Flora Mecklenb.
2 (1844) 16!]).

There is no agreement on the taxonomical status
of C. pocilliformis Bootr. Whereas CLARKE (J.
Linn. Soc. Bot. 36, 1904, 315) did not distinguish
it from C. tristachya, it was reduced to varietal
rank by KUKENTHAL, OHwI, and others, but main-
tained as a distinct species by NELMES and AKIYAMA.
The main difference is to be found in the glumes of
the 3 flowers, which are said to have free margins
in typical C. tristachya. I think the difference is
only gradual, as in the latter the margins of the
glumes are also connate, though at the very base
only.

Glumes with united margins are very rare in the
genus; they are also known in some North
American species belonging to sect. Phyllostachyae
TUCKERM. (see MACKENZ. N. Am. FI. 18, 1935,
174).

Carex tristachya is very near to C. mitrata
FRANCH. & SAv., the main difference being in the
3 spikelets, of which the glumes are infundibuli-
form and the filaments connate in C. tristachya.
But in C. tristachya var. tristachya the margins of
the glumes are connate at the very base only, and
sometimes they are free. Besides, even in var.
pocilliformis, 1 found perfectly free filaments and
anthers, especially in the lower flowers (e.g.
MERRILL 6629). It is very well possible that the
mentioned characters in the ¢ spikelets are not
very important, and that C. tristachya and
C. mitrata have to be considered as subspecies of a
single species. — (Noot.)

12. Section Radicales

(KUK.) NELMES, Reinwardtia | (1951) 389; RaymM. Mém. Jard. Bot. Montréal n. 53
(1959) 54, 75. — Sect. Digitatae Fries subsect. Radicales Kix. Pfl. R. Heft 38

(1909) 480.

Type species: Carex radicalis Bootr.

35. Carex malaccensis C. B. CLARKE, FI. Br. Ind. 6
(1894) 722; J. Linn. Soc. Bot. 37 (1904) 9; Rip.
Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 116; Kix.
Pfl. R. Heft 38 (1909) 289; RipL. Fl. Mal. Pen. 5
(1925) 183; Ne_mes, Reinwardtia 1 (1951) 257. —
Fig. 121.

Rhizome shortly creeping, woody, covered with
brown, readily fraying scales. Stems tufted, slender,
triquetrous, very narrowly winged above, smooth
below, slightly scabrid on the angles above, up to
50 cm by 1-11/, mm. Leaves basal and subbasal,
shorter to longer than the stems, stiff, strongly

154 FLORA MALESIANA [ser. I, vol. 9!

Fig. 129. Carex speciosa KUNTH. a. Habit, x '/,, b. spikelet, lower part 2, upper part d, x 1'/,, c. glume,
d. fruit in utricle, e. fruit, f. young stamens in glume, g. glume, 4. seemingly connate filaments, i. filaments,
all x 13 (WAITz s.n.).

1979 ]

Cy PERACEAE—II (Kern & Nooteboom)

155

revolute when dry, long-attenuate, greyish green
or glaucous, scabrous-asperous in the apical
part, 2-6 mm wide; sheaths brown, membranous
in front, outer ones bladeless. J/nflorescence
racemiform, consisting of 14 remote, subglobose
or pyramidal heads on short (the lowest rather
long), smooth or scabrid peduncles, each head
composed of 1-4 crowded, sessile spikelets. Lower
bracts foliaceous, much exceeding the inflorescence,
not auricled, not or lowest very shortly sheathing,
ultimately horizontally spreading, upper much
reduced. Spikelets androgynous, the 3 part about
as long as or longer than the °, ovoid, patent,
5-10 mm long and wide. Glumes broadly ovate to
oblong-ovate, obtuse, membranous, strongly
nerved, ciliolate above, otherwise glabrous, white
or light brown, 2!/,-3 mm long, the midnerve
excurrent in a stoutish, scabrid, '/,-1 mm long awn.
Utricles trigonous, rhomboid or obovoid, mem-
branous, suberect or patulous, strongly many-
nerved, rather abruptly contracted into the beak,
glabrous, narrowly marginate or winged, scabrid
on the margins at the apex, stramineous, 4—6*/, by
1+/,—2'/, mm; beak linear-conical, flattened, curved
inwards, bidenticulate, with oblique mouth 1-2 mm
long. Nut triquetrous, pyriform, scarcely stipitate,
brown but densely overlain white-scurfy, 2-2!/, by
13/, mm. Style-base pyramidally thickened, per-
sistent on the nut. Stigmas 3.

Distr. Malesia: Malay
Langkawi Is.).

Ecol. On limestone along the sea-shore, on
limestone rocks, at low altitude.

Notes. Like Fimbristylis malayana Ouw1 this
interesting species is apparently endemic in the
Langkawi Is., here exclusively growing on lime-
stone. RIDLEY (/.c.) recorded it for Thailand, but it
is not mentioned for that country either by NELMES
or RAYMOND.

Its nearest ally is C. leucantha ARN. ex BOoTT,
Proc. Linn. Soc. 1 (1845) 257; Trans. Linn. Soc. 20
(1846) 135; Ill. 1 (1858) 10, t. 28, from Ceylon and
India, which differs by the long-sheathing lower
bracts and the hairy utricles.

Peninsula (Kedah,

36. Carex ramosii KUx. in Fedde, Rep. 8 (1910) 8;
Philip. J. Sc. 6 (1911) Bot. 63; MerrR. En. Philip. 1
(1923) 141; Nemes, Reinwardtia 1 (1951) 259;
ibid. 2 (1954) 374. — Fig. 121.

Rhizome shortly creeping, woody, covered with
brown, readily fraying scales. Stems very slender,
erect but often curved, triquetrous, slightly
incrassate just below the inflorescence, smooth, or
slightly scabrid at the top, surrounded below the
leaves by a few, bladeless, brown sheaths, 10-25
cm by !/,-1 mm. Leaves basal, up to twice as long
as the stems, rigid, flat, with revolute margins when
dry, long-attenuate, greyish green, scabrid in the
apical part, 2-5 mm wide. Inflorescence either
a single head of 1-3 sessile spikelets, ovoid or
oblong-ovoid, 1-2 cm long, or spiciform with 1-2

additional subsessile heads near the base of the
stem in the axil of a leafy bract very similar to and
as long as the leaves. Bracts not sheathing, amplexi-
caul by ciliolate auricles, the lowest of the terminal
head foliaceous, 5—10(-25) cm long. Spikelets
androgynous, the ¢ part longer than the 2 but
finally hidden by the fully developed utricles, 6-10
by 5-7 mm. Glumes ovate-lanceolate, obtuse or
acutish, membranous, strongly nerved, glabrous,
stramineous with white margins, 3-4'/, mm long,
the midnerve excurrent in a minutely scabrid,
1-1°/, mm long awn. Utricles trigonous, ellipsoid-
rhomboid, subcoriaceous, suberect, strongly many-
nerved, distinctly winged almost all round, scabrid-
margined, gradually narrowed into the beak,
glabrous, stramineous or brownish, 6-8 by 2-
2?/, mm (wings included); beak compressed, sub-
conical, bidenticulate, with oblique mouth, 3 mm
long. Nut triquetrous, rhomboid-ellipsoid, sessile,
livid, 3 by 1°/,-2 mm. Style-base pyramidally
thickened, persistent on the nut. Stigmas 3.
Distr. Malesia: Philippines (Luzon, Mindanao).
Ecol. In forest at medium altitude.

37. Carex speciosa KUNTH, En. 2 (1837) 504; Mia.
Fl. Ind. Bat. 3 (1856) 348; Boorr, Ill. 1 (1858) 53;
Boeck. Linnaea 40 (1876) 388, incl. var. minor
BoeEcK.; CLARKE, Fl. Br. Ind. 6 (1894) 729; KUK.
Pfil. R. Heft 38 (1909) 481, incl. var. courtallensis
KUk. et var. angustifolia KUk.; CAmMus, Fl. Gén.
I.-C. 7 (1922) 197; Back. Bekn. Fl. Java (em.
ed.) 10 (1949) fam. 246, p. 63; NELMES, Reinwardtia
1 (1951) 390; ibid. 2 (1954) 381; Mém. Mus. Hist.
Nat. Paris n.s. B4 (1955) 160, incl. var. angustifolia;
Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 80;
Dansk Bot. Ark. 23 (1965) 260; KERN in Back. &
Bakh. f. Fl. Java 3 (1968) 488, 492. — C. concolor
NEEs in Wight, Contr. (1834) 125, mon R. Br. 1823.
— C. courtallensis NEES ex BootrT, Ill. 1 (1858) 52,
t. 138, incl. var. angustifolia Bootr; Camus, FI.
Gén. I.-C. 7 (1922) 197, f. 28, 1-8; RAyM. Mém.
Jard. Bot. Montréal n. 53 (1959) 78, f. 19. —
C. longispica Boeck. Allg. Bot. Zeitschr. 2 (1896)
174. — C. speciosa var. abscondita KtxK. Bull.
Jard. Bot. Btzg III, 16 (1940) 318. — C. stenura
NELMES, Kew Bull. (1950) 202; Reinwardtia 1
(1951) 392. — Fig, 121, 129.

Rhizome short, descendent, woody. Stems tufted,
erect or suberect, slender, triquetrous, often
narrowly winged at the top, smooth or finely
scaberulous, 5-50 cm by !/,-1!/, mm, surrounded
below the leaves by fuscous sheaths and fibres.
Leaves subbasal, longer to much longer than the
stems, stiff, flat (but margins revolute when dry),
scabrous on the margins, asperous on the upper
surface towards the long-attenuate apex, greyish or
glaucous-green, 3-10 mm wide. Spikelets 1-3(-4),
5-20 cm distant from one another, lowest some-
times arising from a basal leaf-sheath, androgy-
nous, cylindric, 2-8 cm long, rather laxly 6—18-
flowered, on shortly exserted smooth peduncles,

156

FLORA MALESIANA

[ser. I, vol. 9!

the 2 part 3-7 mm thick, half as long to about as
long as the slender g part. Bracts foliaceous,
usually overtopping the inflorescence, sheathing.
Glumes of the 3 flowers with free margins or rarely
the margins connate in front in the lower 2/,—"/,,
those of the @ flowers triangular-broadly ovate,
rounded to subacute, translucent, many-nerved,
with prominent midrib, muticous, minutely cilio-
late above, long-persistent, 2'/,-3 mm _ long.
Utricles ovoid-ellipsoid or ovoid-pyramidal, tri-
quetrous, coriaceous, suberect, strongly many-
nerved, straight or slightly curved, with whitish-
pubescent margins, greenish-stramineous, 4~7 by
1-21/, mm, spongy at the base, gradually narrowed
into the short, bidenticulate beak. Nut obovoid or
ellipsoid, triquetrous, shortly stipitate, abruptly
shortly beaked, pale to dark brown with promi-
nent pale angles, 3-4 by 11/,-2 mm. Style-base
pyramidally thickened, ciliolate, persistent. Stig-
mas 3.

Distr. Widely distributed in India (from Nepal
to Sikkim, also in S. India), S. China (Yunnan),
Thailand, Indo-China; in Malesia: Sumatra
(Atjeh, E. Coast Res.), Malay Peninsula (Ulu
Kelantan: Bertam, Gua Musang), SE. Borneo
(N. of Bandjermasin, once), and Java (rare in
W. and E., more common in Central Java), also in
Kangean I.

Ecol. Primary forest, often in teak-forest,
distinctly preferring seasonal climatic conditions,
in Borneo on dry serpentine rock (once), at low
and medium altitude, from sea-level up to 1500 m.

Vern. Java: ilat djepun, S.

Notes. According to RAYMOND, /.c., the leaves
in C. speciosa are 11/,-2 mm wide, the number of
spikelets 1-4, and the utricles 4-5 mm long. He
kept C. courtallensis apart from C. speciosa on
account of its broader leaves, the greater number
of spikelets, and the larger utricles. The type-
collection of C. speciosa (WALLICH 3391) does not
answer RAYMOND’s description of this species.
I agree with NeLMes that C. courtallensis is con-
specific with C. speciosa; the specimens from
Sumatra and Java agree very well with the type of
C. courtallensis (WIGHT 991 in K).

The Bornean specimens are remarkable by the
narrow leaves, the long, tail-like g part of the
spikelets, and especially by the infundibuliform
glumes of the ¢ flowers (the last character not
mentioned by NELMgs). I have not seen the Tonkin
specimens NELMES (1955) referred to C. speciosa
var. angustifolia (BooTT) KUK. They have leaves
only 11/,-2 mm wide, and utricles 3!/,-4 mm long.
It is not clear how NELMEs distinguished between
this variety and his C. stenura.

Another member of sect. Radicales from Lower
Burma was described as C. pterocaulos NELMES in
Hook. Ic. Pl. 35 (1947) t. 3467; Mém. Mus. Nat.
Hist. Paris B4 (1955) 159, according to NELMES a
very distinct species because of its widely winged
stems, its sometimes empty bract, and the very

stout 2 part of its spikelets. In some specimens
collected in Thailand, Chiengmai, Doi Chiengdao,
at 1200 m (SLEUMER & SMITINAND 1063, L), the
stems are strikingly winged (4 mm broad) and the
bracts very long and sometimes empty, but the
glumes are only 3-3'/, mm long, the (young)
utricles 4 mm (not 4-5 mm and 6-7 mm respec-
tively); the 2 part of the spikelets is therefore not
stouter than in C. speciosa. I doubt whether
C. pterocaulos is specifically distinct from C.
speciosa.

The polymorphism of the group is badly in need
of special study.

The record of C. speciosa var. angustifolia for
NE. New Guinea, based on CLEMENS 7909a (KUK.
Bot. Jahrb. 70, 1940, 465) is very doubtful for
plant-geographical reasons.

The filaments are often coherent to such a degree
as to give the impression of their being connate.

38. Carex tricephala Boeck. Flora 58 (1875) 263;
Kuk. Pfl. R. Heft 38 (1909) 289; Koorp. Exk. FI.
Java 1 (1911) 205; Kix. Bull. Jard. Bot. Btzg III,
16 (1940) 316; Netmes, Kew Bull. (1950) 190;
Reinwardtia 1 (1951) 256; Mém. Mus. Hist. Nat.
Paris n.s. B4 (1955) 117; KoyAMaA, Contr. Inst. Bot.
Un. Montréal n. 70 (1957) 15, t. 1; RAyM. Mém.
Jard. Bot. Montréal n. 53 (1959) 76; Dansk Bot.
Ark. 23 (1965) 260; KERN in Back. & Bakh. f. FI.
Java 3 (1968) 491. — C. madoerensis CLARKE,
J. Linn. Soc. Bot. 37 (1904) 15; Kew Bull. add.
ser. 8 (1908) 77. — C. thorelii Camus, Not. Syst. 1
(1910) 295; Fl. Gén. I.-C. 7 (1922) 196; NELMEs,
Kew Bull. (1946) 15, 23. — C. plesiocephala TurR.
Kew Bull. (1910) 385; ibid. (1912) 427. — C. his-
pidangula KoyAMA, Nat. Canad. 82 (1955) 200,
t. 1. — Fig. 121.

Rhizome shortly creeping, woody, covered with
the fibrous, fuscous remains of old scales. Stems
slender, triquetrous, narrowly winged just below
the inflorescence, smooth below, antrorsely
scabrous-setulose above, 10-50 cm by 1 mm (up to
2 mm in the winged part), the base clothed with the
fibrous remains of old leaf-sheaths. Leaves basal
and 1-2 cauline, shorter than to much exceeding
the stems, rigid to rather weak, flat, with revolute
margins when dry, long-attenuate, scabrid in the
upper part, light green, 3-10 mm wide; cauline
leaves long-sheathing; ligule short, triangular.
Inflorescence spiciform, consisting of (1—)3(-4)
dense, sessile, ovoid or subglobose spikelets
0-5 cm distant, and 6-10(-12) mm long and wide.
Lower 1-2 bracts foliaceous, usually much exceed-
ing the inflorescence, not sheathing, amplexicaul
by ciliolate auricles, upper much reduced. Spikelets
androgynous, the 3 part about as long as the 9, but
inconspicuous when the utricles are fully developed.
Glumes oblong-ovate to ovate, membranous,
nerved, densely setulose, white, or brownish with
white margins, 2!/,4'/, mm long; midnerve
excurrent in a scabrid, 1/.-1 mm long awn.

1979]

Cy PERACEAE—II (Kern & Nooteboom)

157

Utricles trigonous, ovoid or rhomboid-ovoid,
membranous, patent, slenderly nerved, rather
abruptly narrowed into the beak, densely hispidu-
lous in the upper 2/3, whitish or brownish, 4-6 by
14/,-2'/, mm; beak conic-cylindric, bidentate, with
slightly oblique mouth, 1-2'/, mm long. Nut
triquetrous, ellipsoid, rhomboid, or slightly obo-
void, scarcely stipitate, cinereous, dark brown
on the angles, 2'/,-3 by 1°/,-2 mm. Style-base
pyramidally thickened, persistent on the nut. Stig-
mas 3.

Distr. Yunnan, Upper Burma, Thailand, Indo-

China; in Malesia only known from Madura I. off
NE. Java (type locality !).

Ecol. In grassy fields on heavy calcareous marl,
in muddy places in teak-forests, at low altitude
(up to 200 m), obviously bound to a distinct
seasonal climate and its range apparently coin-
ciding with that of teak. See Netmes 1951, /.c.

Note. Newmes placed this species, together with
C. malaccensis and C. ramosii in sect. Stramentitiae,
but as RAYMOND 1959, /.c., pointed out, it has very
little in common with the other members of that
section.

13. Section Trachychlaenae

DreJER, Symb. Caric. (1844) 9; KUK. Pfl. R. Heft 38 (1909) 415. — Sect. Anomalae
(non CAREY) NELMES, Reinwardtia 1 (1951) 413; Raym. Mém. Jard. Bot. Mon-

tréal n. 53 (1959) 54, 71.

Type species: Carex hispida WILLD. (lectotype).

39. Carex maculata Boott, Trans. Linn. Soc. 20
(1846) 128; Ill. 1 (1858) 9, t. 26; Boeck. Linnaea 41
(1877) 191; Bento. Fl. Austr. 7 (1878) 447;
CLARKE, FI. Br. Ind. 6 (1894) 735; Kix. Pfl. R.
Heft 38 (1909) 427, incl. var. neurochlamys KUK.;
Ouw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936)
434; Kix. Bull. Jard. Bot. Btzg III, 16 (1940) 317,
incl. var. sanguineo-squamata KUx. et f. humilior
Kux.; Ouwt & Koyama, Misc. Rep. Nat. Sc.
Mus. Tokyo n. 5 (1952) 1, t. 1; AKIYAMA, Car. Far
East. Reg. Asia (1955) 120, t. 105, f. 1; YOSHIKAWA,
Ic. Jap. Carex 2 (1958) 206, t. 103; NELMEs, Kew
Bull. (1950) 205; Reinwardtia 1 (1951) 414; Rayo.
Mém. Jard. Bot. Montréal n. 53 (1959) 71; Dansk
Bot. Ark. 23 (1965) 259; KERN in Back. & Bakh.
f. Fl. Java 3 (1968) 489; STEEN. Mt. FI. Java (1972)
pl. 14-5. — C. neurochlamys F.v.M. Fragm. 8
(1874) 258; NeLmes, Kew Bull. (1949) 391; Rein-
wardtia 1 (1951) 413. — C. samoensis BOECK.
Bot. Jahrb. 25 (1898) 588. — C. elibates NELMES,
Kew Bull. (1937) 353; ibid. (1950) 205; Reinwardtia
1 (1951) 416. — C. pruinosa (non BooTT) KUK. Bull.
Jard. Bot. Btzg III, 16 (1940) 317. — C. pruinosa
Boott f. tristigmatosa Back. Bekn. Fl. Java
(em. ed.) 10 (1949) fam. 246, p. 62. — Fig. 121.
Rhizome short, woody. Stems densely tufted,
slender, erect, trigonous, smooth, (5—)15—60 cm by
1-11/, mm, surrounded below the leaves by a few
ferrugineous or brownish, bladeless sheaths or their
fibrous remains. Leaves subbasal, shorter to longer
than the stems, flat with often revolute margins,
greyish green, scabrous towards the long-
attenuated apices, (1-)3-8 mm wide; sheaths pale
to ferrugineous, membranous in front. Lower
bracts sheathing, leaf-like, usually overtopping the
inflorescence. Spikelets (2-)3—5(—6), upper approxi-
mate, lower distant, cylindric, densely flowered;
terminal spikelet 3, sessile, pale, 1-2 mm thick,

lateral ones 9, suberect, dark, 1-4 cm by 3-5 mm,
upper sessile or shortly peduncled, lower on short
to long, firm, smooth peduncles. Glumes ovate or
oblong-ovate, acute or subobtuse, thin, translucent,
often minutely ciliolate, ferrugineous to castaneous
with narrow whitish-hyaline margins and 3-nerved,
pale green central stripe, muticous or scarcely
mucronulate, 2-3 mm long. Utricles ellipsoid,
compressed-trigonous, membranous, distinctly
several-nerved, suberect to patulous, straight,
glabrous but densely ferrugineous-papillose,
scarcely stipitate, 2-3(-4) by 1-13/, mm, sub-
abruptly contracted into a very short, up to '/, mm
long, entire or slightly emarginate beak. Nut
ellipsoid or obovoid, triquetrous, shortly stipitate
and beaked, 11/,-2 by 3/,-1!/, mm, both stipe and
beak usually bent. Style-base oblique, somewhat
thickened. Stigmas 3 (rarely 2 in some fruits).

Distr. Widely distributed from Ceylon and
India through Thailand, China, and Korea to
Formosa, Japan and the Ryu Kyu Is., and through
Malesia to Queensland, New South Wales, New
Caledonia and Samoa; in Malesia: Malay Penin-
sula (Pahang), N. Sumatra (Atjeh), Java (West:
Mt Papandajan; Central: Diéng Plateau; East:
Jang plateau), Lesser Sunda Is. (Flores), Celebes
(Minahassa, Poso), and New Guinea. Distr. map:
MeuseEL, Vergl. Arealkunde 2 (1943) Karte 32c.

Ecol. In swamps, marshes, boggy meadows, wet
mountain heaths, open places in mossy woods,
1600-3500 m.

Vern. New Guinea: sisik, Tomba, toni, ititu,
Mendi, era, teleleme, Onim.

Notes. Very variable. I cannot follow NELMES
in assigning specific rank to C. neurochlamys and
C. elibates. His descriptions are fairly well covered
by that of C. maculata. C. elibates has darker
coloured glumes and often stiffer and narrower

158

FLORA MALESIANA

[ser. I, volags

leaves than typical C. maculata, and for this reason
KUKENTHAL distinguished it as var. sanguineo-
squamata. The other characters given by NELMES
are valueless: in all Carices with a 3 terminal
spikelet this spikelet may bear some 9 flowers; the
very short beak of the utricle in C. maculata is
variable in length, and so it is in C. elibates.
F.v.MUELLER distinguished C. neurochlamys from
C. maculata mainly by the non-papillose, strongly
nerved utricles, a mistake apparently due to
inaccurate observation. According to NELMES the
glumes in C. neurochlamys are usually mucronulate,
not usually so in C. maculata; 1 fail to see any
difference. KUKENTHAL reduced C. neurochlamys
to varietal rank and distinguished it by the stouter

stems, the leaves much overtopping the inflores-
cence, and the many-nerved, less densely papillose
utricles. These characters are not found in the New
Guinea specimens referred to C. neurochlamys by
NELMES.

Carex maculata is closely related to species of
sect. Praelongae (C. phacota and C. pruinosa). One
collection of N. Sumatra (DE WILDE c.s. 13251) has
several fruits with only 2 styles, and becoming as
long as 4 mm, thus exactly resembling the fruits of
C. pruinosa. These fruits occur in the same spikelet
as the ‘normal’ 3-styled fruits. Another deviating
feature in this collection is that some of the utricles
are not glabrous, but minutely scabrid on the
margins. — (Noot.)

14. Section Capitellatae

MeInsu. Act. Hort. Petrop. 18 (1901) 280, 309; OHw1, Mem. Coll. Sc. Kyoto Imp.
Un. B11 (1936) 437; NELMES, Reinwardtia 1 (1951) 404; Raym. Mém. Jard. Bot.
Montréal n. 53 (1959) 17, 18. — Sect. Rarae CLARKE, Kew Bull. add. ser. 8 (1908)
143. — Sect. Unciniaeformes Kix. subsect. Capitellatae (MEINSH.) KUK. Pfi. R.
Heft 38 (1909) 100. — Sect. Extensae Fries subsect. Capitellatae (MEINSH.)
KoyaMa, J. Fac. Sc. Un. Tokyo III, 8 (1962) 202.

Type species: Carex capitellata Botss. & BAL.

40. Carex capillacea Bootrt, Ill. 1 (1858) 44, t. 110;
Boeck. Linnaea 39 (1875) 37; BENTH. FI. Austr. 7
(1878) 438; CLarKE, FI. Br. Ind. 6 (1894) 713; J.
Linn. Soc. Bot. 37 (1904) 7; Philip. J. Sc. 2 (1907)
Bot. 107; Onw1, Mem. Coll. Sc. Kyoto Imp. Un.
B11 (1936) 441; S. T. Blake, J. Arn. Arb. 28 (1947)
101; NELMes, Kew Bull. (1949) 381; Reinwardtia 1
(1951) 405; ibid. 2 (1954) 381; AktyAMA, Car. Far
East. Reg. Asia (1955) 42, t. 6, f. 1 A-D; Yosui-
KAWA, Ic. Jap. Carex 2 (1958) 218, t. 109; RAyYM.
Mém. Jard. Bot. Montréal n. 53 (1959) 18;
Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 206;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 488;
STEEN. Mt. FI. Java (1972) pl. 14-8. — C. simplicis-
sima F.v.M. Fragm. 9 (1875) 191. — C. rara (non
Boott) STAPF, Trans. Linn. Soc. II, Bot. 4 (1894)
246; CLARKE, J. Linn. Soc. Bot. 37 (1904) 7; KUK.
Bull. Jard. Bot. Btzg III, 16 (1940) 313; Back.
Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 64.
— C. rara Boott ssp. capillacea KUK. in Mats.
Index Pl. Jap. 2 (1905) 130, guoad synon.; Pfi.
R. Heft 38 (1909) 102; Philip. J. Sc. 6 (1911)
Bot. 58; MeErRR. En. Philip. 1 (1923) 141; STEEN.
Trop. Natuur 19 (1930) 87, 89 f. 14. — Fig. 121,
130.

Rhizome short, woody. Stems densely tufted,
slender, erect, triquetrous, smooth or slightly
scaberulous just below the inflorescence, 5-50 cm
by 1/,-3/, mm, clothed below the leaves by brown-
ish, bladeless sheaths and their fibrous remains.
Leaves subbasal, much shorter than to slightly

exceeding the stems, flat or canaliculate, smooth or
nearly so, erect, !/,-2 mm wide; sheaths pale,
membranous in front. Spikelet solitary, terminal,
androgynous, ovoid or shortly cylindric, densely
but rather few-flowered, 5-15 mm long, the gd part
c. 1 mm thick, shorter than to about as long as the
2 part, the latter becoming 4-6 mm thick. Glumes
ovate or oblong-ovate, obtuse to acutish, muticous,
translucent, ferrugineous to brown with narrow
paler margins and 3-nerved, pale central stripe,
caducous, 1!/,-3 mm long. Utricles oblong-ovoid or
oblong-ellipsoid, rounded at the base, obscurely
trigonous, subinflated, membranous, finally widely
patent to somewhat reflexed, glabrous, several-
nerved, scarcely stipitate, often minutely reddish
resinous-dotted, (11/,—-)2!/,-31/,(-4) by c. 1 mm,
rather gradually tapering to a very short, subentire
or slightly emarginate beak. Nut ellipsoid or
oblong-ellipsoid, triquetrous, shortly stipitate,
abruptly short-beaked, stramineous to brownish,
11/,-2!/, by c. 1 mm. Style-base slightly thickened.
Stigmas 3.

Distr. From the Himalayan region to Annam,
Manchuria, Korea, Japan, and extending as far
north as Sachalin (var. sachalinensis OHW1);
through Malesia to Australia (New South Wales)
and New Zealand; in Malesia: Sumatra (North:
Atjeh; Central West: G. Singalang), W. Java
(Mt Papandajan), N. Borneo (Mt Kinabalu), SW.
Celebes (Mt Rante Mario), Philippines (Luzon),
and New Guinea.

1979]

Cy PERACEAE—II (Kern & Nooteboom)

159

Ecol. Swamps, wet alpine grasslands, open
seepages, wet borders of small streams, marshy
places in forests, 2000-4000 m.

Vern. New Guinea: ere, tep, Mendi lang.

Notes. Closely related to the Indian C. rara
Bootrt, Proc. Linn. Soc. 1 (1845) 284; Ill. 1 (1858)
44, t. 109. Boortr distinguished C. capillacea from
C. rara by the capillary stems and leaves, the
shorter, ovoid spikelet, the smaller, reflexed,
puncticulate utricles, and the caducous glumes.
Except for the much longer, linear spikelets ($ part
much longer than the 2 one), and the suberect
utricles of C. rara, these characters are unreliable.
The two might possibly better be regarded as
subspecies of a single species.

In typical C. capillacea (var. capillacea) the stems
and leaves are filiform and the utricles 2—2'/, mm
long. The above description comprises also
stouter plants with leaves up to 2 mm wide and

utricles 3-4 mm long, which have been distin-
guished as:

var. sachalinensis (F. ScHmrpDT) OHw1, Mem. Coll.
Sc. Kyoto Imp. Un. B11 (1936) 442; Axryama,
Car. Far East. Reg. Asia (1955) 42, t. 6, f. 1 E-H;
YOSHIKAWA, Ic. Jap. Carex 3 (1960) 386, t. 193. —
C. nana Boott in A. Gray, Mem. Am. Ac. n.s. 6
(1859) 418; Ill. 4 (1867) 139, t. 44, f. 2, non CHAM.
ex STEUD. 1855. — C. uda MAxiM. var. sachalinensis
F. SCHMIDT, Reisen Amur]. (1868) 191. — C. capil-
lacea var. nana FRANCH. Nouv. Arch. Mus. Hist.
Nat. Paris III, 8 (1896) 198. — C. rara ssp. capil-
lacea var. nana KUx. Pfi. R. Heft 38 (1909) 103. —
C. capillacea var. major NELMES, Kew Bull. (1949)
381; Reinwardtia 1 (1951) 406.

Known from Sachalin, Japan, Korea, and
Malesia. The Malesian specimens are often
difficult to refer to one of the varieties.

15. Section Rhizopodae

OuwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 443; NeLMegs, Reinwardtia 1
(1951) 411; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 17. — Sect. Extensae
Fries subsect. Rhizopodae (OHW1) KoyAMaA, J. Fac. Sc. Un. Tokyo III, 8 (1962)

202.

Type species: Carex rhizopoda MAXIM.

41. Carex eremostachya S. T. BLAKE, J. Arn.
Arb. 28 (1947) 99, f. 1A; Netmes, Kew Bull.
(1949) 381; Reinwardtia 1 (1951) 412. — Fig.
121.

Rhizome creeping, slender, 1-2 mm thick,
covered with brown, striate scales. Stems approxi-
mate or loosely tufted, erect or suberect, slender
and rather flaccid, triquetrous, scabrid at the top,
20-45 cm by !/,-1 mm. Leaves subbasal, as long as
to longer than the stems, flat, shortly attenuate,
glabrous, scabrid on margins and nerves towards
the apex, 2-3 mm wide, the lower ones reduced to
brown, entire, bladeless or short-bladed sheaths.
Spikelet 1, terminal, erect, androgynous, oblong-
ellipsoid to shortly cylindric, densely many-
flowered, 10-15 by 5-6 mm, the ¢ part shorter than
the 2 one. Glumes ovate or ovate-lanceolate, acute
or the upper obtusish, thin, glabrous, reddish
brown with 3-nerved, pale, central stripe, 3-4 mm

long, lower mucronate (mucro up to 1 mm),
upper muticous. Urtricles ellipsoid, triquetrous,
angled dorsally, obliquely erect, membranous,
glabrous and smooth, distinctly 4-S-nerved on each
face, pale green, 4*/,-5 by 1'/,-1'/, mm, shortly
stipitate, rather abruptly narrowed into a slender,
linear, bidentate, c. 11/, mm long beak. Nut obo-
void, triquetrous, apiculate, c. 2 by 11/; mm. Style-
base scarcely thickened. Stigmas 3.

Distr. Malesia: W. New Guinea, 9 km NE of
Lake Habbema; only known from the type
collection.

Ecol. Massed on open beaches of small stream
in forest, 2800 m.

Note. Closely allied to C. rhizopoda MAxIM.
from Japan, which is however clearly distinct by its
more flaccid, usually broader leaves, and its longer,
oblong-cylindric spikelet with pale green glumes
much shorter than the utricles.

16. Section Anomalae

[Carey in Gray, Man. Bot. N. Un. St. (1848) 557, pro subsect.]; MACKENZ. N.
Am. FI. 18 (1935) 339. — Japonicae FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris
III, 10 (1898) 107, nomen. — Sect. Tumidae KUx. Pfl. R. Heft 38 (1909) 611;
AKIYAMA, J. Fac. Sc. Hokkaido Imp. Un. V, 2 (1932) 188; NELMEs, Reinwardtia 1
(1951) 394; non MeEINsH. 1901. — Sect. Dispalatae OHW1, Mem. Coll. Sc. Kyoto

160

FLORA MALESIANA

[ser. I, vol. 92

Imp. Un. B11 (1936) 480; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 82. —
Sect. Confertiflorae FRANCH. ex OHWI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936)
474; AktyAMA, Car. Far East. Reg. Asia (1955) 144. — Sect. Molliculae OHWI,
Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 450; KoyAMA, Bot. Mag. Tokyo 72
(1959) 307; Quart. J. Taiwan Mus. 13 (1960) 226. — Sect. Extensae FRIES ser.
Tumidae (KUK.) KoyaMa, J. Fac. Sc. Un. Tokyo III, 8 (1962) 202.

Type species: Carex scabrata SCHWEIN. (lectotype).

42. Carex alopecuroides D. Don, Trans. Linn. Soc.
14 (1825) 332; Prod. Fl. Nepal. (1825) 43; Boott,
Ill. 2 (1860) 88, t. 258; Boeck. Linnaea 41 (1877)
172:

In Malesia only:

var. chlorostachys (D. Don) CLARKE, J. Linn. Soc.
36 (1903) 271 (‘chlorostachya’); KOYAMA, Nat.
Canad. 82 (1955) 199; Contr. Inst. Bot. Un.
Montréal n. 57 (1957) 14; Raym. Mém. Jard. Bot.
Montréal n. 53 (1959) 85; KERN in Back. & Bakh.
f. Fl. Java 3 (1968) 490. — C. chlorostachys
D. Don, Trans. Linn. Soc. 14 (1825) 330; Prod.
Fl. Nepal. (1825) 43; non StTEvEN, 1813. —
C. doniana SPRENG. Syst. 3 (1826) 825; NEEs in
Wight, Contr. (1834) 128; DreJer, Symb. Caric.
(1844) 26, t. 13; S. T. BLAKE, J. Arn. Arb. 28 (1947)
115; Kew Bull. (1950) 204, incl. var. cacuminis
NELMES; NELMES, Reinwardtia 1 (1951) 399; ibid. 2
(1954) 381; AxtyAMA, Car. Far East. Reg. Asia
(1955) t. 161. — C. japonica (non THUNB.) BooTT,
Ill. 2 (1860) 88, t. 257, excl. var. minor Bootrt;
Boeck. Linnaea 41 (1877) 283, excl. syn. SCHKUHR
t. Ww f. 110; CLARKE, FI. Br. Ind. 6 (1894) 736;
BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246,
p. 70. — C. japonica var. chlorostachys KUK. ex
Matsuo. Ind. PI. Jap. 2 (1905) 116; Pfl. R. Heft 38
(1909) 620; Merr. En. Philip. 1 (1923) 139;
Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 212.
— C. subtransversa CLARKE, Philip. J. Sc. 2 (1907)
Bot. 108; Kew Bull. add. ser. 8 (1908) 92; KUx.
Pfil. R. Heft 38 (1909) 614, in nota; Philip. J. Sc. 6
(1911) Bot. 63; Merr. En. Philip. 1 (1923) 142;
S. T. BLAKE, J. Arn. Arb. 28 (1947) 115; NELMEs,
Reinwardtia 1 (1951) 401. — C. japonica var.
mesogyna Kix. Bot. Jahrb. 69 (1938) 265. —
C. japonica ssp. subtransversa (CLARKE) KOYAMA,
J. Fac. Sc. Un. Tokyo III, 8 (1962) 213. — Fig. 121.

Rhizome emitting long, slender stolons covered
with pale brown sheaths. Stems tufted, slender,
erect, very acutely triquetrous to narrowly winged,
smooth or scabrid on the angles above, (2—)30-
90 cm by 1-2 mm, surrounded below the leaves by
a few pale, bladeless sheaths. Leaves subbasal and
1 higher on the stem, shorter to much longer than
the stem, flat, stiffish, long-acuminate, scabrid on
the margins and often asperous on the upper
surface, 2-10 mm wide. Spikelets (2—)3-6, erect,
cylindric, (1!/,-)2-5 cm long, approximate and
fastigiate, or lower 1-2 somewhat distant, sessile,

or lower shortly peduncled; terminal spikelet ¢
(not rarely gynaecandrous), pale, 1-3 mm thick,
lateral ones 2, greenish or stramineous, 4-6 mm
thick. Lower bracts foliaceous, overtopping the
inflorescence, not sheathing, upper much reduced.
Glumes oblong-ovate, acute to obtuse, 2—2!/, mm
long, whitish with 3-nerved, green, central stripe
excurrent in a scabrid, (!/,—)1-17/, mm long awn.
Utricles ellipsoid, obscurely trigonous, subinflated,
membranous, plurinerved, glabrous, ultimately
patent, straight, greenish to stramineous, 3-4 by
1-11/, mm, rather gradually narrowed into a
1-13/, mm long, linear-conic, bidenticulate, smooth
or almost smooth beak. Nut ellipsoid, obovoid, or
oblong-obovoid, triquetrous, 11/,-2 by 3/,-1 mm.
Style straight or slightly bent at the scarcely
thickened base. Stigmas 3, about half as long as
the utricle.

Distr. Nepal, Sikkim, and Khasia to Central
and S. China, Korea, Japan, Ryu Kyu Is., Formosa,
Tonkin, and Annam, and Malesia: Sumatra
(Atjeh; W. Coast: Mt Kerintji), Java (Mt Wajang
in W., Mt Tengger: Ranu Regulo in E.), Central
Celebes (Mt Kambuno), Philippines (Luzon), and
New Guinea.

Ecol. Margins of lakes, riversides, openings in
the mossy forest, also on dry, open slopes, stated
to be a common trackside sedge in New Guinea;
1500-2900 m.

Vern. New Guinea: akele, Lalibu.

Notes. The only collection known from Celebes
(EYMA 1395) was distinguished as C. doniana var.
cacuminis NELMES, which has no taxonomic value;
the dwarfish habit has apparently been caused by
continuous grazing of anoas, the small wild buffalo
of Celebes, Bos (Bubalus) depressicornis.

Both C. alopecuroides and its var. chlorostachys
are treated as varieties of C. japonica THUNB. by
KUKENTHAL, and also recently by KoyAMA, but
C. japonica seems to be distinct by its short-
cylindric, not crowded, peduncled, usually pendu-
lous spikelets, not distinctly awned glumes, and its
very long stigmas. In typical C. alopecuroides the
leaves are usually narrower than in var. chloro-
stachys, the spikelets very slender, and the utricles
somewhat narrower, less inflated, but the two are
very near to each other and connected by inter-
mediates. C. subtransversa CLARKE was described
from N. Luzon, where also C. alopecuroides var.
chlorostachys occurs. I fail to see differences

Cy PERACEAE—II (Kern & Nooteboom)

161

Fig. 130. Carex capillacea Boott. Habit, x '/;
(from Mt Papandajan, VAN STEENIS).

between the two but for the narrower leaves of the
former, which is said to differ also by the suberect—
not patent — utricles; the specimens are not fully
ripe and the degree of reflexion of the utricles

depends upon maturity. KUKENTHAL placed C.
subtransversa under C. brownii and its var. trans-
versa (BOOoTT) KixK., with which it is certainly not
closely related.

43. Carex brownii TUCKERM. En. Meth. Car. (1843)
21 (or 15? ‘brownii’?); Boott, Ill. 4 (1867) 161,
t. 532 (pl. dextrae); Boeck. Linnaea 41 (1877) 151,
incl. var. viridis BOECK.; BENTH. Fl. Austr. 7 (1878)
447; Kix. Pfi. R. Heft 38 (1909) 612, f. 104 A-D;
OxuwIl, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936)
477: S. TsBeAKe; J. Arn: Arb: 28 (1947)-115;
NeELMES, Kew Bull. (1949) 384; Reinwardtia 1
(1951) 398; Koyama, Act. Phytotax. Geobot. 18
(1959) 23; J. Fac. Sc. Un. Tokyo III, 8 (1962) 215;
Phytologia 17 (1968) 409, t. 16. — C. striata R. BR.
Prod. (1810) 243; KuNTH, En. 2 (1837) 458;
STEUD. Syn. 2 (1855) 226; Dreser, Symb. Caric.
(1844) 28, t. 15; non Micux, 1803. — C. nippo-
sinica Onwtl, Act. Phytotax. Geobot. 11 (1942) 255;
AKIYAMA, Car. Far East. Reg. Asia (1955) 144,
t. 133; YOSHIKAWA, Ic. Jap. Carex 2 (1958) 256,
t. 128.

a. ssp. brownii. — Fig. 122.

Rhizome short, without stolons. Stems slender,
erect, triquetrous, smooth or almost so, 25-75 cm
by 1-1!/, mm. Leaves basal and often 1 higher up
the stem, flat, weak, shorter than the stems, long-
acuminate, scabrous on nerves and margins,
3-4 mm wide, lowest ones reduced to bladeless,
reddish-brown sheaths. Spikelets 3—4(-5), erect,
short-cylindric to cylindric, upper approximate,
fastigiate, lowest often more or less distant;
terminal spikelet ¢, or rarely gynaecandrous,
subsessile, pale, (!/,-)1-2!/, cm by 2(-3) mm;
lateral spikelets 2, subsessile or lowest sometimes
on a long, exserted peduncle, short-cylindric,
obtuse, densely flowered, 11/,-2'/.(-3) cm by
5-6 mm. Lowest bract foliaceous, overtopping the
inflorescence, more or less sheathing, upper much
shorter, not or hardly exceeding the inflorescence.
Glumes ovate or oblong-ovate, obtuse, scarious,
1-2 mm long, whitish with 3-nerved green central
stripe excurrent in a scabrous, 1/,-3'/, mm long
awn. Utricles broadly ellipsoid to subglobose,
obscurely trigonous, inflated, membranous,
strongly many-ribbed, glabrous, straight, patent,
dark olive-green to dark brown, contrasted against
the white glumes, 3-4 by 1'/3-13/, mm, suddenly
contracted into a very short, bidenticulate beak.
Nut obovoid, triquetrous, yellowish, 2'/,-2'/, by
11/,-1?/, mm, with a very short, (sometimes
indistinctly) bent stipe and beak. Style-base
slightly thickened. Stigmas 3.

Distr. Japan, Korea, Formosa, E. China
(Yangtze R. valley), Australia (New South Wales,
Victoria), New Zealand; in Malesia: New Guinea
(W. New Guinea: Balim R. valley; Terr. of New
Guinea: W. Highlands, lower Tale valley and

162 FLORA MALESIANA [ser. I, vol. 9!

= VLSSMA "78

\ |

| h
Fig. 131. Inflorescence (nat. size), spikelets and (occasionally) venation (x 2). — 48. Carex michauxiana
BoECK., a. — 18b. C. perakensis CLARKE var. borneensis (CLARKE) Noot., b. — 18a. var. perakensis, c—d.
— 19a. C. turrita CLARKE var. turrita, e-f. — 50. C. maubertiana Boott, g. — 64. C. echinata MurR., h. —

15. C. baccans NEES, i. — 51. C. graeffeana BOECK., j (a SCHODDE 1992, b BROOKE 8561, c-d VAN STEENIS
8277, e-f ANU 15532, g KosTERMANS 14005, 4 DE WILDE c.s. 13323, i ELBERT 62, j SINCLAIR 979").

1979 |

a b C d

Cy PERACEAE—II (Kern & Nooteboom)

e

Fig. 132. Glumes, x 10. — 46. Carex brachyathera OHW1, a. — 49. C. pseudocyperus L. var. fascicularis

(SOLAND. ex Boott) Boott, b. — 52. C. phacota SPRENG., c. — 53. C. pruinosa Boott, d. — 54. C. teres

Boott, e. — 56. C. bilateralis Hayata, f. — 57. C. brunnea THUNB., g (a BRASS 9803, b EyMA 4709,
c JERMY 4634, d VAN STEENIS 4624, e DE WILDE c.s. 13329, f Brass 9515, g EYMA 3862).

Sirunki; E. Highlands, Aiyura, and Kainantu,
Morobe Distr., Langiman R.). Distr. map in Acta
Phytotax. Geobot. 18 (1959) 24.

Ecol. In wet places, pools, drains, etc., also
open grassy area; 1500-2500 m.

Vern. Kariandend, Enga lang.

b. ssp. transversa (BOOoTT) KERN, stat. nov. —
C. transversa Boott, Perry Exp. 2 (1857) 324;
Ill. 4 (1867) 202; FRANcH. & Sav. En. Pl. Jap. 2
(1879) 149, incl. var. dissociata FRANCH. & SAV.;
FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III,
10 (1898) 48, t. 3, f. 1; CLARKE, J. Linn. Soc. Bot.
36 (1904) 314; OHw1, Mem. Coll. Sc. Kyoto Imp.
Un. B11 (1936) 478; AkrtyAma, Car. Far East. Reg.
Asia (1955) 145, t. 134; YosHikAwa, Ic. Jap.
Carex 2 (1958) 258, t. 129. — C. brownii var.
transversa KUK. ex MaAtsum. Index Pl. Jap. 2
(1905) 103; Kux. Pfl. R. Heft 38 (1909) 614. —
C. furusei KoYAMA, J. Jap. Bot. 30 (1955) 135, pro
C. brownii x transversa. — C. brownii var.
dissociata KoyaMa, J. Fac. Sc. Un. Tokyo III, 8
(1962) 215. —.Fig. 122.

Utricles ovoid-ellipsoid, up to 6 mm long,
gradually narrowed into a c. 2 mm long beak.

Distr. Japan, Ryu Kyu Is., Korea, China
(Yangtze valley region); in Malesia: New Guinea
(W. Highlands, Sirunki).

Ecol. In New Guinea near water-course on clay
soil, at c. 2500 m.

Note. The long-beaked utricles are so different
from those of C. brownii s.s., that I prefer to treat
this taxon as a subspecies, not as a variety of
C. brownii.

44. Carex oedorrhampha NeELMeES, Kew Bull.
(1939) 659; ibid. (1949) 384; Reinwardtia 1 (1951)
396; ibid. 2 (1954) 381; RAymM. Mém. Jard. Bot.
Montréal n. 53 (1959) 83; Dansk Bot. Ark. 23
(1965) 261; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 490. — C. tumida Boortrt, Ill. 1 (1858) 66,
t. 181; Boeck. Linnaea 41 (1877) 243; CLARKE,
Fl. Br. Ind. 6 (1894) 741; J. Linn. Soc. Bot. 37
(1904) 16; Kix. Pfl. R. Heft 38 (1909) 615; Bull.
Jard. Bot. Btzg III, 16 (1940) 321, non BEILSCHM.
1850. — C: olivacea (non BootT) KUK. Bot. Jahrb.
70 (Jan. 1940) 467; Bull. Jard. Bot. Btzg III, 16
(Feb. 1940) 321, incl. var. altissima KUK.; BACK.
Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 70,
p.p. — C. oedorrhampha var. arfakiana OHWI, Bot.
Mag. Tokyo 56 (1942) 214. — C. oedorrhampha
var. microcarya NELMES, Kew Bull. (1950) 204;
Reinwardtia 1 (1951) 398. — Fig 122.

Rhizome short, in large tussocks, without
stolons. Stems tufted, slender, erect, triquetrous,
smooth, 50-125 cm by 2-3 mm below. Leaves
subbasal and 1-2 higher up the stem, about as long
as the stem, flat, stiffish, long-acuminate, scabrous
on margins and main nerves, 4-8 mm wide, few
lowest reduced to reddish purple sheaths readily
fraying into thin strips; ligula elongate, up to
11/, cm long. Spikelets 4-6, erect or subcernuous,
upper approximate and fastigiate, lower distant;
terminal spikelet 3, long-linear, subsessile, not or
slightly exceeding the uppermost lateral spikelet,
pale, 2-6 cm long, 1-2 mm thick, lateral spikelets
2, narrowly cylindric, densely flowered, 2!/,-12 cm
long, 4-6 mm thick, upper on shortly, lower on
long-exserted, scabrid peduncles. Lower bracts

164

FLORA MALESIANA

[ser. I, vol. 91

foliaceous, much exceeding the inflorescence, long-
sheathing, upper more or less reduced. Glumes
oblong to ovate-lanceolate, obtuse, translucent,
glabrous or sparsely hispidulous, 13/,-2 mm long,
whitish with 3-nerved, green, central stripe excur-
rent in a scabrid awn 1-2 mm long. Utricles
ellipsoid, membranous, somewhat inflated, pluri-
nerved, glabrous, obliquely erect to subpatent,
olive-brown to fuscous, 3-3/, by 1-11/,; mm,
rather gradually narrowed into a 1—-11/, mm long,
conic-linear, smooth, minutely notched beak which
is often somewhat swollen at or below the middle.
Nut ellipsoid, triquetrous, yellow to brownish,
shortly stipitate, distinctly beaked, (11/,-)2-2!/, by
c. 1 mm. Style straight, thickened at the base.
Stigmas 3.

Distr. E. Himalaya, S. China (Yunnan), N.
Thailand, Tonkin, Assam; in Malesia: Sumatra
(Atjeh; W. Coast Res.: Mt Kerintji), W. Java
(Mt Papandajan), Moluccas (Buru), New Guinea.

Ecol. Wet places in forests, lake margins,
swampy grasslands, 1200-2400 m.

Notes. Boort’s figure, showing a distinct
swelling in the middle of the beak (hence his specific
epithet) is, according to CLARKE, taken from a not
fully ripe collection. In mature utricles the lower
half of the beak is cylindric, somewhat swollen, the
upper conic. In the specimens from Mt Kerintji the
beak is scarcely swollen; they were distinguished as
var. microcarya NELMES.

Besides to C. olivacea the species is closely related
to C. ischnostachya STEUD. from Japan, which is
easily distinguishable by the muticous glumes, but
otherwise very similar.

45. Carex olivacea Boott, Proc. Linn. Soc. 1
(1846) 286; Ill. 1 (1858) 56, t. 149; CLarkg, FI. Br.
Ind. 6 (1894) 741; J. Linn. Soc. Bot. 37 (1904) 15;
KOK. Pfl. R. Heft 38 (1909) 617; Back. Bekn. FI.
Java (em. ed.) 10 (1949) fam. 246, p. 70, p.p.;
NeEvLMes, Reinwardtia 1 (1951) 395; Koyama, Bot.
Mag. Tokyo 72 (1959) 304; J. Fac. Sc. Un. Tokyo
II, 8 (1962) 215; Kern in Back. & Bakh. f. FI.
Java 3 (1968) 490.

ssp. olivacea. — Fig. 122, 133.

Rhizome stout, woody, emitting long, stout
stolons. Stems stout, erect, triquetrous, smooth or
scabrid on the angles above, 50-110 cm by up to
5 mm below. Leaves basal and subbasal, much
exceeding the stems, flat, stiffish, long-acuminate,
scabrous on the margins towards the top, the
broader ones 1-2 cm wide; sheaths stramineous,
somewhat spongy, deeply concave in front, ligule
elongate, up to 5 cm long. Spikelets 4-9 (according
to BootT up to 14), erect, long-cylindric, upper
approximate, fastigiate, lower somewhat distant;
terminal spikelet ¢ (often with some 2 flowers at
the base), peduncled, 51/,-16 cm long and 5 mm
thick, often a second, shorter 3 spikelet added at

Fig. 133. Carex olivacea Bootrt in the Kerintji area,
Westcoast of Sumatra, Rawal Bento, 1400 m
altitude (MEWER 6656, Aug. 1956).

1979]

Cy PERACEAE—II (Kern & Nooteboom)

165

the base, remaining spikelets 2 (but usually with
short 3 apices), very densely flowered, sessile but
lowest sometimes distinctly peduncled, up to 16 cm
by 5-8 mm. Lower bracts foliaceous, much over-
topping the inflorescence, semi-amplexicaul by
dark auricles, upper shorter, usually none sheathing
(lowest sometimes more or less sheathing). G/umes
oblong, obtuse, vinaceous to dark red with
3-nerved, green, 1'/,-3 mm long, central stripe
excurrent in a scabrid awn 3/,-3 mm long. Utricles
obovoid or ellipsoid, membranous, much inflated
when ripe, rugose when dry, slenderly plurinerved,
patent or reflexed, glabrous, olive-brown, 3!/;-
4?/, by 11/,-2 mm, suddenly narrowed into a short,
conic, often recurved, minutely bidenticulate beak
ciliate at the mouth. Nut obovoid or oblong-
obovoid, triquetrous, beaked, stramineous to
yellowish, c. 2 by 1 mm. Style straight, not or
hardly thickened at the base. Stigmas 3.

Distr. NE. India; in Malesia: W. Sumatra
(Mt Kerintji), W. Java (Telaga Bodas, once
collected by H. O. Forses in 1880). The record for

Indo-China (Reinwardtia 1, 1951, 396) refers to a
collection of C. nemostachya StEuD. Distr. map in
Bot. Mag. Tokyo 72 (1959) 304, f. 26 (the occur-
rence in Java erroneously indicated as comprising
the whole western and central part of that island,
the Sumatran locality unknown at the time).

Ecol. On Mt Kerintji in swamp at 1400 m
altitude, at Telaga (= lake) Bodas “in warm water,
1500 m.”

Notes. Closely related to 44. C. oedorrhampha
NELMES, but much stouter, and distinguished from
all other members of its section by its very wide
leaves and very long spikelets.

Ssp. confertiflora (BooTT) KoyAMA, Bot. Mag.
Tokyo 72 (1959) 307; Phytologia 17 (1968) 413,
t. 17 (C. confertiflora Boott in A. Gray, Bot. Jap.
1859, 418; Ill. 4, 1867, 184; C. olivacea var. minor
Kuk. Pfl. R. Heft 38, 1909, 618), from Japan, and
ssp. recurvisaccus (KOYAMA) KoyAMA, /.c. (C. recur-
visaccus KOYAMA, J. Jap. Bot. 15, 1956, 166, f. 2),
from China, Kwantung, differ but slightly from the
typical subspecies.

17. Section Ferrugineae

[TUCKERM. En. Meth. (1843) 12, nomen]; ex BAILEY, Proc. Am. Ac. 22 (1886) 92, as
group; NELMES, Reinwardtia 1 (1951) 409. — Sect. Frigidae Fries subsect. Ferru-
gineae (BAILEY) KUK. Pfl. R. Heft 38 (1909) 559.

Type species: Carex ferruginea SCopP.

46. Carex brachyathera Onwi, Jap. J. Bot. 7
(1934) 190; Mem. Coll. Sc. Kyoto Imp. Un. B11
(1936) 332, f. 4, t. 8 f. 5; AKtyAMA, Car. Far East.
Reg. Asia (1955) 133, t. 118; KoyAMa, J. Fac. Sc.
Un. Tokyo III, 8 (1962) 158, incl. var. brevispiculosa
Koyama. — C. breviculmis var. perciliata (non
KUkK.) RIDL. Trans. Linn. Soc. II, Bot. 9 (1916)
247. — C. tricuspidata Kix. Bot. Jahrb. 70 (1940)
466, incl. var. brevispiculosa KiK. et var. minor
Kux.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 113;
NELMES, Kew Bull. (1949) 384; Reinwardtia 1
(1951) 410. — Fig. 122, 132a.

Rhizome thick, woody, obliquely descending,
sometimes elongated. Stems erect, somewhat
cernuous at the top, triquetrous, smooth or slightly
scaberulous just below the inflorescence, (5S—)20-
70 cm by 1-1!/, mm, at the base covered with a
thick mass of fibrous, brownish remains of leaf-
sheaths. Leaves basal or 1-2 higher on the stem,
shorter than the stem, flat, or involute when dry,
stiffish, asperous, long-attenuate, 1—31/, mm wide.
Spikelets 3-5, single at the nodes, erect to cernuous,
often subfastigiate, rather densely flowered above,
lax-flowered below, 11/,-6 cm long, terminal one
3, subclavate, 1-2 mm thick, long-peduncled,
lateral ones Q, linear-cylindric, 2'/,-3'/, mm thick,
on exserted, slender, smooth or antrorsely scabrid

peduncles. Lower bracts foliaceous, shorter to
slightly longer than their spikelet, long-sheathing,
upper reduced. Glumes oblong, often asymmetri-
cal, truncate or emarginate, thin, brown, whitish
hyaline at the top, 3-4'/, mm long, with strong
midrib excurrent in a hispidulous, up to 1 mm long
awn. Utricles ellipsoid or ellipsoid-fusiform,
obtusely compressed-trigonous, angled ventrally,
membranous, nerveless except for 2 submarginal
nerves, sparsely to rather densely subappressed-
hispid, suberect, stipitate, 3-41/, by c. 1 mm, rather
gradually narrowed into a stout, straight or slightly
bent, bidentate, 1—1!/, mm long beak with oblique,
white-hyaline mouth. Nut ellipsoid, triquetrous,
c. 2 by 1 mm. Style-base pyramidally thickened.
Stigmas 3 (according to NELMES 2 or 3).

Distr. Ryu Kyu Is., Formosa; in Malesia: New
Guinea (Mt Carstensz, Mt Wilhelmina in W., Mt
Sarawaket in E.).

Ecol. In mountain grasslands, on steep rocks,
seepages, sandy margins of streams, 3000-4000 m.

Notes. The New Guinea collections exhibit a
high degree of variability in size, length of bracts,
spikelets, and utricles, probably due to their
growing at high altitude, often in uncongenial
habitats. I do not see any reason to treat them as
varietally distinct from the Formosan plants, as was

166

FLORA MALESIANA

[ser. I, vol. 9!

done by Koyama, on account of the longer bracts
and — according to NELMES — sometimes digynous
flowers.

To judge from its description and excellent

figure, C. drepanorhyncha FRANCH. Pl. David. 2
(1888) 141; Nouv. Arch. Mus. Hist. Nat. Paris III,
9 (1897) 178, t. 4, f. 1; KUxK. Pfl. R. Heft 38 (1909)
563, from Szechuan, is very near to C. brachyathera.

18. Section Sylvaticae

[TUCKERM. En. Meth. (1843) 12, nomen]; BooTT ex MACKENZ. N. Am. FI. 18 (1935)
283; NELMES, Reinwardtia | (1951) 402. — Sect. Hymenochlaenae DREJER subsect.
Debiles (CAREY) KUK. Pfl. R. Heft 38 (1909) 594, p.p.

Type species: Carex sylvatica HUDs.

47. Carex finitima BooTr, Ill. 1 (1858) 44, t. 112;
Boeck. Linnaea 41 (1877) 247; CLarKeE, FI. Br.
Ind. 6 (1894) 736; Kix. Pfl. R. Heft 38 (1909) 598,
f. 101 E-H; Netmes, Kew Bull. (1949) 385, 391;
ibid. (1950) 204; Reinwardtia 1 (1951) 403;
Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 218.
— C. remotiflora HAYATA, Ic. Pl. Form. 10 (1921)
68, f. 45; OHwW1, Mem. Coll. Sc. Kyoto Imp. Un.
B11 (1936) 446; Akiyama, Car. Far East. Reg.
Asia (1955) 155, t. 148, f. 2. — C. fusiformis NEES
var. enervosa KUK. Bot. Jahrb. 70 (1940) 467. —
C. atjehensis Kx. Bull. Jard. Bot. Btzg III, 16
(1940) 314; in Fedde, Rep. 53 (1944) 105 (‘atjehien-
sis’). — Fig. 122.

Rhizome short. Stems tufted, erect, triquetrous,
smooth, (10-)30-90 cm by 1-2 mm below, sur-
rounded below the leaves by a few reddish bladeless
sheaths. Leaves basal and 1-2 higher up the stem,
shorter than to slightly exceeding the stem, flat,
smooth except for the minutely scaberulous mar-
gins, long-attenuate, 2-8 mm wide. Spikelets
4-8(-12 according to Bootrt), linear-cylindric,
terminal one dg or rarely gynaecandrous, peduncled,
11/,-4'/, cm by c. 1 mm (sometimes a smaller
second ¢ spikelet added), remaining ones 9, lax-
flowered, erect or lower cernuous, upper approxi-

mate, lower remote, 2-9 cm by 3-7 mm; peduncles
very slender, smooth or sparsely scaberulous above,
lower long-exserted. Lower bracts foliaceous,
slightly shorter than to much exceeding the in-
florescence, long-sheathing, upper much reduced;
sheaths pale to reddish brown. Glumes oblong-
ovate, acute to very obtuse, 31/,-4(-6) mm long,
translucent, with broad, white margins and
greenish midrib, the latter not reaching the apex
but sometimes excurrent below the apex in a short,
up to 1(-2) mm long awn. Utricles fusiform, tri-
gonous, membranous, with 2 marginal nerves,
otherwise nerveless or obscurely few-nerved,
glabrous, smooth, obliquely erect, shining, light
green, 5—7!/, by 1-2 mm, rather gradually narrowed
into a linear-cylindric, long, smooth beak with
oblique, scarious mouth. Nut ellipsoid or oblong-
ellipsoid, triquetrous, finally dark brown, shortly
stipitate, abruptly beaked, 2!/,-3 by 1-1'/, mm.
Style thickened at the base. Stigmas 3, long,
slender.

Distr. E. Himalaya, S. China (Szechuan, Yun-
nan), Formosa; in Malesia: N. Sumatra (Atjeh:
Gajolands), E. New Guinea.

Ecol. Grassy and bushy slopes, alpine grass-
lands, 2400-3900 m.

19. Section Folliculatae

MACKENz. in Britt. & Brown, Ill. Fl. ed. 2, 1 (1913) 353; N. Am. FI. 18 (1935) 426.
— Sect. Orthocerates Kocu subsect. Folliculatae (MACKENZ.) KOYAMA, J. Fac. Sc.

Un. Tokyo III, 8 (1962) 234.
Type species: Carex folliculata L.

48. Carex michauxiana Boeck. Linnaea 41 (1877)
336; CLARKE, J. Linn. Soc. Bot. 36 (1904) 298;
KUK. Pfil. R. Heft 38 (1909) 705; MACKENZ. N. Am.
Fl. 18 (1935) 427; N. Am. Car. 2 (1940) t. 489;
FERN. in Gray’s Man. Bot. ed. 8 (1950) 375,
f. 742; GLEASON, New Britt. & Brown Ill. Fl. 1
(1952) 358 f. — C. rostrata MicHx, Fl. Bor.-Am. 2
(1803) 173; Boorrt, Ill. 2 (1860) 91, t. 267; non

STOKES, 1787. — C. abacta L. H. BAILEY, Bull. Torr.
Bot. Club 20 (1893) 427.
The Asiatic plants are distinguished as:

var. asiatica (HULTEN) OHWI, Mem. Coll. Sc.
Kyoto Imp. Un. B11 (1936) 491; AkryAMA, Car.
Far East. Reg. Asia (1955) 163, t. 158 f. 2; YosuI-
KAWA, Ic. Jap. Carex 2 (1958) 268, t. 134; KOYAMA,

1979]

CyYPERACEAE—II (Kern & Nooteboom)

167

J. Fac. Sc. Un. Tokyo III, 8 (1962) 235; KERN,
Blumea 13 (1965) 125. — C. michauxiana ssp.
asiatica HULTEN, Kungl. Svenska Vet. Ak. Hand.
5 (1927) 207, f. 14, t. 223; Phytologia 17 (1968) 417,
t. 19. — C. michauxiana f. asiatica (HULTEN)
AKIYAMA, J. Fac. Sc. Hokkaido Imp. Un. 5 (1932)
220, f. 162. — C. dolichocarpa C. A. MEY. ex
Krecz. in Komar. Fl. U.R.S.S. 3 (1935) 458, 623.
— Fig. 122, 131a.

Rhizome short, thick. Stems densely tufted,
stiffly erect, slender, trigonous, smooth, at the base
clothed with the remains of old leaf-sheaths,
20-60 cm by 1-2 mm. Leaves subbasal, shorter to
longer than the stems, firm, flat, long-attenuate,
scabrid towards the apex, light green, 2-5 mm
wide; sheaths tight, white-hyaline ventrally, ligule
as long as wide. Inflorescence consisting of 1
terminal ¢ spikelet and (1—)2-4 9 spikelets. Bracts
long-sheathing, leaf-like, the lower overtopping the
inflorescence; sheaths concave at the mouth.
3 Spikelet (see note) shortly peduncled or subsessile,
few-flowered, 8-20 by 1'/,-2!/, mm; glumes
oblong-ovate, acute, yellowish brown with 3-nerved
green centre and hyaline margins; upper 9 spikelets
subsessile, crowded and sometimes hiding the 3 one
(see note), the lower 1-2 remote, on slender, more
or less exserted, erect, smooth peduncles, sub-
globose or broadly ovoid, 5-—20-flowered, 15-
25 mm long and wide. Glumes ovate, acutish,
hyaline with conspicuously 4—5-nerved centre,
1/,1/, as long as the utricle. Utricles at first
appressed-ascending, soon divergent, lanceolate-
subulate, obtusely trigonous, subcoriaceous, glab-
rous, many-nerved, stipitate, spongy at the base,

shining yellowish green, (8—)12—13 by 1'/,-2 mm,
gradually tapering into the scabrid bidentate beak;
teeth erect, ?/; mm. Nut trigonous with convex
sides, oblong-ovoid, shortly stipitate, shining
yellow, c. 3 by 1'/, mm. Style continuous with the
nut, tortuous, subincrassate at the base. Stigmas 3.

Distr. The typical variety in northeastern N.
America, the var. asiatica in eastern Asia: S. Kam-
chatka, Kuriles, Yezo, Hondo, China (Shanghai);
in Malesia: E. New Guinea.

Ecol. In swamps, bog grasslands, 2250-2650 m.

Vern. Tudik, Mendi lang., koale, Enga lang.

Notes. The differences between the American
plants and the Asiatic ones are but slight, and it is
questionable whether recognition of two geographi-
cal races is justified. In the few American specimens
I could examine the ¢ spikelet does not overtop the
upper @ ones (but this is not rarely also the case in
Asiatic specimens!), the leaves are slightly narrower
(2-3 mm wide), the 2 glumes 3-nerved, and the
utricles in general somewhat smaller. I fail to see
the difference in the relative length of the glumes
mentioned by Koyama. If the detailed description
by MACKENZIE, /.c., refers to American plants only,
the distinction of two varieties seems unjustified, as
the Asiatic plants almost completely fall within the
limits of variation there given.

It is worth noticing that most of the terminal
spikelets in the Mt Giluwe specimens are not
strictly g, but bear some 2 flowers at the top.
Sometimes the ¢ spikelet is either inconspicuous or
absent.

A northern element in the New Guinea moun-
tain flora.

20. Section Pseudocypereae

[TucKERM. En. Meth. (1843) 13, nomen]; ex BAILEy, Proc. Am. Ac. 22 (1886) 69,

as group; Kux. Pfl. R. Heft 38 (1909) 693. — Sect. Orthocerates KOCH subsect.

Pseudocypereae (BAILEY) KoyAMa, J. Fac. Sc. Un. Tokyo HII, 8 (1962) 234.
Type species: Carex pseudocyperus L.

49. Carex pseudocyperus LINNE, Sp. Pl. 2 (1753)
978; Boort, Ill. 4 (1867) t. 451, 452; Boeck.
Linnaea 41 (1877) 321; Kix. Pfil. R. Heft 38 (1909)
695.

In Malesia only:

var. fascicularis (SOLAND. ex Boott) Boortrt, Ill. 4
(1867) 41; Kix. Pfl. R. Heft 38 (1909) 696;
Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 234;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 490. —
C. fascicularis SOLAND. ex Bootrt in Hook. f. FI.
Nov. Zel. 1 (1853) 283; Bootrt, Ill. 1 (1858) 53,
t. 139, 140; S. T. Blake, J. Arn. Arb. 28 (1947) 116;
NELMES, Reinwardtia 1 (1951) 393. — C. pseudo-
cyperus (non L.) R. Br. Prod. (1810) 243;

BenTH. Fl. Austr. 7 (1878) 448; CLarkE, J. Linn.
Soc. Bot. 37 (1904) 16. — Fig. 122, 132b.
Rhizome very short, woody. Stems stout, tufted,
erect, triquetrous with flat or slightly concave
sides, smooth except just below the inflorescence,
surrounded below the leaves by a few bladeless,
somewhat fibrous sheaths, 50-150 cm by 3-4 mm.
Leaves subbasal, longer than the stems, flat, stiffish,
rough-margined, conspicuously septate-nodulose,
long-acuminate, light or yellowish green, 6-10 mm
wide. Spikelets 3-7, usually close together (lowest
1-2 sometimes distant), the terminal 3, peduncled,
3-7 mm thick, the lateral 2, cylindrical, very densely
flowered, 21/,-5 cm long and 7-15 mm thick, the
upper shortly peduncled, the lower on slender,

168

FLORA MALESIANA

[ser. I, volP@?

smooth or scabrid peduncles at length pendulous.
Lower bracts leaf-like, far exceeding the inflores-
cence, scarcely sheathing (occasionally long-
sheathing in distant spikelets), upper subfoliaceous
to setaceous, not sheathing. Glumes oblong or
oblong-spathulate, obtuse, translucent, eventually
ferrugineous, with 3-nerved green centre, ciliate
above, 2—2!/, mm, excurrent into a flat, antrorsely
scabrous awn 2-4 mm long. Utricles ovoid or
ovoid-lanceolate, obscurely trigonous, strongly and
densely many-nerved, coriaceous, glabrous, when
ripe widely spreading or even reflexed, somewhat
inflated, distinctly stipitate (stipe 7/,-1 mm, in some
Australian specimens up to 11/, mm), greenish to
stramineous, 4—5(—7) by 1-11/, mm, tapering above
into a 1'/,-2 mm long, smooth, deeply 2-cleft beak
(teeth c. 1 mm, divergent). Nut obovoid or ellip-
soid, triquetrous, scarcely stipitate, golden or
brown, 1'/,-12/; by 1(-11/3;) mm. Sty/e continuous
with the nut, contorted. Stigmas 3.

Distr. Carex pseudocyperus in the circum-
scription here accepted shows an almost world-
wide distribution. The typical variety is widely
distributed in the northern hemisphere (Eurasia

from W. Europe to Kashmir and Japan, N. Africa,
eastern N. America). Other varieties occur in North
and South America; var. fascicularis in Australia
(Tasmania to Queensland) and in New Zealand, in
Malesia: New Guinea (W. New Guinea: Lake
Habbema, Wissel Lake region; Terr. of New
Guinea: W. Highlands, Kandep valley; Yobobos
grassland area; Lake Inin). Specimens of, this
variety in BM and K are labelled ‘“‘Java, Horsfield”’ ;
recent collections from this island are not extant.

Ecol. In pools and swamps, along lakes, river-
sides; 1750-3225 m.

Vern. Kwai’a’reh, koali, Enga lang., Kepilam.

Notes. Var. fascicularis differs from var.
pseudocyperus in its darker, ferrugineous glumes,
its subinflated utricles with more divergent teeth
to their beaks and longer stipe, and its contorted
style.

The varieties certainly represent geographical
races and might therefore better be considered
subspecies. They are often treated at specific level,
but the differences are so slight that it is here
preferred to follow Boorr (1867) and KUKENTHAL
(1909).

21. Section Occlusae

CLARKE, Kew Bull. add. ser. 8 (1908) 147; RAyM. Mém. Jard. Bot. Montréal n. 53
(1959) 55, 87; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 169. — Sect.
Scleriiculmes NELMES, Kew Bull. (1951) 121; Reinwardtia | (1951) 407;; KoyAMa,
J. Jap. Bot. 29 (1954) 44; Act. Phytotax. Geobot. 16 (1955) 108.

Type species: Carex maubertiana Boott.

50. Carex maubertiana BoorttT, Ill. 1 (1858) 45,
t. 114; CLarkeE, J. Linn. Soc. Bot. 36 (1904) 297;
Camus, Fl. Gén. I.-C. 7 (1922) 200, f. 29, 5-8;
NELMES, Reinwardtia 1 (1951) 408; Raym. Mém.
Jard. Bot. Montréal n. 53 (1959) 87; Koyama, Act.
Phytotax. Geobot. 16 (1955) 39; KERN in Back. &
Bakh. f. Fl. Java 3 (1968) 489. — C. hebecarpa
C. A. Mey. var. lachnosperma CLARKE, FI. Br. Ind.
6 (1894) 747, quoad specim., non C. lachnosperma
NEES. — C. hebecarpa var. maubertiana FRANCH.
Nouv. Arch. Mus. Hist. Nat. Paris III, 10 (1898)
70; KUx. Pfl. R. Heft 38 (1909) 745. — C. hebe-
carpa var. ligulata (non KUK.) BAcK. Bekn. FI.
Java (em. ed.) 10 (1949) fam. 246, p. 69. — Fig. 122,
131g.

Rhizome very short. Stems tufted, erect, tri-
quetrous, smooth, for the greater part hidden by
the leaf-sheaths, 40-60 cm by 2-3 mm. Leaves of
normal length only in the upper half of the stem,
exceeding the inflorescence, flattish to strongly
revolute, stiff, greyish green, asperous above
towards the long-attenuated apices, 3-7 mm wide,
merging above into the foliaceous lower bracts and
below into short-bladed to bladeless, purplish,

rather tight sheaths; ligule obtuse, membranous,
ferrugineous. Spikelets 4-9, erect or suberect,
cylindric, dense-flowered, 11/,-4 cm long, upper
approximate, fastigiate, on included or shortly
exserted peduncles, lower somewhat distant, on
scarcely to long-exserted, scabrid peduncles; ter-
minal spikelet fg, 1-2 mm thick, sometimes a
second, smaller ¢ spikelet added; remaining
spikelets 9, 4-6 mm thick. Lower bracts foliaceous,
exceeding the inflorescence, their sheaths hispidu-
lous especially near the mouth, upper much
reduced. Glumes ovate, obtuse to acutish, trans-
parent, glabrous or hispidulous at the top, pale
with hyaline margins and 3-nerved, greenish
central stripe, covered with resinous flecks and
streaks, muticous or mucronulate, 1'/,-2'/. mm
long. Utricles ellipsoid, trigonous, submem-
branous, obliquely patent, with 2 marginal nerves
displaced on to the dorsal face, densely white-
hispid, broadly stipitate, 3-33/, by 1'/,-1*/, mm,
abruptly narrowed into a c. 1 mm long, bidentate,
finally somewhat upcurved beak. Nut ellipsoid,
triquetrous, c. 2 mm long. Style-base thickened.
Stigmas 3.

1979]

Cy PERACEAE—II (Kern & Nooteboom)

169

Distr. Himalaya, Annam, China (Hupeh); in
Malesia: Sumatra (Atjeh; W. Coast Res.: Mt
Kerintji), W. and E. Java (Priangan; Besuki), E.
Borneo (Berouw: Mt Ilas Mapulu), New Guinea

(Morobe Distr., near Bulolo). Distr. map in Act.
Phytotax. Geobot. 16 (1955) 38.

Ecol. Primary forests, forest-borders, bushy
slopes, 400-2100 m.

22. Section Longispicae

CLARKE, J. Linn. Soc. Bot. 37 (1904) 3; NELMEs, Reinwardtia 1 (1951) 417.
Type species: Carex graeffeana BOECK.

51. Carex graeffeana Boeck. Flora 58 (1875) 123;
CxiarKE, J. Linn. Soc. Bot. 37 (1904) 5; Philip. J.
Sc. 2 (1907) Bot. 107; Kix. Pfl. R. Heft 38 (1909)
403; Philip. J. Sc. 6 (1911) Bot. 62; in Hochr.
Candollea 6 (1936) 433; Bot. Jahrb. 69 (1938) 264;
Merk. Philip. J. Sc. 5 (1910) Bot. 335; En. Philip. 1
(1923) 138; Back. Bekn. Fl. Java (em. ed.) 10
(1949) fam. 246, p. 62; NELMES, Kew Bull. (1938)
109, incl. var. samoensis NELMES; ibid. (1955) 317;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494;
STEEN. Mt. Fl. Java (1972) pl. 14-9. — C. rechingeri
PALLA, Oest. Bot. Z. 57 (1904) 424. — C. philip-
pinensis NELMES, Kew Bull. (1938) 109; ibid. (1949)
385, 392; Reinwardtia 1 (1951) 419; ibid. 2 (1954)
381. — C. exploratorum NELMES, Kew Bull. (1938)
108; Reinwardtia 1 (1951) 418. — C. pandanus
OnwIl, Bot. Mag. Tokyo 56 (1942) 214. — C. euph-
lebia S. T. BiaKke, J. Arn. Arb. 28 (1947) 108,
f. 3C. — Fig. 122, 131].

Rhizome short, woody, forming large clumps.
Stems densely tufted, stoutish, erect, triquetrous,
scabrid on the angles at the top, 30-110 cm by up
to 4 mm below, surrounded below the leaves by
shining brown to blackish red sheaths splitting into
herring-bone-shaped fibres. Leaves basal and
subbasal, often 1-2 higher up the stem, stiff,
subcoriaceous, flat but margins often revolute,
scabrid on the nerves, long-attenuate, greyish or
glaucous-green, much overtopping the stems,
3-12 mm wide. Bracts not sheathing, 2-3 lower
ones foliaceous, semi-amplexicaul and blackish red
auricled at the base, middle ones subfoliaceous,
upper ones reduced to long-awned glumes. Spike-
lets (6—)10-15(—S0), single or binate at the nodes,
upper approximate, fastigiate, subsessile, lower
more distant, on capillary, scabrid, up to 5 cm long
peduncles, more or less nodding, all androgynous
or 1(—2) at the base of the uppermost much smaller
and wholly 3, exceptionally terminal spikelet
wholly ¢, linear-cylindric, (3-)6-13 cm long, very
densely flowered, the ¢ part occupying the upper
1/,—1/,, 1-3 mm thick, the 2 part 4-6 mm thick.
Glumes ovate or oblong-ovate, obtusish, purplish
black, with wide, 3-nerved, greenish central stripe
and very narrow hyaline margins, 11/,—2(—2'/,) mm
long, excurrent in a hispid, short mucro sometimes
1 mm long. Urtricles elliptic or obovate-elliptic,
biconvex or plano-convex, multinerved, longer
than (rarely about as long as) the glumes, finally
divaricate, not stipitate, somewhat acuminate at the

top but scarcely beaked, often puncticulate, some-
times minutely and sparsely setulose at the truncate
or emarginate mouth, 11/,-2!/, by °/,.-11/, mm.
Nut elliptic or obovate, biconvex, brown, 11/,-13/,
by 4/,—1'/, mm. Style short, not thickened towards
the base. Stigmas 2.

Distr. W. Polynesia (Samoa, Fiji Is.) and
Malesia: New Guinea (incl. New Britain), Philip-
pines (Luzon, Negros, Leyte, Mindanao), N.
Borneo (Mt Kinabalu), Lesser Sunda Is. (Flores),
W. Java (Mt Gedeh).

Ecol. On slopes in open places, on open banks
of streams, in peaty alpine grasslands, and in mossy
forest; on Mt Gedeh at 1600-1800 m, in the Philip-
pines at 1500-2200 m (once collected at 400 m), on
Mt Kinabalu at 1200 m, in New Guinea between
800 and 3800 m.

Vern. Jlateum, S; Philippines: a/asas, Buk.,
bagibi, giron, Bag., kigid, Bon., sidak, silak, Ig.

Notes. NeELMes (1938) distinguished between
C. graeffeana (Fiji) with its var. samoensis (Samoa),
C. philippinensis (Java, Philippines, New Guinea),
and C. exploratorum (Borneo), mainly using width
of leaves, number and length of spikelets, and size
of utricles as specific characters. For the New
Guinea specimens two more specific names are
available, C. pandanus OHwi and C. euphlebia
S. T. BLAKE. From NELMEs’s later publications it is
clear that the characters used for discrimination are
by far not so constant as was originally supposed.
There is indeed considerable variation in the
specimens collected in New Guinea or in the
Philippines, and even in those from the only
Javanese locality. The utricles are remarkably
small in the Fiji specimens, but not in those from
Samoa (the latter NeLMes in 1955 no longer
treated as varietally distinct.) Carex exploratorum
is only known from the type collection (Mt
Kinabalu, CLEMENS 34297), in which the much
compressed, sterile, and elongate utricles are
apparently diseased. The few well-developed, nut-
bearing utricles hardly differ from those of the other
Malesian materials. The glumes are larger than
usual, but not until more Kinabalu specimens are
available will it be possible to judge of their
taxonomical value.

On Mt Pulog (Luzon) the lower spikelets are
often branched into a raceme of secondary
spikelets, the total number of spikelets reaching
up to c. 50.

170

FLORA MALESIANA

[ser. I, vol. 9!

23. Section Praelongae

(KUK.) NELMES, Reinwardtia | (1951) 421; RAym. Mém. Jard. Bot. Montréal n. 53
(1959) 52, 55. — Sect. Acutae FRIES subsect. Praelongae KUK. Pfl. R. Heft 38 (1909)

345.

Type species: Carex praelonga CLARKE.

52. Carex phacota SPRENG. Syst. 3 (1826) 826;
NEEs in Wight, Contr. (1834) 126; KUNTH, En. 2
(1837) 420; DREJER, Symb. Caric. (1844) 15, t. 4,
excl. syn. C. notha et C. punctata; BoortrT, Ill. 1
(1858) 63, t. 168; Boeck. Linnaea 40 (1876) 434;
CLARKE, FI. Br. Ind. 6 (1894) 708; J. Linn. Soc.
Bot. 37 (1904) 6 (‘phacodes’); Kix. Pfl. R. Heft 38
(1909) 350, f. 56A—-C; Philip. J. Sc. 6 (1911) Bot.
62; MerRR. En. Philip. 1 (1923) 140; KUx. in Hochr.
Candollea 6 (1936) 431; Bull. Jard. Bot. Btzg III,
16 (1940) 317; Ouw1, Bot. Mag. Tokyo 56 (1942)
214; S. T. Biake, J. Arn. Arb. 28 (1947) 107;
Back. Bekn . FI. Java (em. ed.) 10 (1949) fam. 246,
p. 62; NELMEs, Reinwardtia 1 (1951) 426; ibid. 2
(1954) 382; Raym. Mém. Jard. Bot. Montréal
n. 53 (1959) 55; Dansk Bot. Ark. 23 (1965) 257;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494;
STEEN. Mt. Fl. Java (1972) 48a, pl. 14-7.—
C. lenticularis D. Don, Trans. Linn. Soc. 14 (1824)
331; Prod. Fl. Nepal. (1825) 43; non Micnx, 1803.
— C. platycarpa Hocust. ex STEUD. Syn. 2 (1855)
214. — C. hexasticha REINW. ex Mia. FI. Ind. Bat.
3 (1857) 353, p.p. — C. pruinosa var. aristata
O. K. Rev. Gen. Pl. 2 (1891) 748. — Fig. 122, 132c.

Rhizome short. Stems tufted, slender, erect,
triquetrous, smooth or sparsely scaberulous just
below the inflorescence, 20-120 cm by 1-3 mm,
surrounded below the leaves by a few ferrugineous
to dark brown, bladeless sheaths split into fine,
sometimes reticulate fibres. Leaves subbasal and
occasionally 1-2 higher up the stem, shorter to
much longer than the stems, stiff, with strongly
revolute margins, gradually attenuated, 3-8 mm
wide. Spikelets 4-6, rarely more, subapproximate
or the lowest more distant, cylindric, 2—8(—10) cm,
terminal wholly ¢ or with a few @ flowers, erect,
11/,-41/. mm _ thick, remainder androgynous
($ part short) or wholly 9, 4-6 mm thick, lower
usually nodding on slender, smooth or scabrid,
long peduncles. Lower bracts foliaceous, much
overtopping the inflorescence, upper ones much
smaller, none sheathing, but dark brown auricled
at the base. Glumes oblong, obtuse, truncate, or
bilobed-emarginate, thinly membranous, ferru-
gineous to castaneous, or pale with reddish
flecks, 11/,-3 mm long, with 3-nerved central stripe
excurrent in a wide, scaberulous-margined awn
1/,-2 mm long. Utricles elliptic, obovate, or sub-
orbicular, compressed-biconvex, membranous,
nerveless, narrowly marginate, glabrous, densely
beset with ferrugineous or reddish, raised glandu-
lar papillae, obliquely erect, shortly stipitate,
21/,-3(—31/2) by 11/,-2!/, mm; beak extremely short,

entire. Nut pyriform to suborbicular, compressed-
biconvex, broadly stipitate and beaked, 1°/,-2 by
11/,-13/, mm. Style thickened at the base. Stigmas 2.

Distr. Ceylon, Himalayan region, N. Thailand,
S. & E. China, Korea, Formosa, Ryu Kyu Is., and
Japan; in Malesia: Sumatra, Java, Lesser Sunda
Is. (Flores), Philippines (Luzon), N. Celebes, and
New Guinea.

Ecol. In open damp places, swampy grassland,
1500-2700 m.

On Mt Diéng (Java) VAN STEENIS (1972, /.c.)
observed in the shallow crater marsh Telaga
Pangonan that it formed in a stand of Scirpus
mucronatus L. large, solid, hummocky tussocks,
75 cm high and 50 cm thick, similar as do C. pani-
culata L. and the grass Molinia caerulea (L.)
MoeNCH. in Europe and C. secta Bootr (‘nigger-
head’) in New Zealand. This peculiar habit is
obviously for a large part developing with a
fluctuating water-level. — (Ed.)

Vern. New Guinea: int, koole, kwajare, Enga,
tsineme, Mendi, gogowe, Kapauku, yaguogufa,
Okapa, tsiri ku, Upper Kangel valley.

53. Carex pruinosa Boott, Proc. Linn. Soc. 1
(1845) 255; Trans. Linn. Soc. 20 (1846) 131; Ill. 1
(1858) 65, t. 174; STEuD. Syn. 2 (1855) 213; Mia.
FI. Ind. Bat. 3 (1856) 352; CLARKE, Fl. Br. Ind. 6
(1894) 709; J. Linn. Soc. Bot. 34 (1898) 111; ibid.
37 (1904) 7, excl. syn.; KUK. Pfl. R. Heft 38 (1909)
352; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam.
246, p. 62, excl. f. tristigmatosa BACK.; NELMES,
Reinwardtia 1 (1951) 428; Koyama, Bot. Mag.
Tokyo 72 (1959) 300, 306; RAyM. Mém. Jard. Bot.
Montréal n. 53 (1959) 56; Dansk Bot. Ark. 23
(1965) 258; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 494. — C. hexasticha Retnw. ex Mia. FI.
Ind. Bat. 3 (1857) 353, p.p. — C. pruinosa f. sub-
mutica O. K. Rev. Gen. Pl. 2 (1891) 748. — Fig.
123, 132d.

Closely related to 52. C. phacota SPRENG., from
which it is distinguished by the following characters:

Glumes lanceolate or elliptic, acute to obtusish
(but not truncate-emarginate), muticous or (the
lower ones) excurrent in a short awn up to 1 mm
long. Utricles slightly larger, 3-4 mm long, dorsally
slenderly 2—5-nerved, ventrally 1-—3-nerved, the
glandular papillae usually whitish. Spikelets more
approximate, fastigiate, and suberect to subcernu-
ous, 3/,—-5'/, cm long. Leaves greyish green to glau-
cous, 2-6 mm wide; sheaths not splitting into fibres.

Distr. Assam, Annam, NE. Thailand; in
Malesia: Java (West: several localities in Priangan;

1979 ]

CyYPERACEAE—II (Kern & Nooteboom)

171

Central: Diéng plateau; East: Jang plateau). In the
Leyden Herbarium there is a sheet labelled
“Sumatra, WartTz’’, but Warrz probably did not
visit this island. See Fl. Males. I, 1 (1950) 554.
KUKENTHAL’s record for the Moluccas (Buru)
refers to a collection of C. phacota; see there. The
rather different ssp. maximowiczii (MIQ.) KUK.
(C. maximowiczii MiQ.) in Japan and Korea; see
Koyama, /.c., with map.

Ecol. Swamps, damp grassy places, along
streams, 1500—2500 m.

Notes. According to BACKER /.c., 52. Carex
phacota and C. pruinosa would be connected by
intermediates, but I have not seen any transitional
forms and I find the two clearly distinct.

Carex pruinosa f. tristigmatosa BACK. belongs
to 39. C. maculata Bootr.

54. Carex teres BooTr, Ill. 1 (1858) 62, t. 167, excl.
utriculo imo; Boeck. Linnaea 40 (1876) 393;
CLARKE, FI. Br. Ind. 6 (1894) 707; J. Linn. Soc.
Bot. 34 (1898) 110; Kux. Pfl. R. Heft 38 (1909) 348;
KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494. —
C. teres var. spathulata Ktx. Bull. Jard. Bot.
Btzg III, 16 (1940) 316. — C. phacota (non SPRENG.)
KUwx. /.c. — C. spathulata NeLMES, Kew Bull.
(1950) 207; Reinwardtia 1 (1951) 423. — C. kemi-
riensis NELMES, Kew Bull. (1950) 206; Reinwardtia
1 (1951) 422. — C. petecticalis NELMES, Kew Bull.
(1950) 205; Reinwardtia 1 (1951) 424. — Fig. 123,
132e.

Probably loosely tufted on a woody, ascending
rhizome. Stems erect, stiff, triquetrous, smooth,
50-120 cm by 2-3 mm, at the base surrounded by
bladeless to short-bladed, brownish to dark
sheaths fraying into fine reticulate fibres. Leaves
subbasal, crowded in the lower 1/, of the stem,
flattish with revolute margins, stiff, 4-7(-11) mm
wide. Spikelets 4-8, cernuous, upper subapproxi-
mate, fastigiate, lower more separated, gynaecan-
drous usually with short ¢ base, or lower wholly
2, cylindric, dense-flowered but often laxer at the
base, 3—6(—8) cm long, the 2 part 5-9 mm thick, the
3 part 2-3(—5) mm, upper on short, lower on long,
smooth peduncles. Lower bracts foliaceous, much
to little exceeding the inflorescence, upper setaceous
to glumiform, none sheathing but dark-auricled at
the base. Glumes elliptic-oblong or oblong-obovate,
subobtuse to very obtuse, truncate or bilobed,
21/,-3'/. mm long, dark red with wide, 3-nerved,
pale central stripe excurrent in a flat, more or less

hispidulous awn up to 13/, mm long. Utricles ovate
to elliptic, plano-convex, membranous, obscurely
to distinctly 3—S-nerved on each face, straight,
patulous, stramineous, usually densely covered
with purplish flecks and minutely granular-
puncticulate, scarcely stipitate, gradually beaked,
3—4(-4"/,) by 11/,-2 mm;; beak short, '/,—3/,mm long,
entire or slightly emarginate. Nut broadly elliptic or
suborbicular, compressed-biconvex, 1'/,—2!/, mm
long, brown, shortly stipitate and beaked. Style
slightly thickened at the base. Stigmas 2, short.

Distr. E. Himalaya; in Malesia: Sumatra
(Atjeh: Mt Kemiri, Mt Losir, Senubong Mts;
W. Coast: Mt Ophir, Mt Singgalang, Mt Kerintji),
W. Java (Mt Papandajan).

I have not seen KERR 21031 from Laos, Pu Bia,
the type collection of C. kerrii NELMES, Kew Bull.
(1939) 304; ibid. (1946) 7, 28; Mém. Mus. Hist.
Nat. Paris n.s. B4 (1955) 175; Raym. Mém. Jard.
Bot. Montréal n. 53 (1959) 56. From the description
I infer that it is not specifically distinct from
C. teres.

Ecol. In marshes, damp mountain meadows and
heaths, in ericoid forest, in Sumatra between 2100
and 3500 m, in Java at 2300 m.

Notes. Variable, particularly as to the size of
spikelets and utricles, and the nervation of the lat-
ter, but I do not see how to distinguish between the
three “‘endemic species” into which NELMEs split
up the rather scanty Malesian materials. According
to him C. spathulata differs from C. teres princi-
pally in its stouter and shorter spikelets, C. petecti-
calis from C. spathulata by its longer inflorescence
(11-13 versus 7-10 cm), longer spikelets (3-6
versus 2—41/, cm), and shorter utricles (23/,-31/,
versus 31/,-41/, mm) with shorter beak. Carex
kemiriensis, only known from a single collection,
is said to differ by the densely papillose utricles, a
difference I am unable to find.

KUKENTHAL thought the Sumatran and Javan
plants to be varietally distinct from the continental
Asian ones by the reticulate-fibrous basal sheaths,
the shorter spikelets, the spatulate glumes, and
slightly larger utricles. However, in the type
collection of C. teres the basal sheaths are distinctly
fibrous-reticulate and the glumes variable in shape,
just like in the Malesian plants (see also BooTT’s
figure!).

Provisionally I prefer to include both the Indian
and Malesian materials in the variable C. teres, a
mountain species with a broken-up area.

24. Section Carex
Sect. Vulgares (ASCHERS.) NELMES, Reinwardtia | (1951) 429.

Type species: Carex acuta («) L.

55. Carex gaudichaudiana KUNTH, En. 2 (1837)
417; CLarKE, J. Linn. Soc. 37 (1904) 6; KUK. Pfi.
R. Heft 38 (1909) 312; Bot. Jahrb. 69 (1938) 264,

incl. var. humilior KGK.; OHw1, Bot. Mag. Tokyo
56 (1942) 215; S. T. BLake, J. Arn. Arb. 28 (1947)
107; Koyama, Act. Phytotax. Geobot. 18 (1959)

172

FLORA MALESIANA

[ser. I, vol. 91

22, f. 28. — C. caespitosa (non L.) R. Br. Prod.
(1810) 243. — C. vulgaris FR. var. gaudichaudiana
Bootr, Ill. 4 (1867) 169, t. 567; BENTH. FI. Austr. 7
(1878) 442. — C. vulgaris (non FR.) F.v.M. Fragm.
8 (1874) 257. — C. lacerans Kux. Pfi. R. Heft 38
(1909) 326; NeLMEs, Kew Bull. (1949) 385, 392;
Reinwardtia 1 (1951) 430. — Fig. 123.

Rhizome emitting short to rather long, horizon-
tal stolons clothed with reddish, ribbed scales.
Stems erect, triquetrous, scabrid above, 3-40
(-75) cm by 1-1!/, mm. Leaves subbasal, often
longer than the stem, somewhat rigid, flat or canali-
culate, scabrid on the margins, 11/,-4 mm wide,
with involute margins when dry, the lower reduced
to reddish to blackish red sheaths. Spikelets 3-6,
approximate to rather distant, cylindric, terminal
3, 1-4 cm by 2-3 mm, peduncled, sometimes a
second smaller one immediately under it also dg,
remainder 9 (or sometimes with a few ¢ flowers at
the top), sessile or lowest very shortly peduncled,
suberect, (1—)2-4(-6) by 3-4(-6) mm, densely
flowered. Lower bracts foliaceous, usually over-
topping the inflorescence, not sheathing, dark-
auricled at the base, upper much reduced. Glumes
ovate or oblong-ovate, obtuse to rather acute,
reddish to blackish red, with a central 3-nerved
pale stripe, 2-31/, mm long, the midrib sometimes
very shortly excurrent. Utricles elliptic, ovate, or
ovate-lanceolate, compressed, plano-convex,
obliquely erect, submembranous, strongly 5—7(-9)-
nerved on the dorsal, 3—5-nerved on the ventral
face, glabrous, green, becoming brownish, often
minutely reddish-puncticulate especially towards
the apex, scarcely stipitate, subabruptly beaked,

2-3(—31/,) by 1-1'/, mm; beak short, entire or
bidenticulate. Nut elliptic, obovate, or suborbicular,
compressed, biconvex or plano-convex, shortly
stipitate and apiculate, brown, 11/,-2 mm long.
Style not or scarcely thickened at the base. Stigmas
De

Distr. Australia (Tasmania to Queensland) and
New Zealand to E. China and Japan; in Malesia:
throughout New Guinea. Distr. maps: DURIETZ,
Act. Phytogeogr. Suec. 13 (1940) 219; Koyama,
Act. Phytotax. Geobot. 18 (1959) 24.

Ecol. Open marshes, edges of lakes, marshy
banks of streams, boggy alpine grasslands,
1450-3800 m.

Vern. New Guinea: kwajare, lebandili, Enga,
taua-tane, Tari.

Notes. Very similar to C. nigra (L.) REICH., a
common European and N. American species, and
sharing with it an extreme polymorphism. Carex
gaudichaudiana is characterized by the strongly
nerved, more distinctly beaked utricles, and the
lower bracts usually exceeding the terminal
spikelet.

The type of C. gaudichaudiana is from Australia.
The Asiatic plants have been distinguished as var.
thunbergii (STEUD.) Kix. Pfl. R. Heft 38 (1909)
313. — C. thunbergii STEUD. Flora 29 (1846) 23;
Syn. 2 (1855) 221 (type from Japan).

The differences between the Australian plants
and the Asiatic ones are but slight; see KOYAMA, /.c.

Carex appendiculata (TrAutTv.) KUx. Pfl. R.
Heft 38 (1909) 338 from E. Siberia, according to
Koyama also only a variety of C. gaudichaudiana,
is C. nigra (L.) REICH.

25. Section Graciles

[TUCKERM. En. Meth. (1843) 10]; ex KUK. Bot. Jahrb. 27 (1899) 516, quoad basion. ;
OuwIi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 464; NeLMgs, Reinwardtia 1
(1951) 353; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 52, 57; KOYAMA, J. Fac.
Sc. Un. Tokyo III, 8 (1962) 161. — Sect. Hymenochlaenae DReJER subsect. Graciles

(KUK.) KUk. Pfl. R. Heft 38 (1909) 599.
Type species: Carex gracilis R. BR.

56. Carex bilateralis HAyATA, Mat. Fl. Form.
(1911) 380; Ic. Pl. Form. 6 (1916) 127, f. 40 e-;
OuwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936)
471, p.p.; AkryAMA, Car. Far East. Reg. Asia
(1955) 105, t. 84, f. 2; Koyama, J. Fac. Sc. Un.
Tokyo III, 8 (1962) 163. — C. subteinogyna OHWI,
Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 469,
f. 17 & t. 15, f. 22; AxtyAMA, Car. Far East. Reg.
Asia (1955) 102. — C. spathaceo-bracteata KUx.
Bot. Jahrb. 70 (1940) 466; NeLmes, Reinwardtia 1
(1951) 356. — C. acrophila S. T. Biake, J. Arn.
Arb. 28 (1947) 114; NELMEs, Kew Bull. (1949) 382;
Reinwardtia 1 (1951) 354. — C. asperinervis
Koyama, Act. Phytotax. Geobot. 16 (1955) 6, t. 3,

f. U-V & f. W (ut C. subteinogyna). — Fig. 123,
132f.

Rhizome very short, woody. Stems tufted,
slender, erect, trigonous, smooth except on the
angles above, 20-90 cm by 3/,—11/, mm, surrounded
below the leaves by shining, fuscous to blackish,
bladeless sheaths tending to split in front into
reticulate fibres. Leaves shorter to longer than the
stems, rigid, conduplicate, keeled, often flexuous
at the long-attenuate top, scabrid, 1—2'/, mm wide
when flattened out. Inflorescence with 4-6 fascicles
of (1-)2-3(-5) spikelets, narrow, 5-15 cm long;
fascicles approximate or lower somewhat distant.
Spikelets erect (or some possibly slightly cernuous),

1979]

linear-cylindric, simple, rather densely flowered,
androgynous (2 part considerably longer than to
subequalling the 3 part), 1-4 cm long, upper
sessile or subsessile on included peduncles, lower
on included to long-exserted, smooth or scaberu-
lous peduncles. Lower bracts setaceous, shorter to
longer than their fascicles but usually much shorter
than the inflorescence, suddenly widening into a
spathaceous, strongly nerved, reddish brown base
with membranous margins clasping the base of the
spikelets, with ampliate, short or long sheaths;
upper bracts much reduced. Glumes slightly
shorter than utricles, oblong-lanceolate, acute to
very obtuse, translucent, glabrous or very sparsely
hispidulous on the midrib towards the apex,
3-nerved in the centre, ferrugineous to fuscous
with rather wide, whitish-hyaline margins, muti-
cous or apiculate, 4-6 mm long. Utricles narrowly
elliptic, plano-convex (occasionally trigonous),
membranous, slenderly multinerved, suberect,
reddish brown, whitish-hispidulous on the margins
in the upper half and often on the nerves, 31/,—5'/,
(-6!/,) by 1-11/, mm, contracted below into a stout,
stipe-like base '/,-1 mm long, rather gradually
narrowed into a bidenticulate, 11/,-13/, mm long
beak. Nut elliptic, oblong-elliptic, or oblong-
obovate, compressed-biconvex (occasionally
trigonous), not stipitate, shortly beaked, brown,
13/,-21/,. by 1-1'/, mm. Style-base slightly
thickened. Stigmas 2 (occasionally 3), 3-7 mm long.

Distr. Malesia: New Guinea (Carstensz Mts,
Lake Habbema, Mt Wilhelmina, Star Mts in W.,
Morobe Distr., Mt Sarawaket in E.).

Ecol. Alpine grasslands, marshy hollows, high
mountain ridges, (2100?—)3000-4050 m.

57. Carex brunnea THUNB. FI. Jap. (1784) 38;
ScHKUHR, Riedgr. 2 (1806) 16, t. Xx f. 111;
Boeck. Linnaea 39 (1875) 145; CLARKE, FI. Br.
Ind. 6 (1894) 705; J. Linn. Soc. Bot. 37 (1904) 5;
Philip. J. Sc. 2 (1907) Bot. 107; Kux. Pfi. R. Heft
38 (1909) 599; in Fedde, Rep. 8 (1910) 8, incl. var.
subteinogyna KUK.; Philip. J. Sc. 6 (1911) Bot. 63;
Camus, Fl. Gén. I.-C. 7 (1922) 194; Merr. En.
Philip. 1 (1923) 137; Ow, Mem. Coll. Sc. Kyoto
Imp. Un. Bll (1936) 466; Netmes, Kew Bull.
(1950) 201, incl. var. dolichocarpa NELMES; Rein-
wardtia 1 (1951) 357; Kew Bull. 2 (1955) 309;
RAYMOND, Mém. Jard. Bot. Montréal n. 53 (1959)
58; KoyAMA, Micronesica 1 (1964) 109, incl. var.
meyenii (NEES) KOYAMA; KERN in Back. & Bakh.
f. Fl. Java 3 (1968) 493. — C. gracilis R. BR. Prod.
(1810) 242, non Curt. 1777; KUNTH, En. 2 (1837)
513; Boort, Ill. 1 (1858) 59, t. 154-156; BENTH.
Fl. Austr. 7 (1878) 442. — C. meyenii NEES, Nova
Acta Nat. Cur. 19, Suppl. 1 (1843) 123; Krauss,
Pac. Sc. 4 (1950) 264-267, f. 8-9. — C. hattoriana
NAKAI ex TUYAMA, Bot. Mag. Tokyo 49 (1935)
508, t. 15; Onw1, Mem. Coll. Sc. Kyoto Imp. Un.
B11 (1936) 466; Axtyama, Car. Far East. Reg.
Asia (1955) 103, t. 82, f. 1; KoyAMA, J. Fac. Sc. Un.

Cy PERACEAE—II (Kern & Nooteboom)

[73

Tokyo III, 8 (1962) 164. — C. kanehirae Ouw1,
Act. Phytotax. Geobot. 8 (1939) 67; J. Jap. Bot. 18
(1942) 138. — C. spadiceo-vaginata OHWI, Bot.
Mag. Tokyo 56 (1942) 215. — C. teinogyna (non
Boott) BAcK. Bekn. Fl. Java (em. ed.) 1 (1949)
fam. 246, p. 61. — C. buruensis NELMES, Kew Bull.
(1950) 201; Reinwardtia 1 (1951) 360; ibid. 2 (1954)
379. — C. megacarpa KoyAMA, Bot. Mag. Tokyo
69 (1956) 210, f. 2. — Fig. 123, 132g.

Rhizome very short, woody. Stems tufted,
slender, erect, triquetrous, smooth, or scaberulous
on the angles above, (10—)30-100 cm by '1/,-
1!/, mm, surrounded below the leaves by dull
brown bladeless sheaths tending to split in front
into reticulate fibres. Leaves subbasal, shorter to
longer than the stems, rigid, flattish or condupli-
cate, rarely subfiliform, usually asperous above,
gradually attenuated towards the apex, (1-)2-6 mm
wide. Inflorescence with (2-)4-8 fascicles of 2-7
spikelets (rarely all the spikelets solitary), narrow,
erect or more or less nodding, c. 5-50 cm long,
upper fascicles or spikelets approximate and some
fastigiate, lower rather distant, one of the spikelets
at each node (especially at the lower ones) usually
longer than the others and with some smaller
spikelets branching from it. Spikelets erect or
suberect, cylindric or narrowly cylindric, rather
densely to rather loosely flowered, androgynous
(2 part usually much longer and thicker than the
6 part, 2-4 mm thick), 1-4'/, cm long, upper
sessile or subsessile on included or shortly exserted
peduncles, lower on long-exserted, smooth or
scaberulous peduncles. Lower bracts foliaceous or
subfoliaceous, longer than their fascicles but usually
shorter than the inflorescence, long-sheathing,
upper bracts much reduced. Glumes from much
shorter than to almost as long as the utricles, ovate
to ovate-lanceolate, acute to obtuse, glabrous,
obsoletely nerved but distinctly 3-nerved in the
centre, muticous or apiculate, rarely an awn to
2 mm present, light castaneous, sometimes with
whitish-hyaline margins, 2-4(-5) mm long. Utricles
ovoid to oblong-elliptic, plano-convex, mem-
branous, prominently multinerved, shortly whitish
setulose at least on the margins, suberect, cinna-
momeous to castaneous, cuneately tapering to a
1/,-1 mm long stipe, subabruptly beaked, 21/,-5(-6)
by 1-1!/, mm; beak !/,-2 mm, bidenticulate. Nut
ovate or oblong-ovate, compressed-biconvex,
scarcely stipitate and beaked, stramineous,
11/,-2'/, by 1-11/, mm. Style-base _ slightly
thickened. Stigmas 2 (or rarely 3), up to about as
long as the utricle.

Distr. Widely distributed from Madagascar and
the Mascarene Is. to Ceylon, India, Farther India,
China, Japan, the Hawaiian Is., New Caledonia,
Australia (New South Wales and Queensland); in
Malesia: New Guinea, Moluccas (Buru), Philip-
pines (Luzon, Bohol, Negros, Mindanao), Celebes,
Lesser Sunda Is. (Lombok, Sumbawa, Flores),
NW. Borneo (Sarawak), Java (a few localities in

174

FLORA MALESIANA

[ser. I, vol. 9?

W., Central & E.), N. Sumatra (Pajakumbuh;
Gajolands: Ketambe).

Ecol. Primary forest, mossy forest, exposed
ridges, grassy slopes, (80—)800—2800(—3100) m, in
Lombok in monsoon forest scrub on dry calcareous
soil, 200-400 m, in W. Sumatra on limestone
hills.

Vern. Philippines: selak, Ig., tayalid, Bag.

Notes. As will be seen from the above des-
cription C. brunnea in the sense accepted here is
extremely polymorphic, which is to be expected in
so wide-ranging a species. Especially Japanese
taxonomists have split it up into several micro-
species and infraspecific taxa among which I do not
see sharp distinctions. To me it is even very doubt-
ful whether 59. C. teinogyna and 56. C. bilateralis,
both closely related to C. brunnea, but which I have
decided to treat provisionally as separate species,
can be upheld as such in future.

Typical C. brunnea, described from Japan, has
small, broadly ovate, c. 2'/, mm long utricles sub-
tended by distinctly shorter glumes. Malesian
specimens approaching this have only been found
in N. Luzon.

According to KOYAMA (1962) the greater part of
the Malesian specimens belong to C. hattoriana
NAKAI ex TUYAMA, occurring from Bonin and
Formosa through Malesia to Australia. To him it
is quite distinct from C. brunnea by the strikingly
large utricles 4—41/, mm long, the longer glumes,
and the larger ¢ part of the spikelets.

Carex spadiceo-vaginata Onwi, from New
Guinea, is said to differ from C. brunnea by its
looser spikelets, sparsely setulose, 3 mm long
utricles with longer beak.

The type of C. buruensis NELMES is a very delicate
plant with almost filiform leaves and reduced
inflorescences, which may be due to the habitat.
Essential differences with C. brunnea I cannot find.
It is connected with broader-leaved specimens of
C. brunnea by a collection from New Guinea,
which to NELMES “‘represents a new variety or per-
haps a new species’.

I share the opinion of Koyama that C. meyenii
NeEgEs (C. brunnea ssp. meyenii (NEES) KOYAMA)
differing from typical C. brunnea by the lanceo-
late, 3-4'/, mm long, long-beaked utricles setu-
lose only on the margins, falls within the vari-
ability of C. brunnea, but refrain from maintaining
it as a subspecies. I also agree with KOYAMA
in the reduction of C. kanehirae Onwi from
Micronesia, to C. brunnea. I expect that several
other ‘species’ described from Japan and the
Pacific will have to be reduced to the polymorphic
C. brunnea.

Carex brunnea var. subteinogyna Kix. (non C.
subteinogyna Oxuwt), from the Philippines was
described as having looser spikelets, lighter
coloured, longer, acuminate glumes 5 mm long,
and very long-beaked and sparsely setulose utricles
with longer stipe. NeELMES (1950) altered this

circumscription considerably so as to cover also
specimens from Celebes and New Guinea, and
described moreover a var. dolichocarpa NELMES
with 5—5%/, mm long utricles from Java. The latter
variety is undoubtedly the same as C. megacarpa
Koyama. I fail to see how to draw a line between
the two varieties.

The collection DE WILDE 14068 from the
Gajolands often has 3 styles and accordingly
trigonous (fertile) nuts, and the terminal spikelets
are wholly g. This might be a hybrid with 20.
C. verticillata. — (Noot.)

58. Carex longipes D. Don, Trans. Linn. Soc. 14
(1825) 329; DresJER, Symb. Caric. (1844) 24, t. 10;
Mia. FI. Ind. Bat. 3 (1856) 347; Boorr, Ill. 4 (1867)
190; Boeck. Linnaea 40 (1876) 376; CLARKE,
Fl. Br. Ind. 6 (1894) 704; J. Linn. Soc. Bot. 34
(1898) 108; ibid. 36 (1903) 295; KUx. Pfl. R. Heft
38 (1909) 603; Bot. Jahrb. 70 (1940) 467, incl. var.
ramosa KuxK.; BACK. Bekn. Fl. Java (em. ed.) 10
(1949) fam. 246, p. 61; NELMES, Reinwardtia 1
(1951) 361; RAym. Mém. Jard. Bot. Montréal n. 53
(1959) 58; KERN in Back. & Bakh. f. Fl. Java 3
(1968) 493; STEEN. Mt. FI. Java (1972) pl. 14-6. —
Fig. 123, 134.

Rhizome very short. Stems tufted, slender, erect,
trigonous, smooth except on the angles above,
15-80 cm by 1-1!/, mm, surrounded below the
leaves by the fibrous, brownish remains of old
leaf-sheaths. Leaves subbasal, rarely one higher up
the stem, shorter to somewhat longer than the
stem, flat, scabrid, 1'/,-6 mm wide. Inflorescence
very lax, with up to 7 spikelets, 10-40 cm long.
Spikelets single at the nodes, erect or the lower
cernuous, cylindric, simple or the lower 1-3
branched near the base, lax-flowered, androgynous
(2 part much longer than the very short ¢ part),
rarely wholly 2, 1-3'/,(—4'/.) cm by 4-6 mm, upper
approximate, sessile, or subsessile on shortly
exserted peduncles, lower distant on filiform,
minutely scabrid, usually long-exserted peduncles,
the lowest often near the base of the stem. Lower
bracts foliaceous, shorter than inflorescence,
long-sheathing, upper much reduced. Glumes
3-4 mm, shorter than the utricles, ovate-lanceolate
to lanceolate, acute or subtruncate to bilobed-
emarginate, pale ferrugineous with whitish hyaline
margins, glabrous, with darker 3-nerved central’
stripe, the midnerve excurrent in a_ stoutish,
straight, antrorsely scabrid, up to 5 mm long awn
often overtopping the utricle. Uftricles elliptic,
compressed, biconvex, membranous, dorsally
strongly 6-9-nerved, less distinctly nerved ventrally,
glabrous and smooth or rarely the margins setulose,
straight, obliquely erect, scarcely stipitate, pale
green, fully ripe yellowish to light brown, shining,
subabruptly long-beaked, 5—7 by 11/,-2 mm; beak
sparsely hispid above, 2-3 mm long, deeply biden-
tate (teeth '/,—3/, mm). Nut broadly elliptic to ovate,
biconvex, stipitate, abruptly beaked, 2!/.-3 by 1*/.-

1979]

Fig. 134. Carex longipes D. Don. a. Habit, x 1/3,
b. young fruit with utricle in axil of bract, c. fruit
in utricle, d-e. fruits, all x 7 (VAN STEENIS 6788).

Cy PERACEAE—II (Kern & Nooteboom)

175
13/, mm. Style distinctly thickened at the base,
subpersistent. Stigmas 2, shorter than the utricle.

Distr. Nepal and India to China (Hupeh) and
Indo-China (Tonkin); in Malesia very rare: Java
(a few localities in W., Central and E.), Celebes
(Menado, Poso, top of G. Lumut), and NE. New
Guinea (Sattelberg).

Ecol. In forests, along forest-trails, on swampy
mountain meadows, 1500-2200 m.

59. Carex teinogyna Boortr, Ill. 1 (1858) 60, t. 158;
Boeck. Linnaea 39 (1875) 145; CLARKE, FI. Br.
Ind. 6 (1894) 705; KUK. Pfl. R. Heft 38 (1909) 602,
f. 102 F-H (‘teiogyna’), incl. var. scabriculmis
KUxK.; Bull. Jard. Bot. Btzg III, 16 (1940) 320,
excl. pl. jav.; NELMES, Reinwardtia 2 (1954) 378;
Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 146;
Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 59;
Koyama, Bot. Mag. Tokyo 72 (1959) 307; J. Fac.
Sc. Un. Tokyo III, 8 (1962) 162. — C. scabriculmis
Ounwt, Act. Phytotax. Geobot. 2 (1933) 27; Mem.
Coll. Sc. Kyoto Imp. Un. B11 (1936) 468; AkI-
YAMA, Car. Far East. Reg. Asia (1955) 102, t. 80;
YOSHIKAWA, Ic. Jap. Carex 2 (1958) 246, t. 123. —
C. brunnea (non THunsB.) NELMES, Kew Bull.
(1950) 200. — Fig. 123.

Rhizome very short, woody. Stems tufted,
slender, erect, triquetrous, smooth or scabrid on
the angles above, 20-60 cm by ?/,-1 mm, sur-
rounded below the leaves by spadiceous sheaths
ultimately fraying into persistent fibres. Leaves
subbasal, shorter to slightly longer than the stems,
flat to conduplicate, stiff, long-attenuate, asperous
especially in the upper part, 2-3(-4) mm wide.
Inflorescence with 3-6 fascicles of 1-3 spikelets,
narrow, 10-20 cm long, upper fascicles approxi-
mate, lower more distant. Spikelets erect, mostly
simple, rarely branched at the base, lax-flowered,
androgynous (@ part longer than the 3 one), upper
on scarcely, lower on shortly to rather long-
exserted, smooth or scabrid peduncles, up to
3 cm long. Lower bracts foliaceous, longer than
their fascicles but usually shorter than the in-
florescence, long-sheathing, upper much reduced.
Glumes about as long as utricles, oblong-
lanceolate, acuminate, acute, translucent, glabrous,
ferrugineous, sometimes with — pale-hyaline
margins above, muticous or with an awn up
to 2 mm long, 3!/,-5 mm long. Utricles elliptic,

plano-convex or compressed-biconvex, mem-
branous, slenderly multinerved, suberect,
castaneous, whitish appressed-hispidulous,

cuneate-stipitate, subabruptly beaked, 31/,-4'/2
(-5) by 1-1/3 mm; stipe '/;1/, mm; beak 1'/.-
2 mm, bidentate. Nut elliptic to oblong-elliptic,
compressed-biconvex, not stipitate, shortly beaked,
brown, 2(-2?/2) by 1(-14/,) mm. Sty/e-base slightly
thickened. Stigmas 2, very long (7-12 mm),
flexuous, persistent.

Distr. Assam, Upper Burma, S. China (Hunan),
Tonkin, Annam, Japan (Honshu, Shikoku,

176 FLORA MALESIANA [ser. I, vol. 9!

Ecol. Rocky riverbanks, ravines, 200-1150 m.
Note. Sometimes the glumes of the ¢ flowers
cup-shaped, the margins connate in front.

Kyushu), Korea (Quelpaert); in Malesia: N.
Sumatra (Atjeh: Leuser; Gajolands: Mt Kemiri
and Sangir Valley).

26. Section Paludosae

[Friges, Fl. Scan. (1835) 190, pro grege; TUCKERM. En. Meth. (1843) 14; O. F.
LANG, Linnaea 24 (1851) 618]; BAILEY, Proc. Am. Ac. 22 (1886) 74, as group;
KUK. Pfl. R. Heft 38 (1909) 730. — Subsect. Lacustres CAREY in Gray, Man. (1848)

561. — Sect. Tumidae MEINsH. Act. Hort. Petrop. 18 (1901) 283, 376.
Type species: Carex paludosa GOODEN.

Insufficiently known

60. Carex sp.

Rhizome woody, emitting stout stolons covered
with pale sheaths. Stem rather stout, triquetrous,
smooth, leafy, c. 60 cm by 3 mm. Leaves basal,
overtopping the stem, long-attenuate, flat or
somewhat folded lengthwise, septate-nodulose,
glaucous-green, subcoriaceous, c. 5 mm wide;
ligule lanceolate, acute, c. 1!/, cm long; margins
smooth below, scabrid above; lower sheaths
strongly septate-nodulose, not fibrous, stramineous
or light brown. Inflorescence erect, consisting of 4
spikelets. Terminal spikelet 3, c. 11/, cm long,
2 mm wide, lateral spikelets 2, erect, cylindrical,
densely flowered, the upper 2 approximate, sub-
sessile, the lowest distant on a smooth, 1!/, cm long
peduncle. Bracts foliaceous overtopping the
inflorescence, not or scarcely sheathing. Glumes of
the 2 flowers ovate, deeply emarginate at the top,
pale with purplish margins, c. 3 mm long, the

strong midrib excurrent in a firm, smooth or
scabrid awn about as long as the glume; glumes of
the ¢ flowers similar, more oblong, purplish with
pale centre. Utricles young, lanceolate, glabrous,
many-nerved, 5—5!/, by 1'/, mm, rather gradually
narrowed into the stout, straight, bidentate beak
with straight, !/,; mm long teeth. Style straight,
ciliolate, c. 2 mm, not thickened at the base.
Stigmas 3, longer than the style.

Distr. Malesia: New Guinea: Western High-
lands, Sirunki, swamp near Nanguris village, in
fast flowing very deep water course (water depth
approx. 230 cm), at c. 2500 m, 14 Sept. 1962:
WALKER ANU 691 (CANB).

Note. Only a single, immature specimen was
collected. The plant is related to the Eurasian
C. riparia CurT., from which it differs by the
narrower leaves, the small, single ¢ spikelets, and
the deeply incised g and 2 glumes, and to the
Japanese C. rugulosa KUK. (not seen).

II. Subgenus Vignea

(BEAUV.) CLARKE, FI. Br. Ind. 6 (1894) 700; KUx. Bot. Jahrb. 27 (1899) 495; Pfl. R.
Heft 38 (1909) 111; Netmes, Reinwardtia | (1951) 431. — Vignea BEAUV. in

Lestib. Ess. Fam. Cyp. (1819) 22.
Type species: Carex arenaria L.

27. Section Divisae

[CuRIsT, Bull. Soc. Bot. Belg. 24 (1885) 18, nomen]; (KUK.) KUx. Pfl. R. Heft 38
(1909) 119; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 234. — Sect.
Capituligerae KUK. subsect. Divisae CHRIST ex KUK. Bot. Jahrb. 27 (1899) 500.

Type species: Carex divisa Hubs.

61. Carex duriuscula C. A. Mey. Mém. Ac. St.
Pétersb. 1 (1831) 214, t. 8; KuntTu, En. 2 (1837)
373; OnwI, Mem. Coll. Sc. Kyoto Imp. Un. B11
(1936) 234. — C.. stenophylla WAHLENB. var.
duriuscula TRAutv. Act. Hort. Petrop. 10 (1887-

89) 537; Kix. Pfl. R. Heft 38 (1909) 121. — Carex
sp. NELMES, Kew Bull. (1949) 386; Reinwardtia 1
(1951) 446.

Rhizome creeping, slender, c. 1 mm_ thick,
clothed with brown sheathing scales. Stems erect

1979]

Cy PERACEAE—II (Kern & Nooteboom)

177

or curved, trigonous,’ smooth, or scaberulous
below the inflorescence, 7-15 cm by c. !/, mm.
Leaves subbasal, shorter than the stems, rigid,
curved, circinnate at the apex, canaliculate to con-
volute, smooth or scaberulous towards the apex,
c. 1 mm wide. Spikelets 4-7, androgynous (¢ part
about as long as the Q), sessile, approximate,
elliptic-lanceolate (very young), subdense-flowered,
5-7 mm long, forming a more or less oblong head
1-1!/, by c. !/, cm. Bracts glumiform, the lower
long-awned. Glumes oblong-lanceolate or ovate-
lanceolate, acutish, very thin, translucent,
castaneous with broad whitish-hyaline margins,
31/,-4 mm long, the midrib not extending to the
apex to very shortly excurrent. Utricles (very
immature) elliptic, plano-convex, nerveless, gla-
brous, scabrid on the margins in the upper half,
3-4 by c. 1 mm, tapering above into a short beak

with oblique mouth. Stigmas 2, about as long as the
utricle.

Distr. Siberia, N. Mongolia and Manchuria; in
Malesia: New Guinea, Lake Habbema (BrRAss
9235) in W., Mt Victoria (LAE 61670) and Kondo,
Mt Hagen (NGF 43535) in E.

Ecol. In New Guinea in sunny bog and alpine
grassland, 2400-3225 m.

Note. NELMES, /.c., supposed this New Guinea
collection to be close to C. arenicola F. Scumipt,
but I agree with S. T. BLAKE, J. Arn. Arb. 28 (1947)
116, that it is evidently allied to C. stenophylla
WAHLENB. of the northern hemisphere. The resem-
blance to small, narrow-leaved specimens of the
latter is surprising. I have seen extra-Malesian
material of C. duriuscula; the BRAss collection very
well matches the collection I saw of this species,
which is apparently very near to C. stenophylla.

28. Section Paniculatae

[KUNTH, En. 2 (1837) 389, nomen]; MEINSH. Act. Hort. Petrop. 18 (1901) 281,
313; KUkK. Pfl. R. Heft 38 (1909) 174; NeL_mgs, Reinwardtia | (1951) 432. — Sect.
Muricatae Fries subsect. Paniculatae KUNTH ex KUK. Bot. Jahrb. 27 (1899) 515.

Type species: Carex paniculata L.

62. Carex appressa R. Br. Prod. (1810) 242;
KUunrTH, En. 2 (1837) 389; KUNZE, Suppl. Riedgr.
(1840-50) 45, t. 11; Boorrt, Ill. 1 (1858) 46, t. 119,
120; Boeck. Linnaea 39 (1875) 99; KUx. Pfl. R.
Heft 38 (1909) 178, f. 29 E-J; Bot. Jahrb. 69 (1938)
261; Ouwi, Bot. Mag. Tokyo 56 (1942) 214;
S. T. BLake, J. Arn. Arb. 28 (1947) 101; NELMEs,
Kew Bull. (1949) 386, 392; Reinwardtia 1 (1951)
432; Kew Bull. 2 (1955) 318. — C. paniculata
(non L.) BENTH. FI. Austr. 7 (1878) 440. — Fig.
123.

Rhizome short, stout, woody, forming very
dense clumps. Stems densely tufted, erect, triquet-
rous (often very acutely so), rigid, scabrid on the
angles above (see note), 30-180 cm by 2-4 mm,
surrounded below the leaves by spadiceous, dark-
nerved, bladeless sheaths and their fibrous remains.
Leaves in the lower '/,—'/, of the stem, shorter to
longer than the stem, very stiff, flat to conduplicate,
long-acuminate, with very scabrous margins, pale
green, 3-10 mm wide. Inflorescence a slender,
oblong-cylindric, contracted, spike-like panicle,
5-25 by 1-2 cm; branches numerous, erect, often
appressed or even partly adnate to the stems,
upper crowded, lower approximate or slightly
distant. Bracts inconspicuous, setaceous, lower
sometimes as long as the branches, upper reduced
to glumes. Spikelets very numerous, sessile,
androgynous, ovoid or ovoid-lanceolate, few-
flowered (¢ and @ parts about equal in length),
4-8 mm long. Glumes ovate, acute, thin and

translucent, with ciliolate margins, otherwise
glabrous, slenderly nerved, ferrugineous to casta-
neous with hyaline margins, 2-3 mm long, the
midrib often excurrent in a short mucro up to
1/, mm long. Utricles ovate, plano-convex, with
obtuse margins, coriaceous, 6—-12-nerved dorsally,
3-6-nerved ventrally, glabrous, distinctly setulose-
margined above, subpatent, stramineous to dark
brown, rounded at the spongy-thickened base,
shortly stipitate, subabruptly beaked, 2'/,-31/, by
11/,-2 mm; beak 1/,-1 mm long, bidentate,
grooved on the back, with slightly oblique mouth.
Nut ovate to obovate, compressed-biconvex,
broadly stipitate, beakless or shortly beaked,
11,-13/, by c. 1 mm. Stigmas 2.

Distr. Widespread in Australia, also in New
Zealand and New Caledonia; in Malesia: New
Guinea (Arfak Mts, Lake Habbema in W. and
Western Highlands, Finisterre Range, Mt Sara-
waket, in E.).

Ecol. Open marshes, shores of lakes, alpine
meadows, 1900-3225 m, on Mt Sarawaket as low
as 900 m.

Vern. Kwajare, Enga.

Note. The stems are sometimes smooth or
almost so:

var. virgata (SOL. ex Boott) KUK. Pfl. R. Heft 38
(1909) 179 (C. virgata Sov. ex Boott in Hook. f.
Fl. Nov. Zel. 1 (1853) 282; Ill. 1 (1858) 46, t. 121,
122; Borck. Linnaea 39 (1875) 98.) — Only known
from New Zealand and Tasmania.

178 FLORA MALESIANA [ser. I, vol. 9!

29. Section Multiflorae

[KUNTH, En. 2 (1837) 387, nomen]; (CAREY) BAILEY, Proc. Am. Ac. 22 (1886) 135,
as group; KUK. Pfl. R. Heft 38 (1909) 142; NeLMeEs, Reinwardtia 1 (1951) 433;
Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 19. — Sect. Vignea BEAUV. subsect.
Multiflorae CAREY in Gray, Man. Bot. N. Un. St. (1848) 540. — Sect. Muricatae
Fries subsect. Multiflorae KUK. Bot. Jahrb. 27 (1899) 514.

Type species: Carex multiflora MUEHL. ex WILLD.

63. Carex nubigena D. Don, Trans. Linn. Soc. 14 Candollea 6 (1936) 430; Back. Bekn. Fl. Java
(1825) 326; Nees in Wight, Contr. (1834) 120; (em. ed.) 10 (1949) fam. 246, p. 60; NELMEs,
KuNTH, En. 2 (1837) 385; Boorrt, Ill. 1 (1858) 1, | Reinwardtia 1 (1951) 434, excl. specim. Sumatrae;
t. 2; Boeck. Linnaea 39 (1875) 90; CLARKE, Fl. Koyama, Bot. Mag. Tokyo 72 (1959) 302; KERN in
Br. Ind. 6 (1894) 702; J. Linn. Soc. Bot. 37 (1904) Back. & Bakh. f. Fl. Java 3 (1968) 494; STEEN. Mt
5, incl. var. fallax CLARKE; KUK. Pfl. R. Heft 38 FI. Java (1972) pl. 14-1. — C. fallax Steup. [in
(1909) 145, incl. var. fallax Kwtx.; in Hochr. Zoll. Syst. Verz. (1854) 60]; Syn. 2 (1855) 189;

Fig. 135. Carex nubigena D. Don in tufts around a shallow depression (sawahan), probably the site of an

old silted-up crater, covered with a heavily deer-grazed, very short turf of herbs and grasses subject to

frost in the dry season; background some scattered Casuarina junghuhniana Miq. Mixed with tussocks of
Pennisetum alopecuroides (L.) Spr. East Java, Mt Jang, c. 2000 m altitude (VAN STEENIS, 1936).

(1875) 57; Onw1, Mem. Coll. Sc. Kyoto Imp. Un.
Bll (1936) 247 (var. franchetiana Onwt). —
Fig. 123, 135.

Rhizome short, woody, forming dense tufts.
Stems erect, slender but rigid, smooth or slightly
scaberulous just below the inflorescence, obtusely
trigonous, 20—60(—100) cm by 1-3 mm, clothed
below the leaves by brownish to pale, bladeless
sheaths and their fibrous remains. Leaves subbasal,
shorter to longer than the stems, rigid, canaliculate
to conduplicate, with scabrid margins, 1!/,-3 mm
wide. Inflorescence pyramidal to oblong, head-like
or spike-like, 1-3(-S) cm by 7-13 mm. Spikelets
5-10(-15), crowded, or lowest 1-2 sometimes
slightly separated, ovoid to subglobose, sessile,
androgynous (with very few ¢ flowers), 5-10 by
5-7 mm. Lower 2-3 bracts foliaceous, mem-
branous-margined at the base, erect or curved,
lowest much exceeding, other equalling to exceed-
ing the inflorescence, upper much reduced to
glume-like, none sheathing. G/umes ovate to
oblong-lanceolate, acutish, very thin, translucent,
ferrugineous to whitish hyaline, with brownish to
greenish, 3-nerved central stripe, 21/,-31/, mm long,
with a mucro up to 1 mm long. Utricles ovate or

Cy PERACEAE—II (Kern & Nooteboom)

179

ovate-lanceolate, plano-convex, membranous,
strongly many-nerved on both faces, winged,
glabrous, obliquely erect, greenish to brown,
spongy-thickened at the base, shortly stipitate,
subgradually beaked, 3'/,-4'/, by 1'/,-2 mm; beak
c. 1'/, mm, serrulate-margined, dorsally grooved,
bidentate. Nut elliptic to suborbicular, biconvex,
broadly stipitate, shortly beaked, 1'/,-13/, by
4/,-1 mm. Style-base not or scarcely thickened.
Stigmas 2 (according to Bootr occasionally 3).

Distr. From Ceylon, S. India and the Himalaya
to China (Hupeh, Yunnan), Formosa and Japan;
in Malesia: Java (Central: Diéng Plateau; East:
Mts Kawi, Tengger-Semeru & Jang). Distr. map
in Bot. Mag. Tokyo 72 (1959) 302, f. 23 (the area
in Malesia should be restricted to Central and
East Java!).

Ecol. Marshy places, along streams, according
to BACKER 1600-3000 m, sometimes gregarious. On
Mt Jang deer feed on this species of which the leaf-
bases have a sweet taste, as in Gahnia javanica
(VAN STEENIS, /.c.).

Note. The immature collection LAE 65208
from New Guinea (W. slope of Mt Kenive,
9° 10’ S and 147° 45’ E) might possibly be referred
to this species. — (Noot.)

30. Section Stellulatae

KUNTH, En. 2 (1837) 399; OHwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 253;
NELMES, Reinwardtia | (1951) 441. — Sect. Elongatae KUNTH sensu KUK. Pfl. R.

Heft 38 (1909) 226, p.p.

Type species: Carex stellulata GOODEN.

64. Carex echinata Murr. Prod. Stirp. Gotting.
(1770) 76; Boecxk. Linnaea 39 (1875) 124; BENTH.
Fl. Austr. 7 (1878) 439. — C. stellulata GOODEN.
Trans. Linn. Soc. 2 (1794) 144; Kx. Pfl. R. Heft 38
(1909) 228. — C. nubigena (non Don) KUk. Bull.
Jard. Bot. Btzg II, 16 (1940) 314. — C. perileia
S. T. BLAKE, J. Arn. Arb. 28 (1947) 102; NELMEs,
Kew Bull. (1949) 386, 392; Reinwardtia 1 (1951)
441. — C. gajonum NELMES, Kew Bull. (1952) 84;
Reinwardtia 2 (1954) 382. — C. omiana FRANCH.
& SAv. var. perileia KOYAMA, Bot. Mag. Tokyo 69
(1956) 211. — Fig. 123, 131h.

Rhizome short, forming dense tufts. Stems
slender, erect, obtusely trigonous below, more
acutely so above, smooth or slightly scaberulous
just below the inflorescence, 15-90 cm by 1-2 mm.
Leaves subbasal, shorter to longer than the stems,
canaliculate-conduplicate, long-attenuate, scabrid
on the margins above, 1—3 mm wide, the lower ones
reduced to pale to castaneous, bladeless sheaths.
Inflorescence ovoid to oblong, head-like or spike-
like, 11/,-3'/,(-41/.) cm long. Spikelets 3-8,
gynaecandrous (¢ flowers very few), ellipsoid,
obovoid or ovoid to subglobose, sessile, dense-

flowered, approximate or lowest somewhat distant,
5-10 by 5-8 mm, finally squarrose by the widely
spreading utricles. Bracts glumiform, the lowest
with a setaceous awn, others more shortly aristate,
or indistinguishable from the glumes or the lower
bracts foliaceous, overtopping the inflorescence.
Glumes ovate, acute, thin, wholly pale or brownish
with wide whitish-hyaline margins, 3-nerved in the
centre, 2!/,-31/, mm long. Utricles elliptic or ovate-
lanceolate to broadly ovate, rounded to cardate at
the base, plano-convex, membranous, several-
nerved on both faces, glabrous, smooth, straight
or slightly recurved, widely patent to reflexed
when mature, spongy-thickened at the base,
scarcely stipitate, (3-)4-5(—-S?/,) by (1-)1?/,-2 mm,
yellowish green to brownish, subgradually nar-
rowed into a bidenticulate beak more or less scabrid
on the margins or rarely wholly smooth and with a
dorsal split with brown overlapping margins. Nut
oblong-ovoid, plano-convex, scarcely stipitate and
beaked, brownish, 2—2'/, by 1-1!/, mm. Style-base
slightly thickened. Stigmas 2.

Distr. N. America, Eurasia to Australia and
New Zealand; in Malesia: N. Sumatra (Gajolands:

180

FLORA MALESIANA

[ser. I, vol. 9!

Mts Kemiri, Losir, Bandahara) and New Guinea
(Arfak and Lake Habbema in W., and many
mountains in E.).

Ecol. Peat swamps, wet alpine grasslands,
marshy lake shores, 1850-3600 m, locally often
abundant.

Vern. New Guinea: kisis, pemp, Papua, Mendi
lang., koali, Enga lang., armemséna, kul, Mani-
kiong lang.

Notes. In the wide sense here accepted C. echi-
nata is a widely spread species. The characters used
for differentiating the numerous microspecies
described in sect. Stellulatae are far from reliable;
they mainly refer to the width of the leaves, the size
of the utricles, the scabridity of their margins and
the intensity of their nervation. For N. America
MacKeEnzigE, N. Am. FI. 18 (1931) 99-114 recog-
nized c. 20 spp. which can hardly be maintained.
I have not seen Japanese materials of the section,
but to judge from the descriptions and figures
C. basilata Ouwi, Act. Phytotax. Geobot. 11
(1942) 258; YosHIkAWwA, Ic. Jap. Carex 3 (1960)
296, t. 148 [C. muricata (non L.) Onwi1, Mem. Coll.
Sc. Kyoto Imp. Un. B11 (1936) 253; AktyAMA, Car.
Far East. Reg. Asia (1955) 64, t. 31] is hardly
different from the European plants, and C. omiana
FRANCH. & SAv.; OHwI, /.c. 254; AKIYAMA, /.c.
t. 32 with its lanceolate utricles less scabrid mar-
gins must be very near to the New Guinean speci-
mens.

Carex perileia S. T. BLAKE was based on a
specimen with a single fruiting culm, and distin-
guished from C. echinata by its narrower leaves and
its longer utricles with relatively larger beak deeply

split on the back with entirely smooth margins.
Several additional collections have shown that size
and scabridity of the utricles are very variable.
Sometimes the utricles are not longer and hardly
less scabrid than in European materials. In typical
C. echinata the uppermost spikelet is seemingly
long-stalked by the relatively large number of ¢
flowers at its base and sometimes almost wholly 3,
in the New Guinean specimens also this spikelet is
sessile as there are so few ¢ flowers that it has a
wholly 2 appearance. This may be the same in some
Australian forms of C. echinata, as according to
BENTHAM, /.c., there are very few ¢ flowers at the
base of the spikelets, sometimes even none.

Whether C. perileia represents a special New
Guinea race cannot be decided without a critical
study of the whole section or of at least the
Australasian and E. Asian representatives; in my
opinion it is not specifically distinct.

Carex gajonum NeELMES, from N. Sumatra
(Gajolands: Mts Losir and Kemiri), was distin-
guished because the lower bracts are foliaceous,
much overtopping the inflorescence, and the
utricles being broader, cordate at the base, and
obliquely erect instead of widely patent when
mature. In some Sumatran plants, however, the
lower bracts are setaceous, in some New Guinean
ones they are foliaceous, overtopping the inflores-
cence. In the collection DE WiLpE 13323, from
Mt Bandahara, the ripe utricles are patent as in
true C. echinata, making the spikelets squarrose.
Furthermore I compared several utricles, and al-
though they are generally broader in Sumatra,
there is no constant difference. — (Noot.)

31. Section Elongatae

KunTH, En. 2 (1837) 402; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 256;
KUK. Pfl. R. Heft 38 (1909) 226, p.p.; NELMES, Reinwardtia 1 (1951) 438.

Type species: Carex elongata L.

65. Carex remota LINNE, Amoen. 4 (1759) 293;
Boeck. Linnaea 39 (1875) 129; KUx. Pfl. R. Heft
38 (1909) 233.

The typical subspecies is widely distributed in
Europe, extending to N. Africa and W. Asia; in
Malesia two other subspecies occur:

a. ssp. alta (Bootr) KUK. Pfl. R. Heft 38 (1909)
234, incl. var. brizopyrum KOK.; BACK. Bekn.
Fl. Java (em. ed.) 10 (1949) fam. 246, p. 60; KERN
in Back. & Bakh. f. Fl. Java 3 (1968) 494; STEEN.
Mt. Fl. Java (1972) pl. 14-4. — C. alta Boorrt,
Proc. Linn. Soc. 1 (1845) 254; Trans. Linn. Soc. 20
(1846) 130; Ill. 1 (1858) 59, t. 153; Mia. FI. Ind.
Bat. 3 (1855) 347; Boeck. Linnaea 39 (1875) 126;
CLARKE, FI. Br. Ind. 6 (1894) 707; J. Linn. Soc. 37
(1904) 6; NeLmes, Reinwardtia 1 (1951) 438. —

C. brizopyrum Kunze, Suppl. Riedgr. (1840-50)
169, t. 43. — C. remota var. rochebrunii CLARKE,
J. Linn. Soc. 37 (1904) 6, p.p. (quoad ZOLLINGER
3192). — C. craspedotricha NELMES, Kew Bull.
(1939) 657; ibid. (1946) 28; RayM. Mém. Jard. Bot.
Montréal n. 53 (1959) 19; Dansk Bot. Ark. 23
(1965) 252. — ? C. imbricata KUx. in Hand.-Mazz.
Symb. Sin. 7 (1936) 1260, f. 38 n. 2. —? C. squamata
Krecz. Not. Syst. Herb. Inst. Bot. Ac. Sc. URSS 9
(1946) 196. — Fig. 123.

Rhizome very short, woody. Stems densely
tufted, stiff to rather weak, erect or suberect,
smooth, 15-80(—120) cm by 1—1!/, mm, surrounded
at the base by brown, fibrous remains of old leaf-
sheaths. Leaves in the lower 1/; of the stem, shorter
to longer than the stems, flat, scabrid on the
margins especially towards the long-attenuated

1979]

Cy PERACEAE—II (Kern & Nooteboom)

181

apex, 1'/,4 mm wide; sheaths long, mouth
concave in front. Spikelets 5—-18(—24) in an up to
c. 15 cm long, spiciform inflorescence, sessile,
obliquely erect, upper densely crowded, lower
separated to their own length from one another,
lowest 1-2 often distant, ellipsoid to ellipsoid-
cylindric, densely flowered, gynaecandrous with
only a few ¢ flowers, 5—15 by 3-5 mm. Lower bracts
foliaceous, far exceeding the inflorescence, not
sheathing, upper glumiform. Glumes ovate or
oblong-ovate, acute, thin and translucent, finely
nerved, whitish with greenish 3-nerved centre,
2-3 mm long, usually excurrent in a short, up to
1/, mm long mucro. Utricles ellipsoid or ovoid-
ellipsoid (often in outline broadest above the
middle because of the wings), obliquely erect,
plano-convex, membranous, slenderly nerved on
both sides in the lower centre, glabrous, winged in
the upper '/,-*/4,, yellowish green to brownish,
scarcely stipitate, subabruptly beaked, 27/,-3 by
1-11/, mm; wings varying in width, denticulate-
ciliate; beak c. 1/, mm long, bidentate with
slender, straight teeth. Nut ellipsoid or ovoid,
biconvex, shortly stipitate and beaked, 11/3—17/, by
2/,>/; mm. Style slightly thickened at the base.
Stigmas 2.

Distr. Insufficiently known; according to
literature from India (Himalaya) to Central China;
C. craspedotricha Netmes from Thailand un-
doubtedly belongs here. The type collection of both
C. alta Boott and C. brizopyrum KUNZE are from
Java; in Malesia only known from Java (from
Mt Patuha in W. to Jang in E.).

Ecol. In moist or swampy grasslands, along
water-courses, damp forest-borders, 1500-2200 m;
once collected at 1150 m.

b. ssp. rochebrunii (FRANCH. & SAv.) KUK. Pfl. R.
Heft 38 (1909) 234; Kern in Back. & Bakh. f. FI.
Java 3 (1968) 494. — C. rochebrunii FRANCH. &
SAV. En. Pl. Jap. 2 (1879) 126, 555; NeELMEs, Kew
Bull. (1946) 29. — C. remota var. rochebrunii
CLARKE, FI. Br. Ind. 6 (1894) 707. — C. monopleura
Krecz. Not. Syst. Herb. Inst. Bot. Ac. Sc. URSS 7
(1937) 35; NELMES, Kew Bull. (1950) 208; Rein-
wardtia 1 (1951) 440.

Usually slenderer than ssp. alta, with narrower
leaves. Inflorescence up to 10 cm long, with 3-9
spikelets; upper spikelets not rarely sterile. Glumes
3/,-4 mm long. Utricles lanceolate, ventrally
nerveless or nearly so, narrowly winged, 4—4!/, mm
long. Nut elliptic-oblong, c. 2 by 1 mm.

Distr. India (Sikkim), China, Japan, Formosa;
in Malesia: Sumatra (Atjeh: G. Leuser, and W.:
Mt Kerintji) and W. Java (Mt Papandajan).

Ecol. Marshy places in forests, along rivers and
streamlets, 1700-2700 m.

Notes. Ssp. rochebrunii is very near to the
European ssp. remota, mainly differing by the
narrow, lanceolate, narrowly winged utricles and
the oblong nuts. In my opinion the differences
between C. rochebrunii (described from Japan) and
C. monopleura (based on Sikkim material) are too
slight to justify specific separation.

In its extreme tall form, with relatively dense
inflorescence and broadly winged utricles, ssp. alta
gives the impression of being a separate species, but
slender, weak plants occur (they were distinguished
as var. brizopyrum by KUKENTHAL), and the width
of the wings of the utricles is variable.

KUNZE, /.c., in describing C. brizopyrum, stressed
the importance of the bristly appendage of the con-
nective, but this isalso found in European C. remota.

32. Section Heleonastes

KUNTH, En. 2 (1837) 393; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 261;

NELMES, Reinwardtia 1 (1951) 443.

Type species: Carex heleonastes EHRH.

66. Carex curta GOODEN. Trans. Linn. Soc. 2
(1794) 145; ScHkunR, Riedgr. 1 (1801) 43, t. C,
f. 13; KuNTH, En. 2 (1837) 403; S. T. BLAKE,
J. Arn. Arb. 28 (1947) 101; NeLMeEs, Kew Bull.
(1949) 386; Reinwardtia 1 (1951) 443; YosHIKAWA,
Ic. Jap. Carex 1 (1957) 32, t. 16. — C. canescens
(non L.) Boorrt, Ill. 4 (1867) 154, t. 496; Borck.
Linnaea 39 (1875) 122, excl. var. 8; BENTH. FI.
Austr. 7 (1878) 439; CLARKE, FI. Br. Ind. 6 (1894)
706; KUxK. Pfl. R. Heft 38 (1909) 216, f. 35 C-E;
Oxuwil, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936)
268; AKIYAMA, Car. Far East. Reg. Asia (1955) 60,
t. 26. — Fig. 123.

Rhizome short, forming dense tufts. Stems
slender, erect, triquetrous, slightly scaberulous just

below the inflorescence, 10-40(-60) cm by I1-
11/, mm, surrounded below the leaves by light
brown, withered leaf-bases. Leaves subbasal,
shorter to longer than the stems, flat to condupli-
cate, grey-green, scabrid on the margins above,
long-attenuate to the triquetrous apex, 2—3(-5) mm
wide. Inflorescence oblong, spike-like, 2—3'/,(-6) cm
long. Spikelets 4-8, gynaecandrous (3 part few-
flowered, inconspicuous), ovoid, ellipsoid, or sub-
cylindric, sessile, dense-flowered, approximate or
contiguous, 6-9 by 4-5 mm. Bracts glumiform,
shortly aristate, the lowest rarely subulate to
subherbaceous. Glumes ovate, acute, very thin,
whitish, 3-nerved in the centre, 2—2'/, mm long, the
midrib sometimes slightly excurrent. Utricles ovate

182

or ovate-elliptic, biconvex, membranous, several-
nerved on both faces, scarcely marginate, glabrous
or slightly papillose at the apex, golden yellow,
densely whitish puncticulate, straight, suberect,
very shortly stipitate, 2-2'/, by 1-1'/, mm, scarcely
beaked; mouth minutely bidenticulate. Nut elliptic
to broadly ovate, plano-convex or biconvex,
scarcely stipitate, abruptly beaked, light brown,
c. 1'/, by 1 mm. Style-base not thickened. Stigmas
2

Distr. Widely spread in N. America, extra-
tropical S. America, Eurasia (also Kashmir and
Japan), and SE. Australia (New South Wales,
Victoria); in Malesia: New Guinea (Lake Habbema
in W., Neon basin 15 km NNE of Woitape in E.).

Ecol. Plentiful in open grassland on wet sandy
soil and on marshy flats, 2800—c. 3225 m

Note. For the correct name of this species, which
has commonly been called C. canescens L., see
E. S. MARSHALL, J. Bot. 45 (1907) 365; S. T. BLAKE,
J. Arn. Arb. 28 (1947) 101; NELMeEs, Reinwardtia |
(1951) 444.

Doubtful (Sect. Vigneastra)

Carex subfilicina OHw1, Bot. Mag. Tokyo 56
(1942) 213; Netmes, Reinwardtia 1 (1951) 446
(under doubtful species). — I did not see any
collections of this species. From the description it
is possibly conspecific with either C. filicina NEES
or C. cruciata WAHLENB. — (NOootT.)

Excluded

Carex arnottiana NEES ex DREJER, Symb. Caric.
(1844) 16, t. 5; Boeck. Linnaea 40 (1876) 436. —
This species is endemic in Ceylon. BOECKELER,
l.c., cited it also to occur in “‘Java, alt. 6000 ped.
(Arnott)’’. As the annotation for the type specimen
is exactly matching this and ARNOTT never collected
in Java, this record is based on a miswritten label.

Carex cladostachya WAHLENB. Vet.-Ak. Handl.
24 (1803) 149; Boeck. Linnaea 40 (1876) 361; KUK.
Pfil. R. Heft 38 (1909) 267. — This Central Ameri-
can species was recorded by BOECKELER, /.c., also
from the Philippines on the strength of CUMING
625. This specimen must certainly belong to
CumING’s American collections, which were
separately numbered from the Philippine ones.

Carex dimorpholepis StEuD. Syn. 2 (1855) 214.
— C. cernua Bootrt, Ill. 4 (1867) 171, t. 578, non
J. F. Gmev. 1791. — The old specimens of this
species in the Leyden Herbarium annotated “‘Arch.
Ind. ? Leg. ?” in all probability do not originate
from the Malayan Archipelago, as they would be
the only Malesian collection extant. C. dimor-
pholepis is known from India, Upper Burma, and
China to Korea and Japan. See NELMES, Kew Bull.
(1950) 208.

FLORA MALESIANA

[ser. I, vol. 9!

Carex divulsa STOKES var. javanica NELMES, Kew
Bull. (1950) 208; Reinwardtia 1 (1951) 436. — See
below under C. pairaei.

Carex erythrolepis Kux. Pfl. R. Heft 38 (1909)
628; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam.
246, p. 65. — “Java, Hillebrand in herb. musei
berol. sine indicatione loci.’”’ Wrongly localized;
see Fl. Males. I, 1 (1950) 232. The name must be
referred to the synonymy of C. wahuensis C. A.
Mey., an endemic of the Hawaiian Islands. See
NELMES, Kew Bull. (1950) 203.

Carex haenkeana Presi, Rel. Haenk. 1 (1828)
205; STEUD. Syn. 2 (1855) 241; Mia. FI. Ind. Bat. 3
(1857) 354; F.-ViLL. Nov. App. (1882) 310. —
“Habitat in insulis Philippinis.’’ This is C. pseudo-
cyperus L. var. haenkeana (PRESL) KUK. Bot.
Jahrb. 27 (1899) 550; Pfil. R. Heft 38 (1909) 696, a
S. American plant. The record attributed to the
Philippines was based on an erroneously localized
Malaspina Expedition specimen. See MERRILL,
En. Philip. 1 (1923) 142.

Carex lindleyana NEES in Wight, Contr. (1834)
121; Boeck. Linnaea 40 (1876) 362; KUx. Pfi. R.
Heft 38 (1909) 280. — This is a species from the
Deccan Peninsula and Ceylon. BOECKELER recorded
it besides from the Philippines on the strength of an
unnumbered collection of Cuminc. If correctly
identified this certainly rests on an erroneous
localisation; it might have been collected in Ceylon
by CuMING himself.

Carex pairaei F. SCHULTZ var. javanica NELMES,
Kew Bull. (1950) 208; Reinwardtia 1 (1951) 437.

Carex divulsa Stokes and C. muricata L.
(= C. pairaei F. SCHULTZ) were recorded by
Netmes from Java, the only ones from outside
Europe, both based on a single specimen, that of
C. divulsa even on a single culm. The scanty
materials were alleged to have been collected by
Riw_ey during his Java tour on Mt Papandajan.

However, there is no doubt that the specimens
were mislocalized, possibly by using drying paper
to which remnants of rambles in Europe adhered.
The Carex flora of the easily accessible Mt Papan-
dajan is well known, e.g. by the intensive search
for Carices by VAN STEENIS.

Similar records opposing all rules of plant distri-
bution are those of Elisma natans for Java, Linaria
alpina for the Malay Peninsula, and Scheuchzeria
palustris for Sumatra (see Taxon 5, 1956, 157).
NEES, in litt. Dec. 17, 1953, admitted that the
records must be due to an error.

Carex scabrifolia STEUD. Syn. 2 (1855) 237; Mia.
Fl. Ind. Bat. 3 (1857) 354. — “Carex nr 67 et 83.
Herb. Zollinger. Java.’ The type is not from Java,
but from Japan, Decima in Nagasaki. See ZOLL.

1979] Cy PERACEAE—II (Kern & Nooteboom) 183

Syst. Verz. 1 (1854) 60; Kix. Pfl. R. Heft 38 (1909) Carex typhoides Bory; Hassk. Cat. Bog. 296;
737; KoyAMaA, J. Fac. Sc. Un. Tokyo III, 8 (1962) Mua. FI. Ind. Bat. 3 (1857) 352. — This is C. bor-
248. — HUBERT WINKLER 2097, wrongly distributed bonica LAMK from the Mascarenes; certainly not
as C. scabrifolia, belongs to C. baccans NEES. occurring in Malesia.

29. UNCINIA

Pers. Syn. Pl. 2 (1807) 534; Boeck. Linnaea 41 (1877) 339; CLARKE, J. Linn. Soc.
Bot. 20 (1883) 389; KUx. Pfl. R. Heft 38 (1909) 50; NELMeEs, Kew Bull. (1949) 140;
HAMLIN, Dom. Mus. Bull. 19 (1959) 1; BALGooy, Pac. Pl. Areas 3 (1975) 320,
map 209; Noort. Blumea 24 (1979) 511. — Fig. 137a-d.

Perennial, monoecious herbs, glabrous (or with hispid utricles). Stems central,
tufted or approximate on a more or less creeping rhizome, erect or ascendent,
sharply trigonous to subterete, striate, smooth, or scabrid below the inflorescence.
Leaves narrowly linear, flat or involute, more or less scabrid on margins and nerves;
basal sheaths bladeless, often disintegrating into fibres. Inflorescence a single,
terminal spikelet; 3 part above, shorter than the lower 2 part. Glumes spirally
arranged, ovate to oblong, concave, persistent or caducous, all flower-bearing, the
lowest often produced into a setaceous to foliaceous bract. — 3 Flowers naked,
consisting of (1—)2-3 stamens with linear (or dilated, New World spp.) filaments and
linear anthers; connective shortly produced. — 92 Flowers naked, enclosed in a
bottle-shaped, obtusely trigonous organ (utricle, perigynium) which is closed up to
the truncate top, glabrous (in all Mal. and Austr. spp.) or hispid; style incrassate at
the base; stigmas 3, exserted from the utricle. Rachilla (see note) reduced to a rigid
bristle below the nut and produced far beyond the mouth of the utricle, hooked at
the top. Nut trigonous.

Fig. 136. Range of the genus Uncinia Pers. (from VAN BALGooy, Pac. PI. Areas 3, 1975, 320, map 209)

184 FLORA MALESIANA [ser. I, vol. 9!

Distr. A genus of probably Antarctic origin, with wide southern distribution; from the extreme south
of S. America including the Falkland Is., along the Andes to Mexico and Jamaica; islands in the southern
parts of the Atlantic and of the Indian Ocean; from Tasmania through E. Australia northwards to
Malesia, Mt Pulog in the Philippines being the most northern extension; highly developed in New
Zealand and neighbouring islands, from there one species extending to Hawaii. Fig. 136.

Note. Subdivision of the genus. KUKENTHAL divided Uncinia into two subgenera, Uncinia (‘ Eu-Uncinia’)
and Pseudocarex, the latter containing only U. kingii Bootr from Antarctic S. America, in which the
rachilla is but shortly hooked. Subg. Uncinia is divided into two sections which are to some extent also
geographically defined: sect. Uncinia, mainly Australian with few species in S. America, and sect. Platyan-
dra which is exclusively American.

KEY TO THE SPECIES

1. Glumes persistent. Stems sharply trigonous, scabrid beneath the inflorescence . . . . 1. U. riparia
1. Glumes caducous, when young often an abscission layer visible. Stems obscurely trigonous, smooth

1. Uncinia riparia R. Br. Prod. (1810) 241;
Bootr in Hook. f. Fl. Tasmania 2 (1860) 102,
t. 152 f. B; BENTH. FI. Austr. 7 (1878) 434; CLARKE,
J. Linn. Soc. Bot. 20 (1883) 392, excl. var.; KUK.
Pfi. R. Heft 38 (1909) 63, excl. var.; S. T. BLAKE,
J. Arn. Arb. 35 (1954) 234; Noort. Blumea 24
(1979) 513. — Carex riparia (R. Br.) Por. in
Lamk, Enc. Méth. Suppl. 3 (1813) 282. — U. sclero-
phylla NELMES, Kew Bull. (1949) 143. — U. ohwiana
KoyaMa, Bot. Mag. Tokyo 69 (1956) 214, f. 6.

Rhizome more or less creeping. Stems approxi-
mate on the rhizome, very slender, sharply trigo-
nous, scabrid in the upper half, (10—)40-75 cm by
1/,-1 mm. Leaves from slightly shorter to slightly
longer than the stems, rigid, flat or canaliculate,
long attenuate, scabrous on margins and nerves in
upper half, 11/,-3(-?4) mm wide; basal sheaths
bladeless or short-bladed, fuscous. Spikelet
narrowly linear, often very loosely flowered,
ebracteate or with a filiform bract usually not over-
topping the inflorescence, 3—7(—15) by 2-5 mm, the
3 part few-flowered, 1—11/,(-2) cm long. Glumes
persistent, oblong-ovate, acute, muticous, rigid,
stramineous with broad 3-nerved green centre and
sometimes brown margins, (4—)5—6(—?8'/,) mm
long, the midrib not reaching the apex. Utricles
slightly exceeding the glumes, erect, linear-oblong
or linear-lanceolate, compressed-trigonous, gla-
brous and smooth, fine-nerved, stramineous,
6-7 by c. 1 mm, at the base subgradually narrowed
into a c. 11/, mm long stipe, at the apex into a
c. 1'/, mm long, compressed-conical beak with
narrow, hyaline mouth. Nut narrow ellipsoid.

Distr. New Zealand, Tasmania, SE. Australia
(Victoria, Upper Hume R. and Mt Kosciusko,
according to BENTHAM, /.c.; no specimens seen); in
Malesia: New Guinea (West Irian, summit of
Mt Wilhelmina; Papua New Guinea).

Ecol. In shaded places, usually between moss on
the floor of the mossy or subalpine forest, 3000-
4100 m, above 4000 m also in grassland with
shrubs. Fi. fr. Jan.—Dec.

Note. Whether the glumes are persistent or
caducous can often only be seen in old inflores-

2. U. compacta

cences. It is not impossible that hybrids occur with
U. compacta, thus giving more variability and
plants in which the glumes fall very late. In these
plants the sharply triquetrous culms become more
or less rounded and smooth instead of scabrous
(not found in Malesia).

2. Uncinia compacta R. Br. Prod. (1810) 241;
Boott in Hook. f. Fl. Tasmania 2 (1860) 103;
F.v.M. Fragm. 8 (1874) 152; BENTH. FI. Austr. 7
(1878) 434; CLarKE, J. Linn. Soc. Bot. 20 (1883)
395; CHEESEMAN, Man. New Zeal. FI. (1906) 800;
KUk. Pfl. R. Heft 38 (1909) 65; CHEESEMAN, Man.
New Zeal. Fl. ed. 2 (1925) 245; LourTEIG, Bull.
Com. Nat. Fr. Rech. Antarct. (1968) 25; Noort.
Blumea 24 (1979) 515. — Carex compacta Porr. in
Lamk, Enc. Méth. Suppl. 3 (1813) 282. — U. rupes-
tris RAOUL, Ann. Sc. Nat. III, 2 (1844) 117; Bootr
in Hook. f. Fl. Nov. Zel. 1 (1853) 286; CLARKE,
J. Linn. Soc. Bot. 20 (1883) 392; Kix. Pfl. R. Heft
38 (1909) 64, incl. var. capillacea KUK.; HAMLIN,
Dom. Mus. Bull. 19 (1959) 39. — U. filiformis
Boott in Hook. f. Fl. Nov. Zel. 1 (1853) 286;
HAMLIN, Dom. Mus. Bull. 19 (1959) 43. — U. ner-
vosa BooTt in Hook. f. Fl. Tasmania 2 (1860) 102.
— U. riparia R. Br. var. stolonifera KUK. & STEEN.
Bull. Jard. Bot. Btzg III, 13 (1934) 213. — U. riparia
(non R. Br.) Onwt, Bot. Mag. Tokyo 56 (1942)
213. — U. subtrigona NELMES, Kew Bull. (1949)
144. — Fig. 137a-d.

Plant laxly to densely cespitose or with short
rhizome and stems densely tufted, erect; sometimes
stems decumbent, forming new tufts; culms slender,
rarely more than 1 mm thick, obscurely trigonous,
smooth, 5-45 cm. Leaves shorter or longer than
stems, flat, involute, convolute or conduplicate,
sometimes (var. nervosa) plano-convex and then
often canaliculate, long attenuate, scabrous on
margins and nerves at least in upper half, '/,-3 mm
wide, the tip mostly rather acute, triquetrous,
rarely flat or plano-convex and blunt (var. nervosa);
basal sheaths bladeless, brown. Spikelets nar-
rowly oblong, loosely to very densely flowered,
sometimes bracteate, (1—)1!/,-5!/. cm by 2!/,-

1979] Cy PERACEAE—II (Kern & Nooteboom) 185

Fig. 137. Uncinia compacta R. BR. var. nervosa CLARKE. a. Habit, nat. size, b. leaftip, x 7. — U. compacta
R. Br. var. alpina Noor. c. Habit, nat. size, d. utricle, x 10. — Exocarya scleroides (F.v.M.) BENTH.
e. Flower diagram (a-b ANU 7161, c-d ANU 7289).

186

FLORA MALESIANA

[ser. I, vol. 9!

20 mm, the ¢ part ?/,-1'/, cm long. Glumes
caducous, when young the abscission line often
already visible, oblong-ovate or lanceolate, acute,
muticous, at least the margins hyaline, greenish or
brown, with 3-nerved central stripe but often with
several more nerves, 4-6(-8 mm in the Antarctic
Is.) by 1.6-3 mm. Utricles shorter to slightly longer
than glumes, obliquely erect to patent, lanceolate
elliptic or ovoid, with 2 conspicuous submarginal
nerves, sometimes slenderly nerved towards the
base or striate when these nerves are prolonged,
3!/,-6(—7) by 1-2 mm, at the base contracted into a
1-11/, mm long stipe, at the apex (gradually)
narrowed into a 1-2 mm long beak. Nut ellipsoid.

Distr. Australia (Victoria, New South Wales,
Tasmania), New Zealand, Amsterdam I., Kergue-
len, Crozet, Marion, Gough and Tristan da Cunha;
in Malesia: Philippines (Luzon: Mts Pulog &
Banahao; Mindanao: Mt Apo), N. Borneo
(Mt Kinabalu), Celebes (Latimodjong Mts), New
Guinea.

Ecol. In the tropics in the high mountains,
2000-4300 m, outside the tropics in temperate and
cold climates, in open places and in forest.

Note. The second character in the key to the
species differentiating U. compacta from U. riparia
breaks partly down in some specimens from Vic-
toria and New South Wales; these specimens have
a sharply triquetrous and (interruptedly) scabrous
stem.

KEY TO THE VARIETIES

1. Spikelets c. 1 cm long. Leaves strongly circin-
nate towards the apex. Usually small plants
c. var. alpina
1. Spikelets 11/,-S'/, cm. Leaves not or less circin-
nate. Plants often larger.

2. Leaves flat (always in New Guinea), involute,
convolute or conduplicate with acute, tri-
gonous or triquetrous tip. . a. var. compacta

2. Leaves plano-convex with flat, or plano-
convex blunt tip. b. var. nervosa

a. var. compacta.

Spikelets 11/,-51/, cm. Leaves flat, involute,
convolute or conduplicate. Leaf tip acute, trigo-
nous or triquetrous, undulate or (somewhat)
circinnate.

Distr. As the species.

Ecol. In forest, 2000-3700 m. Fi. fr. Jan.—Dec.

b. var. nervosa CLARKE, J. Linn. Soc. Bot. 20 (1883)
395. — U. nervosa Bootrt in Hook. f. Fl. Tasmania
2 (1860) 102; HAMLIN, Dom. Mus. Bull. 19 (1959)
50. — Fig. 137a—b.

Spikelets 1'/,-3 cm. Leaves plano-convex,
usually canaliculate, sometimes some of the leaves
convolute. Leaf tip flat, or plano-convex, blunt.

Distr. Australia (New South Wales, Kosciusko
area and Tasmania), New Zealand; in Malesia:
New Guinea.

Ecol. In open places, mostly in grassland,
3000-4026 m. Fi. fr. June—Nov.

c. var. alpina Noort. Blumea 24 (1979) 519. —
Fig. 137c-d.

Spikelets c. 1 cm long. 2? Flowers c. 5; utricles
3'/,4 mm. Leaves involute, strongly circinnate
towards the triquetrous or trigonous apex.

Distr. Malesia: New Guinea (Mt Wilhelm and
Mt Giluwe).

Ecol. In exposed places, often temporarily
covered by snow, also on solifluction terraces,
3770-4350 m. Fi. fr. Nov.—April.

ADDENDA

7: 452 Add to A. SUBFAMILY CyPEROIDEAE I. Tribe Hypolytreae after 6. Paramapania: 6a. Exocarya.
Change in the SYNOPTICAL KEY TO THE MALESIAN GENERA fork 6, second lead, into:

6. Hypogynous scales 2 or 4.
6’. Hypogynous scales 2.
6’. Hypogynous scales 4.

. 7. Hypolytrum
6a. Exocarya

7: 454 Change in the KEY TO THE GENERA fork 19, first lead, into:
19. Hypogynous scales 2 or 4. Stigmas 2. Inflorescence paniculate.

19’. Hypogynous scales 2, either of them with a stamen in its axil .

19’. Hypogynous scales 4, stamens 3

. 7. Hypolytrum

6a. Exocarya

6a. EXOCARYA

BENTH. in Hook. Ic. Pl. 3 (1877) pl. 1206. — Fig. 137e.

Leaves situated throughout the stem. Inflorescence an aemubeliaee panicle, the
lower 2-3 bracts long, foliaceous. Spikelets small, the upper 1 or 2 flowers bisexual,
the other 2-3 flowers male. Glumes imbricate, the lower ones often empty. Flowers

187

1979] Cy PERACEAE—II (Kern & Nooteboom)

compressed, the 2 outer hypogynous scales subopposite, folded, transverse and
sharply keeled, the 2 inner ones flat or concave, parallel with the glumes. Stamens
3. Style broadened towards the base, with 2 filiform stigmas. Nut exserted from the
glumes, crowned by the persistent style-base.

Distr. Monotypic. Australia (Queensland, New South Wales); in Malesia: E. New Guinea

(Ferguson I.).

1. Exocarya scleroides (F.v.M.) BENTH. in Hook.
Ic. Pl. 3 (1877) pl. 1206; F. M. BarLey, Queensl.
FI. 6 (1902) 1777; S. T. Biaxe, Proc. R. Soc.
Queensl. 54 (1943) 72. — Cladium scleroides
F.v.M. Fragm. 9 (1875) 12. — Scleria ustulata
F. M. BaILey, 3rd Suppl. Syn. Queensl. Fl. (1890)
81. — E. montivaga Domtn, Bibl. Bot. XX, Heft 85
(1915) 484. — Fig. 137e.

Stems from a creeping rhizome, to c. 1 m high,
foliaceous, 3-angled. Leaves flat, to 6!/, mm broad,
tapering into long subulate points; sheaths close,
with minutely fimbriate mouth. Jnflorescence an
umbellate panicle, varying in size, but often very
large, to 20 cm @, the longest rays up to 15 cm,
pedicels filiform. Involucral bracts several, folia-
ceous, the longest scarcely as long as the inflores-
cence. Spikelets dark brown, narrowly obovate,
c. 4 mm long. Lower glumes empty, outer ones
very short, c. 1 mm, gradually passing into the
c. 2 mm long flowering ones. Hypogynous scales
as long as the glumes. Nut much exserted, ovoid-
oblong (or globose: ‘E. montivaga’), c. 4 mm long,

smooth, the remains of the spikelet forming a small
tuft at its base. Style-base black, larger than the
ovary at the time of flowering but not enlarged
afterwards.

Distr. E. Australia (New South Wales, Queens-
land); in Malesia: Papua New Guinea (once
collected in Ferguson I.).

Ecol. In forest, 720 m. Fi. fr. Nov.

Although sometimes rather abundant, its
occurrence in Australia is very sporadic over its
rather extensive geographical range. It seems to be
usually a constituent of some of the less densely
closed forests.

A notable feature is that only a small propoftion
of the spikelets produce mature nuts (BLAKE, /.c.).

Note. This plant belongs to the tribe Hypoly-
treae and is obviously intermediate between
Paramapania and Hypolytrum, differing from the
former in the presence of only 2 interior, flat
hypogynous scales, while Hypolytrum lacks those
scales. The stomata are tetracytic, as in Lepironia
and Scirpodendron.

Excluded
Eriophorum comosum (WALL. in Roxb.) NEEs in Wight, Contr. (1837) 110; Mia. Fl. Ind. Bat. 3 (1856)

330; STEEN. Bull. Jard. Bot. Btzg III, 13 (1933) 200.

MIQUEL cited this to occur in Malaya (Penang I.); on what grounds is uncertain. It must be a mystifica-
tion or mislocalisation of specimens. CLARKE (1893) did not mention this locality in Fl. Br. Ind.

In Herbarium Bogoriense VAN STEENIS, /.c., found specimens of E. comosum, said to have been collected
in Karimata I. (off W. Borneo), mixed with specimens of Machaerina rubiginosa. This was interpreted as
an unintentional mixture; E. comosum certainly does not occur in the low Karimata I.

Eriophorum filamentosum Boeck. Bot. Jahrb. 5 (1884) 506 was based on a GRIFFITH collection credited

to have been collected in Malaya.

CLARKE (FI. Br. Ind. 6, 1893, 664) reduced this to Xerotes leucocephala R. Br., of which Xerotes
filamentosa A. CUNN. msc. is a synonym. This is not a Cyperaceous plant, but belongs to Lomandra of the
Liliaceae sens. lat., a genus which occurs outside Australia only in New Guinea. It is hard to believe that
BOECKELER made such an error. This may be a slip of the pen by CLARKE.

The identity of BOECKELER’s type should still be checked; but no Eriophorum has ever been found in

Malaya.

LILIACEAE—I (J. P. Jessop, Adelaide)!

Herbs, usually glabrous, with perennial underground stems (corms, bulbs, tubers,
or rhizomes) in all Mal. spp. Aerial stems usually herbaceous and annual, erect or
climbing. Leaves simple, caespitose and basal, sometimes distichous, if cauline
usually alternate, generally linear to lanceolate or oblanceolate especially when
basal, but sometimes shorter and broader (to ovate) when cauline, usually sessile
(in Asparagus and Petrosavia reduced to non-photosynthetic scales), usually with
parallel venation. Stipules 0. Inflorescence terminal or axillary, usually racemose
(less often at least partly umbellate) or flowers solitary, usually bracteate. Flowers
bisexual (except, in Mal., Asparagus cochinchinensis and Astelia alpina), usually
actinomorphic. Perianth segments almost invariably 6 in two more or less similar
or less often distinctly dissimilar whorls of 3, petaloid, connate or free, the outer
whorl sometimes saccate at the base. Stamens 6, inserted on receptacle or perianth;
filaments connate or free, rarely forming a corona-like ring attached to the
perianth; anthers basifixed or dorsifixed, rarely sessile, usually 2-celled, extrorse to
introrse or rarely dehiscing by an apical pore. Ovary usually superior, of 3 (usually
fused) carpels; styles 1 or 3, simple or 3-branched; locules usually 3 (1 in Trica-
listra); ovules 1 to numerous, placentation axile, rarely basal or parietal, usually in
2 rows. Fruit usually a loculicidal or septicidal capsule or berry, rarely the ovary
wall ruptured by the developing seed which develops unprotected by a fruit,
perianth caducous or persistent. Seeds with copious fleshy or cartilaginous endo-
sperm.

Distribution. About 180 genera with approximately 3500 spp., distributed all over the world,
especially in the temperate regions of Asia, Australasia and Africa, but relatively poorly repre-
sented in South America (13 genera).

In Malesia 22 genera, with a total of 31 spp., no genus being represented by more than two
species. The only genus endemic to the region is the Malayan genus Tricalistra whose separation
from Tupistra is, however, somewhat uncertain. Most other genera are represented in Malesia
by a minority of their species, exceptions being Gloriosa and Peliosanthes, which are probably
both monotypic, and Petrosavia, which consists probably of two species.

The genera can roughly be arranged into three geographical groups.

Old World genera are Asparagus, Chlorophytum, Dianella, Gloriosa and Iphigenia, among
which Chlorophytum is mainly Africa-centred, and Dianella mostly Australasian.

Northern hemisphere genera, especially from the Far East, Sino-Himalayan, are the following:
Aletris, Disporopsis*, Disporum, Lilium*, Liriope*, Ophiopogon, Peliosanthes, Petrosavia,
Tricyrtis*, and Tupistra. Of these four, provided with an asterisk, are found only in Malesia in
the Philippines, and Tupistra only in Malaya and Sumatra. All of them are absent from East
Malesia. Most of their species occur in the montane zone, testimony of their subtemperate
ecology.

Australasia-derived genera are Arthropodium, Astelia, Caesia, Schelhammera, Thysanotus, and
Tricoryne. Their occurrence in Malesia is confined to New Guinea, except for Thysanotus
chinensis BTH. which is found through Malesia as far as Thailand and S. China. Their ranges are
sometimes wider in Austral regions, as Arthropodium and Caesia occur also in the Malagasy
area, and Caesia also in South Africa, while Astelia ranges widely from Mauritius to the southern
Pacific islands and the Falkland Islands. Except for Astelia, which occurs in Malesia only in the

(1) Gratefully acknowledging a stipend from C.S.I.R., Pretoria, for work on this revision during the
period July 1973 to June 1974, at the Rijksherbarium, Leyden, when on long-leave from Rhodes Univer-
sity, Grahamstown, South Africa.

With bibliographic co-operation of the General Editor.

(189)

190 FLORA MALESIANA [ser. I, vol. 9!

alpine zone of New Guinea, all the species of the genera of this southern group are bound to
lowland drought habitats.

Ecology. Of the 22 genera 13 are integrated in lowland to montane everwet-forest conditions.
Astelia alpina is a high-altitude cushion plant which sometimes plays a significant role in the
alpine bogs of New Guinea. The eight other genera, Arthropodium, Asparagus, Caesia, Gloriosa,
Iphigenia, Schelhammera, Thysanotus, and Tricoryne, are constituents of areas subject to a
seasonal climate. Consequently, the ranges of Asparagus racemosus, Gloriosa superba, Iphigenia
indica and Thysanotus chinensis show in Malesia the usual disjunctions of drought-preferring
plant species. They are predominantly grassland or open savannah plants at low altitudes. Most
of them are Australasia-centred.

Except for Astelia papuana, species of these Liliaceae sensu stricto do not form major con-
stituents or natural communities.

Dispersal. The great majority of Liliaceae spread and reproduce vegetatively by the branching
of their subterranean axes. In most species this appears to be a slow process, with the branches
often not extending more than a few centimetres in a year. It may, however, result in fairly dense
monospecific stands, for example, in Astelia and Liriope.

Fruits are generally capsules or berries. In the former, dispersal mechanisms do not usually
result in the removal of seeds to any great distance, although wind and water can contribute
significantly.

Birds are probably the most efficient vectors over longer distances. Several genera have
fleshy fruits and two (Ophiopogon and Peliosanthes) have a fleshy coating to the seeds, which are
exposed through rupture of the ovary wall. Mammals may also disperse the seeds by eating the
fruits. Liliaceae seeds in other areas are known to be carried by ants if there is a substance
attractive to ants (often oil bodies) in the testa or fruit. Specific data on the Malesian species have
not, however, been found.

For a study of the structure and relationships of the seeds, see HUBER, Die Samenmerkmaie
und Verwandtschaftsverhdltnisse der Liliifloren, Mitt. Bot. Staatssamml. Miinch. 8 (1969)
219-538.

Cytology. Liliaceae, because of the usually large size of their chromosomes, because of the
ease with which material can often be obtained at the stages of division required for study, and
because many species are in cultivation, have been fairly well studied cytologically. Chromosome
numbers, and even basic chromosome numbers vary widely sometimes within, as well as between,
genera. At least six different somatic numbers have, for example, been reported for Disporum,
based on x = 6, 7, 8, 9 and 11. Other genera, for example, Asparagus (x = 10) and Dianella
(x = 8), have relatively stable basic numbers, although polyploidy may be common.

Taxonomy. The family Liliaceae, in the sense of BENTHAM & HOOKER and of KRAUSE in
E. & P. Nat. Pfl. Fam. is a very large and rather heterogeneous one including possibly as many
as 3500 species. Many more recent authors have attempted to distribute these species over a larger
number of families. In this treatment the family delimitation of HUTCHINSON (Families of flower-
ing plants, ed. 3, 1973) has been adopted, with two modifications: the inclusion of Petrosavia
which HUTCHINSON placed in its own family, and of the naturalised Nothoscordum which
HUTCHINSON placed in the Amaryllidaceae. It is very doubtful if the family is more naturally
defined by excluding a number of genera represented in Malesia, as HUTCHINSON has done,
but this has been followed here as much for the convenience of dealing with smaller families as
for any conviction that these families have any botanical significance. Table 1 indicates the
genera retained in the Liliaceae and the families to which other genera, sometimes included in the
Liliaceae, were ascribed by HUTCHINSON. These families have also been included in the key to the
genera of Malesian Liliaceae.

Much work remains to be done on the relationships of Liliaceous genera and HUTCHINSON’S
work, although the most recent, is probably no better than KRAUSE’s. HUTCHINSON’s placing of
Ophiopogon and Peliosanthes in separate tribes is, for example, almost certainly unjustified.

Uses. Several species of Liliaceae native to Malesia have been taken into cultivation as
garden ornamentals, for example of the genera Dianella, Gloriosa, Lilium, Liriope, and Ophio-
pogon. Other uses are, however, rather few. Several genera, for example, Asparagus, Gloriosa,
Ophiopogon, have been used in traditional medicines but they have not contributed to modern

1979 } LILIACEAE—I (Jessop) 191

Table 1. Malesian Liliaceous genera in the classification by KRAUSE (1930), first column, and by HUTCHIN-
SON (1973), third column.

Subfamily I. Melanthioideae
if 2. Petrosavieae — Petrosavia (Petrosaviaceae)
IIe 6. Uvularieae — Schelhammera Would
ay ees vularieae
I: 7. Tricyrteae — Tricyrtis Tricyrtideae
I. 8. Anguillarieae — Iphigenia Iphigenieae
Subfamily III. | Asphodeloideae
Ill. 11. Asphodeleae — Chlorophytum
2 ines Asphodeleae
Ill. lla. Asphodelinae — Arthropodium Asphodeleae
— _ Tricoryne Johnsonieae
— Caesia Asphodeleae
Ill. 11g. Dianellinae — Stypandra :
sae Dianelleae
Ill. 17. Lomandreae = pete } (Xanthones hacnceces
Subfamily IV. = Allioideae — Nothoscordum (Amaryllidaceae)
(introduced)
Subfamily V. Lilioideae — Lilium Tulipeae
Subfamily VII. Dracaenoideae
VII. 27. Dracaeneae — Cordyline
D (Agavaceae)
— Dracaena
— Astelia Milliganieae
Subfamily VIII. Asparagoideae
VIII. 28. Asparageae — Asparagus Asparageae
VIII. 29. Polygonatae a= ee : } Pe hyeonaiee
— _ Disporopsis
VIII. 30. Convallarieae — ee } Uspidiurene
— Tricalistra
Subfamily IX. | Mondoideae — Liriope Ophi ae
— Ophiopogon ETB Oe
— Peliosanthes Peliosantheae
Subfamily X. Aletroideae — Aletris Narthecieae
Subfamily XI. Luzuriagoideae — Luzuriaga
— Geitonoplesium (Philesiaceae)
— Eustrephus
Subfamily XII. Smilacoideae — Rhipogonum
— Smilax (Smilacaceae)
— Heterosmilax

192 FLORA MALESIANA [ser. I, vol. 9!

medicine. Others have edible underground parts (e.g. Arthropodium) or fruits (e.g. Astelia), but
none is probably of great significance.

Notes. A very large number of exotic Liliaceae are recorded to be or have been cultivated in
gardens in Malesia. They have been treated elaborately, with keys for their identification by
C. A. BACKER in his ‘Handboek voor de Flora van Java’, part 3 (1924), in Dutch, and by C. A.
BACKER & R. C. BAKHUIZEN VAN DEN BRINK Jr in their ‘Flora of Java’, volume 3 (1968), in
English.

Only one exotic, Nothoscordum inodorum, has been introduced and has run wild in West Java;
this has been incorporated in the treatment.

Almost all drawings were made by Mr. L. DuTKiEwicz, Adelaide.

KEY TO THE GENERA
including the families sometimes segregated from Liliaceae sensu lato

1. Leaves reduced to small, non-photosynthetic scales.
2.-Stem: branched bearme ereen'cladodes “Yates. ] 4: . . Sees eenols St Sessa Acnarrans
2. Stem simple. Small echlorophyllous saprophyte .. . . . . 4 Petrosavia
1. Leaves usually well-developed, always green (sometimes absent when the plant i is in flower).
3. Anthers 1-celled. Plants Leberg) woody and often prickly climbers (Heterosmilax, Rhipogonum,

Smilax) . AIREY 0, Teas pee. Wes oes: Sit ose: eeu eee. eee Smilacaceae
3. Anthers 2-celled. Plants erect or - herbaceous, not prickly, ‘rarely woody climbers.

4. Plants dioecious.

5. Panicle glabrous or scaberulous. Flowers clustered at all or most nodes (Lomandra)

Xanthorrhoeaceae
5. Panicle silvery-scaly. Flowers solitary ateach node ............... .22. Astelia
4. Flowers bisexual.
6. Foliage leaves caespitose, usually basal or radical.
7. Inner perianth segments fringed... 2.9. 5. 2 ss 3 ee te eo ee Oe VSaNO GIs
7. Perianth segments not fringed.
8. Inflorescence a simple umbel. Bulbous . . . .... . . 23. Nothoscordum
8. Inflorescence a raceme, spike or panicle. Plant rhizomatous.
9. Flowers sessile. Anthers sessile or subsessile on the perianth.
VOW TU tanCADSUISR ay oe oe ssctewsiee ses ee, a ie tiw’o xopacielipeer voy 0 ho, |B ace pa A OES LEREIES
10. Fruit fleshy.
11. Style distinct. Stigma simple or indistinctly lobed. . ....... .. . 17. Tupistra
11. Style absent. Stigmas distinctly3 ...... bee ee es ed Onme I CRHISERST

9. Flowers pedicelled. Filaments usually well- developed.
12. Seeds fleshy and exposed soon after the commencement of their development.

13. Filaments free of one another. Corona absent ...... . 2 ss ee ES liniope
13. Filaments connate or anthers sessile on a staminal corona.
14. Anthers borne on a distinct staminal corona. ....... . ... . 20. Peliosanthes
14. Anthers borne on connate filaments ............... . 19. Ophiopogon

12. Seeds retained in the fruit until mature.
15. Anthers dorsifixed.
16 Ovaryenalt-inferiorn, =)..4. 4 90. 5 4. oo ae See en so ce ae
16. Ovary fully superior.
17. Fruit a capsule. Stems to 5 cm long and 2-3 mm diameter (Romnalda) . Xanthorrhoeaceae

17. Fruit a berry. Stems woody and usually long (Dracaena, incl. Pleomele) . . Agavaceae
15. Anthers basifixed.
SE cuitarberrys. st. eee. | heen Teiges. et Se eee ee . . . . 9. Dianella

18. Fruit a capsule.
19. Base of anthers with a papillose appendage. Pedicels 12-20 mm. Outer perianth segments

distinctly broader than innenonesi 2.) eee on ee . . . 6. Arthropodium
19. Base of anthers without appendages. Pedicels 3-12 mm. Perianth segments equal.
20. Perianth spirally twisting after anthesis, blue ........... > « #. Caesia

20. Perianth segments persistent, but not twisted after flowering, white to green
8. Chlorophytum
6. Foliage leaves distributed at intervals along the stem.
21. Plant with scaly bulb. Flowers 10-25cmlong .......... J. 4)

1979] LILIACEAE—I (Jessop) 193

21. Plant with a rhizome, corm or tuber. Flowers up to at most 9 cm.
22. Stem(s) woody. Venation reticulate (Eustrephus, Geitonoplesium). . .... . Philesiaceae
22. Stem(s) herbaceous. Venation parallel.
23. Style simple, or if 3 lobed the filaments forming a corona.
DAS eruitra berry. Inilonescence aipanicle |) 6) ays ee 3 erhcns las ee 9. Dianella
24. Fruit consisting of 1-3 nutlets. Inflorescence consisting of umbels each on a winged peduncle
11. Tricoryne
23. Style branched, sometimes to the base.

25. Flowers large, 5-9 cm. At least some leaves ending in a coiled tendril . . . 1. Gloriosa
25. Flowers smaller, up to 3 cm. Leaves never ending in a tendril.
26. Filaments expanded to form acorona. Fruita berry ......... 15. Disporopsis
26. Filaments free, sometimes connivent.
Dime rilitva Dery; Bile OR en oes ek Gone ee ee aes 14. Disporum

27. Fruit a capsule.
28. Pedicels not articulated. Anthers dorsifixed.

29 Rootstock a rhizome. Stylelbranchesibifide ys eee 2. Tricyrtis
29 sRootstocka\ corm. style branches)simple. ys) 4 4 ee 3. Iphigenia
28. Pedicels articulated. Anthers basifixed ............. 5. Schelhammera

1.GLORIOSA

LINNE, Sp. Pl. (1753) 305; Gen. Pl. ed. 5 (1754) 144; Baker, J. Linn. Soc. Bot. 17
(1879) 457; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 266; HuTcH. Fam.
FI. Pl. ed. 2, 2 (1959) 606; FIELD, Kew Bull. 25 (1971) 243; Lilies and other Liliaceae
1973 (1972) 93. — Methonica ToURN. ex CRANTZ, Inst. rei herb. 1 (1766) 474. —
Fig. 1, 2.

Climbing, or less often erect, glabrous herbs. Rhizome perennial, tuberous,
horizontal; roots fibrous. Aerial stem annual, moderately branched; the branches
spreading or erect-spreading at the base. Leaves cauline, alternate, opposite or in
whorls of 3 (4), flat, with many veins and a strong midrib, sessile, entire, lanceolate
to ovate, slightly amplexicaul, obtuse at the base, narrowing gradually to an apical,
coiled tendril (sometimes lacking tendrils in erect forms); basal leaves lacking a
lamina or tendril, ensheathing the stem. Pedicels solitary, axillary in the axils of
vegetative leaves, not articulated, cernuous. Perianth segments connate at the base,
many-veined, subequal, reflexed or rarely spreading with a longitudinal papillose
furrow in the basal + third of the adaxial surface. Filaments filiform, attached to
the receptacle; anthers dorsifixed, linear-oblong, extrorse. Ovary superior, sessile,
oblong-ovoid to oblong-obovoid, 3-celled, 1 cell slightly shorter than the other 2;
ovules axile, numerous; style filiform with 3 stigmatic branches, reflexed or spread-
ing from the attachment to the ovary. Capsule coriaceous, septicidal; seeds globose;
perianth persistent but withering as the capsule enlarges.

Distr. Widespread in tropical and southern Africa, Madagascar, India, Burma and SE. Asia, as far as
West Malesia.

Ecol. Usually climbing in bushes, in habitats ranging from savanna to forest.

Note. Stated by FieLp (1971, 1972) to be monotypic. In Malesia only a single indigenous species has
ever been recognized, although other species have been described elsewhere.

1. Gloriosa superba LINNE, Sp. Pl. (1753) 305; Nutt. Pl. (1927) 436; Back. Onkr. Suiker. (1928)
Baker, Fl. Cap. 6 (1897) 525; BACK. Trop. Natuur 189, Atlas (1933) t. 201; GAGNeP. FI. Gén. I.-C. 6
3 (1914) 117, tab. col.; Rw. Fl. Mal. Pen. 4 (1924) (1934) 807; H. Perrier, Fl. Madag. fam. 40 (1938)
338; Back. Handb. FI. Java 3 (1924) 50; Petcu, 135; Sitva, Ceyl. J. Sc. sect. A, 12 (1945) 155;
Ann. R. Bot. Gard. Perad. 9 (1925) 243; HEYNE, SANTAPAU, J. Bomb. Nat. Hist. Soc. 46 (1946) 202;

FLORA MALESIANA [ser. I, vol. 9!

Fig. 1. Gloriosa superba L. Botanic Garden Singapore, Febr. 1952 (Photogr. M. R. HENDERSON).

1979 ]

HEND. Mal. Gard. Pl..4 (1951) 10; Mal. Wild
Flow. Monoc. (1954) 178, f. 104; Hottr. MAHA
Mag. 15 (1958) 75 (hybrids); BAcK. & BAKH. f. FI.
Java 3 (1968) 85; HutcH. & Daz. Fl. W. Trop.
Afr. ed. 2, 3 (1968) 106; Fie_p, Kew Bull. 25 (1971)
243. — Methonica superba (L.) CRANTZ, Inst. rei
herb. 1 (1766) 474; ZoLv. Syst. Verz. 1 (1854) 66;
Nat. Tijd. N. I. 14 (1857) 149; Mia. FI. Ind. Bat. 3
(1859) 550. — G. virescens LINDL. Bot. Mag. (1825)
t. 2539. — Fig. 1, 2.

Stems usually climbing to c. 2 m (rarely to 6 m),
less often erect; green. Leaves (including the tendril)
8-17'/,(-25) by (13/,-)1'/,4(C4'/2) cm; tendril
usually less than 1 cm long. Pedicels 4!/,-19 cm.
Perianth segments narrowly elliptic, with undulate
or crisped margins, 5—7(-9) by 3/s-1'/,(-3) cm
broad, yellow or red, often (perhaps always in
Asia) yellow or green towards the base at first but
becoming red throughout later. Filaments spread-
ing, 2'/,-5 cm long; anthers 7-10 mm long. Ovary
8-15 mm long; style including the filiform, 3-7
(-12) mm long style branches 3'/,-5'/, cm long.
Capsule 4-10 by c. 1'/,-2 cm. Seeds vivid-red or
orange-red, with a fleshy testa, c. 5 mm @, tardily
falling.

Distr. Tropical and southern Africa, Madagas-
car, India to Indo-China; in Malesia: Java (also
Madura and Kangean Is.), S. Celebes, and all
Lesser Sunda Is.

Ecol. Brushwood, hedges, teak-forest, only in
regions subject to a strong dry season, from near
the beach and dunes up to c. 300 m altitude (very
rarely 600 m), locally common. It is not native in
Sumatra and Borneo, and probably not in con-
tinental Malaya. This disjunction in its range
between continental SE. Asia and Central South
Malesia is clearly caused by its drought preferring
ecology. Fi. fr. Jan.—Dec.

Uses. Commonly grown as a garden ornamental.
The tuber is said to be poisonous (through
colchicin), but only slightly so as was tested by
BoorsMA (BACKER, 1914; HEyngE, 1927). Fig. 2.

Vern. Klimlelie, D, flame lily, superb lily, E;
Java: kembang djonggrang, k. kuku matjan, k. sung-

LILIACEAE—I (Jessop)

195

sang, M, dongkél sungsang, mandalika, pa(n)tjing
towo, J, katongkat, S, mand(h)alika, Md, Balin.;
Lesser Sunda Is.: enatba, sikal, Dawan lang.,
Timor.

Note. In sterile state easily distinguished from
another climbing monocot with coiled apical leaf-
tendrils, Flagellaria indica L., by absence of a
leaf-sheath.

Fig. 2. Gloriosa superba L. Old rhizome with scar,

the apex with a new tuber emitting roots and a

vertical shoot. The two triangular elongations of

the new tuber will grow later into new rhizomes;

x 1/,. Botanic Garden, Bandung, 1952. Dug up by
L. VAN DER PUL.

2. TRICYRTIS

WALL. Tent. FI. Nap. (1826) 61, t. 46; KRAUSE in E. & P. Nat. Pfi. Fam. ed. 2, 15a
(1930) 269; HutcH. Fam. FI. Pl. ed. 2, 2 (1959) 608, nom. gen. cons. — Compsoa
D. Don, Prod. Fl. Nep. (1825) 50. — Fig. 3.

Erect puberulous or glabrescent herbs. Rhizome short, creeping. Aerial stem
annual, simple or branched. Leaves cauline, alternate, flat, with several veins and a
strong midrib, sessile, entire, lanceolate to ovate, with a sheathing base. Jnflores-
cence terminal, racemose, simple or branched, or flowers in the axil of vegetative
leaves. Pedicels solitary, not articulated. Perianth segments free or very shortly
united, equal or subequal but the outer three saccate at the base, erect to spreading.

196 FLORA MALESIANA [ser. I, vol. 9!

SER

fea AMANENAABY Gay yer Tt

Fig. 3. Tricyrtis imeldae GUTIERREZ. a. Habit, * '/4, b. flower, nat. size, c. outer perianth segment, d.
inner perianth segment, e. gynoecium, all x 11/3, f. fruit, nat. size, g. seed, x 12 (Redrawn from GUTIER-
REZ, Philip. J. Sc. 103, 1974, 3, fig. 1).

1979] LILIACEAE—I (Jessop) 197

Filaments flattened, more or less connivent, free of the perianth; anthers dorsifixed,
versatile, oblong, extrorse. Ovary superior, sessile, oblong, 3-celled, 3-angled;

ovules axile, numerous; style columnar; with 3 spreading or recurved bifid
branches. Capsule septicidal. Seeds oblong or ovoid.

Distr. Possibly c. 20 spp., largely in Japan, also in Manchuria, Korea, throughout China to the Hima-
layas, Taiwan, and North Malesia: Philippines.

Notes. HUTCHINSON placed this genus in the tribe Tricyrtideae possibly with a South African genus
Sandersonia as the only other member. KRAUSE placed it in the Tricyrteae with the closely related, possibly
congeneric, Brachycyrtis. The genus appears to be taxonomically rather isolated, but probably closest to
Gloriosa of the Malesian genera.

A thorough revision of the species is badly needed.

1. Tricyrtis imeldae GUTIERREZ, Philip. J. Sc. 103
(1974) 171, f. 1. — Fig. 3.

Stems erect to 70 cm high, unbranched, puberu-
lous at first. Leaves thick and fleshy when fresh but
becoming membranous when dried, acute, the
lower narrowly elliptic-lanceolate, with cuneate
base, the upper broadly lanceolate to elliptic, with
cordate base, (6-)12-16 by (3—)4—-5 cm, glabrous
except for the main veins beneath. Inflorescence a
terminal bifurcate raceme c. 18-20 cm, puberulous;
pedicels 3-5 mm, puberulous. Flowers greenish-
white with purple spots inside, to over 3 cm long,
infundibuliform, glabrous, segments linear-oblong
to oblong spathulate. Filaments 16-18 mm long;
anthers 3 mm long, yellowish-brown. Ovary 10 mm
long; style 8 mm, its branches 8 mm long, purple,

spreading, tuberculate on the inner surface. Fruit
c. 25-30 by 4-6 mm. Seeds flat, oblong, c. 2 mm
long.

Distr. Malesia: S. Philippines (Mindanao:
Tasaday, Cotabato), reported to be rather rare;
only known from the type.

Ecol. Primary forest, along stream at c. 1300 m.
Fl. Aug.

Vern. Philippines: amutmagiso, Tasaday.

Notes. I have only seen the type and have not
been able to add to the description by GUTIERREZ.

He compared the species with the Formosan
T. stolonifera, from which it chiefly differs in the
shape and colouring of the perianth segments.
Close study of more material and variability is
needed to check the specific difference.

3. IPHIGENIA

KUNTH, En. Pl. 4 (1843) 213; Baker, J. Linn. Soc. Bot. 17 (1880) 450; KRAUSE in
E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 272; H.PeRRIER, FI. Madag. fam. 40
(1938) 136; HuTcH. Fam. FI. Pl. ed. 2, 2 (1959) 612; OBERMEYER, Kirkia 1 (1961)
84, nom. gen. cons. — Aphoma Rar. FI. Tellur. 2 (1836) 31, nom. rejic. — Fig. 4.

Erect, glabrous herbs. Corm enclosed in dry leaf bases; roots fibrous. Aerial stem
annual, unbranched. Leaves cauline, with tubular, ensheathing bases; the lowest 1
or 2 often with a poorly developed lamina; the others decreasing in size from the
lowest towards the uppermost, the uppermost often approaching the lowest bracts
in size, c. 4-7; lamina flat, glabrous, sessile, entire, linear to lanceolate, acute.
Flowers solitary, in a few-flowered cluster or in a raceme terminating the stem.
Pedicels solitary, erect-spreading, not articulated. Perianth segments free, ++ equal,
spreading or reflexed, few-veined. Filaments flattened, attached to the receptacle;
anthers dorsifixed or basifixed, extrorse. Ovary superior, sessile, ovoid, oblong or
obovoid, 3-celled; ovules co, axile; styles 3, (free or) fused at the base only.
Capsule globose, cylindrical or ellipsoid, loculicidal. Seeds globose or angular;
perianth deciduous.

Distr. Southern and tropical Africa (5 spp.), Madagascar (2 spp.), SE. Asia (4 spp.), of which one species
extends through Malesia to Australia, and New Zealand.
Ecol. Usually in open grassland, sometimes in damp areas.

198

FLORA MALESIANA

[ser. I, vol. 9}

1. Iphigenia indica (L.) A. GRAY ex KUNTH, En.
Pl. 4 (1843) 213; Mia. FI. Ind. Bat. 3 (1859) 552;
Bru. Fl. Austr. 7 (1878) 31; BAKeEr, J. Linn. Soc.
Bot. 17 (1880) 450; E.v.M. Descr. Not. 6 (1885) 18;
Hook. f. Fl. Br. Ind. 6 (1892) 357; BAILEY, Queens].
Fl. 5 (1902) 1641; Laut. Bot. Jahrb. 50 (1913) 292;
Merr. En. Philip. 1 (1922) 202; Back. Handb. FI.
Java 3 (1924) 51; Krause, Bot. Jahrb. 59 (1925)
548; BAcK. & BAKH. f. Fl. Java 3 (1968) 85. —
Melanthium indicum LiINnE, Mant. 2 (1771) 226. —
Anguillaria indica (L.) R. Br. Prod. (1810) 273;
WALL. Pl. As. Rar. 3 (1832) 37, t. 259. — Fig. 4.

Corm usually + globose, 5-10 mm @. Leaves
linear-lanceolate, often with a single conspicuous
vein and 2-8 rather inconspicuous ones, the longest
c. 10-40 cm long, up to 6 mm wide, the shortest
often less than a quarter of the length of the longest;
lower leaf bases sometimes pubescent, glandular
pubescent or scabrid, especially on the veins.
Flowers 1-3. Pedicels erect or erect-spreading
540 mm. Perianth segments + spreading,
narrowly oblanceolate (inner whorl sometimes
narrower than the outer), 5-9 by up to 2 mm, dark-
brown or red-brown, reddish, purplish or white,
described by BACKER (1924) as having a green keel
and apex. Filaments linear, up to half as long as the
perianth, green basally, brown distally; anthers
dorsifixed, c. 1/,-1 mm long. Ovary obovoid to
ovoid, c. 11/,-2'!/, mm long; styles recurved,
c. 1 mm. Capsule c. 10-20 mm long; seeds
c. 1'/, mm ©.

Distr. Ceylon, India to Thailand, S. China, in
Malesia rare: N. Sumatra (Lake Toba), Java
(Indramayu in W, Surabaja and Madura I. in E),
the Lesser Sunda Is. (Timor and Wetar), Philip-
pines (Luzon, Mindanao), New Guinea, and
Australia (W. Australia, Northern Territory and
Queensland).

Ecol. A rather uncommon species of open, often
poor grassiand, always under seasonal climatic
conditions. In Java only in the lowland but else-
where also in the hills, in N. Sumatra at c. 1000 m.
Flowers are reported by BACKER (1924) to occur
for a short period during the rainy period (Dec.—
Jan.) in Java, with fruit ripening in March, after
which the aerial parts soon wither and disappear.
Elsewhere other flowering dates have been noted
from July to August, N of the equator, where
seasons are reversed.

b

Fig. 4. Iphigenia indica (L.) A. GRAY ex KUNTH.
a. Habit, nat. size, b. flower, c. fruit, both x 2
(Partly after WALLICH, PI. As. Rar. 3, 1832, t. 259).

4. PETROSAVIA

Becc. Nuov. Giorn. Bot. Ital. 3 (1871) 7, t. 1; Riv. J. Str. Br. R. As. Soc. n. 24
(1891) 170; KRAusE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 256; HutTcH. Kew
Bull. (1933) 156; STEEN. Trop. Natuur 23 (1934) 52; NAKAI, J. Jap. Bot. 17 (1941)
191; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 546. — Protolirion RipL. Ann. Bot. 9
(1895) 45; Groom, /.c.; RIDL. Fl. Mal. Pen. 4 (1924) 322; KRAUSE in E. & P. Nat.

199

1979 | LILIACEAE—I (Jessop)

Pfl. Fam. ed. 2, 15a (1930) 257, f. 87; NAKAI, J. Jap. Bot. 17 (1941) 191. — Miyoshia
MAKINO, Bot. Mag. Tokyo 17 (1903) 144; NAKAI, J. Jap. Bot. 17 (1941) 191. —
Fig. 5.

Erect, glabrous, saprophytic, pale yellow or cream coloured, herbs lacking
chlorophyll. Rhizome slender, simply or sparsely branched, bearing alternate,
often imbricate scale-leaves. Aerial stems | or less often up to 3 or more, un-
branched. Leaves scale-like, spiral, sessile, entire; the base usually partly embracing
the stem, rather well-spaced. Raceme terminal, usually simple, sometimes corym-
bose. Pedicels solitary, with 0-2 alternate bracteoles near the centre or in the lower
half (often concealed by the subtending bract), not articulated, in the axils of bracts
resembling the scale-leaves. Perianth segments erect-spreading, with a single vein,
cream-coloured to yellow; the outer 3 distinctly inserted outside the inner 3, even
in the open flower, and narrower and shorter. Filaments sublinear attached to the
receptacle or to the base of the perianth segments; anthers basifixed, introrse or
dehiscing laterally. Ovary superior or semi-inferior; the 3 carpels free above the
receptacle; styles 1 on each carpel, capitate or subcapitate; ovules attached to ven-
tral surface, numerous, in 3-c. 6 rows. Fruit dry, dehiscing along the ventral suture;
the 3 segments spreading. Seeds numerous, ovoid, with longitudinal ridges;
perianth persistent.

Distr. Japan (Prov. Mino), China (Kwangsi and Taiwan), Indo-China (Tonkin), in Malesia: Malaya,
N. half of Sumatra, Borneo (Sabah, Sarawak), and Central Celebes.

In addition to the two species from Malesia and Japan, a third has been described from Tonkin and
S. China (Kwangsi): Petrosavia sinii (KRAUSE) KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 257;
GacneP. Fl. Gén. I.-C. 6 (1934) 802, f. 78 (10-13); (Anonymous) Icon. Corm. Sin. 5 (1976) 424, f. 7677. —
Protolirion sinii KRAUSE, Notizbl. Berl.-Dahl. 10 (1929) 806.

It is highly probable that this is a synonym of P. sakuraii and would thus neatly fill a gap in the range of
that species.

Ecol. Saprophytes on the forest flora, in Malesia in the hills at 1000-2000 m.

Notes. This genus has been placed in the Liliaceae (Liliales) by KRAUSE (1930), the Petrosaviaceae
(Alismatales) by HUTCHINSON (1959) and the Miyoshiaceae (Miyoshiales) by NAKAI (1941). Both in being
saprophytic and in having 2- or multi-seriate ovules it is anomalous in either the Liliales (sensu HUTCHIN-
SON) or the Alismatales. ERDTMAN (Pollen Morph. & Pl. Taxon., Angiosp., 1952, 235) described the pollen
as 1-sulcoidate which is unknown in the Scheuchzeriaceae or Alismataceae (Alismatales) but present,
although uncommon, in the Liliaceae. M. Y. STANT (Bot. J. Linn. Soc. 63, 1970, Suppl. 1, 147) investigated
the anatomy of P. stellaris and found it to be indistinguishable from that in the (saprophytic) Triuridaceae
(Triuridales). Although here retained in the Liliaceae it is admitted that further investigation may show it
to be better placed elsewhere.

The roots lack root-hairs but contain an endotrophic mycorrhiza (Groom, 1895).

KEY TO THE SPECIES

1. Inflorescence corymbose; pedicels all arising close to the peduncle apex, usually 10 mm or more long
1. P. stellaris

1. Inflorescence racemose; pedicels spaced along the peduncle, usually 8 mm or less long 2. P. sakuraii

1. Petrosavia stellaris BEcc. Nuov. Giorn. Bot.
Ital. 3 (1871) 8, t. 1; Rw. J. Str. Br. R. As. Soc.
n. 24 (1891) 171; Groom, Ann. Bot. 6 (1892) 380;
HutcuH. Kew Bull. (1933) 156; STEEN. Trop.
Natuur 23 (1934) 52, f. 12 right; NaKat, J. Jap.
Bot. 17 (1941) 191; HeEnp. Mal. Wild Flow.
Monoc. (1954) 178, f. 103; STANT, Bot. J. Linn.
Soc. 63 (1970) Suppl. 1, 147, anat. — Protolirion
paradoxum Rip_. Ann. Bot. 9 (1895) 56; Groom,

l.c. 45, pl. 3; Rip. Mat. Fl. Mal. Pen. Monoc. 2
(1907) 87; Fl. Mal. Pen. 4 (1924) 322, f. 195;
NAKAI, J. Jap. Bot. 17 (1941) 191. — Fig. 5a.
Aerial stems (4-)6-11 cm high. Scale-leaves of
rhizome ovate, c. 2-5 mm long, their base amplexi-
caul, often forming a closed sheath. Scale-leaves of
aerial stem narrowing gradually to the acute apex,
3-6 mm long, their base partly embracing the stem.
Raceme simple, corymbose, with 1-12 flowers.

FLORA MALESIANA

[ser: I, vols

Fig. 5. Petrosavia stellaris BEcc. a. Habit, nat. size

(Redrawn from HUTCHINSON, Fam. FI. Pl. 2,

1959, fig. 347). — P. sakuraii (MAKINO) J. J. SMITH

ex STEEN. b. Habit, nat. size, c. flower, d. flower in

fruit, both x 10 (Redrawn from MAKINO, Bot.
Mag. Tokyo 17, 1903, pl. 5).

Pedicels all arising close to the apex of the stem,
usually less than 1 mm apart, (5—)10—16(—-20) mm.
Outer perianth segments ovate to lanceolate, acute,
1-2 by c. !/, mm. Inner perianth segments ovate,
+ acute, 2!/,-31/, by 11/,-2 mm. Filaments c.2 mm;
anthers less than !/, mm long. Ovary + superior;
styles up to 1 mm long. Capsule segments 3-4 mm
long. Seeds c. 3/, mm long.

Distr. Malesia: Sumatra (West Coast Res.),
Malay Peninsula, Borneo (Sarawak, Sabah), and
Central Celebes (Masamba: Mt Kambuno).

Ecol. Recorded by RIDLey (1924) “in dry woods
at the foot of Dacrydiums” and by EyMA (EYMA
1305, L) from ‘forest, rather dark, without
undergrowth’. Other records refer to sandy forest,
mossy forest and among bamboos. Recorded
between 100 and 1000 m. F/. usually Febr.—Sept.

2. Petrosavia sakuraii (MAKINO) J. J. SMITH ex
STEEN. Trop. Natuur 23 (1934) 52. — Miyoshia
sakuraii MAKINO, Bot. Mag. Tokyo 17 (1903) 144,
pl. 5; /.c. 208. — Protolirion miyoshia-sakuraii
MAKINO, Bot. Mag. Tokyo 17 (1903) 208, nomen
err. et provis., illeg.; PILG. in E. & P. Nat. Pfl. Fam.
Nachtr. 3 (1908) 44, f. 8, ditto. — Protolirion
sakuraii (MAKINO) DANDY, J. Bot. 69 (1931) 53. —
Fig. 5b-d.

Aerial stems (5—)10-21(-27) cm high. Scale-
leaves of rhizome ovate, c. 2-5 mm long, their base
amplexicaul, often forming a closed sheath. Scale-
leaves of aerial stem narrowing gradually to the
acute apex, 4-6 mm long, their base partly embrac-
ing the stem or rarely completely amplexicaul.
Raceme simple or with few-flowered branches
towards the base, elongate, with (3—)6—25(—30)
flowers. Pedicels of mature flowers usually at least
2 mm apart, 3-8 mm. Outer perianth segments
ovate to lanceolate, acute, 1!/,-2 by +/.-1 mm.
Inner perianth segments ovate, + acute, 2-3 by
11/,-13/, mm. Filaments c. 2 mm; anthers less than
1/, mm long. Ovary superior to semi-inferior; styles
up to 1 mm. Capsule segments 2-3 mm long. Seeds
c. 3/, mm long.

Distr. Japan (Prov. Mino), Formosa, Burma
and Malesia: northern half of Sumatra (Gajo-
lands; Westcoast Res.).

Ecol. Along forest paths and on flat forest
ridges, 1000-2000 m. F/. March-July, Nov.

5. SCHELHAMMERA

R. Br. Prod. (1810) 273; Bru. Fl. Austr. 7 (1878) 31; BAKER, J. Linn. Soc. Bot. 17
(1879) 466; BAILEY, Queensl. Fl. 5 (1902) 1642; MAIDEN & BETCHE, Cens. N.S.W.
PI. (1916) 40, as Schellhammera; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930)
266; Hutcn. Fam. FI. Pl. ed. 2, 2 (1959) 606, as Schelhammeria; nom. gen, cons. —

Fig. 6.

1979] LILIACEAE—I (Jessop) 201

Erect, mainly glabrous herbs. Rhizome rather thick; roots fibrous. Aerial stems
annual, simple or with 1-3 branches, often slightly swollen at the nodes; branches
erect. Leaves cauline, alternate, flat, with many veins and usually a strong midrib,
sessile or shortly petioled, lanceolate to ovate, often at least partly amplexicaul,
obtuse or cuneate at the base, acute at apex. Leaves on rhizome and lower parts of
aerial stems and branches scale-like. /nflorescence terminal, consisting of a sessile
or peduncled umbel or of a single flower. Pedice/s straight, articulated or not.
Perianth segments free, equal, spreading to shallowly campanulate, with several
usually inconspicuous veins. Filaments flattened, tapering from the base, attached
to the base of the perianth segments; anthers basifixed, linear-oblong, extrorse.
Ovary superior, sessile, obovoid, globose or oblong, often fairly deeply 3-lobed,
3-celled; ovules axile, few (c. 4-12) per locule; style filiform, with 3 deeply divided
branches. Capsule somewhat fleshy.

Distr. Three spp. in eastern Australia, one of which also in East Malesia: New Guinea.
Ecol. Most records suggest a preference for rain-forest, but also recorded in scrub and on open slopes
in the lowland and hills.

1. Schelhammera multiflora R. Br. Prod. (1810)
274; F.v.M. Descr. Not. 4 (1876) 73; Bru. FI.
Austr. 7 (1878) 32; Laur. Bot. Jahrb. 50 (1913)
292; HALL. f. Nova Guinea 8 (1914) 989; KRAUSE,
Bot. Jahrb. 59 (1925) 548. — Fig. 6.

Stems 10-40 cm, rarely minutely and sparsely
pubescent. Leaves 4-8 by 1-3 cm, usually minutely
ciliate on the margin and veins towards the base;
petiole usually absent, rarely up to 5 mm. Scale-
leaves ovate to lanceolate, dry, sessile, amplexicaul,
usually 5-10 mm long. Pedicels usually 5—30,
articulated at base of flower, 1-3 cm. Perianth
segments obovate, acute, with the sides turned up
to form a gutter-shaped structure in which the
anthers are partly enclosed, swollen on the abaxial
surface at the base, 4'/,-8 by c. 2 mm, white.
Filaments c. 3 mm; anthers c. 2 mm long, brown or
black. Ovary obovoid, 6-lobed, c. 11/; mm long;
style 3-4 mm, divided for at least half its length;
branches adhering to one another rather firmly at
first, later recurving; ovules few (c. 2-4) per
locule. Fruit and seeds not seen.

Distr. Australia (Queensland) and E. Malesia:
S. New Guinea (Merauke area: Okaba; Fly R.
area).

Ecol. In lowland grassland on open slopes, sub-
ject to a strong or distinct dry season, in Queens-
land also in open forest, obviously a rare species,
below 400 m. F/. March—Sept.

Note. The two further Australian species are
S. undulata, which has larger solitary flowers, and Fig. 6. Schelhammera multiflora R. BR. a. Habit,
S. pedunculata, which has peduncled umbels. x 1/,, b. flower, c. capsule, both x 2.

202 FLORA MALESIANA [ser. I, volNoe

6. ARTHROPODIUM

R. Br. Prod. (1810) 276; BAKegr, J. Linn. Soc. Bot. 15 (1876) 351; Bru. Fl. Austr. 7
(1878) 55; BAILEY, Queensl. Fl. 5 (1902) 1629; Ewart, FI. Vict. (1930) 292; KRAUSE
in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 286; BLAcK, FI. S. Austr. 1 (1943) 193;
PAYENS, Nova Guinea n.s. 8 (1957) 388; SCHLITTLER, Mitt. Bot. Mus. Un. Zitirich
207 (1957) 6, map; HuTcH. Fam. FI. Pl. ed. 2, 2 (1959) 598. — Dichopogon KUNTH,
En. Pl. 4 (1843) 622; BAKgr, J. Linn. Soc. Bot. 15 (1876) 318; Bru. Fl. Austr. 7
(1878) 58; BAILEY, Queens]. Fl. 5 (1902) 1631; Ewart, FI. Vict. (1930) 291;
KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 285; BLAck, FI. S. Austr. 1
(1943) 193; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 598. — Fig. 7.

Stemless herbs. Rhizome very short. Roots fibrous or somewhat fleshy, often
producing well-defined, distant tubers. Leaves basal, linear to lanceolate, gradually
expanding towards the base to form (apparently dry and membranous) open
sheaths, sometimes ciliate or with bristles along the margins; veins of the sheaths
sometimes persisting on an outer zone of fibres. Inflorescence a raceme or panicle.
Pedicels \—3(—5)-nate, in the axils of bracts and usually associated with a few
smaller bracts also apparently in the axil of the principal bract, articulated usually
in the distal half, rather long and filiform. Perianth segments usually free, the inner
whorl broader, with 3 or less often 5 veins, spreading. Filaments linear-filiform,
usually attached to the base of the perianth segments; anthers basifixed, oblong to
linear, often strongly arcuate, dehiscing laterally or introrsely; papillose or pubes-
cent appendages always present, usually adnate to the filaments and often also to
the anthers. Ovary superior, sessile, subglobose to ellipsoid, 3-celled; ovules axile,
2-10 in each locule; style filiform, simple, minutely capitate. Capsule dehiscing
loculicidally ; perianth segments adhering, marcescent, not twisting after flowering.
Seeds angular.

Distr. Madagascar (1 sp.), New Zealand (2 spp.), New Caledonia (1 sp.), and c. 9 spp. in Australia, of
which one extends into Malesia: New Guinea (South Papua).

Ecol. Usually in open grassland or open woodland, often at rather low altitudes (below 250 m).

Species have been described as having chocolate or vanilla scents.

Notes. PAYENS (1957) made a strong case for combining Dichopogon with Arthropodium. Nevertheless,
authors (e.g. BURBIDGE and Gray, FI. A.C.T., 1970) have continued to separate these genera principally
on whether the papillose or pubescent staminal appendages are adnate to the filaments (in Arthropodium)
or to the anthers (in Dichopogon). An examination of species represented at Leyden and Adelaide has
convinced me that these two genera cannot be separated. A particularly significant form is A. neo-
caledonicum in which the appendages are attached to both filaments and anthers such that it would be
difficult to assign this species to a genus. In most of the material of Dichopogon the appendages are also

distinctly adnate to the filaments as well as to the anthers. Dichopogon strictus and Arthropodium capillipes,
although placed in separate genera, have almost indistinguishable stamens.

1. Arthropodium strictum R. Br. Prod. (1810) 276; Roots bearing distant tubers c. 1-1'/, cm long.
F.v.M. Descr. Not. 6 (1885) 17; Laur. Bot. Jahrb. Leaves 3-12, suberect, sublinear but narrowing
50 (1913) 292; Krauss, ibid. 59 (1925) 548; PAYENS, gradually towards the apex and sometimes also
Nova Guinea n.s. 8 (1957) 390; EICHLER, Suppl. towards the basal sheath where they again become
Fl. S. Austr. (1965) 83. — Dichopogon strictus broader, glabrous or minutely ciliate on the mar-
(R. Br.) BAKER, J. Linn. Soc. Bot. 15 (1876) 319; gins, 20-45 cm by 3-7 mm; veins of leaves not
Bru. Fl. Austr. 7 (1878) 58; BAILEY, Queensl. Fl. 5 forming fibrous sheaths at base of plant. Peduncle
(1902) 1631; Ewart, FI. Vict. (1930) 291; simple or more often with 1-4 ascending branches,
GARDNER, En. Pl. Austr. Occ. (1931) 18; BLAcK, (20—)30—60(—110) cm high. Bracts rarely up to 6 cm
Fl. S. Austr. 1 (1943) 194. — Fig. 7. long at the base of the lowest branch, but usually

1979] LILIACEAE—I (Jessop) 203

5-15 mm long, ovate to lanceolate, narrowing
gradually to the apex, partly scarious. Pedicels
solitary or less often 2- or 3-nate, erect-spreading,
12-20 mm. Perianth segments purple, 9-14 by
c. 3-3'/, mm. Filaments c. 11/,-2 mm; anthers
3-5 mm long; appendages less than 1 mm long,
free of or only shortly adnate to filaments. Ovary
globose-ellipsoid, 11/4 mm long; ovules 8-10 in
each locule; style 6-7 mm. Capsule subglobose,
c. 5mm @, with several seeds; perianth marcescent
or rarely persistent.

Distr. Australia (all states but not yet from the
Northern Territory) and Malesia: SE. New Guinea
(Port Moresby area).

Ecol. Open grassland and open woodland at
low altitude, subject to a long dry season.

Vern. Chocolate lily, E (Australia).

Note. PAYENS (1957) stated that the perianth
segments are connate for 1 mm. This was not con-
firmed by me for material he had examined.

Fig. 7. Arthropodium strictum R. Br. a. Habit,

x 1/,, b. flower, x 2, c. withered flower, x 3,

d. gynoecium, X 7, e. anther, x 10(C. R. ALcocK
2875, S. Australia).

7. CAESIA

R. Br. Prod. (1810) 277; BAKER, J. Linn. Soc. Bot. 15 (1876) 357; Bru. Fl. Austr. 7
(1878) 46; BAKER, Fl. Cap. 6 (1897) 400; BaImLey, Queensl. Fl. 5 (1902) 1632;
Ewart, FI. Vict. (1930) 289; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930)
288; BLACK, FI. S. Austr. | (1943) 192; PHiLiips, Gen. S. Afr. Fl. Pl. ed. 2 (1951)

204 FLORA MALESIANA [ser. I, vol. 9!

1002; PAYENS, Nova Guinea n.s. 8 (1957) 383; SCHLITTLER, Mitt. Bot. Mus. Un.
Ziirich 207 (1957) 6, map; HuTCH. Fam. FI. Pl. ed. 2, 2 (1959) 596; OBERMEYER,
Bothalia 11 (1973) 122. — Fig. 8.

Stemless herbs. Rhizome very short. Roots fibrous, fleshy or tuberous. Leaves
basal, subulate to linear, expanded at the base to form a sheath. Inflorescence a
raceme or more often a panicle. Pedice/s |—3-nate, in the axils of bracts and usually
associated with a few smaller bracts also apparently in the axil of the principal
bract, articulated usually in the distal half. Perianth segments free or shortly con-
nate, subequal, 3-veined, spreading. Fi/aments filiform or linear, glabrous, attached
to the receptacle or to the base of the perianth segments; anthers basifixed, oblong,
dehiscing introrsely. Ovary superior, sessile, globose or subglobose, 3-celled;
ovules axile, 2 in each locule; style filiform, simple, minutely capitate. Capsule
dehiscing loculicidally; perianth segments adhering, marcescent, twisting spirally
after flowering. Seeds angular or globose, usually solitary in each locule.

Distr. Madagascar (1 sp.), South Africa (2 spp.; Cape Province) and Australia (7-9 spp.), of which one
species also occurs in Malesia: S. Papua New Guinea (W. Distr.: Wassi Kussa area).

Ecol. Usually in the open, but also in savanna.

Note. Generically this group is regarded as well-defined by the spirally-twisting perianth and 2-ovuled

locules. The South African monotypic genus Nanolirion BTH. (1883) was formerly distinguished by having
a 1-3-flowered inflorescence (PHILLIPS, 1951), but OBERMEYER (1973) placed it under Caesia.

1. Caesia setifera BAKER, J. Linn. Soc. Bot. 15
(1876) 359; Bru. FI. Austr. 7 (1878) 47; Ewart &
Davies, Fl. North. Terr. (1917) 71; PAYENs, Nova
Guinea n.s. 8 (1957) 384, f. 1. — Fig. 8.

Roots bearing distant spindle-shaped tubers
c. 1-1'/, cm long. Leaves 2-4, suberect, filiform,
glabrous, c. 10-25 cm long, c. 1 mm broad; veins of
leaves forming fibrous sheaths at base of plant.
Peduncle thin and wiry, usually with 1 or 2 ascend-
ing branches, (17—)25-45 cm high. Bracts bearing
branches or pedicels in their axils, rarely up to Smm
long, ovate to lanceolate, partly scarious. Pedicels
1-6-nate, usually erect-spreading, (3—)5-10 mm.
Perianth segments blue, linear-oblong, 6-8 by
1-1!/, mm. Filaments 3-4 mm (the outer up to 1 mm

longer than the inner); anthers yellow, c. 3/,-
3/,mm long. Ovary c. 3/,-1 mm long; style 4-5 mm.
Capsule subglobose, deeply 3-lobed, c. 3 mm @.

Distr. Australia (Queensland and Northern
Territory); in Malesia: S. Papua New Guinea:
Western District, Wassi Kussa area: Morehead,
Weam, Arufi, Tarara.

Ecol. In the Wassi Kussa area scattered on open,
grass-sedge plains on thin sand over clay, in
maintained savanna grassland, in savanna with
Melaleuca and Acacia, on wet flats in savanna
forest, on sour grey soils, at very low altitudes sub-
ject to a long dry season. F/. July-Aug., Dec.—Jan.

A field note recorded that usually one flower is
Opening in sequence.

Fig. 8. Caesia setifera BAKER. a. Flower after anthesis, with persistent twisted perianth, b. sepal inside,
c. stamen, all x 6, d-e. anther, dorsal and lateral, x 15 (a Brass 8599, b-e BRASS 8560).

1979] LILIACEAE—I (Jessop) 205

8. CHLOROPHYTUM

Ker-GAWL. Bot. Mag. 27 (1808) t. 1071; BAKER, J. Linn. Soc. Bot. 15 (1876) 321;
KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 284; POELLN. Ber. Deut. Bot.
Ges. 61 (1943) 126; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 598; Dress, Baileya 9
(1961) 29; OBERMEYER, Bothalia 7 (1962) 690; BACK. & BAKH. f. Fl. Java 3 (1968)
86; CHAROENPHOL, Thai For. Bull. 7 (1973) 67; PANIGRAHI, Kew Bull. 30 (1975)
563; MARAIS & REILLY, Kew Bull. 32 (1978) 653.

Perennial stemless herbs. Rhizome horizontal, often very short. Roots fibrous, or
fleshy or tuberous. Leaves basal, usually rosulate, linear to lanceolate, often with a
fimbriate margin, expanding towards the base to form a sheath. Inflorescence a
raceme or a panicle. Pedice/s 1-6-nate, in the axils of bracts and associated with a
few small bracts also apparently in the axil of the principal bract, articulated
usually near or below the middle. Perianth segments free, subequal, with 3 or 5
veins, most frequently spreading or reflexed. Filaments filiform, glabrous or
papillate, attached to the receptacle; anthers basifixed, linear-oblong, introrse.
Ovary superior, sessile or shortly stipitate, globose or obovoid, 3-lobed, 3-celled;
ovules axile, 2 or more in each locule; style filiform, simple, minutely capitate.
Capsule dehiscing loculicidally; perianth segments adhering, marcescent, not
twisting after flowering. Seeds flat, suborbicular.

Distr. Especially in Africa (mainly southern and tropical), Madagascar and Asia (especially India)
to Australia (2 spp.); in West Malesia two non-endemic species.

The total number of species is estimated by KRAuSE (1930) at c. 100 and by OBERMEYER (1962) at
nearly 300.

Ecol. Species of Chlorophytum occur in a wide range of habitats from coastal to montane regions.
They grow in many soil types and in rock crevices, and in open grassland and dense forest.

Notes. OBERMEYER (1962) recorded that in species with fascicled flowers plants do occasionally occur
in which the flowers are borne spirally on a distinct but very short lateral branch of the inflorescence.
This supports the theory that fascicled flowers and their associated bracts represent an abbreviated lateral
shoot.

She also discussed the difficulty encountered in separating Chlorophytum from Anthericum L. (1753).
She was only able to find a single character on which these genera could always be separated: the seeds of
Chlorophytum are flat, but of Anthericum angular and smaller. Several other characters were found to be
generally, but not universally, of value in separating them.

KEY TO THE SPECIES

1. Perianth segments 3-5 mm long. Anthers less than 1 mm long. Leaves usually 4-8 mm broad
1. C. laxum
1. Perianth segments 7-12 mm long. Anthers 4-5 mm long. Leaves usually 10-50 mm broad
2. C. malayense

1. Chlorophytum laxum R. Br. Prod. (1810) 277; Roots often bearing tubers c. 1-5 cm long (absent
Hook. f. Fl. Br. Ind. 6 (1892) 336; BAcK. Handb. in material I have seen from Malesia and stated by
Fl. Java 3 (1924) 52, incl. f. javanicum (HassK.) BACKER & BAKHUIZEN VAN DEN BRINK f. to be
Back.; GAGNEP. FI. Gén. I.-C. 6 (1934) 804; BAcK. absent in Javanese plants). Leaves 4-12 or rarely
& BAKH. f. Fl. Java 3 (1968) 86; HutcH. & Daz. more, linear to lanceolate, usually channelled
Fl. W. Trop. Afr. ed. 2, 3 (1968) 100. — C. laxi- especially towards the base, with a rather promi-
florum Baker, J. Linn. Soc. Bot. 15 (1876) 328, nent midrib; suberect, glabrous, 10-30(-50) cm by
nom. illeg. — Nolina javanica Hassk. Tijd. Nat. 4-8(-12) mm, rarely forming fibrous sheaths
Gesch. Phys. 10 (1843) 120; Mia. Fl. Ind. Bat. 3 round the base of the plants; the outer arcuate and
(1859) 554. recurved; the inner often straight. Peduncle straight

206

FLORA MALESIANA

[ser. I, vol. 9!

or flexuous, unbranched or rarely with a single
branch, (5—)15—35(—60) cm. Lower bracts 3-12 mm
long, usually lanceolate-acuminate, partly scarious.
Pedicels solitary or, less often, 2-nate, often erect-
spreading but recurving in fruit, 3-12 mm.
Perianth segments white or ‘greenish white’, linear-
oblong, 3-5 by c. 1 mm. Filaments (1'/,-)2-3
(-4) mm long (the outer often longer than the
inner); anthers up to !/, mm long. Ovary c. 1 mm
long, obovoid or globose; style less than 2 mm;
ovules 2 per locule. Capsule usually obovoid, less
often globose or depressed-globose, 3-lobed,
5-10 mm long.

Distr. Tropical Africa (e.g. Senegal, Liberia,
Ghana, Nigeria, Zambia, Ethiopia), S. Asia (e.g.
India, Indo-China, Thailand), China (incl.
Hainan); in Malesia: N. Sumatra (Eastcoast Res.),
Malay Peninsula, W. and Central Java, and SE.
Borneo (Bandjermasin), extending to N. Australia
(Queensland and Northern Territory).

Ecol. Recorded from a variety of localities
including rock crevices and in sandy soils, but
usually a species of shady places (including bamboo
forest, deciduous forest and a coconut grove),
usually below 1000 m. F/. Jan.—Dec.

Notes. BACKER (1924) and BACKER & BAK-
HUIZEN VAN DEN BRINK /. (1968) treat the Javanese
plants as belonging to forma javanicum (HASSK.)
BACK. They give no reason for this decision and in
the absence of a monograph of this species I would
prefer not to uphold this form.

Extra-Malesian synonyms are omitted from the
references.

2. Chlorophytum malayense RIDL. Fl. Mal. Pen. 5
(1925) 341; PANIGRAHI, Kew Bull. 30 (1975) 565. —
C. orchidastrum (non LINDL. Bot. Reg. 10 (1824)
t. 813) RrpL. Mat. Fl. Mal. Pen. Monoc. 2 (1907)
92; Fl. Mal. Pen. 4 (1924) 327; GaGnep. Fl. Gén.
I.-C. 6 (1934) 806; CHAROENPHOL, Thai For. Bull. 7
(1973) 67.

Roots sometimes bearing tubers 3-4 mm or more
long. Leaves 3-10, sublinear or indistinctly petioled
and with a lanceolate lamina, channelled towards
the base, with a rather prominent midrib, suberect,
glabrous, (10—)25-60 by (*/,-)1-5(-10) cm, never
forming fibrous sheaths round the base of the plant.
Peduncle glabrous or less often glandular pubes-
cent, with 0-~ branches (30—)40—50(?-90) cm high.
Lower bracts up to 9 cm long, usually narrowly
lanceolate-acuminate, usually partly scarious; the
lowest usually sterile. Pedicels usually 2-3-nate,

less often solitary, erect-spreading, not recurving
in fruit, 3-10 mm. Perianth segments white, elliptic
to linear-oblong, 7-12 by 2-3 mm. Filaments
occasionally minutely pubescent, 4-5 mm (not
always similar in length but neither whorl regularly
longer than the other); anthers 4-5 mm long.
Ovary c. 11/,-2 mm long, obovoid or globose; style
3-4(-8, GAGNEPAIN, 1934) mm; ovules 4-6 per
locule. Capsule globose, strongly emarginate and
very deeply 3-lobed, c. 5-8 mm @.

Distr. Indo-China, Thailand and Malesia:
Malay Peninsula (Perak and Perlis).

Ecol. This is usually a forest (including bamboo,
oak, pine) species, often associated with limestone,
50-1500 m. F/. Jan.—Dec.

Notes. GARRETT (in sched.) recorded that the
plant is night-flowering in Thailand.

RIDLEY (1925) gave the new name C. malayense
to what he had previously identified as C. orchid-
astrum LINDL. However, he gave no characters by
which these species could be separated.

In the neighbouring territories of Thailand and
Indo-China CHAROENPHOL (1973) and GAGNEPAIN
(1934) have continued to recognize C. orchidastrum,
but Iam following PANIGRAHI (1975) who considers
that C. orchidastrum sensu stricto is confined to
Africa, that the Indian material should be referred
to C. nimmonii and C. glaucum, and that the SE.
Asian material is C. malayense. Our species can,
according to PANIGRAHI, be recognized inter alia by
drying greenish brown or glaucous, in the leaves
not being distinctly petioled and 3-5 cm broad, in
the scape being up to 50 cm long and shorter than
the leaves and in the bracts being up to 9 cm long.
Few collections have been made of C. malayense
and further field work is needed toconfirm its status.

C. longissimum Ripv. (J. Str. Br. R. As. Soc.
n. 49, 1907, 209) was described from Trang,
Peninsular Thailand, close to the Malesian border.
It closely resembles C. malayense. CHAROENPHOL
distinguished these species on whether the inflores-
cence is erect and sometimes branched (C. malay-
ense) or trailing on the ground and never branched
(C. longissimum). The type of C. longissimum has
not been seen, but specimens at Kew identified as
such, and agreeing with the type description, are
possibly sufficiently characterized by these inflores-
cence characters to retain it as a distinct taxon.
Whether the differences are sufficient for specific
rank must be left to future examination; it is
provisionally kept distinct. It was not discussed
by PANIGRAHI.

9. DIANELLA

LAMK [Encycl. 2 (1786) 276, nom. inval.] ex Juss. Gen. Pl. (1789) 47; Baker, J.
Linn. Soc. Bot. 14 (1875) 574; Bru. Fl. Austr. 7 (1878) 13; BAck. Handb. FI.
Java 3 (1924) 53; Krause in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930)

1979} LILIACEAE—I (Jessop) 207

295; Se Mitt. Bot. Mus. Un. Ziirich 163 (1940) 256; Blumea 6 (1948) 200;
Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 606. — Rhuacophila BL. En. Pl. Jav. (1827)
13. — Fig. 9.

Erect or climbing glabrous herbs. Rhizome short to stoloniferous. Roots fibrous.
Leaves usually cauline and distichous or basally rosulate, linear, distally dorsiven-
tral, in the lower parts sometimes laterally compressed and keeled and often form-
ing a closed sheath at the base. Inflorescence a panicle. Pedicels solitary or few,
usually in the axils of bracts, articulated immediately below the flower. Perianth
segments free, 3-7-nerved, spreading or recurved, equal or subequal. Filaments
filiform or linear, often swollen in the distal half, glabrous, attached to the recep-
tacle or the inner whorl attached to the perianth segments; anthers basifixed,
linear to oblong, dehiscing by an apical pore or by a slit which is initiated in an
apical pore. Ovary superior, more or less sessile, globose, 3-celled; ovules axile,
4-8 in each locule; style filiform, simple, minutely capitate. Fruit a berry, usually
shiny blue; perianth segments adhering, marcescent, not twisting after flowering;
base of style persistent. Seeds globose or angled.

Distr. Continental Africa (one record), Madagascar, through India into China, throughout Malesia
and Australia to New Zealand and the Pacific Islands (New Caledonia, Sandwich Is., Norfolk and Fiji).

Estimates of the number of species vary widely; there are possibly 20-30 spp. in all.

Ecol. Both in forest and in more open localities.

KEY TO THE SPECIES

1. Leaf bases strongly compressed and keeled. Leaves usually with minute teeth or prickles along the
margin and abaxial surface of the midrib. Fertile bracts differing considerably in size from even the

smaller leaves. Filaments strongly thickened at the apex under the anther. .. . .

1. D. ensifolia

1. Leaf bases obtuse in section, not keeled. Leaves always smooth. Fertile bracts grading into the leaves.

Filaments widened about the middle

1. Dianella ensifolia (L.) DC. in Redouté, Liliac. 1
(1802) t. 1, cf. Coops in Bosser c.s. Fl. Mascar. Lil.
(1978) 32; Mia. FI. Ind. Bat. 3 (1859) 560; HEMsL.
Rep. Chall. Bot. 1, 3 (1884) 201; RipL. Mat. FI.
Mal. Pen. Monoc. 2 (1907) 92; Laut. Bot. Jahrb.
50 (1913) 293; HALL. f. Nova Guinea 8 (1914) 995;
Merr. En. Born. (1921) 114; En. Philip. 1 (1922)
203; Ripi. Fl. Mal. Pen. 4 (1924) 329; KRAUSE,
Bot. Jahrb. 59 (1925) 551; MeRR. Contr. Arn. Arb.
8 (1934) 18; Wi_tp, Kew Bull. 8 (1953) 251;
M. R. HENDERSON, Mal. Wild Flow. Monoc.
(1954) 186, f. 110; Back. & Baku. f. Fl. Java 3
(1968) 87; STEEN. Mt. FI. Java (1972) t. 28-1. —
Gladiolus odoratus indicus RuMPH. Herb. Amb. 5
(1747) 185, t. 73. — Dracaena ensifolia LINNE,
Mant. (1767) 63. — Dianella nemorosa LAMK,
Encycl. 2 (1786) 276, nom. inval.; Tabl. Enc. 2
(1792) 388, t. 250, nom. illeg.; SCHLITTLER, Mitt.
Bot. Mus. Un. Zirich 163 (1940) 256; Blumea 6
(1948) 209, incl. many forms, /.c. 216-223. —
Dracaena ensata THUNB. Diss. Bot. Drac. (1808) 4.
— Dianella montana BL. En. Pl. Jav. 1 (1827) 12;
Hassk. Tijd. Nat. Gesch. Phys. 11 (1844) 180; Pl.
Jav. Rar. (1848) 114; Mia. Fl. Ind. Bat. 3 (1859)
560; Back. Handb. Fl. Java 3 (1924) 54. —

2. D. javanica

D. odorata [RUMPH.] BL. En. PI. Jav. 1 (1827) 13;
HAL. f. Nova Guinea 8 (1914) 996; Merr. Int.
Rumph. (1917) 136; Krause, Bot. Jahrb. 59 (1925)
550; SCHLITTLER, Mitt. Bot. Mus. Un. Zurich 163
(1940) 258. — D. revoluta (non R. BR.) SCHAUER,
Nov. Act. Ac. Nat. Cur. 19 (1843) Suppl. 1: 445;
Laut. Bot. Jahrb. 50 (1913) 293; MerR. En.
Philip. 1 (1922) 203. — D. bancana Mia. FI. Ind.
Bat. Suppl. (1861) 610; BAck. Handb. Fl. Java 3
(1924) 54. — D. caerulea (non Sims, Bot. Mag. 15,
1801, t. 505) Merr. Philip. J. Sc. 2 (1907) Bot. 266;
ibid. 5 (1910) Bot. 337; HALL. f. Nova Guinea 8
(1914) 993; Gress, Arfak (1917) 100; Merr. En.
Philip. 1 (1922) 202; KrAusE, Bot. Jahrb. 59 (1925)
552; HoLtuuts, Blumea 5 (1942) 167. — D. robusta
Evmer, Leafl. Philip. Bot. 5 (1913) 806. — D. bam-
busifolia HALL. f. Nova Guinea 8 (1914) 995, t. 182;
Krause, Bot. Jahrb. 59 (1925) 550. — D. flabellata
Hatt. f. lic. 997, t. 183; Krause, Ic. 549. —
D. carinata HALL. f. l.c. 999, t. 186; KRAUSE, /.c.
550. — D. parviflora Zipp. ex HALL. f. l.c. 998,
t. 184; Krause, /.c. 551. — C. albiflora HALL. f.
l.c. 998, t. 185; Krause, /.c. 551. — D. monophylla
HALL. f. lc. 1000, t. 188; Krause, /.c. 551. —
D. serrulata HAL. f. /.c. 1000, t. 187; KRAUSE,

FLORA MALESIANA

Hy
&
. ¥
a
ee
Te

[ser. I, vol. 9!

Fig. 9. Dianella javanica (BL.) KUNTH, on Mt Kaba, S. Sumatra (Photogr. DE Voocp).

lic. 549. — D. parviflora Riv. J. Fed. Mal. St.
Mus. 6 (1915) 186. — D. pullei KRAUSE, Nova
Guinea 14 (1924) 175; Bot. Jahrb. 59 (1925) 553. —
D. ledermannii Krause, Bot. Jahrb. 59 (1925) 549.
— D. monticola Krause, I.c. 553. — ? D. levis (non
R. Br.) C. T. Wuirte, Proc. Linn. Soc. N.S.W. 51
(1926) 298. — D. sparsiflora SCHLITTLER, Mitt. Bot.
Mus. Un. Zurich 163 (1940) 262. — D. ensata
(THUNB.) R. J. HENDERSON, Taxon 26 (1977) 136.

Stem 0-1 m high, unbranched, rarely with a few
branches. Rhizome horizontal, moderately
branched. Leaves basal, scattered along the stem or
in a terminal rosette, distichous, with a sheathing
lower part, (25—)30—60(-100) cm by (4-)8-30 mm;
above the base keeled and with the sides of the
lamina becoming firmly appressed to one another
and fused to form an isobilateral portion; distally
with a dorsiventral linear or linear-lanceolate

1979]

LILIACEAE—I (Jessop)

209

lamina which is sometimes absent from the lower
leaves, almost always with minute serrations or
prickles along the midrib on the lower surface,
with a conspicuous midrib and numerous smaller
veins. Inflorescence exceeding the leaves, lax or
with short terminal branches often c. 1-2 cm long,
bearing up to 20 pedicels. Lower bracts usually
narrowly linear-lanceolate and bilaterally com-
pressed above the basal sheath like the leaves;
bracts subtending pedicels 1-4(-7) mm long or
rarely absent. Pedicels 4-15(-22) mm. Perianth
segments blue, white, lilac or yellow, spreading,
(4-)5—8(-9) mm long. Filaments often more than
half as long as the perianth segments, filiform or
narrowly linear, white or yellow with a yellow or
orange, glabrous swelling below the anther. Ovary
green, 11/.-2 mm long; ovules 4 in each locule;
style green, white or blue, 4-6 mm long. Fruit
shiny blue, 6—8(—11) mm @. Seeds several, 3-4 mm
long.

Distr. Continental Africa (WiLp, 1953),
Madagascar, continental Asia (Himalayas, Burma,
Thailand, Indo-China) to southern China (Yunnan,
Hainan), Japan and Formosa, throughout Malesia
to Australia (Northern Territory, Queensland, New
South Wales), Tasmania, New Zealand and many
Pacific islands.

Ecol. A highly adaptable species, occurring in
habitats ranging from open grassland to primary
forest, from sea-level to over 3000 m. Fl. Jan.—

Dec.
Vern. Malaya: benjuang, satagit, senjuang,
siak-siak jantan; Sumatra: (akar) tu(n)daun,

mentuntil, tengari, ukop, Banka, siak-siak, Riouw,
sieuh, Djambi, sitanggit, Batak, sesiah, Pasemah,
sitangie, Indragiri; Java: djamaka, d. putih, suliga,
S, tegari, J; Borneo: labeh-labeh, Dusun Penam-
pang dial., angkup-angkup, Bokan dial., tembalong
tipoh, Dusun dial.; Philippines: ablaas, Bag., bariu-
bariu, oyon-dyon, P.Bis., dufigau, Ig., hogangan,
If.; Moluccas: mariuii, Talaud; New Guinea:
suruma; bururl, tirambaramp, Mendi, buru-buru,
Biak dial., moalengen, Aiome, tanglenu, Wigote,
Wapi lang., sinda, Mumuni, Orokaiva lang., tsiri
kande kande, Hagen Subdistr., bonkaige, Sinasina
lang., Nimai dial., pfifiriki, Kutubu lang., kili-kili,
Bembi, kikipatia, Koroba Subdistr., sabetari,
Rawa, kilina, Kaigorin, sarpeim, Miwaute, Wapi
lang., pengeh-pengeh, Maipa, Mekeo lang.,
baibigehgi, Utukap, Miniafi lang.

Notes. BACKER (1924, 1968) recognized two
species, D. montana and D. bancana, to include the
material from Java, which he distinguished mainly
on flower colour, venation of petals, and inflores-
cence form. However, the large number of inter-
mediates makes this separation impracticable.

SCHLITTLER (1940) recognized 3 spp. in Malesia
(D. odorata, D. nemorosa and D. sparsiflora), but
in 1948 he reduced these to a single one under the
name D. nemorosa, with 24 forms in Malesia.

Being a common species over a large area,

D. ensifolia is exceptionally well represented in
herbaria and, despite its variability, I doubt that
even with intensive field work distinct subspecific
taxa can be defined.

2. Dianella javanica (BL.) KUNTH, En. Pl. 5 (1850)
52; Mia. FI. Ind. Bat. 3 (1859) 561; HALL. f. Nova
Guinea 8 (1914) 995; Merr. En. Born. (1921) 114;
En. Philip. 1 (1922) 203; Ripv. Fl. Mal. Pen. 4
(1924) 328; Back. Handb. FI. Java 3 (1924) 53;
Krause, Bot. Jahrb. 59 (1925) 549; Merr. Contr.
Arn. Arb. 8 (1934) 19; ScHLITTLER, Blumea 6
(1948) 206, incl. f. stenophylla, alba et rubra
SCHLITTLER, /.c. 208; Mitt. Bot. Mus. Un. Ziirich
207 (1957) 6, 11, map, 25; BAck. & BAKH. f. FI.
Java 3 (1968) 87; STEEN. Mt. Fl. Java (1972) t. 28-2.
— Rhuacophila javanica BL. En. P|. Jav. (1827) 14;
Rip. J. Linn. Soc. Bot. 42 (1914) 166; J. Fed. Mal.
St. Mus. 6 (1915) 186. — Rhuacophila celebica
BL. En. Pl. Jav. (1827) 14. — Eustrephus javanicus
(BL.) D. Dretr. Syn. Pl. 2 (1840) 1117. — Eustre-
phus celebicus (BL.) D. Dietr. /.c. — D. celebica
(BL.) KUNTH, En. Pl. 5 (1850) 45; Mia. Fl. Ind.
Bat. 3 (1859) 561. — D. austro-caledonica SEEM.
FI. Vit. (1868) 312; Laur. Bot. Jahrb. 50 (1913) 294.
— Fig. 9.

Stem always present, up to 2 m high, unbranched
or branched. Rhizome ‘horizontal, moderately
branched. Leaves concentrated towards the ends
of the branches, distichous, 12—35(-40) by 3/,-
2!/, cm, with a sheathing but not closed base which
is continuous with the lamina, lacking a compressed
and fused intermediate portion, lacking serrations
or prickles; midrib usually more conspicuous than
the many other veins. Inflorescence lax, exceeding
the leaves. Bracts subtending branches of the
inflorescences grading gradually in size into the
foliage leaves, up to 25 by 3 cm, ovate to lanceolate
not bilaterally compressed; bracts subtending the
pedicels '/,-3(-5) mm long or absent. Pedicels
6-20 mm. Perianth segments blue (usually pale),
white to yellow or lilac, spreading, (6—)8-12 mm
long; outer whorl strongly cucullate at the apex.
Filaments usually less than half as long as the
perianth segments, white or yellow, filiform, usually
with a distinct papillose fusiform swelling about
the middle. Ovary green, c. 2 mm long; ovules
c. 8-10 in each locule; style white or green 2-5 mm.
Fruit green or yellow at first, sometimes (at least)
becoming black, ellipsoid, 8-15 mm long, with up
to 10 seeds. Seeds c. 1'/, mm @.

Distr. Throughout Malesia, also in New Cale-
donia and Ile des Pins, and Fiji Is. (Viti Levu).

Ecol. Grows in a wide variety of habitats,
including mossy forests, thickets, forest borders, on
narrow open ridge-crests, in debris of craters,
among rocks, on stream banks and in exposed
places, locally common, (1000—)1500-3000 m. FI.
Jan.—Dec.

Vern. Java: hadjéra, S; Philippines: kallawad,
If., apilug, sapilau, urdya, \g., talobatub, Bon.

210 FLORA MALESIANA [ser. I, vol. 9"

Fig. 10. Thysanotus tuberosus R. BR. a. Habit, x 1/4, b. flower, x 5, c. sexual organs, x 10, d. fruit in
persistent perianth, x 5. — T. chinensis BTH. e. Habit, x '/, (a-d BRASS 6517, e VAN ROYEN & SLEUMER
5632).

211

1979] LILIACEAE—I (Jessop)

10. THYSANOTUS

R. Br. Prod. (1810) 282; BAKER, J. Linn. Soc. Bot. 15 (1876) 334; Bru. Fl. Austr. 7
(1878) 36; Riv. FI. Mal. Pen. 4 (1924) 328; KRAUusE in E. & P. Nat. Pfl. Fam. ed.
2, 15a (1930) 285; Ewart, FI. Vict. (1930) 288; BLAcK, FI. S. Austr. 1 (1943) 190;
PAYENS, Nova Guinea n.s. 8 (1957) 386; HUTCH. Fam. FI. Pl. ed. 2, 2 (1959) 598,
nom. gen. cons. — Chlamysporum SALIsB. Parad. Lond. (1808) t. 103. — Halongia
JEANPLONG, Act. Bot. Ac. Sc. Hung. 16 (1970) 293, f. 1-6. — Fig. 10.

Stemless herbs. Rhizome horizontal, very short. Roots fibrous or tuberous.
Leaves basal, rosulate, filiform or linear, glabrous, expanding towards the base to
form a sheath. Inflorescence a raceme, panicle or umbellate, erect or twining.
Pedicels 1—7-nate, in the axils of bracts and associated with a few small bracts also
apparently in the axil of the principal bract, or umbellate, articulated often in the
median third. Perianth segments free, 3—S-nerved, spreading; outer whorl green;
inner whorl usually blue to purple or violet, fringed. Stamens usually 6, rarely 3.
Filaments linear, glabrous, attached to the perianth segments, bent over to one side
of the ovary (BLACK, 1943); 3 often shorter; anthers basifixed, linear, curved de-
hiscing introrsely. Ovary superior, sessile, oblong to globose, 3-celled; ovules axile,
2 in each locule; style filiform, simple, minutely capitate. Capsule dehiscing locu-
licidally; perianth segments adhering, marcescent, twisting after flowering; style
persistent. Seeds angled, with a fleshy white or orange strophiole.

Distr. Up to 30 spp., all in Australia (all states, but especially Western Australia); in Malesia: 2 of
these spp., one in New Guinea and another throughout Malesia as far as Thailand, Indo-China and
southern China.

Ecol. A wide range of mostly open habitats from grasslands and sandy heaths to open forests, including
both Eucalyptus savanna and pine forest.

Notes. The fleshy appendage of the seed has been referred to as a caruncle (e.g. BLACK, 1943), but it
appears to be derived from the funicle rather than from the testa and I am, therefore, using the term
strophiole as done by PAYENS (1957).

The term ‘cluster’ is sometimes used in preference to umbel as in some material the inflorescence appears
umbellate but in other material the pedicels arise from a short but distinct axis.

KEY TO THE SPECIES

1. T. chinensis

1. Roots lacking tubers. Inflorescence a simple umbel :
. 2. T. tuberosus

1. Roots with tubers. Inflorescence a panicle, the branches terminated ‘by umbels

Mal. — Halongia purpurea

1. Thysanotus chinensis BTH. Fl. Hongk. (1861) 372;
BaKER, J. Linn. Soc. Bot. 15 (1876) 337; HALL. f.
Nova Guinea 8 (1914) 994; Merr. En. Philip. 1
(1922) 202; Krause, Bot. Jahrb. 59 (1925) 548;
GARDN. En. Pl. Austr. Occid. (1931) 18; ScHLIT-
TLER, Mitt. Bot. Mus. Un. Ziirich 207 (1957) 6,
map; PAYENS, Nova Guinea n.s. 8 (1957) 386;
STEEN. Blumea 20 (1972) 433. — T. chrysantherus
F.v.M. [in Bru. Fl. Hongk. (1861) 372, nomen]
Fragm. 5 (1866) 202; Bru. Fl. Austr. 7 (1878) 40;
Naves, Nov. App. (1880) 266; ViDAL, Phan.
Cuming. (1885) 153; BAILey, Queens]. Fl. 5 (1902)
1629. — Chlamysporum chrysantherum (F.v.M.)
O. K. Rev. Gen. Pl. 2 (1891) 708. — T. siamensis
RD. J. Str. Br. R. As. Soc. 2. 59 (1911) 209; FI.

Pen. 4 (1924) 328.
JEANPLONG, Act. Bot. Ac. Sc. Hung. 16 (1970) 296,
f. 1-6. — Fig. 10e.

Roots fibrous. Leaves c. 5 to numerous, erect,
often shallowly channelled, usually 15-40 cm, up
to c. 1 mm wide. Peduncle about as long as the
leaves, straight, unbranched, terminating in a
2-6(-12)-flowered cluster. Bracts with scarious
margins, 2-4 mm long. Pedicels 1-2 cm, articulated
in the basal third, erect in bud, usually spreading
or recurving in fruit. Outer perianth segments
(? always) green, with a scarious, white margin,
6-9 by 2-3 mm; inner segments pale blue to purple
or light violet, with inturned usually fringed
margins, 6-9 by 3-5 mm. Filaments 1'/.-2 mm,

212 FLORA MALESIANA fser. I, vol. 9!

S

Fig. 11. Tricoryne platyptera Rcus. a. Habit, x */4, b. flower, x 2, c. withered flower, < 4, d. stamen,
x 20, e. fruit, x 5 (NGF 38690, PULLEN 7090).

1979]

LILIACEAE—I (Jessop)

213

the outer whorl shorter than the inner; anthers
11/,-3 mm long, the outer shorter than the inner.
Ovary c. 1 mm long; style c. 3-4 mm. Capsule
oblong, 4-5 mm long. Seeds c. 1/, mm long.

Distr. Australia (Western Australia and Nor-
thern Territory); in Malesia: New Guinea (West
New Guinea and Papua), SE. Moluccas (Aru Is.:
Tranggan), Celebes (Masamba), Philippines
(Luzon, Mindanao), Lesser Sunda Is. (Flores),
onto continental SE. Asia: S. Peninsular Thailand
(Setul), Indo-China (Tonkin), S. China (incl.
Hong Kong).

Ecol. Open grassland, in grassy pine forest,
most frequently in open savanna, in New Guinea
of Melaleuca, etc., in places subject to a moderate
to strong dry season, from close to sea-level up to
1600 m. F/. Dec.—Aug.

2. Thysanotus tuberosus R. Br. Prod. (1810) 282;
BAKER, J. Linn. Soc. Bot. 15 (1876) 335; Bru. FI.
Austr. 7 (1878) 41; Ewart, Fl. Vict. (1930) 289;
GARDN. En. Pl. Austr. Occid. (1931) 18; Brass, J.
Arn. Arb. 19 (1938) 190; BLack, FI. S. Austr. 1
(1943) 191; PAYENs, Nova Guinea n.s. 8 (1957)
387; Burs. & Gray, FI. A.C.T. (1970) 102. —
Chlamysporum tuberosum (R. Br.) O. K. Rev.
Gen. PI. (1891) 708. — Fig. 10a-d.

Roots partly fibrous, bearing spindle-shaped
distant tubers 10-25 mm long. Leaves 2-6, erect,
linear to terete, 20-50 cm by up to c. 1 mm broad.
Peduncle about as long as the leaves or up to
50% longer, usually with 1-6 branches; main axis

and branches each terminating in a 2-6(—20)-
flowered cluster; 1—3(—8) flowered fascicles of
flowers usually also present in the axils of bracts
along the main axis and branches. Bracts usually
with scarious margins, the lowest 3—60(—80) mm
long; the upper shorter. Pedicels 7-17 mm, articula-
tion usually below the centre, erect to spreading.
Outer perianth segments purple with a scarious pale
margin, 7-14 by 2-3 mm; inner segments purple
with inturning fringed margin, 7-15 by 21/,-5 mm.
Filaments 1—2 mm, those of the outer whorl often
shorter than the inner; anthers 2-4 mm long; outer
often shorter than the inner. Ovary 1-1'/, mm long;
style (2-)3—5 mm. Capsule oblong, c. 5-6 mm long.
Seeds up to 2 mm long.

Distr. Australia (all states except Tasmania)
and S. Papua New Guinea (Western District:
Wassi Kussa; Mabaduan).

Ecol. Common grass associate in lowland
savanna forest on poorly drained flats, subject toa
distinct dry season. F/. Dec.—April. In Australia
found in exposed localities up to 500 m.

Notes. VAN ROyYEN recorded in Queensland
that the outside of the flowers was white with a
green midrib and the anthers dark purplish green.

PAYENS (1957) recognized two varieties separable
on flower size. Both occur in Australia but only
var. parviflora BTH. in Papua. The material avail-
able is insufficient to justify recognition of these
varieties.

Six other synonyms based on extra-Malesian
material are omitted from the synonymy.

11. TRICORYNE

R. Br. Prod. (1810) 278; BAKER, J. Linn. Soc. Bot. 15 (1876) 361; Bru. Fl. Austr. 7
(1878) 50; BAILEY, Queens]. Fl. 5 (1902) 1636; Ewart, FI. Vict. (1930) 287;
KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 287; BLAck, FI. S. Austr. 1
(1943) 192; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 602. — Fig. 11.

Erect, glabrous or variously hairy herbs; stems green, terete, angled or flattened
and leaf-like. Rhizome horizontal or erect, often very short; roots thick or fibrous.
Leaves basal and/or cauline, more or less linear or reduced to scales, alternate,
often amplexicaul at the base. Inflorescence of terminal umbels surrounded by
small scarious bracts and outer larger, sometimes leaf-like bracts. Pedicels articu-
lated just below the flower. Perianth segments free or shortly connate, equal or sub-
equal, with 3 or 5 (less often 7) prominent veins, spreading. Filaments filiform, with
a dense tuft of hairs in the distal part, attached to the receptacle; anthers basifixed,
linear to oblong; introrse. Ovary superior, sessile, deeply 3-lobed, 3-celled; ovules
basal, 2 in each locule; style filiform, simple, minutely capitate. Fruit consisting of
(1-)3 indehiscent nutlets; perianth segments adhering, marcescent, twisting spirally
after flowering. Seeds subglobose.

Distr. In Australia (all states) 6 spp., of which one in Malesia: S. Papua New Guinea.

Ecol. Low altitude plant. BAILEY recorded most of the Queensland species from ‘sandy shores’ and
specimen annotations suggest a preference for sandy soils in grassland or savanna.

214

FLORA MALESIANA

[ser. I, vol. 9

Note. The terms ‘stem’ and ‘leaves’ give some difficulty as the aerial shoot might be better regarded as
an inflorescence often bearing leaf-like bracts rather than as a vegetative shoot with leaves.

1. Tricoryne platyptera Rcup. Syst. Pflanzenk.
(1871) 72; Bru. Fl. Austr. 7 (1878) 51; BAILEy,
Queens]. FI. 5 (1902) 1636; Compr. Cat. Queens].
Pl. (1913) 559, f. 539; Brass, J. Arn. Arb. 19 (1938)
190; PAYENS, Nova Guinea n.s. 8 (1957) 385. —
T. pterocaulon BAKER, J. Linn. Soc. Bot. 15 (1876)
363. — Fig. 11.

Virgate, erect or subscandent herb. Stems and
branches strongly flattened (winged), terete towards
the base of the plant, glabrous, 25-80 cm high;
flattened portions with a prominent midrib,
c. 2-8 mm broad. Rhizome short; roots rather thick.
Leaves cauline, narrowly triangular, scale-like, up
to 12(-25) mm long. Bracts 14 mm long. Pedicels

Perianth segments yellow, oblong or elliptic-
oblong, with a scarious margin, 6—8(—10) by up to
c. 3 mm; those of the outer whorl usually with
(3-)5(—7) veins; those of the inner whorl slightly
narrower, with 3 veins. Filaments 2'/,-3 mm,
yellow; anthers c. 1-11/, mm long. Ovary c.+/, mm
long; style 3!/,-4 mm. Nutlets green.

Distr. Australia (tropical Queensland) and
Malesia: S. Papua New Guinea (Western District:.
Wassi Kussa area), Thursday I.

Ecol. In New Guinea in savanna with Melaleuca
and Acacia, in lowland savanna-woodland on sour
grey soil and in grass of creek flats, subject to a
strong dry season. F/. Dec.—Jan., July-Aug.

usually 5-12(-15) in number, (2-)4-12 mm. In N. Queensland found up to 950 m.

12. LILIUM

LINNE, Sp. Pl. (1753) 302; Gen. Pl. ed. 5 (1754) 143; Baker, J. Linn. Soc. Bot. 14
(1875) 225; ELtwes, Monogr. genus Lilium (1880); Witson, Lilies of E. Asia
(1925); KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 329; DRYSDALE
Woopcock & STEARN, Lilies of the World (1950); SEALY, Kew Bull. 5 (1950) 273;
Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 609.

Erect, usually glabrous herbs. Bulb scaly; roots thick, produced from below the
bulb and in some species from the base of the aerial stem. Aerial stem annual,
erect, usually unbranched. Leaves cauline, alternate or verticillate, linear or
broadly flattened, usually sessile, sometimes with bulbils in their axils. Flowers
solitary and terminal, or in a racemose inflorescence of solitary flowers in the axils
of the often leaf-like bracts. Pedicels erect to cernuous, not articulated. Perianth
segments free, + equal, infundibuliform or campanulate, sometimes clawed,
sometimes adhering and forming a tube, spreading or recurving distally, with a
nectariferous longitudinal furrow towards the base. Filaments filiform or subulate,
attached to the receptacle or to the base of the segments; anthers dorsifixed, linear,
introrse, versatile. Ovary superior, sessile, oblong to obovoid, 3-celled; ovules
axile, co; style terete sometimes clavate, 3-lobed. Capsule coriaceous, loculicidal,
perianth deciduous. Seeds flat.

Distr. Widespread with possibly 80 spp. in the temperate regions of North America, Europe and Asia
(also Taiwan); in Malesia: Philippines.

Ecol. The species occupy a wide range of habitats, including open areas and woods.
Note. Many species are of horticultural importance.

KEY TO THE SPECIES

1. L. longiflorum
2. L. philippinense

1. Leaves 6-15 mm broad. Nectariferous furrow on perianth segments glabrous
1. Leaves 2-4 mm broad. Nectariferous furrow on perianth segments papillose

1. Lilium longiflorum THunB. Trans. Linn. Soc. 2
(1794) 333; ELwes, Monogr. genus Lilium (1880)
t. 7; MeRR. En. Philip. 1 (1922) 204; Back. Handb.
Fl. Java 3 (1924) 62; Witson, Lilies of E. Asia

(1925) 23; DryspALE Woopcock & STEARN,
Lilies of the World (1950) 253; Hatus. Mem. Fac.
Agric. Kagoshima Un. 5 (1966) 62, err. longifolium;
BACK. & BAKH. f. Fl. Java 3 (1968) 90.

1979]

Bulb usually subglobose, often yellowish, up to
6'/, cm @; scales closely imbricate. Stem 30-90
(-100) cm high, smooth, green, producing roots
above the bulb. Leaves alternate, 20—40(—60),
sessile, lanceolate or oblong-lanceolate, attenuate,
with up to 20 or more veins of which up to 7 are
usually more conspicuous, the largest on each plant
up to 15 by 1'/, cm, usually much smaller distally.
Flowers often solitary, but up to at least 7, white,
horizontal, (12-)15—20 cm long, infundibuliform,
the tube (in dried material) 9-12 mm @. Perianth
segments not clawed, reflexed distally; nectari-
ferous furrow glabrous. Filaments filiform above,
linear below, in dried material 9-14 cm; anthers
7-12 mm long; pollen yellow. Style 8-12 cm;
stigma deeply 3-lobed. Capsule c. 4-6 cm long.

Distr. China, Japan and Taiwan; in Malesia:
Philippines: Batan Is. and Y’ami (N of Luzon),
cf. MERRILL and HATUSIMA.

Native country not exactly known, possibly
naturalized over part of its range.

Ecol. Open grassy slopes at low altitude.

Vern. Easter lily, E; teppo-yuri, Japan; Philip-
pines: vonitan, Iv.

Note. Stated by DryspDALE WoopDcock &
STEARN to be commercially the most important
species of Lilium, with numerous cultivated forms.

2. Lilium philippinense BAKER, Gard. Chron. (1873)
1141, f. 243; J. Linn. Soc. Bot. 14 (1875) 228

LILIACEAE—I (Jessop)

pA Ne,

(‘philippense’); Curtis, Bot. Mag. III, 32 (1876)
t. 6250; ELwes, Monogr. genus Lilium (1880) t. 3;
Me_rr. Philip. J. Sc. 5 (1910) Bot. 337; En. Philip.
1 (1922) 204; Witson, Lilies of E. Asia (1925) 20;
DrysDALE Woopcock & STEARN, Lilies of the
World (1950) 311.

Bulb subglobose, whitish, c. 3'/,4 cm @; scales
unknown. Stem 30-90 cm high, smooth, green or
mottled with purple, producing roots above the
bulb. Leaves alternate, 30-40, linear, attenuate,
with up to 7 veins of which 1 or 3 are more con-
spicuous, 8-17 cm by 2-4 mm. Flowers 1 or 2,
white, with green and reddish outside towards the
base, horizontal, (10—)14-25 cm long, infundibuli-
form, the tube (in dried material) 8-12 mm @.
Perianth segments not clawed, spreading distally;
nectariferous furrow papillose. Filaments linear in
dried material, 13-17 cm; anthers 5-15 mm long;
pollen yellow. Style 10'/,-16 cm; stigma deeply
3-lobed. Capsule c. 5 cm long.

Distr. Taiwan and Malesia: Philippines: North
Luzon (Bontoc, Benguet and Pangasinan Prov.).

Ecol. Open grassy slopes in the pine region,
1100-2300 m. Fi. May, fr. Oct.

Vern. Philippines: kanyon, IIk.,
soyasoi, tubtubkau, \g., tuktukpau, Bon.

Note. Except for the presence of papillae on the
nectariferous furrow hard to distinguish in flower
from L. longiflorum.

luplupak,

13. ASPARAGUS

LINNE, Sp. PI. (1753) 313; Gen. Pl. ed. 5 (1754) 147; BAKer, J. Linn. Soc. Bot. 14
(1875) 594; Bru. Fl. Austr. 7 (1878) 17; Baker, Fl. Trop. Afr. 7 (1898) 425;
KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 362; Hutcu. Fam. FI. Pl. ed. 2,
2 (1959) 608; Jessop, Bothalia 9 (1966) 31; BACK. & BAKH. f. Fl. Java 3 (1968) 92.
— Asparagopsis KUNTH, Abh. K. Ak. Wiss. Berlin (1842) 35.

Climbing or erect, glabrous or pubescent, softly woody plants usually with
bisexual flowers, rarely dioecious. Rhizome perennial; roots mostly thick and often
tuberous. Aerial stems annual or perennial, usually much-branched; branches
sometimes similar to the main stems and sometimes morphologically distinct.
Leaves cauline, alternate, scale-like, usually brown and at least partially scarious,
often with a spine from the abaxial surface; leaves of the rhizome scale-like, lacking
a spine. Leaf-like structures (cladodes) solitary or fascicled, flat, angled or terete,
arising in the axils of cauline leaves but sometimes absent from flower-bearing
branchlets. Pedicels solitary or fascicled in the axils of the cauline leaves, articu-
lated. Perianth segments free or minutely connate, equal or subequal, 1-veined,
white or nearly so, often with a green longitudinal band on the abaxial surface,
spreading or less often refiexed. Filaments flattened, attached to the perianth seg-
ments; anthers dorsifixed, oblong, introrse. Ovary superior, + sessile, 3-celled;
ovules axile, 2 to few; style usually filiform with 3 short branches, less often divided

216 FLORA MALESIANA [ser. I, vole?

nearly to the base. Fruit usually a red 1- to few-seeded berry; perianth not usually
persistent. Seeds globose or partly angled.

Distr. Widespread in Europe, Africa, Madagascar, and Asia; in Malesia 2 spp., one of which is wide-
spread in the Old World and the single one known from Australia.

There are probably fewer than 100 spp., but the taxonomy of the genus is poorly understood.

Ecol. Species have a preference for arid areas in the open and for savanna, but some occur in damp
forests.

Notes. The morphology of the spines has been discussed by CusseT & TRAN (Bull. Soc. Bot. Fr. 113,
1966, 151). The nature of the leaf-like organs (cladodes) is controversial. They are most frequently treated
as axillary structures, i.e. modified branches (e.g. KAUSSMANN, Bot. Stud. 3, 1955), and my own work
confirms this. However, ARBER (Monocotyledons: a morphological study, 1925) believed that in a few
species they were in fact leaves, while SCHLITTLER (Bot. Jahrb. 79, 1959, 428) concluded that they are leaves
in all species.

KEY TO THE SPECIES

1. Flowers unisexual, borne at normal vegetative nodes. Cladodes usually flat, less often triquetrous

1. Flowers bisexual, usually on branches lacking cladodes. Cladodes triquetrous . .

1. Asparagus cochinchinensis (LouR.) MERR. Philip.
J. Sc. 15 (1919) Bot. 230; En. Philip. 1 (1922) 206;
GaGNEP. FI. Gén. I.-C. 6 (1934) 780; MERR.
Comm. Lour. (1935) 108; MAKINo, Ill. Fl. Japan
(1954) 735. — Melanthium cochinchinense Lour.
Fl. Coch. (1790) 216. — A. lucidus LINDL. Bot. Reg.
30 (1844) Misc. 29; BAKER, J. Linn. Soc. Bot. 14
(1875) 605; MerrR. & ROLFE, Philip. J. Sc. 3 (1908)
Bot. 96, incl. var. dolichocladus MERR. & ROLFE.

Plant dioecious. Stems glabrous, erect, procum-
bent or climbing, shallowly ridged; branches
numerous, uSually simple, arising singly, less than
10(-30) cm long, spreading or erect-spreading,
often straight. Roots with distant, elongate tubers.
Scale-/leaves with scarious portion less than
5(-8) mm long; spines absent on branches, poorly
developed or up to 5 mm long on main axes.
Cladodes present on branches and towards the ends
of stems, flat, or 3-angled, 1-3-nate, linear-arcuate,
5-15(-40) by !/,-1!/. mm. Pedicels 1- or 2(—3)-nate,
arising from the axils of scale-leaves subtending
cladodes, articulated near or above the middle,
2-3(-6) mm long. Perianth segments yellow-green,
pale green or white, spreading similar, oblong-
elliptic, c. 2-3'/, mm long. — ¢$ Flowers: filaments
shorter than the perianth segments, cuneate from a
broad base; anthers nearly 1 mm long (one
specimen); ovary rudimentary. — @ Flowers:
filaments c. half the length of the perianth seg-
ments; anthers rudimentary; ovary obovoid,
11/,-2 mm long; style c. 1/, mm, with 3 stigmatic
ridges; ovules 2 per locule. Berry green when ripe,
c. 4-7 mm @. Seeds 1-4, globose or angled,
c. 2-3 mm @.

Distr. Korea, Japan, Ryu Kyu Is., S. China,
Indo-China, Taiwan; in Malesia: Philippines
(N. Luzon: Benguet Prov.), one record (LOHER
1928), probably from above 1200 m.

Ecol. There are several records from within the
spray zone on coral or limestone substrates; also in

1. A. cochinchinensis
2. A. racemosus

bush up to at least 200 m, but no ecological details
known from the Philippines.

Notes. The recorded flower colours may be mis-
leading. It is possible that the segments are white
with a green or yellow-green band.

GAGNEPAIN recorded that the flowers were uni-
sexual. I have seen insufficient material to determine
whether the plants are always dioecious or to con-
firm that the flowers are always functionally
unisexual, but in all flowers I examined one sex
appeared to be rudimentary.

2. Asparagus racemosus WILLD. Sp. Pl. 2 (1799)
152; BAKER, J. Linn. Soc. Bot. 14 (1875) 623; Bru.
Fl. Austr. 7 (1878) 17; ENGL. Bot. Jahrb. 7 (1886)
448; Rip. Fl. Mal. Pen. 4 (1924) 331; Back.
Handb. Fl. Java 3 (1924) 72; HeyNe, Nutt. Pl.
(1927) 444; H. Perrier, Fl. Madag. fam. 40
(1938) 21; STEEN. Atlas Trop. Nederl. (1938) map
7°; Jessop, Bothalia 9 (1966) 72; HutcH. & DALz.
Fl. W. Trop. Afr. ed. 2, 3 (1968) 93; Back. &
BAKH. f. Fl. Java 3 (1968) 93. — A. dubius DECNE,
Nouv. Ann. Mus. Paris 3 (1834) 363; Herb. Timor.
Descr. (1835) 35; SPAN. Linnaea 15 (1941) 476,
added A. penduliflorus Ziep., nomen, in syn. —
Asparagopsis decaisnei KUNTH, En. Pl. 5 (1850)
103, nom. illeg.; ZOLL. Syst. Verz. 1 (1854) 67; Mia.
FI. Ind. Bat. 3 (1859) 562; RIDL. in Forbes, Wand.
(1885) 520. — Asparagopsis schoberioides KUNTH,
En. Pl. 5 (1850) 70; IL’In, Fl. U.S.S.R. 4 (1968)
328. — Asparagopsis javanica KUNTH, En. Pl. 5
(1850) 100; ZoLL. Syst. Verz. 1 (1854) 67; Mia. FI.
Ind. Bat. 3 (1859) 562.

Plant with bisexual flowers. Stems glabrous,
usually climbing, up to 2-3 m high or more,
smooth or grooved, in their lower part un-
branched; branches numerous and branched;
solitary; final branches usually 1-4-nate, up to
5(-10) cm long, spreading or ascending, straight.
Roots with distant, elongate tubers. Scale-/eaves

1979]

LILIACEAE—I (Jessop)

217

with scarious portion up to 5(-10) mm long and
spine usually absent on final branches, up to
5(-10) mm long. Cladodes present mainly on
branches and towards the ends of stems, triquet-
rous, linear-crenate, 1-—3(-4)-nate, (7—)10—25
(-40) mm long, rarely over 1 mm broad. Pedicels
1- or 2-nate, usually on branches 2-6 mm long
lacking cladodes, less often on normal branches,
articulated usually near the middle, 3-5 mm long.
Flowers bisexual, scented. Perianth segments white
with a green band, spreading similar oblong to
obovate-oblong, 2-3(-4) mm long. Filaments
shorter than or about the same length as the
perianth segments; anthers c. 0.2-0.3 mm long.
Ovary obovoid, c. 11/, mm long; style c. '/; mm,
with 3 stigmatic ridges or 3 short branches; ovules
2 per locule. Berry red when ripe, c. 4-6 mm @.
Seeds 1-3, globose or angled, c. 2-3 mm @.

Distr. Widespread in Africa including the
southern Cape, Guinea and Madagascar, through
southern Asia into China, in South Malesia and
the northern states of Australia; in Malesia: Malay
Peninsula (Langkawi, on limestone rocks), Java
(in the western half only on the Northcoast, in E on
both sides; also Madura and Kangean Is.), Lesser
Sunda Is. (Lombok, Sumba, Sumbawa, Flores,
Timor), and SE. Moluccas (Tenimber Is.).

The range in Malesia is distinctly disjunct on
both ends which is due to its drought preference;

it is absent from the Sundaland rain-forest core.

Ecol. In Malesia a distinct drought-loving plant
and bound to the seasonal areas, in Java to the
zone with at most 20 rainy days in the 4 driest
consecutive months of the year, mostly in coastal
areas, in sunny thickets and on dunes, in monsoon
forest with Bambusa spinosa, Acacia leucophloea,
etc., predominantly in the lowland, but ascending
to c. 1150 m (BACKER). F/. mostly Aug.—Jan.

Vern. Christusdoorn, D; bek bun, Chinese;
Java: sangga langit, J; Timor: niesie saub, nénésan;
Tenimber Is.: skikirie, Saumlak.

Notes. There are many synonyms from Africa,
Asia and Australia, but none are known to have
been used for the Malesian area.

A. racemosus is closely allied to A. cochinchinen-
sis and can only be distinguished by a combination
of characters. In Malesia they are geographically
separated but until a revision of the genus, at least
in Asia, is undertaken the taxonomy of this group
must remain uncertain.

Excluded

According to MERRILL (En. Philip. 1, 1922, 206)
Naves (Nov. App. 1880, 264) has credited A. decli-
natus L. and A. racemosus WILLD. to the Philippine
flora, but both were apparently admitted on
erroneous identifications.

14. DISPORUM

SALISB. Trans. Hort. Soc. 1 (1812) 331; D. Don, Trans. Linn. Soc. Lond. 18 (1841)
513; BAKER, J. Linn. Soc. Bot. 14 (1875) 588; Hook. f. Fl. Br. Ind. 6 (1894) 359;
Back. Handb. FI. Java 3 (1924) 73; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, l5a
(1930) 368. — Lethea Norona, Verh. Bat. Gen. ed. 1, 5 (1790) Art. 4, 2, nomen. —
Drapiezia BL. En. Pl. Jav. (1827) 8. — Fig. 12.

Glabrous rhizomatous herbs. Aerial stems annual, erect, branched. Leaves
cauline; the lower brown and scale-like with a sheathing base; the others alternate
becoming opposite distally, flat, with many veins, petioled, usually ovate, obtuse at
base. Peduncles terminating vegetative branches, not articulated, bearing 1-6
nutant flowers in an umbel. Perianth segments free, equal or subequal, erect or
distally recurved, several-veined, usually saccate or spurred at the base. Filaments
linear to lanceolate, attached to the receptacle or the base of the perianth; anthers
dorsifixed, linear-oblong, extrorse. Ovary superior, sessile, ovoid to obovoid,
3-celled; ovules axile, 2 per cell; style filiform, with 3 recurved stigmatic branches.
Fruit a 1—3-seeded berry.

Distr. Approximately 30 spp. currently recognized in the western U.S.A. and Canada, Japan, China,
northern Deccan Peninsula, through Thailand to West Malesia: Malay Peninsula, Sumatra, Java, and
Bali.

In my opinion it is highly unlikely that more than half of these will be recognized after critical revision.

Ecol. Typical forest plants.

Note. BACKER /.c. pointed to the peculiar sympodial structure of the stem.

218

FLORA MALESIANA

[ser. I, vol. 91

1. Disporum cantoniense (LourR.) MERR. Philip. J.
Sc. 15 (1919) 229; MERR. Comm. Lour. (1935) 109.
— Fritillaria cantoniensis Lour. Fl. Coch. (1790)
206. — Uvularia chinensis KER-GAWL. Bot. Mag.
(1806) t. 916. — D. pullum Sauiss. Trans. Hort.
Soc. 1 (1812) 331; Hassk. Pl. Jav. Rar. (1848) 105;
Mia. FI. Ind. Bat. 3 (1859) 552; BAKER, J. Linn.
Soc. Bot. 14 (1875) 589; Hook. f. Fl. Br. Ind. 6
(1892) 360; RipL. J. Fed. Mal. St. Mus. 4 (1909)
82, incl. var. multiflorum RIDL.; Koorp. FI. Tyjib. 1
(1922) 47; RIDL. Fl. Mal. Pen. 4 (1924) 338; HEND.

Mal. Wild Flow. Monoc. (1954) 185, f. 109;
CHAROENPHOL, Thai For. Bull. 8 (1974) 89. —
Drapiezia multiflora BL. En. Pl. Jav. (1827) 8;
(1974)89. — Drapiezia multiflora BL. En. Pl. Jav.
(1827) 8; JUNGH. Java ed. 2 (neerl.) 1 (1853) 522;
ZOLL. Syst. Verz. 1 (1854) 66, incl. var. albiflora
ZoL_. — D. leschenaultianum D. Don, Proc. Linn.
Soc. 1 (1839) 45; Trans. Linn. Soc. 18 (1841) 518;
Merr. Contr. Arn. Arb. 8 (1934) 19. — D. hors-
fieldii D. Don, Proc. Linn. Soc. 1 (1839) 45
(WALLICH 5088D). — Streptopus multiflorus (BL.)

Fig. 12. Disporum cantoniense (Lour.) MERR. a. Habit, x 3/4, 6. flower of the spurred form, c. ditto of the
non-spurred form, both x 2!/,, d. gynoecium, e. stamen, both x 5, f. fruit, x 11/, (drawn from various
collections).

1979]

LILIACEAE—I (Jessop)

219

D. Dietr. Syn. Pl. 2 (1840) 1121. — D. multiflorum
(BL.) D. Don, Trans. Linn. Soc. 18 (1841) 518;
Mia. FI. Ind. Bat. 3 (1859) 552. — D. calcaratum
D. Don, Trans. Linn. Soc. 18 (1841) 516; BAKEr,
J. Linn. Soc. Bot. 14 (1875) 588; Hook. f. Fl. Br.
Ind. 6 (1892) 359; CHAROENPHOL, Thai For. Bull.
8 (1974) 89. — Uvularia multiflora (BL.) KUNTH,
En. Pl. 4 (1843) 207. — D. chinense (KER-GAWL.)
O. K. Rev. Gen. Pl. 2 (1891) 708; BAck. Handb.
Fl. Java 3 (1924) 73; Doct. v. LEEUWEN, Verh.
Kon. Ak. Wet. A’dam sect. II, 31 (1933) 147;
BACK. & BAKH. f. FI. Java 3 (1968) 94. — Fig. 12.

Stems erect, often 45-80 cm high, up to 2!/, m,
branched only in the upper half; branches erect-
spreading, becoming flexuose distally. Roots
fibrous or slightly thickened but not tuberous.
Leaves usually ovate, less often oblong or lanceo-
late, rarely suborbicular, acuminate, 6'/,-17 by
2!/.-9 cm, with 5-9 veins prominent on the lower
surface and numerous finer veins; petiole to 8 mm.
Inflorescence an umbel of 2-7 flowers terminating a
short branch which arises opposite a leaf and bears
a single sometimes somewhat reduced leaf;
peduncle 0-3'/, cm; pedicels 1-4 cm. Perianth
segments lanceolate to oblanceolate, acute, keeled
below and usually saccate or less often with a spur
to 5 mm long at the base, white, greenish to
purple, 1—2(-21/,) cm long. Filaments linear to
lanceolate, often less than half as long as the

perianth; anthers bright yellow, 2!/,-4 mm long.
Berry bluish black, 7-10 mm @. Seeds c. 3 mm @.

Distr. SE. Asia from the northern Deccan
Peninsula to southern China and Japan; in
Malesia: Malay Peninsula (Perak, Pahang) and
throughout Sumatra, Java, and Bali.

Ecol. In both primary and secondary forest,
rarely in the open, (700—)1000-2550 m; especially
common in W. Java. Fl. Jan._Dec. DOCTERS VAN
LEEUWEN /.c. observed the flowers to be proto-
gynous; self-pollination is possible but bumblebees
also regularly visit the flowers.

Vern. Java: baradja_ lintang, kamalakian,
kibeunteur areuj, (ki)tamiang, lili leaweung, radja
lintang, tangkal milon, S, égér égér, glinggangan,
lenguk, srintil, tombagan, J; N. Sumatra: si-
demsapo, sumbul sumbul, Karo-Batak, kayu
(si-mar)soma-soma, S.

Notes. Variation in the length of the spur has
been used in segregating species, but BACKER
recognized the continuity of this gradation in
Javanese material. Some Javanese plants have
spurs as long as those in typical ‘D. calcaratum’.

Branching is partly sympodial. Short terminal
axes bear the umbel. Continuation of growth of the
aerial shoot is by a branch arising in the axil of the
second leaf below the umbel. The node separating
two leaves associated with the inflorescence is
sometimes so short that the leaves appear opposite.

15. DISPOROPSIS

Hance, J. Bot. 21 (1883) 278; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930)
370; NaKAl, J. Jap. Bot. 12 (1936) 147; CHANG & Hsu, Taiwania 19 (1974) 64. —
Fig. 13.

Glabrous rhizomatous herbs. Aerial stems annual, erect, unbranched. Leaves
cauline, alternate, flat, with many veins, petioled, entire, ovate to oblong, obtuse or
subcuneate at the base; brown scale-like leaves on the rhizome, less often also at
the base of the aerial stems. Flowers solitary in the axils of the leaves, nutant.
Pedicels articulated. Perianth campanulate; segments 3-5, fused, equal, viscid,
saccate at the base. Filaments expanded to form a corona attached to the perianth;
anthers dorsifixed, sagittate, introrse. Ovary superior, sessile, ovoid, 3-celled ; ovules
axile, 4-6 per cell; style short and thick, with simple or 3-lobed stigma. Fruit a
1—5-seeded berry.

Distr. Probably 4 or 5 spp. from Thailand, Indo-China, southern China, Taiwan; in Malesia: Philip-

pines.
Ecol. Forests, usually in the mountains.

1. Disporopsis fusco-picta HANCE, J. Bot. 21 (1883)
278. — Disporum pullum (non SA.IsB.) MERR.
Philip. J. Sc. 1 (1906) Suppl. 182. — Disporum
luzoniense MeErRR. Philip. J. Sc. 5 (1910) Bot. 338;
En. Philip. 1 (1922) 206. — Fig. 13.

Stems erect, 25-45 cm. Roots fibrous or slightly

thickened but not tuberous. Leaves 6-9, ovate to
oblong, obtuse or subcuneate at the base, acumi-
nate, 6-121/, by 21/,-5 cm, with usually 7 veins
prominent on the lower surface and numerous finer
veins; petiole (3—)5-12 mm. Flowers in the axils of
the lower leaves. Pedicels 1-2!/, cm. Perianth seg-

220 FLORA MALESIANA [ser. I, vol. 9!

Fig. 13. Disporopsis fusco-picta HANCE. a. Habit, x 1/3, b. flower, c. ditto, laid open showing corona and
stamens, both x 2!/,, d. fruit, x 2 (a—c STEINER 2147, d PNH 7471).

ments fused below middle, 13-15 by 2-3 mm, lan-
ceolate, obtuse, shallowly saccate at the base, white
with (always ?) dull purple inside. Corona attached
close to perianth sinus, c. 4 mm long, its lobes
emarginate. Anthers sessile on the corona, c. 1 mm
long, very shallowly sagittate. Berry becoming
purple, blue or black, c. 1 cm @. Seeds c. 4 mm @.

Distr. Southern China, Taiwan, in Malesia:
Philippines (N. Luzon: Lepanto, Bontoc and
Benguet Prov.).

Ecol. Mossy forest, 1700-2500 m, with one
record of association with (the secondary pyro-
genous savanna of) Pinus insularis. Fl. rarely
recorded: May-June, or later.

16. TRICALISTRA

RIDL. J. Fed. Mal. St. Mus. 4 (1909) 83; Fl. Mal. Pen. 4 (1924) 330. — Tupistra
sensu HUTCH. Fam. FI. Pl. ed. 3 (1973) 749, in part. — Fig. 14d.

Stemless herbs. Rhizome horizontal, woody; roots thick and fleshy. Leaves large,
basal, caespitose, petioled, expanding towards the base to form a sheath, with a
strong main vein and numerous finer veins. Inflorescence a spike. Flowers numerous,
each in the axil of a bract. Perianth segments 6, fleshy, fused for about half their
length, campanulate, equal. Stamens 6, attached to the perianth; anthers sub-
sessile, dorsifixed, oblong, dehiscing introrse-laterally. Ovary superior, sessile,
subcylindrical, 3-lobed, 1-celled, containing (2 or) 4 discord ovules but with traces
of two further carpels; stigmas 3, hippocrepiform, sessile. Fruit a drupe, green when
unripe, globose, 1-seeded.

Distr. Monotypic. Malesia: Malay Peninsula.

Note. Regarded as a synonym of Tupistra by HUTCHINSON and recognized, but with some doubt, by
AIRY SHAW (Willis Dict. ed. 8, 1973). The only consistent difference is that Tricalistra has no style, con-

sistently 6 stamens, and a 3-lobed stigma, a set of characters of equal standing as those separating other
genera in the Aspidistreae.

4 (1909) 83; Fl. Mal. Pen. 4 (1924) 331. — Fig. 14d.

Scale-like leaves several, sessile, lanceolate to
5 cm long. Foliage /eaves few, broadly lanceolate
to oblanceolate, acuminate, cuneate at the base;
dark green, chartaceous, 30-40 cm Jong when in
flower, lengthening in fruit, 8-11 cm _ broad;
petiole poorly differentiated, sometimes winged,
12-20 cm. Inflorescence 12-15 cm long when in
flower, lengthening in fruit, with 25-35 flowers.
Bracts ovate, obtuse, caducous, to 3 mm long.
Perianth fleshy, c. 5 mm long, lobes as long as the
tube, recurved, ovate, acute, dull ochreous yellow.

2271

tube, thecae divaricate at base, less than 1 mm
long. Ovary c. 3 mm long. Fruit to c. 2 cm long.

Distr. Malesia: Malay Peninsula (Pahang:
Cameron Highlands).

Ecol. One collection annotated ‘on rocks in open
places’, 1000 m. F/. Nov. (one record), fr. April.

Note. Syntypes collected by RmpLey (13692) are
in SING and K. Rww_ey referred to the leaf having
6 veins. However, leaves on both type specimens
have at least 100 of which about 13 are more
conspicuous than the others. The only other collec-
tion (Nur SF 32725) agrees with the type.

17. TUPISTRA

Ker-GAwL. Bot. Mag. 39 (1814) t. 1655; BL. Tijd. Nat. Gesch. Phys. 1 (1834) 67;
Baker, J. Linn. Soc. Bot. 14 (1875) 580; Hook. f. Fl. Br. Ind. 6 (1892) 324;
KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 372. — Fig. 14a-c.

Stemless herbs. Rhizome tuberous or horizontal, thick and woody; roots thick
and fleshy. Leaves large, basal, scattered or distichous, usually distinctly petioled,
expanding towards the base to form a sheath, with a strong main vein and numer-
ous finer veins. /nflorescence a spike. Flowers numerous, each in the axil of a bract.
Perianth segments 6 or 8, fleshy, fused, campanulate, equal. Anthers sessile or sub-
sessile, inserted in tube, dorsifixed, oblong or ovoid, introrse. Ovary superior,
sessile, subglobose or not externally differentiated from style, 3(—4)-celled; ovules
2 in each locule; style cylindrical; stigma large, peltate or capitate, entire or
variously lobed. Fruit a globose berry, usually 1-seeded; seeds turgid; perianth

persisting below the fruit.

Distr. Eastern Himalayas to southern China; in Malesia: Malay Peninsula and Sumatra.

The type of T. squalida (the type species) was described, undoubtedly in error, from Amboina.
About 25 spp. have been described, but many of these should probably be reduced to synonymy.
Ecol. Most frequently recorded from dry evergreen forest, often in deep shade and often from near

streams.

Syst. It appears to me that in the Aspidistreae too many small genera are distinguished on insignificant

characters.

Note. The Malesian species have been insufficiently collected for convincing taxonomic judgements to

be made.

KEY TO THE SPECIES

1. Spike to 17 cm long; style and ovary up to 7 mm long; stigma usually 11/,.-3 mm @ .

1. T. violacea

1. Spike at least 20 cm long; style and ovary at least 7!/, mm long; stigma at least 4"/, mm @

1. Tupistra violacea RipL. J. Str. Br. R. As. Soc.
n. 41 (1904) 35; Mat. Fl. Mal. Pen. Monoc. 2
(1907) 93; Fl. Mal. Pen. 4 (1924) 330.

Leaves few, elliptic to oblanceolate, acute or
acuminate, cuneate at the base; lamina 50-70 by
T'/,-13 cm; petiole well-defined, to 40 cm. Inflores-
cence ascending to 17 cm long when in flower,
lengthening in fruit, with 30-40 flowers. Bracts
somewhat amplexicaul ovate, c. 5 by 3-8 mm.

2. T. grandis

Perianth segments violet; the tube 5-6 mm long,
41/,-6 mm wide; lobes darker than tube, oblong to
ovate, 4-5!/, by 3-4 mm. Anthers sessile, attached in
tube or throat, c. 1-1!/, mm long. Style and ovary
white, spotted violet, 3'/,-7 mm long; stigma
11/,-4 mm @, obscurely lobed.

Distr. Malesia: Malay Peninsula (Perak:
Bujong Malacca; Penang Highlands); probably
also in Thailand.

222 FLORA MALESIANA [ser. I, vol. 9!

~ %

= v ,
Wig,
VK)

Fig. 14. Tupistra grandis Rwv. a. Habit, x '/, b. flower from outside, c. flower laid open, both x 2. —
Tricalistra ochracea RiDL. d. Flower, halved, showing gynoecium, 3 tepals, to which stamens are attached,
x 10 (a-c LORZING 8753, d RIDLEY 13692).

1979]

LILIACEAE—I (Jessop)

Ecol. Mountain forest, apparently rare and no
collections made since 1901 have been seen. Fi.
March, Dec.

Note. This species closely resembles T. squalida,
the type species of the genus, from the Himalayas.
Further study is needed to confirm that they should
be kept separate. In the absence of recent collec-
tions much of the information here is taken from
Ripiey (1924). The scale-like leaves recorded
for T. gracilis are likely to occur in T. violacea,
but the basal part of the plant has not been
preserved.

2. Tupistra grandis RID. J. Bot. (1900) 73; Mat.
Fl. Mal. Pen. Monoc. 2 (1907) 93; Fl. Mal. Pen. 4
(1924) 330; B. M. ALLEN, Mal. Nat. J. 19 (1966)
303. — T. perakensis Nicuots. Ill. Dict. Gard.,
Cent. Suppl. (1901) 722. — Fig. 14a-c.

Leaves few, surrounded by several sessile lanceo-
late scale-like leaves to 25(-40) cm long, elliptic or
sublinear to oblanceolate, acute or acuminate,
cuneate at the base; lamina (50-)70-150 by 6—
14 cm; petiole often winged and poorly defined,
to 40 cm. Inflorescence erect, 20-50 cm long when
in flower, lengthening slightly in fruit, with up to
100 or more flowers, with a strong smell. Bracts
somewhat amplexicaul, ovate, 5-10 by 2-6 mm.
Perianth segments violet to dark purple; tube
5-11 mm long, 5-10 mm wide; lobes darker than
tube, oblong to ovate, 4-14 by 3-5 mm. Anthers
sessile, attached near top of tube, 11/,-2 mm long.
Style and ovary white, 7'/,-16 mm long; stigma
4'/,-12'/, mm @, peltate, flat to biconvex, some-
times rugulose or irregularly lobed.

Distr. Malesia: Malay Peninsula (Perak,
Kelantan, Pahang, Selangor, Langkawi) and N.
Sumatra (Tapanuli Res.: Karo Highlands);
probably also in Thailand.

Ecol. In Malaya records suggest that it usually
occurs in wet rocky places on limestone. In
Sumatra it grows in forest at 600-1225 m, but in

Malaya there is a record of 150 m in Perak. Fi.
Jan.—Dec.
Vern. N. Sumatra: singkut antu, Karo-Batak.
Note. The Sumatran specimens tend to have
longer spikes and larger flowers than the Malayan
material. No characters have, however, been found
on which to base taxonomic separation.

Excluded

Tupistra singapureana [WALL. Cat. n. 5195];
Baker, J. Linn. Soc. Bot. 14 (1875) 581; Hook.
f. Fl. Br. Ind. 6 (1892) 325, was omitted from the
genus by S. Kurz (J. As. Soc. Beng. 44, ii, 1875,
199) and has indeed appeared not to belong to
Liliaceae. It was referred by RIDLEY (Mat. Fl. Mal.
Pen. Monoc. 1, 1907, 232) to Neuwiedia curtisii
Rotre and by RoLFe (Kew Bull. 1907, 412) to
Neuwiedia singapureana (BAKER) ROLFE.

According to DE VoGEL (Blumea 17, 1969, 331)
= Neuwiedia zollingeri RCHB. var. singapureana
(BAKER) DE VoGEL (Orchidaceae).

Tupistra squalida KErR-GAWL. Bot. Mag. 39
(1814) t. 1655; E>warps, Bot. Reg. (1823) t. 704;
LoppiGces, Bot. Cab. 6 (1821) t. 515; Br. Tijd.
Nat. Gesch. Phys. 1 (1834) 67, t. I1IC; Mia. FI.
Ind. Bat. 3 (1859) 569; BAKER, J. Linn. Soc. Bot. 14
(1875) 580; cf. Hook. f. Fl. Br. Ind. 6 (1892) 324,
in nota sub T. nutans. — Rhodea tupistra SCHULT.
Syst. 7, 2 (1829) 173, nom. illeg.

The provenance of this species, the type of the
genus, was given as ‘Amboyna’. This is certainly
erroneous. It was described from a cultivated plant
in the nurseries of LoppIGEs, and was obviously in
the former century cultivated in several botanic
gardens. The more curious it is that its proper
identity and native country remains more or less
uncertain.

BAKER /.c. reduced T. nutans WALL. (Bot. Reg.
t. 1333) from India to this species, but HOOKER f.
kept these two entities apart.

18. LIRIOPE

Lour. Fl. Coch. (1790) 200; L. H. Bamey, Gent. Herb. 2 (1929) 3; KRAUSE in
E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 376; WANG & TANG, Act. Phytotax. 1
(1951) 331; Hume, Baileya 9 (1961) 135. — Ophiopogon (non KER-GAWL.) KUNTH,
En. Pl. 5 (1850) 297, in part. — Fig. 16d.

Stemless or shortly caulescent, rhizomatous herbs. Roots thick, sometimes with
tuberous swellings. Leaves usually basal, petioled or subpetioled linear to oblanceo-
late or lanceolate, many-nerved. Inflorescence a spike or raceme. Flowers solitary
or fascicled in the axil of each bract. Perianth segments fleshy, campanulate, free or
with a broad tube and short lobes, equal, with 1 vein, violet or white. Anthers
6-8, subsessile or pedicels short, attached near base of perianth, dorsifixed, in-
trorse. Ovary superior, sessile, ovoid to subglobose, 3—-4-celled; ovules axile, 2 in

224 FLORA MALESIANA [ser. I, vol. 9!

if Uh KN
WANN Sitmas
I f AK ) ZEN ;
DAN
), N

K

Fig. 15. Ophiopogon caulescens (BL.) BACK. Habit, with seed, x !/4. In forest above mountain garden
Tjibodas, Febr. 1936.

1979] LILIACEAE—I (Jessop) 225

each locule; style short and thick; stigma capitate or peltate. Ovary wall rupturing
early in the growth of the seeds which are therefore exposed during most of their
development. Seeds black, globose or slightly elongate, with fleshy testa; perianth

persistent.

Distr. About 5 spp., in Japan, China, Indo-China and North Malesia: N. Philippines.
Note. Some species are widely grown as ornamentals.

1. Liriope graminifolia (L.) BAKER, J. Linn. Soc.
Bot. 17 (1879) 499; Merr. En. Philip. 1 (1922) 207;
Hume, Baileya 9 (1961) 150. — Asparagus gramini-
folius LINNE, Sp. Pl. ed. 2 (1762) 450. — Dracaena
graminifolia (L.) LINNE, Syst. Nat. ed. 12 (1767)
275. — ? L. spicata Lour. Fl. Coch. (1790) 201;
L. H. BatLtey, Gent. Herb. 2 (1929) 33; MerR.
Comm. Lour. (1935) 109; Hume, Baileya 9 (1961)
150, 152, 158. — ? Ophiopogon spicatus (LourR.)
KER-GAWL. Bot. Reg. 7 (1821) t. 593; NAveEs,
Nov. App. (1880) 264. — Mondo graminifolia
(L.) Komz. Tokyo Bot. Mag. 40 (1926) 333. —
L. muscari [non (DECNE) L. H. BAILEY] HATus.
Mem. Fac. Sc. Kagoshima Un. 5, 3 (1966) 62. —
Fig. 16d.

Rhizome horizontal, slender, moderately woody.
Roots bearing distant tubers. Leaves basal, caespi-
tose, linear to narrowly linear-oblanceolate,
minutely denticulate on the margins, the central
vein sometimes conspicuously larger than the
others, expanded to form membranous wings
towards the base, 25-90 cm long, 2-9 mm broad.
Peduncle erect, (12—)30—50 cm long, shorter than
leaves. Bracts deltoid, to 4 mm long. Pedicels

2-S-nate, 2-12 mm long, articulated at the base of
the flower. Leafy shoots occasionally produced
from axil of bracts on inflorescence. Perianth
segments free, 3!/,4 mm long, violet. Filaments
to 2 mm long; anthers c. 1 mm long. Seeds oblong
(perhaps only when young) or globose, c. 5 mm
long.

Distr. Japan, China and North Malesia:
Philippines (Batan Is.; Mindoro and Luzon).

The paucity of collections suggests that this
species is rare in the Philippines.

Ecol. Open slopes at c. 1400 m (MERRILL, /.c.),
but obviously in the Batan Is. at low altitude.

Notes. The differences between L. graminifolia
and L. spicata are not clear. HUME depended for
their separation largely on the conspicuous mem-
branous basal wings to the leaves of the former,
associated with quantitative characters of the
leaves and inflorescence. L. graminifolia is likely to
remain the correct name for the Philippine species
whether L. spicata is treated as a synonym or not.

L. muscari (DECNE) L. H. BAILey (Gent. Herb.
2, 1929, 35) differs according to HUME by caespitose
habit and stiffer and wider leaves (8—26 mm).

19. OPHIOPOGON

KER-GAWL. Bot. Mag. 27 (1807) t. 1063; Hook. f. Fl. Br. Ind. 6 (1892) 267; RIDL.
Fl. Mal. Pen. 4 (1924) 326; RoprRIGUEZz, Bull. Soc. Bot. Fr. 75 (1928) 997; FI.
Gén. I.-C. 6 (1934) 655; BACK. & BAKH. f. FI. Java 3 (1968) 95, nom. gen. cons. —
Mondo ADANS. Fam. 2 (1763) 496; FARWELL, Amer. Midland Nat. 7 (1921) 41;
L. H. Barley, Gentes Herb. 2 (1929) 17; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2,
15a (1930) 377; OHwi in Fedde, Rep. 36 (1934) 45. — Flueggea RicH. Neues J.
Bot. 2 (1807) 8 (‘Fluggea’); BAKER, J. Linn. Soc. Bot. 17 (1879) 500. — Chloopsis
BL. En. Pl. Jav. (1827) 14; Hassk. Flora 34 (1851) 481. — Fig. 15, 16a-c.

Stemless herbs. Rhizome sometimes very short; roots fibrous or fleshy, sometimes
tuberous. Leaves basal, linear or petioled with a broad lamina, expanded at the base
to form a conspicuous scarious sheath. /nflorescence a raceme. Pedicels 1—several-
nate, in the axils of bracts and usually associated with a few smaller bracts also
apparently in the axil of the principal bract, articulated usually in the distal half.
Perianth segments free, equal, with 1 vein, spreading or campanulate, white or
violet. Filaments often connate, short, thick, glabrous, attached to the base of the
perianth segments or to the receptacle; anthers basifixed, linear-oblong, dehiscing
introrsely. Ovary superior to inferior, 3-celled; ovules basal, 2(—6) in each locule;

226

FLORA MALESIANA

[ser. I, vol. 91

style columnar, minutely 3-lobed. Ovary wall rupturing early in the growth of the
seeds which are therefore exposed during most of their development. Seeds blue,
often globose, with fleshy testa; perianth often wholly or partly persistent.

Distr. India through to southern China to Thailand, Indo-China, Japan and Taiwan; in Malesia:
Malay Peninsula, Sumatra, Java, Borneo, and Philippines.
About 70 spp. of Ophiopogon have been described, but I doubt whether more than a third of these

should be recognized.
Ecol. In forest.

Notes. RIDLEY recognized O. malayanus, O. intermedius and O. prolifera from the Malay Peninsula.
No fertile material has been seen from this area and the only specimens seen (CORNER SF 37872, K & L,
and SOEPADMO & MAHOoMUuD 1214, K) do not differ significantly from O. caulescens.

No Malayan material identified as O. intermedius or O. prolifera has been seen.

RipD_ey characterized O. prolifera by having the filaments connate, and distinguished O. intermedius
from O. malayanus on its larger flowers (12 mm wide). He did not give a comparable figure for O. malaya-

nus but recorded that the segments were 3 mm long.

The descriptions could all be of varieties of O. caulescens, falling within the morphological range known
for that species from Java, except that the shortest perianth recorded for Java material is 4 mm long
(almost 50% greater than the length recorded for O. malayanus).

O. malayanus has also been recorded from the Philippines and Borneo (MERRILL, 1922).

Material from continental Asia of O. intermedius resembles O. japonicus, but no material has been seen

from the Malay Peninsula.

KEY TO THE SPECIES

1. The broadest leaves on each plant more than 4 mm broad. Rhizome well-developed. Lowest bracts

12-25 mm long

1. O. caulescens

1. The broadest leaves on each plant less than 3 mm broad. Rhizome poorly developed. Lowest bracts

5-10 mm long

1. Ophiopogon caulescens (BL.) BACK. Handb. FI.
Java 3 (1924) 74; VAN HELTEN, Med. Alg. Proefsta-
tion Landb. n. 16 (1924) 49 (‘gauliscens’); RIDL. J.
Bot. 43 (1925) Suppl. 122; Doct. v. LEEUWEN,
Verh. Kon. Ak. Wet. A’dam sect. IT, 31 (1933) 252;
BACK. & BAKH. f. Fl. Java 3 (1968) 95. — ? O. inter-
medius D. Don, Prod. Fl. Nep. (1825) 48; Ript. J.
Fed. Mal. St. Mus. 4 (1909) 81, incl. var. macran-
thum RIDL.; Fl. Mal. Pen. 4 (1924) 327. — Chloop-
sis caulescens BL. En. Pl. Jav. (1827) 14; Zo.t.
Syst. Verz. 1 (1854) 67; Mia. FI. Ind. Bat. 3 (1859)
553. — Chloopsis acaulis Bu. En. P|. Jav. (1827) 14;
Mia. Fl. Ind. Bat. 3 (1859) 553. — ? O. prolifera
LINDL. J. Hort. Soc. 1 (1846) 76; Mia. FI. Ind.
Bat. 3 (1859) 568; RipL. Mat. Fl. Mal. Pen. Monoc.
2 (1907) 91. — Fluggea wallichiana KUNTH, En. PI.
5 (1850) 303. — O. wallichianus (KUNTH) Hook.
f. Fl. Br. Ind. 6 (1892) 268; Ripi. J. Fed. Mal. St.
Mus. 8 (1917) 118. — O. malayanus RiD-. J. Str.
Br. R. As. Soc. n. 41 (1904) 34; Mat. Fl. Mal. Pen.
Monoc. 2 (1907) 91; Merr. En. Born. (1921) 115;
En. Philip. 1 (1922) 207; Contr. Arn. Arb. 8 (1934)
19. — Mondo malayanum (RIDL.) FARWELL,
Amer. Midland Nat. 7 (1921) 42. — O. japonicus
(non (L. f.) KER-GAWL.) Koorp. FI. Tjib. 1 (1922)
47. — O. acaulis (BL.) Ripi. J. Bot. 63 (1925)
Suppl. 122. — Fig. 15, 16b.
Rhizome rather woody and well-developed,
sometimes supported by several thick prop-roots;
roots thick but not bearing tubers. Leaves numer-

2. O. japonicus

ous, linear to linear-oblanceolate, often slightly
arcuate, subacute or narrowly obtuse, often with a
prominent midrib, glaucous (waxy), or with
glaucous stripes, on the lower surface, (15—)25-
55(-65) cm by 3-10(—22) mm. Peduncle filattened,
10-35(-46) cm long. Flowers 3-12, + secund.
Lower bracts 12-25 mm long. Pedicels solitary or,
less often, 2-nate, spreading or erect-spreading,
articulated often near the centre, 3-8 mm. Perianth
segments white or violet, oblong or ovate- or
elliptic-oblong, usually obtuse, the inner often
slightly smaller than the outer, free segments
(5-)7-7'/, by 2-3 mm. Anthers (2-)4-51/, mm long;
filaments connate at the base, c. 1/,-2 mm, up to
1 mm broad at the base. Ovary inferior or semi-
inferior; style terete, linear-obconic, (4'/,—)5—
7(-9) mm, simple or minutely trifid; ovules 2 per
locule. Seeds up to 6, glossy blue, globose or
slightly ovoid or ellipsoid, 4-8 mm long.

Distr. Continental SE. Asia and West Malesia:
Malay Peninsula (also Langkawi), Sumatra, all
over Java, Sabah (Kinabalu area), S. Philippines
(Sulu Is.: Jolo).

Ecol. Generally a forest species reported from
‘rain forest’ and ‘primary forest’. Also recorded
from screes, (650—)1000-2000 m. Fi. Jan.—Dec.
DOCcTERS VAN LEEUWEN /.c. stated that self-pollina-
tion is the rule.

Vern. Java:
Murut dial.

suket alank, J; Sabah: ryran,

synonymous with O. caulescens, it will become the
correct name.

2. Ophiopogon japonicus (L. f.) KER-GAWL. Bot.
Mag. 27 (1807) t. 1063; Merr. Philip. J. Sc. 1
(1906) Suppl. 35; ibid. 5 (1910) Bot. 338; En.
Philip. 1 (1923) 207; Hume, Baileya 9 (1961) 142. —
Convallaria japonica LINNE f. Suppl. (1781) 204. —
Mondo japonicum (L. f.) FARWELL, Amer. Mid-
land Nat. 7 (1921) 42. — O. merrillii MASAM. Bull.
Soc. Bot. Fr. 84 (1937) 90. — Fig. 16a.

Rhizome poorly developed; roots fibrous with
distinct tubers usually less than 11/, cm long and
4 mm broad (not present in all herbarium material
even where roots are well represented). Leaves
numerous, linear, usually more or less straight,
acute or subacute, usually minutely denticulate on
vein and margin, usually with a distinct midrib,
glaucous (waxy) on the lower surface, 18-60 cm by
11/,-3(—3!/.) mm. Peduncle (10—)12-35 cm. Lower

LILIACEAE—I (Jessop)

22]

bracts 5-10 mm long. Pedicels solitary, spreading
or erect-spreading, articulated near or below the
middle, 2'/,-3 mm. Perianth segments white or
violet, oblong-elliptic or ovate, obtuse, the inner
slightly smaller than the outer, free segments,
31/,4 by 2-3 mm. Anthers 2-2'/, mm long;
filaments connate at the base, c. '/, mm long and
up to !/, mm broad at the base. Ovary inferior or
semi-inferior; style terete, sublinear, c. 3 mm,
simple; ovules 2 per locule. Seeds up to 6, glossy
blue, globose or slightly ellipsoid, c. 4 mm long.

Distr. Japan and North Malesia: Philippines
(Luzon: Benguet Prov.; Mindanao: Mt Apo).

Ecol. Chiefly in mossy forest, rather common in
the Mountain Province, 850—2400(—-2900) m. Fi.
May-July, fr. Dec.—Jan.

Vern. Philippines: Jangigit, Ig., takadu, Bon.,
uli-uli, Bag.

Note. A commonly cultivated garden plant in
Java and other places in Malesia, especially for
lining borders, but never flowering in the lowland.

20. PELIOSANTHES

Anpr. Bot. Repos. 10 (1810) t. 605; HAssk. Tijd. Nat. Gesch. Phys. 10 (1843) 121
(‘Piliosanthes’); BAKER, J. Linn. Soc. Bot. 17 (1879) 503; Hook. f. Fl. Br. India 6
(1892) 265; RmpL. Fl. Mal. Pen. 4 (1924) 323; RopriGuEz, Fl. Gén. I.-C. 6 (1934)
668; Jessop, Blumea 24 (1976) 141. — Teta Roxs. [Hort. Beng. 1814, 24, nomen]
Fl. Ind. ed. Carey 2 (1832) 165. — Bulbisperma REINW. ex BL. Cat. (1823) 59,
nomen. — Bulbospermum BL. En. Pl. Jav. (1827) 15. — Lourya BAILL. Bull. Soc.
Linn. Paris 1 (1888) 743. — Neolourya RODRIGUEZ, Bull. Mus. Hist. Nat. Paris II,
6 (1934) 96. — Fig. 16e-g.

Erect, stemless, perennial herbs. Rhizome usually very short and horizontal;
roots thick. Leaves basal, usually distinctly petioled, the blade sublinear to ovate or
obovate, many-nerved. Peduncles flattened, at least at the base, erect, usually soli-
tary. Pedicels and inflorescence surrounded at the base by scarious scale-like leaves.
Inflorescence a simple raceme. Pedicels articulated. Flowers 1-6-nate in the axils of
each bract. Perianth segments fleshy, campanulate or subglobose, fused below,
equal, 1-veined, white, green, blue, violet or purple. Anthers sessile, attached to a
short annular tube (corona) arising from the perianth, introrse. Ovary superior to
inferior, 3-celled; ovules basal, 2-5 in each locule; style simple, conical to cylindri-
cal; stigma capitate or undifferentiated. Ovary wall rupturing early in the growth
of the seeds which are therefore exposed during most of their development. Seeds
blue, ellipsoid to pyriform, with fleshy testa; perianth often persistent.

Distr. Monotypic, widespread in continental SE. Asia, from the southern Deccan to NE. India and
southern China, in West Malesia: Malay Peninsula, Sumatra, Java, Lesser Sunda Is. (Sumbawa) and
Borneo.

Ecol. Usually in forest, from the lowland to the mountains.

Note. In the past far too many species were described. For an account of the taxonomic problems on
specific delimitation, see Jessop (1976).

228

FLORA MALESIANA

[ser. I, vol. 9!

We

Fig. 16. Ophiopogon japonicus (L. f.) KER-GAWL. a. Habit, x 1/4. — O. caulescens (BL.) BACK. b. Flower,

x 2.— Ophiopogon sp. c. Mature seeds, « 2. — Liriope graminifolia (L.) BAKER. d. Flower, x 4. —

Peliosanthes teta ANDR. ssp. humilis (ANDR.) Jessop. e. Habit, x !/,, f. flower, perianth shown reflexed to

reveal corona, x 3, g. LS of flower to show attachment of corona and position of ovary, x 7 (a,c

SWINBURNE 5.n. ex Hort. Adelaide, b after STEEN. Mt. Fl. Java pl. 28: 3b, dlargely after E. & P. Nat. Pfi.
Fam. ed. 2, 15a, fig. 153 ‘O. spicata’, e-g SORENSEN c.s. 2960).

1. Peliosanthes teta ANDR. Bot. Repos. 10 (1810)
t. 605; BAKER, J. LINN. Soc. Bot. 17 (1879) 505;
Hook. f. Fl. Br. Ind. 6 (1892) 265; RipL. Mat. FI.
Mal. Pen. Monoc. 2 (1907) 88; Fl. Mal. Pen. 4
(1924) 323; RopriGuez, Fl. Gén. I.-C. 6 (1934)
669; Jessop, Blumea 24 (1976) 154.

For synonyms see under the subspecies.

Leaves (2-)4-8(-12); leaf-blades (7!/,-)121/,-
47'/, by 14/,-81/,(-11'/,) cm; petioles (4-)7!/,-
50 cm long, slightly compressed. Peduncles to
35(-75) cm high; lower sterile bracts 0-4(-15), to
15(-40) mm long; fertile bracts to 15(—-30) mm long,
smaller towards the apex of the raceme. Flowers
1-6-nate. Pedicels 1-6(-10) mm long, enlarging
after flowering; articulation usually close to flower.
Perianth segments suborbicular to linear, 11/,—
6(-8) mm long. Corona forming a disk c.3-4mm @,
entire or 6-toothed. Anthers usually rather closely
adpressed to the style, c. 1/,-2 mm long. Ovary
most frequently semi-inferior; style often 3- or

6-ridged or fluted, */,-1(-2) mm long. Seeds up to
10-12 mm long.

Distr. Tropical SE. Asia; in Malesia: Malay
Peninsula, Sumatra, Java, Lesser Sunda Is.
(Sumbawa), Borneo.

Ecol. Primarily in wet evergreen forest, from
0-3000 m above sea-level. A few records indicate
that dry areas are also occasionally occupied,
possibly in wet enclaves. The subspecies appear to
grow in similar habitats.

Note. Grown as a garden ornamental or pot
plant.

a. ssp. teta. — Cf. Jessop, Blumea 24 (1976) 155. —
Teta viridiflora Rox. Fl. Ind. ed. Carey 2 (1832)
165. — P. teta var. mantegazziana PAMP. Nuovo
Giorn. Bot. Ital. n.s. 11 (1904) 151; Bull. R. Soc.
Toscana Ortic. III, 10 (1905) 50, f. 11. — P. mante-
gazziana (PAMP.) PAMP. Nuovo Giorn. Bot. Ital.
n.s. 13 (1906) 138; Merr. & Quis. Philip. J. Sc. 82

LILIACEAE—I (Jessop)

229

ad

~
: "seem,

Fig. 17. Range of Peliosanthes teta ANDR. ssp. teta (broken line), and ssp. humilis (ANDR.) Jessop (even
line).

(1953) 323. — P. graminea RiD-. J. Str. Br. R.
As. Soc. n. 59 (1911) 207. — P. teta var. angusti-
folia Riv. /.c. — P. tonkinensis WANG & TANG,
Bull. Fan Mem. Inst. Biol. Peiping, Bot. 7 (1936)
83.

Pedicels 2-6-nate. Leaf index 2—24(—34). Flowers
usually green, rarely blue. Anthers c. 0.5-0.6 mm
long.

Distr. India to southern China; in Malesia:
Malay Peninsula (Pahang, Selangor, Penang and
Langkawi Is.). Fig. 17.

b. ssp. humilis (ANDR.) Jessop, Blumea 24 (1976)
155. — P. humilis ANDR. Bot. Repos. 10 (1811)
t. 634; Mia. FI. Ind. Bat. 3 (1859) 568. — Bulbi-
Sperma ovigera REINW. ex BL. Cat. (1823) 59,
nomen. — Bulbospermum javanicum BL. En. PI.
Jav. (1827) 15; Zoxx. Syst. Verz. 1 (1854) 68. —
P. javanica (BL.) Dietr. Syn. Pl. 2 (1840) 1123;
Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 121;
Pl. Jav. Rar. (1848) 116; Mig. Fl. Ind. Bat. 3
(1859) 568; Back. Handb. Fl. Java 3 (1924) 75;
BAcK. & BAKH. f. Fl. Java 3 (1968) 95; STEEN. Mt.
Fl. Java (1972) t. 28-4. — P. violacea WALL. ex
BAKER, J. Linn. Soc. Bot. 17 (1879) 504; HeENb.
Mal. Wild Flow. Monoc. (1954) 184, f. 108. —

Lourya campanulata BAIL. Bull. Soc. Linn. Paris 1
(1888) 743. — P. albida BAKER, Bot. Mag. 116
(1890) t. 7110; Hook. f. Fl. Br. Ind. 6 (1892) 267;
Rip. Mat. Fl. Mal. Pen. Monoc. 2 (1907) 90;
Merr. En. Born. (1921) 115; FiscHER, Kew Bull.
(1932) 183. — P. viridis Riv. J. Str. Br. R. As.
Soc. n. 31 (1898) 95. — P. lurida Rw . I.c. 95. —
P. grandifolia Riv. I.c. 97. — P. stellaris Riv.
l.c. 97. — P. parviflora RIL. ibid. n. 61 (1912) 61.
— P. sumatrensis RiDv. J. Fed. Mal. St. Mus. 8
(1917) 118. — P. sessiliflora Ripv. /.c. 118. —
P. hypogyna RIDL. ibid. 10 (1920) 121. — P. monti-
cola Ripv. /.c. 155. — P. campanulata (BAILL.)
RODRIGUEZ, Bull. Mus. Hist. Paris II, 6 (1934)
96. — Fig. 16e-¢g.

Pedicels solitary in the axil of each bract. Leaf
index 2-10. Flowers sometimes green, but often
white, blue, violet or purple. Anthers c. !/,-2 mm
long.

Distr. India to southern China; in Malesia:
Malay Peninsula (throughout, incl. the Langkawi,
Penang, Singapore and Tioman Is.), Sumatra
(throughout, incl. Simalur and Billiton Is.), Java
(throughout), Lesser Sunda Is. (Sumbawa) and
Borneo (Sarawak, Southeast, Sabah, incl. the
Anambas and Karimata Is.). Fig. 17.

230 FLORA MALESIANA [ser. I, vol. 91

21. ALETRIS

LINNE, Sp. Pl. (1753) 319; Gen. Pl. ed. 5 (1754) 149; Amoen. Acad. 3 (1756) 11
(‘Alethris’); Hook. f. Fl. Br. Ind. 6 (1892) 264; FRANCHET in Morot, J. de Bot. 10
(1896) 178, 195; KRAusE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 378; HARA, J.
Jap. Bot. 42 (1967) 312. — Metanarthecium Max. Bull. Ac. St. Pétersb. 11 (1867)
438; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 260. — Meta-aletris
Masao. Trans. Nat. Hist. Soc. Formosa 28 (1938) 46. — Fig. 18, 19.

Erect, stemless, rhizomatous herbs. Roots fibrous. Leaves basal, linear or lanceo-
late, sessile, the veins of the decayed bases persisting as fibres at the base of the
plant. Inflorescence a raceme or spike. Flowers solitary in the axils of the bracts,
with a single bracteole. Pedicels not articulated. Perianth segments connate at the
base, equal, with three (often indistinct) veins, ascending or reflexing, glabrous or
pubescent, white or pink. Filaments subulate, shorter than and attached to perianth;
anthers dorsifixed, oblong to ovoid, dehiscing introrsely. Ovary half-inferior,
3-celled; ovules axile, numerous; style simple or minutely 3-lobed. Fruit a capsule;
perianth persistent. Seeds oblong.

Distr. North America (few spp.) and eastern Asia, from Japan to southern China (also Taiwan) and
the Himalayas, in Malesia: only on the high mountains of the Philippines (N. Luzon), Sabah (Mt Kina-
balu) and N. Sumatra (Gajolands and Westcoast Res.).

There are possibly fewer than 15 spp., mostly in Asia, but HARA /.c. estimates the number at c. 30.

Ecol. Elfin forest and mossy mountain forest, but mostly in open, sometimes damp situations in sedge-

or grasslands, crevices of rocks, and mountain heaths, locally often common, between (1000-)1600 and
3250 m.

KEY TO THE SPECIES

5 toe 1. A. foliolosa
oo vietsy ie 2. A. spicata

1. Perianth glabrous. Leaves mostly spreading, usually at least 4 mm broad
1. Perianth glandular-pubescent. Leaves usually erect, less than 4 mm broad

1. Aletris foliolosa STAPF, Trans. Linn. Soc. II, 4
(1894) 240; Docr. v. LEEUWEN, Trop. Natuur 9
(1920) 98, fig.; MERR. En. Born. (1921) 115; Doct.
vy. LEEUWEN, Verh. Kon. Ak. Wet. A’dam sect. II,
31 (1933) 251, f. 66 (‘foliosa’). — A. rigida STAPF,
Trans. Linn. Soc. II, 4 (1894) 241; Gras, J. Linn.
Soc. Bot. 42 (1914) 164; Merr. En. Born. (1921)
115. — ? Liriope brachyphylla MeErRR. Philip. J.
Sc. 2 (1907) Bot. 266; En. Philip. 1 (1922) 206. —
? Metanarthecium brachyphyllum (MERR.) MASAM.
Bull. Soc. Bot. Fr. 84 (1937) 18. — A. sumatrana
Masao. /.c. 18. — Meta-aletris sumatrana (MASAM.)
Masao. Trans. Nat. Hist. Soc. Formosa 28 (1938)
46. — Meta-aletris rigida (STAPF) MASAM. I.c. 46.
— Fig. 18a-—d, 19.

Leaves glabrous, ascending at first but usually
spreading when mature and finally recurved, to
6(-10) cm by 4-7(-10) mm;; veins several, close to
one another. Peduncles 1 or more, 6-65 cm high,
glabrous, rigid, with 1-6 narrow-lanceolate, sterile
bracts to 2 cm long; fertile bracts 4-20, lanceolate,
5-10 mm long; bracteoles similar to bracts but
shorter. Pedicels 0-5 mm long in flower, sometimes
over 1 cm in fruit, usually expanding gradually to

the ovary. Perianth segments white, pink, yellow
or brownish, glabrous, shortly connate, arising
near the middle of the ovary, oblong, obtuse,
21/,-31/, by c. 1/.-3/, mm. Anthers yellow to red,
1/,-1 mm long; filaments arising c. 1/; mm from
base of perianth, the base conspicuously decurrent,
1/,-2 mm long. Ovary 3-lobed, 2-2'/, mm long,
ellipsoid or obovoid; style 1-11/, mm, simple or
minutely 3-lobed, caducous. Capsule ellipsoid or
ovoid, to 7 mm long.

Distr. Malesia: Sabah (Mt Kinabalu) and
North Sumatra (Gajolands and some volcanoes of
the Westcoast Res., especially Mt Singalang),
possibly also in the Philippines (Mindoro: Mt
Halcon).

STapPr’s (1894) record from Malaya seems to be
erroneous.

Ecol. Elfin and mossy forest, but mostly in the
open mountain heath, both on stony ground and
in boggy places, between moss-cushions, 2000—
3450 m. Fl. (Jan.—_)March-Sept.(—Dec.).

In West Sumatra DocTERS VAN LEEUWEN /.c.
observed that the flowers are proterogynous and
self-pollination is the rule, all flowers setting fruit.

1979 ]

231

Se
WS

Bo
¢
4

5%

Fig. 18. Aletris foliolosa STAPF. a. Habit, x 1/,, b. flower, x 5, c. tepal with stamen, x 10, d. dehisced
fruit, x 4. — A. spicata (THUNB.) FRANCH. e. Habit, x 1/4, f. flower, x 71/2, g. fruit, x 71/,(a—-d SCHIFFNER
1700, e CLEMENS 9178, f—g JAcoBs 7426).

Notes. A. foliolosa and A. rigida were both
Originally described from Borneo, differing prin-
cipally in size, but, as pointed out by several collec-
tors in field notes (e.g. SINCLAIR and SLEUMER), the
range of intermediates is such that the characters
STAPF used to separate them break down. The
smaller form tends to occur at higher altitudes
(above 2700 m) and does not occur in Sumatra.

Plants from Sumatra often differ from those
from Borneo in having longer pedicels, but this
character is too variable to be of taxonomic
significance.

Liriope brachyphylla MeErR. is known only from
the type collection from Mt Halcon in Mindoro
(Philippines: MERRILL 5710). This collection falls
within the above description of A. foliolosa, being

232 FLORA MALESIANA [ser. I, vol. 9!

somewhat atypical in that the leaves appear softer
and the flowers fewer (up to 7 per inflorescence)
than usual. Further material is needed to decide on
the identity of the Philippine material with
certainty.

2. Aletris spicata (THUNB.) FRANCH. in Morot,
J. de Bot. 10 (1896) 199; Merr. Philip. J. Sc. 1
(1906) Suppl. 182; ibid. 5 (1910) Bot. 338; En.
Philip. 1 (1922) 207. — Hypoxis spicata THUNB.
Fl. Jap. (1784) 136. — A. japonica LAMBERT, Trans.
Linn. Soc. 10 (1811) 407, non THuNB. — Fig. 18e-g.

Leaves glabrous, usually erect, thinner than in
A. foliolosa, usually 7-20 cm by 1-3 mm; veins 3,
well-spaced. Peduncles 25-60 cm, glandular
pubescent at least in the upper part, rigid, with
6-15 narrow-lanceolate sterile bracts to 5 cm long;
fertile bracts 30-70, lanceolate, 3-8 mm long;
bracteoles similar to the bracts but shorter;
pedicels 0-1 mm, not greatly elongating in fruit,
rather abruptly expanding to the base of the ovary.
Perianth segments white, sometimes pink towards
the tip, glandular pubescent, connate to one-third
of their length, arising near the apex of the ovary,
oblong, subacute, 2!/,-3 by c. 1 mm. Anthers
orange, to c. '/, mm long, subsessile, attached near
middle of perianth. Ovary c. 2 mm long, oblong to
obovoid; style c. 1 mm, minutely 3-lobed. Capsule
obovoid, 5 mm long.

Distr. Japan, southern China, Taiwan and
N. Malesia: Philippines (N. Luzon: Abra, Benguet
and Bontoc Prov.).

Ecol. Among grasses above (1000-)1600-
Fig. 19. Aletris foliolosa StaPF. In low vegetationon 2300 m, usually in pine forest, but often in open
old lavas; Sumatra Westcoast Res., Mt Singalang, _ places. F/. Jan.—Dec.

c. 2800 m altitude (W. MEIER, 1956). Vern. Philippines: salenganga, lg.

22. ASTELIA

BANKS & SOLAND. ex R. Br. Prod. (1810) 291, nom. gen. cons.; BTH. Fl. Austr. 7
(1878) 11; KRAusE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 360; Skortss. Kong].
Svenska Vet. Ak. Handl. III, 14, 2 (1934) 1-106; VAN BaLGooy, Pac. Pl. Areas 2
(1966) 86, map 47; Moore & EpaGar, FI. New Zeal. 2 (1970) 27-38. — Funckia
WILLD. Mag. Ges. Naturf. Fr. Berlin 2 (1808) 19, nom. rejic. — Fig. 20.

Stemless or short-stemmed, dioecious herbs. Rhizome usually well-developed;
roots fibrous. Leaves rosulate, 3-ranked, linear to lanceolate, forming a closed
sheath at the base. Inflorescence a panicle; peduncles 3-angled. Pedicels solitary in
the axils of bracts, not articulated. Perianth segments connate, |- or 3-nerved, erect,
spreading or reflexed, the outer often larger than the inner. Filaments filiform or
somewhat flattened, attached to the perianth segments; anthers dorsifixed, ovoid,
introrse. Ovary superior, sessile, depressed-globose to ellipsoid, 1- or 3-celled;
ovules few to numerous, axile or parietal; style thick or absent. Fruit a berry;
perianth persistent. Seeds ovoid or angled, glossy.

1979

LILIACEAE—I (Jessop)

233

Distr. About 20-30 spp. in the Pacific (13 in New Zealand) including Australia (Victoria, New South
Wales and Tasmania), Mauritius and the Falkland Is.; in E. Malesia: New Guinea.

Ecol. Some species are epiphytic in forests; others grow on the ground or on rocks, usually in wet
areas. Several occur in bogs and may contribute to peat formation. They are to be found from sea-level
in the south but only at alpine altitudes near the equator.

1. Astelia alpina R. Br. Prod. (1810) 291; Bru. FI.
Austr. 7 (1878) 11; F.v.M. Trans. R. Soc. Vict. 1, 2
(1889) 35; Burkitt, Kew Bull. (1899) 113;
Ropway, Tasm. Fl. (1903) 212; Laur. Bot.
Jahrb. 50 (1913) 298; Krause, Bot. Jahrb. 59
(1925) 559; Skottss. Kongl. Svenska Vet. Ak.
Hand. III, 14, 2 (1934) 26. — A. papuana SkorTTssB.
lic. 29, incl. f. minor Skottss. l.c. 30; HooGt.
Blumea Suppl. 4 (1958) 235. — Fig. 20.

Erect herb, forming dense cushions. Leaves
linear to narrowly lanceolate, often with a silvery
film on the upper surface and densely covered by
silvery scales on the lower surface or glabrescent,
often with red margins, 3-13(-40) cm by 3-7
(—25) mm, with 1-3 major veins and several minor;

leaf bases persistent and forming a silvery sheath
round the rhizome. — ¢ Panicle densely covered
by silvery scales, 2-8 cm long, with up to 3 simple
branches subtended by bracts to 4(-12) cm long.
Pedicels 3-6(-15) mm. Perianth segments green,
pale yellow or brown, spreading or reflexed,
21/,4 mm long, scaly on the outer surface.
Filaments 3/,-11/, mm; anthers c. 1/, mm long.
Gynoecium sterile, green, 11/,-2'/, mm long. —
2 Panicle similar to the male but shorter, 1-5 cm
long, bracts to 3(—7) cm long. Pedicels 2—-4(-10) mm.
Perianth segments green, pale yellow or brown,
erect, 6—7'!/,(—9) mm long, glabrescent on the outer
surface. Staminodes 3/,-1!/, mm long. Gynoecium
c. 6 mm long, ovoid; style not differentiated from

Fig. 20. Astelia alpina R. BR. a. Habit, 3 plant, x '/,, b. 3 flower, x 5, c. 2 flower, x 2!/,, d. fruit, x 2
(a—b BRAss 10332, c HOOGLAND & PULLEN 5782, d Brass 10217).

234

FLORA MALESIANA

[ser. I, vol. 9!

ovary. Berry green, becoming bright red when ripe,
ovoid or ellipsoid, c. 8-13 mm long. Seeds numer-
ous, c.2 mm @.

Distr. Australia (Tasmania, Victoria, southern
New South Wales) and E. Malesia: New Guinea
(from Mt Wilhelmina in W through the highlands
of Papua New Guinea to Mt Albert Edward in
E).

Ecol. Alpine and subalpine grasslands forming
tussocks to large solid cushions in bogs, sometimes
forming almost pure communities in very wet areas
(e.g. in Pindaunde Valley), 3225-4500 m. Fig. 21.

Vern. Papua New Guinea: maunz, Habi’inz
dial., pangjubank, Enga lang., pogwe, tudik, Mendi
lang., waiamaia, Chimbu, whyameya, Kundiawa
Subdistr.

T
150

4 4 4 4 4 4 4 m — J

1 4 4 ~. 4 1 a

Fig. 21. Range of the genus Astelia (after VAN BALGOoy, in Pac. Pl. Areas 2, 1966, 87).

23. NOTHOSCORDUM

KunrtH, En. Pl. 4 (1843) 457, nom. gen. cons.; KRAUSE in E. & P. Nat. Pfl. Fam. ed.
2, 15a (1930) 322; BACK. Handb. FI. Java 3 (1924) 61; TRAuB, Plant Life 10 (1954)
123; BACK. & BAKH. f. FI. Java 3 (1968) 132.

1. Nothoscordum inodorum (W. Ait.) G. NICHOLS.
Ill. Dict. Gard. 2 (1885-89) 457; A. & G. Syn.
Mitt. Eur. Fl. 3 (1905) 167; BAck. Handb. FI. Java
3 (1924) 62; Back. & SLoot. Handb. Theeonkr.
(1924) 89, fig.; TRAUB, Plant Life 10 (1954) 127;
BAcK. & BAKH. f. FI. Java 3 (1968) 132. — Allium
inodorum W. Air. Hort. Kew. 1 (1789) 427. —
Allium fragrans VENT. Jard. Cels. (1800) t. 26. —
N. fragrans KUNTH, En. PI. 4 (1843) 461.

Erect, inodorous, glabrous herb with a subter-
ranean coated bulb. Leaves radical, linear, flat,
somewhat glaucous, shorter than the peduncle,
15-45 by '/,-11/, cm. Umbels borne on an erect,
20-70 cm long peduncle, 6-17-flowered. Flowers
fragrant. Pedicels 11/,-5 cm. Tepals basally shortly
connate and green, 1—1', cm, persistent, the seg-
ments oblong, 1-nerved, white, whether or not
with a purple median streak. Stamens 6, inserted

1979]

on the base of the perianth; filaments ligulate with
a subulate top; anthers medifixed. Ovary oblong-
obovoid, sessile, 3-celled, each cell with ~ ovules;
style filiform; stigma small. Capsule obovoid, mem-
branous, loculicidally 3-valved, 8-9 mm long.
Seeds several, black, often containing more than |
embryo.

Distr. Native of subtropical North America,

LILIACEAE—I (Jessop)

235

Mediterranean), in Java sometimes cultivated as
an ornamental, escaped from the Tjibodas Botanic
Garden in West Java on Mt Gedeh.

Ecol. Locally naturalized, in W. Java (Priangan)
sometimes occurring in great numbers in fields,
tea-estates, and along roadsides; 1000-1500 m. FI.
Jan.—Dec. Easily propagating by bulbils and seed,
difficult to eradicate.

often cultivated and naturalized (e.g. in the Vern. Java: babawangan, S.

Excluded

Astelia novoguineensis KRAUSE, Bot. Jahrb. 59 (1925) 559 is according to SKOTTSBERG (Bot. Jahrb. 65,
1932, 260) = Helmholtzia novoguineensis (KRAUSE) SKOTTSB. (Philydraceae).

Laxmannia sessiliflora DECNE, Herb. Timor. Descr. (1835) 35, t. 16; SPAN. Linnaea 15 (1841) 477; Mia.
Fl. Ind. Bat. 3 (1859) 66; Bru. Fl. Austr. 7 (1878) 66. This was said by MIQUEL to occur both in West
Australia and Timor, but the latter addition is a slip of the pen. Cf. STEEN. Bull. Jard. Bot. Btzg III, 17
(1948) 463.

Veratrum malayanum JACK, Mal. Misc. 1 (1820) 25; in Hook. Bot. Misc. 2 (1831) 74. — Veratronia
malayana (JACK) Mia. FI. Ind. Bat. 3 (1859) 553 is, according to Kurz (Flora 56, 1873, 224) and MERRILL
(cf. Fl. Males. I, 4, 1951, 249) = Hanguana malayana (JACK) MERR. (Flagellariaceae).

Xerotes arenaria R. Br. and X. longifolia R. BR. were cited by MIQueL (FI. Ind. Bat. 3, 1859, 248) to
occur in Java, obviously both based on specimens collected by ZOLLINGER on Mt Salak, in West Java.
Cf. ZOLLINGER, Syst. Verz. 1 (1854) 66. According to Kurz (Flora 56, 1873, 224) these are misidentifica-
tions for Hypolytrum, assumedly H. nemorum (VAHL) SPRENG. (Cyperaceae).

my ie ‘3 1 : 7
bn . ' ‘en Pe efron os ° rr

wii -: : Fk ; a8 Zz
ae Py eM VT he ee Ve : y ; wwe .
BAA Oe ty Mus, Ae cae, j hid are Vs a We nd
: ae at Ss ieee 1 i a ) 4) eae ee n oo i ab ie, it! 10 yore =)
/ viv = 7 Pore er thi S ” a ee i . 4 =a?
i ae aks : 4 NAD es sth ? m9) ay “Toe
ee rd ENTE CSE RS CMR
. , ee

Nes a, it ae hr

“lo awry on Par 1G. Hacer nk; eae .
anit aes a ow he Bra. vy : 9 the. oi cal
Siar fixe, i". dues | 66 ceculhh pa len cand el aati
ae r, “Si.gS ta Op cehey Weteeas
5-70 bom 7

DIPTEROCARPACEAE
(P.S. Ashton, Arnold Arboretum, Harvard University )!

Small or large resinous usually evergreen trees, usually buttressed, and often (if
large trees) with flaky or fissured bark. Some or most parts with a tomentum of
fascicled hairs, or sometimes single hairs, unicellular or multicellular glandular
hairs, or multicellular, short or long lobed or peltate hairs. Leaves alternate,
simple, margin entire or sinuate, not crenate, terminating + abruptly at the + pro-
minent geniculate petiole, penninerved (in Dryobalanops and some Hopea nerves
co, dense and slender), often with domatia in axils between nerves and midrib or
along midrib and (rarely) nerves; tertiary nerves scalariform or reticulate. Stipules
paired, large or small, persistent or fugaceous, leaving small to amplexicaul scars.
Inflorescence paniculate, racemose, rarely cymose, + regularly, rarely irregularly,
branched, terminal or axillary; bracts and bracteoles paired, small or large,
persistent or fugaceous. Flowers secund or distichous, bisexual, actinomorphic,
scented, nodding. Ca/yx persistent, 5-merous; 2—5 sepals usually greatly enlarging
into wing-like lobes in fruit; sepals either free to base, imbricate in bud, remaining
so or becoming valvate in fruit, or fused at base, forming a cup or tube + enclosing
the fruit, adnate to or free from it. Corolla 5-merous, contorted, base connate or
free, usually partially or entirely unicellular hairy. Stamens 5-110, 1—3 verticillate
or irregular, hypogynous or subperigynous, centrifugal; filaments compressed or
filiform, free or connate, frequently cohering with petals on falling; anthers erect,
2-celled with (2—)4 pollen sacs, introrse or laterally dehiscent; tapetal cells
binucleate, pollen grains 2-celled at anthesis; connective with short or prominent
appendage. Ovary superior or semi-inferior, 3-, rarely 2-, locular; style + thick-
ened at base into a stylopodium, entire or trifid towards apex; stigma obscure or
prominent, 3- or 6-lobed. Ovules 2(—3) in each loculus, axile, pendulous or
laterally anatropous, bitegmatic with ventral raphe and superior micropyle. Fruit
indehiscent, l-seeded; with woody pericarp and persistent+aliform sepals.
Embryo-sac development of Polygonum type: endosperm of the nuclear type,
embryo development normal, ripe seeds with or more usually without endo-
sperm; cotyledons equal or more usually unequal and with one more or less
enclosing the other, laminar or fleshy, entire or lobed, enclosing the radical.
Germination epigeal or hypogeal; pericarp splitting irregularly or along 3 sutures.

Distribution. The newly described monotypic genus Pakaraimaea MAGUIRE & ASHTON (1977),
locally found in the south of former British Guyana, makes the family pantropical, confined to the
lowlands and hills of the tropics below 1800 m. Fig. 2. This genus represents a distinct subfamily
Pakaraimoideae.

The second subfamily, Monotoideae is represented in Africa and Madagascar, with some 36 spp.
of Monotes A.DC. and a few species of Marquesia GILG (cf. BANCROFT, 1935).

Subfamily Dipterocarpoideae, comprising 13 genera and some 470 spp. ranges from the Seychelles
through Ceylon (where a proportionally large diversification exists) to the south of Peninsular
India, and then to East India, Bangladesh, Burma, Thailand, Indo-China, to continental S. China
(Yunnan, Kwangsi, S. Kwantung, Hainan) and through Malesia southeastwards to the D’Entre-
casteaux Is. off S.E. Papua (not in New Britain and New Ireland), and northwards to the Batan Is.
north of Luzon, Philippines.

(1) With some co-operation of the General Editor for the general chapters.
Drawings by Miss R. van Crevel (details) and Mr. J. van Os (habit) were made under supervision of Dr. M.
Jacobs.

pie ¥ |

238 FLORA MALESIANA (ser. lL volg@-

Fig. 1. Characteristic habit of the large Dipterocarpaceae with high unbranched bole and huge dome-shaped
crown: Shorea rubella ASHTON, from Brunei (Photogr. ASHTON).

1982] DIPTEROCARPACEAE (Ashton) 239

Fossils do not significantly extend subfamilial range in Asia and Malesia, but they do essentially
so in East Africa. Fig. 2.

In Malesia 10 genera with 386 spp. occur, predominantly in the humid non-seasonal areas, absent
only from the seasonal area encompassing the Lesser Sunda Is. east of Sumbawa as far as the
Tenimber Is.

The local species diversity of these genera is very uneven, with a tendency to decline eastwards, as
is shown by the total number of species per island. Fig. 3-4. See also Fig. 19, 28, 43, 65, and 79.

Three of the 10 genera are endemic in Malesia, viz the monotypic genus Upuna in Borneo,
Neobalanocarpus in Malaya (& Pattani adjoining Kelantan in N. Malaya), and the genus
Dryobalanops (7 spp.) on the Sunda shelf (Sumatra, Borneo, Malaya); the 7 others Malesia shares
with continental Asia, and Ceylon (except Anisoptera and Parashorea). A further three are endemic
to Southern India, Ceylon and the Seychelles.

Four genera range widely through Malesia and also have species (mostly few) in East Malesia
(Celebes, Moluccas, and New Guinea), viz Anisoptera (11 spp., 10 in Malesia), Vatica (65 spp., 55 in
Malesia), Hopea (102 spp., 84 in Malesia), and Shorea (194 spp., 163 in Malesia).

Of the remaining three Cotylelobium (6 spp., 3 in Malesia) is known in Malesia only from the
Sunda shelf islands, while Dipterocarpus (69 spp., 53 or 54in Malesia) and Parashorea (14 spp., 10in
Malesia) occur on the Sunda shelf islands, but also in the Philippines.

Some of the Malesian genera formerly had in the Tertiary a wider distribution, e.g. Dryobalanops
occurred in West Java and Southern India, Dipterocarpus in N.E. Africa, and Anisoptera (now only
from Chittagong and Burma southeastwards) in India.

It is noteworthy that there are hardly any clear disjunctions in the generic ranges (apart of course
from seas separating adjacent islands), the exception being that of Cotylelobium, with 1 sp. in
Ceylon and further from S. Thailand to West Malesia which stems obviously from extinction.
Vateria and its close ally Vateriopsis are confined to Ceylon and the Seychelles respectively; this
huge oversea gap must be ascribed to ancient geomorphological processes.

Species ranges. | have (1972) discussed the relationship between the ecological and geographical
ranges of dipterocarps, and speculated on their evolutionary significance. Patterns of Malesian
dipterocarp distribution can be summarised as follows:

Widespread species. These form 4 principal categories: (1) The first, those which occur both in
seasonal and non-seasonal zones, often tend to be gregarious in the former and occur there on a
wider range of substrates; but in the latter they become local, scattered, and confined to deep fertile
soils where leaching is least apparent, especially on basic and intermediate igneous rocks, calcareous
shales, and around the base of limestone hills. Shorea assamica, S. guiso, Dipterocarpus gracilis, D.
hasseltii, D. kerriiand Anisoptera costata serve as examples. — (2) The second category are those of
wide West Malesian distribution in the non-seasonal zone, often including the Philippines; these
occur on the granite of the Sunda core but spread through the region on other igneous rocks and on
the deep clay soils of the shale and phyllite ridges of the great sedimentary geosynclines. — (3)
Thirdly are those of freely drained yellow/red soils prone to drought and of moderate to low
fertility, mostly of coastal hills and islands though some extend up inland ridge-tops; coastal N.W.
Borneo is the present centre of distribution, but a relict distribution which frequently includes
eastern coastal Malaya, and less frequently Riouw, Perak (W. Malaya), S. Borneo and rarely E.
Sumatra suggests a more extensive former continuity of this habitat. — (4) Lastly are the minority
of riparian species, many of which may be of recent origin, being rapidly dispersed by freshwater by
means of their floating fruits.

What is said about the coherence of generic ranges holds also for species ranges which rarely show
disjunct distributions (Dipterocarpus retusus, the continental D. turbinatus GAERTN. f., and Shorea
roxburghii are notable exceptions), though several Ceylon species are vicarious with others in
Malesia and similar distant vicarism occurs on either side of Wallace’s line.

Endemicity. Dipterocarpaceae show a high rate of endemicity in the non-seasonal humid tropics
not reflected in the more seasonal parts of their range. Fig. 4. This may be ascribed to their poor fruit
dispersal in a windless climate, allowing easy isolation by natural barriers such as quite small river
valleys (ASHTON, 1969, 1972), and rapid edaphic specialisation. Endemicity, both local and
island-wide, is greatest in New Guinea (73% of 15 spp.) and Borneo (59% of 267 spp.), and high in
the Philippines (465°% of 45 spp.); in Sumatra (113% of 96 spp.) and Malaya (19% of 156 spp.) it is

240 FLORA MALESIANA [ser. I, vol. 92

Fig. 2. Range of the Dipterocarpaceae: Dipterocarpoideae (line and 2 fossil sites in E. Africa), Monotoideae (2
genera, Afro-Madagascan, interrupted line, dots Monotes, squares Marquesia), Pakaraimoideae (monotypic
genus in northern South America).

Fig. 3. Density map of Dipterocarpaceae in Malesia, total number of species in each island.

Fig. 4. Density map of Dipterocarpaceae in Malesia, segregated into endemics (above the hyphen) and
non-endemics (below the hyphen).

surprisingly comparatively low. Explanations may possibly be sought in the central position within
the Sunda shelf and the relative edaphic uniformity of Malaya and Sumatra, the prevailing
archipelagic condition of the Philippines with only intermittent connections with Sundaland during
the late Tertiary and Quaternary period, and the youthful physiography and hence edaphic
diversity of New Guinea and Borneo. The close proximity of all Borneo to, and frequent absorption
into, Sundaland and its central position in western Malesia have probably also led to periodic
enrichment by immigration followed by local evolution in ensuing periods of isolation.
Literature: ASHTON, Biol. J. Linn. Soc. | (1969) 155; Proc. 2nd Aberdeen/Hull Symp. Mal. Ecol.

1982] DIPTEROCARPACEAE (Ashton) 241

(1972) 35; BANCROFT, Am. J. Bot. 22 (1935) 511, f. 1 (map); MAGuiIRE & ASHTON, Taxon 26 (1977)
341.

Fossils. Fossil data of the family have been surveyed by R.N. LAKHANPAL (1974). Fossil wood
and leaf impressions which have been identified with the Dipterocarpaceae, and which anatomically
apparently differ little from present day genera, have been recorded from both East Asia and Africa.
The winged specimens, apparently fruits, called Calycites, from the Upper Cretaceous strata of the
east coast of the United States, and compared by BERRY (1914) to Dipterocarpaceae, and Woburnia,
a similar structure from the Lower Cretaceous of Bedfordshire (England) similarly compared by
Stopes (1912) are both considered by BANCROFT (1933) to be too incomplete for determination.
SCHWEITZER (1959) who studied and surveyed the records of fossil wood of Dipterocarpaceae
considered, however, that the Woburnia wood is distinctly Dipterocarpaceous and does not differ
very much from recent woods. WOLFE c.s. (1975) correctly pointed out that on the Woburnia site no
other samples could be found and concluded, it seems correctly, that the specimen must have been
mislocalised.

Rasky (1956) claimed to have found a flower in the Upper Miocene of Budapest, referred to
Monotes (CHANDLER, 1964: 81). WEYLAND (1964) mentioned that fruit fragments from the
Oligocene of the Zevengebergte and the Miocene of Oehningen would belong to the fossil Monotes
macranthus (HEER) WEYLAND. KIRCHHEIMER (1957) was earlier of the opinion that these remains are
too inadequate to warrant inclusion in Monotoideae. The only authenticated fossil fruit is that of
Dipterocarpus verbeekianus, recorded by HEER (1874, 1883) from the Tertiary in Sumatra.

Leaf impressions. HEER (/.c.) recorded Dipterocarpus leaf impressions from the Tertiary in
Sumatra; later authors have confirmed his conclusions. Crié (1888) recorded Pliocene leaf
impressions of a possible Dipterocarpus from Java, while SCHUSTER (1911) identified Pleistocene
Javanese impressions as Hopea fagifolia (H. sangal) and Vatica lanceaefolia BL. EDwARDs (1923)
recorded the modern Philippine species Anisoptera thurifera, Shorea guiso and S. polysperma from
the Pliocene in Luzon. The records of dipterocarp leaves from the Tertiary of Labuan, North
Borneo by GEYLER (1887) are not considered by BRANDIS (1895) or EDWarbs (/.c.) to merit
inclusion. WOLFE (1972) claimed to have found an Eocene (para-)tropical flora in Alaska, in which
also leaf impressions of Anisoptera occurred. Such a geographically remote record must require
substantiation by more unequivocal material.

Wood. KRAUSEL (1922, 1926) has recorded undoubted dipterocarp woods from the Tertiary
(claimed then to be Miocene) of southern Sumatra and West Java, which have been associated by
DEN BERGER (1923, 1927) with Shorea and Dryobalanops, his D. spectabilis and D. javanica being the
only records of that genus from Java. KRAUSEL’s Caesalpinioxylon palembangense is considered by
DEN BERGER, and later KRAUSEL himself, to be Shoreoxylon. Dipterocarpoxylon annamense of
CoLani (1919), from the Tertiary of Annam, is only tentatively accepted by EDwarps. The
specimen of Dipterocarpoxylon described by HOLDEN (1916) from Burma possesses uniseriate rays,
unlike all Asiatic genera, and does not bear secretory canals throughout the wood; hence the
identity is insufficiently grounded, but PRAKASH (1973) has subsequently confirmed the fossil genus
from Burma. Shoreoxylon has been identified from the Tertiary of Assam by EybE (1962), while
Miocene fossil woods from the east coast of Southern India have been identified by NAVALE (1962)
as Anisopteroxylon, Dipterocarpoxylon, Hopeoxylon, and Shoreoxylon and by him and AWASTHI
(1971) as Dryobalanoxylon. Fossil wood from Quaternary sites, c. 10,000 years old, from the Siwalik
hills, N. India, and from W. Bengal have been identified with Anisoptera by GHOSH & GHOSH (1958),
and GHOSH & KAZmI (1958).

SCHWEITZER (1959), who made a major study of fossil wood of Dipterocarpaceae, gave maps of
fossil sites. It is interesting that he identified fossil wood from Timor as Shoreoxy/on — an island
where the family does not now occur — and that he recorded Dryobalanoxylon from several places
where the genus is at present absent, viz Cambodia, S. Sumatra, W. Java, and Ambon in the
Moluccas.

It is agreed that no fossil wood is found older than the Miocene.

Pollen. J. MULLER’s studies of pollen (1970) indicate that the thin walls do not preserve well, and
the lack of distinctive sculpturing or other diagnostic characters combines to limit their value in the
fossil record. It is noteworthy that he found the pollen percentage proportionally always low, even
in stands on peat completely dominated by Shorea albida where it comprised only a few percent.

242 FLORA MALESIANA [ser. I, vol. 9?

This is against expectation, and the reason is unclear. MULLER suggested that much pollen is
devoured by thrips, but then the persistent exines should be found.

CHANDLER (1964: 36) mentioned (with a question mark) Oligocene pollen from the London Clay
Flora, but she appeared far from assured of this identification as she remarked later (/.c.: 56) that she
found the apparent absence of Dipterocarpaceous fossils strange. The identity of the pollen record
(by MA KHIN SEIN) is very much doubted by J. MULLER.

MULLER (1964) found pollen of Dipterocarpus and Dryobalanops in the Tertiary of Borneo (Brunei),
the first genus being represented from the Oligocene (MULLER, 1970), the latter from the Miocene.

The fossil record of Dipterocarpaceae in Asia and Malesia rests therefore on the wood fossils of
several genera in India and West Malesia from the Tertiary (mostly Miocene and later) and the
pollen records of the same age, but also on the presence of pollen of Dipterocarpus already in the
Oligocene of Borneo.

Northeast African fossils. Fig. 2. In 1933 BANCROFT described Dipterocarpoxylon africanum from
the Tertiary beds near Mt Elgon (Kenya). In the same year CHIARUGI described three species of
Dipterocarpoxylon from the ?Plio-Pleistocene beds of Somaliland which SCHWEITZER (1959) later
considered to be identical with D. africanum BANCROFT. The fossil woods collected by WEYLAND
(1964) from the volcanic tuffs of Mt Elgon, of uncertain date but probably of late Tertiary and
pre-Pleistocene times, also undoubtedly represent dipterocarp material and must be associated with
the Asiatic subfamily and show strong evidence of being congeneric with Dipterocarpus, supporting
CHIARUGI's identification. In 1935 SEwarD had described a few leaf impressions from the Nubian
sandstone of Egypt as Dipterocarpophyllum humei and D. zeraibense; the age of the strata being
uncertain but probably Tertiary. Finally LAKHANPAL (1974) mentioned that Prof. Y. LEMOIGNE had
told him that he had found a dipterocarp wood in an Upper Tertiary deposit in Ethiopia to be
described in a future paper.

Concluding, it is clear that Dipterocarpoideae were present in East Africa at least in the Upper
Tertiary. It is difficult to imagine that they would have been derived from the Indian subcontinent,
as according to geophysical theory this rafted block of land had by then hardly joined the Asian
plate while, moreover, Dipterocarpus could hardly have migrated through the desert zones of S.W.
Asia. Thus, even the poor fossil record presents formidable implications. The alternative is that
Dipterocarpoideae were already earlier represented in N.E. Central Gondwanaland. It is a pity that
the older fossil record in S.W. Asia and N.E. Africa is so scant, as this area is vital for understanding
exchange of the Gondwana element and tropical S.E. Asia.

Literature: AWASTHI, Palaeobotanist 18 (1971) 229; BANCROFT, Foren. Forhandl. 55 (1933) 59;
Am. J. Bot. 22 (1935) 164; DEN BERGER, Verh. Geol.-Mijnb. Gen. Ned. & Kol. 7 (1923) 143; Bull.
Jard. Bot. Btzg III, 8 (1927) 495; Berry, U.S. Geol. Surv. prof. paper (1914) 84; BRANDis, J. Linn.
Soc. Bot. 31 (Nov. 1895) 243; CHANDLER, London Clay Flora, Suppl. London, 1961, The Lower
Tertiary Floras of Southern England. IV. Summary & survey of findings in the light of recent
botanical observations (1964) 36, 56; CHIARUGI, Palaeontographica Hal. 32, n. 1 (1933) 106;
COLANI, Bull. Serv. Geol. Indochine 6, n. 3 (1919) 2; Criz, Samml. Geol. Reichsmus. Leiden I, 15
(1888) 1; Eowarps, Geol. Mag. 60 (1923) 159, 409; Eypg, Palaeobotanist 11 (1962) 115; GEYLER,
Vega-Exp. Vetensk. iskttagelser 4 (1887) 473; S.S. GHosH & A.K. GHOSH, Sci & Cult. 24 (1958) 238;
S.S. GHOSH & KAzMI, Sci. & Cult. 23 (1958) 485-487; Heer, Abh. Schweiz. Palaeontolog. Ges.
(1874); Neue Denkschr. Allg. Schweiz. Ges. Gesammt. Naturw. 28 (1883) 1; HOLDEN, Rec. Geol.
Surv. India 17 (1916) 47; KIRCHHEIMER, Die Laubgewachse der Braunkohlzeit (1957) 432; KRAUSEL,
Verh. Geol.-Mijnb. Gen. Ned. & Kol. 5 (1922) 231; Proc. Kon. Ak. Wet. A’dam, sect. sc., 25 (1922)
9; Leid. Geol. Meded. 2, n. 1 (1926) 1; LAKHANPAL, Birbal Sahni Inst. Palaeobot. Spec. Publ. |
(1974) 30-39; MuLLer, Abstr. 10th Int. Bot. Congr. Edinb. (1964) 271; Biol. Rev. 45 (1970) fig. 5
facing p. 435; NAVALE, Palaeobotanist 11 (1962) 66; PRAKASH, Palaeobotanist 20 (1973) 48-70;
RASKY, Fossil plants from the marl formation of the environs of Budapest; Féldt. K6zl. Budap. 86
(1956) 167-179; ScHUsTER, Abh. K. Bayer. Ak. Wiss. M.-Ph. KI. (1911) 25; SCHWEITZER,
Palaeontographica 105B (1959) 1-66; SEwARD, Leaves of Dicotyledons from the Nubian Sandstone
of Egypt; Min. Finance Surv. Dept. Egypt (1935) 1-21; Stopes, Phil. Trans. R. Soc. B 203 (1912) 75;
WEYLAND, Lehrbuch der Palaobotanik ed. 2 (1964) 455; WoLFE in Graham (ed.), Floristics and
Paleofloristics of Asia and Eastern North America (1972) 207-208, pl. 3, f. 3; WOLFE c.s. Ann. Mo.
Bot. Gard. 62 (1975) 819.

1982] DIPTEROCARPACEAE (Ashton) 243

Ecology. Overall range. Confined to tropical climates with a mean annual rainfall exceeding
1000 mm, and/or a dry season of less than 6 months. Most species do not occur above c. 1000 m
where an important floristic transition occurs; a few are +entirely confined above 700 m. Above
1500 m records are spurious: a Shorea sp. in N. Sumatra at 1700 m and Hopea beccariana BURCK in
Brunei at 1750 m. The subfamily geographical range is divisible into three climatic zones:

Savanna zone. A limited number of Indo-Burmese species are fire-tolerant: Shorea robusta, the
Indian Sal, S. roxburghii and the Indo-Burmese S. obtusa, S. siamensis, Dipterocarpus obtusifolius,
D. tuberculatus and D. intricatus. They are distinguished by their thick bark (the latter 3 are the only
fissured members of their genus) and are the only dipterocarps to be deciduous for a more or less
prolonged period during the dry season. They form single species or codominant stands on the
well-drained plains, and dry hills (occasionally to 1400 m) of Central and East India, Burma,
Thailand and Indo-China. Generally the fire-climax savanna woodlands (Dry Dipterocarp forests
of CHAMPION, 1936) in which they occur grow on infertile, yellow skeletal hill soil and laterite
bearing red soils in the plains in areas with less than 2000 mm mean annual rainfall and 3-5 months
dry season; these forests do not exist in Malesia, but Shorea siamensis and Dipterocarpus obtusifolius
occur, on dry rocky headlands and old secondary forest (schima-bamboo forests of SYMINGTON,
1943) respectively in Perlis, N.W. Malaya, where there are regularly 1-2 consecutive dry months at
the end of each year. In the foothills of the sandstone mountains of Arakan and southern Cambodia
the Dry Dipterocarp forests themselves extend, probably following the advent of man-induced fire,
into areas with a dry season of but |—2 months and more than 2000 mm rainfall, on porous yellow
podsolic sands; here they are floristically impoverished, and of the dipterocarps only D. intricatus
and D. obtusifolius remain. They flower annually shortly before, during, or after leaf fall during the
dry season, and the ripe fruit fall after the coming of the following rains. Regeneration is abundant
in disturbed forest; the saplings are cut back annually by fire and drought, the roots often becoming
deep and extensive before a permanent leader is established. Mature trees sucker readily in response
to damage. At the more humid climatic and edaphic ends of their range these species become more
shortly deciduous or evergreen.

Malesian species occurring in the seasonal evergreen forest zone (there are three categories, species
of wide distribution, i.e. occurring in both seasonal and aseasonal evergreen forest, species confined
to seasonal forest marked with *, and species occurring on drought-prone sites in the aseasonal
tropics marked with +):

ANISOPTERA ferrea* farinosa*
costata forbesii* gratissima*
scaphula* glabrifolia* guiso
thurifera gregaria* henryana*

COTYLELOBIUM griffithii hypochra*
melanoxylon helferi* laevis

DIPTEROCARPUS iriana montigena*
baudii* malibato negrosensis
costatus * novoguineensis palosapis
dyeri* odorata* polita
gracilis pedicellata polysperma
grandiflorus * philippinensis roxburghii*
hasseltii pierrei* selanica*
kerrii* plagata siamensis*
littoralis* sangal VATICA
obtusifolius* ultima* bantamensis*
philippinensis* PARASHOREA cinerea*
retusus malaanonan flavovirens*

HOPEA stellata* lowii
acuminata SHOREA mangachapoi*
bilitonensis* assamica odorata
cagayanensis* contorta pachyphylla*
celebica* falcifera rassak

244 FLORA MALESIANA [ser. I, vol. 9?

Seasonal evergreen forest zone. The vast majority of dipterocarps are therefore confined to areas
where the mean annual rainfall exceeds 2000 mm. The presence of even a short but regular dry
season, which in N.W. Malaya, for instance, hardly exceeds a month, has a profound influence on
both number and kinds of species present. 62 Malesian species occur where there is a regular season
of at least one month with less than 100 mm rainfall, 25 appear to be confined there, and a further 9
extend into the aseasonal zone only on sandy, coastal and skeletal ridge soils prone to water stress:
these figures exaggerate the species diversity of dipterocarps in any one region, for all but 7 of the
species occur in only one of the three Malesian seasonal regions: the north-west of the Malay
Peninsula and northern Sumatra, seasonal parts of the Philippines, and the Moluccas and New
Guinea. The relative floristic poverty of dipterocarps in seasonal evergreen forest is reflected in the
flora in general: STAMP (1925) cited merely “at least 1000 tree species’ in these forests 1n all Burma; an
estimated 400 species occur in the seasonal evergreen forests on the southern flanks of the
Cardamom mountains in southern Cambodia; by contrast 2,500 occur in the non-seasonal parts of
lowland Peninsular Malaya. No estimates of the total species diversity of the seasonal evergreen
forests in Malesia exist. Such areas are widespread in the Philippines, S. Celebes, some local parts of
the Moluccas, and S. New Guinea. It is noteworthy that no fire-resistant or deciduous species of
dipterocarps occur in Malesia, such as are known from certain tracts in continental S.E. Asia with
Dry Dipterocarp forest.

East of Sundaland the dipterocarp flora becomes increasingly impoverished. See Fig. 3, 4, 19, 28,
43,65, and 79. The fact that even in the non-seasonal New Guinean lowlands only a single section of
Hopea has undergone an ebullition of speciation suggests that the poverty of dipterocarp species
east of Wallace’s line is in part due to probably geologically recent immigration there; clearly
Makassar Straits must have been a formidable barrier, although it is, at its narrowest point only a
mere 125 km wide, and southwards not exceeding 275 km. Two features underline this: firstly that
Borneo possesses over 250 species and Celebes only 8, which can never be ascribed to climatic
difference between Borneo and Celebes; secondly, of these 8 species there are 2 endemic (Hopea
celebica and Vatica flavovirens}, 2 are shared with the Moluccas (Hopea gregaria, Shorea
montigena); Shorea assamica 1s a wide, from Assam to Malaya and Sumatra, with ssp. philippinensis
in the Philippines, and ssp. koordersii in Celebes, the Moluccas and Philippines, and some doubtful
sterile specimens in S.E. Borneo (represented in aseasonal parts of Borneo by the vicariant S.
agamii); Anisoptera costata ranges from Burma to Borneo and Mindanao (one record); A. thurifera
occurs in the Philippines, Celebes, Moluccas, and New Guinea; the last, Vatica rassak, ranges from
Borneo to New Guinea, but is also found in the Sulu Archipelago joining Borneo and the
Philippines just north of Celebes. None of these species which occur both west and east of Wallace’s
Line is not also found in the Philippines, suggesting that they spread along the Philippines over
Wallace’s Line and avoided Makassar Straits by marching northwards around it. In the south a
single species, Dipterocarpus retusus transgresses Wallace’s Line from Bali through Lombok to
Sumbawa; if the fossil record of Shorea in Timor is correct too, a southern route eastwards via Java
and the Lesser Sunda Islands, avoiding Makassar Straits, might also be imagined.

The tendency towards gregariousness persists into the seasonal evergreen forest zone, but is there
more pronounced on impoverished soils and in areas probably colonised in geologically recent
times. An example of the former are the Dipterocarpus costatus dominated forests on the
impoverished sandstone soils of Arakan and southern Indo-China; this species also occurs between
700-1100 m in the mountains of S.E. Asia, but in non-seasonal Malaya ceases to be gregarious. This
differential in gregariousness occurs also with several other widespread species, including
Anisoptera costata, A. scaphula, A. thurifera, Dipterocarpus gracilis, Shorea assamica, S. contorta, S.
guiso, and S. hypochra. In eastern Malesia several species are gregarious (see VAN SLOOTEN, 1952).
The majestic stands of kayu bapa (Shorea selanica) on the hills of Buru below 1000 m and on the Sula
Is. are celebrated. Shorea assamica ssp. koordersii is gregarious in Celebes, the Sula Is. and Obi; S.
montigena becomes locally gregarious in Buru; Anisoptera thurifera is gregarious both in the
seasonal zone of the Philippines and in New Guinea where it is grouped along the crests of
steep-sided ridges below 800 m over large areas; in New Guinea Hopea forbesii is also often
gregarious in the south-east.

Dipterocarp populations in these seasonal evergreen forests flower more or less regularly
annually too, from November to March in the Indo-Burmese region including N.W. Malaya (e.g.

1982] DIPTEROCARPACEAE (Ashton) 245

BurGeEss, 1972) and also the Philippines. Flowering, and even more fruit set, is heavier in some years
than others; not all individuals flower; in Thailand less than half the trees in a stand that are of
flowering age normally flower in any one season and a minority of these flower in two consecutive
seasons (SMITINAND’s and personal observations). The developing fruit are much infected by beetle
larvae while parakeets favour the ripe fruits. In Dipterocarpus many seasons may consequently pass
without a single seedling becoming established. Germination is immediate on falling. Several
species are shortly deciduous in seasonal evergreen forests including some, such as Dipterocarpus
gracilis and Anisoptera costata which are evergreen in non-seasonal western Malesia, but none are
deciduous for a prolonged period.

Mountains higher than 1000 m along the coasts and around the head of the great valleys of the
seasonal Far East, even at latitudes higher than the normal range of lowland evergreen forests,
collect cloud and comparatively humid though still seasonal conditions prevail on their slopes. The
species occurring at these latitudes in the Indo-Burmese region and Java, Dipterocarpus costatus, D.
gracilis, and D. retusus, occur also in the mountain forests of Malaya.

Non-seasonal humid zone. As this is the zone to which the great majority of Asian, including
Malesian, dipterocarps are confined a review of the biological characteristics of the family here must
prelude considerations of climatic and edaphic ranges of the species and their overall role in the
forest communities.

Reproductive biology. A most important characteristic of the family in the non-seasonal zone is its
flowering behaviour. Flowering does not occur annually, but at +irregular intervals and then of
varying intensity though gregariously, species from most or all genera flowering in a single season
(this includes those species that occur, and flower annually, in seasonal evergreen forests) (Woop,
1956; McC ure, 1966; Mepway, 1972; COCKBURN, 1975). In a heavy flowering nearly all species in
an area may flower, and the majority of individuals; more frequently probably less than half the
individuals flower, though critical observations of this kind have only been made by BurRGEss in the
semi-gregarious species Shorea curtisii (1972). Gregarious flowering may occur in a single river
valley or throughout a region as large as N.E. Borneo. The 1955 general flowering in Sabah was
observed by Woop, when over 2/3 of the 200 species then known in that state were collected fruiting,
and when the only area not to experience a general flowering was the extreme south-east. His
description summarises the main features seen in dipterocarp flowerings elsewhere in this zone. All
genera flowered concurrently over a period of a few weeks in a single area, though there was a slight
delay west of the Crocker range compared with the east, and one of as much as two weeks towards
the upper altitudinal limits both in exclusively montane species (some of which flowered poorly or
not at all) and in species of wide ecological amplitude. At the height of the flowering in Sepilok
Forest Reserve, Sabah, ‘the ground appeared to be carpeted in snow and the scent of the flowers
pervaded the jungle’. Detailed observations showed great variation in the period during which
anthesis occurred between closely related species; in some sections of Shorea it continued over a
period from May until as late as August in the lowlands and November in the mountains. ASHTON
(1969) observed that anthesis of individual species in Shorea sect. Richetioides in Andulau Forest
Reserve, Brunei hardly overlapped, being sequential through the flowering season. CHAN &
APPANAH (1980) have demonstrated, by meticulous phenological study of numbered wild trees, that
species which flower early do so within 15 days, but flowering of the last species is over a period of to
25 days. Woop also observed that some, such as Dryobalanops lanceolata, flowered heavily over a
short period, whereas in the taxonomically isolated Shorea smithiana anthesis occurred over a very
long one; in S. curtisii stands it may continue over several months (BuRGEss, 1972). Among the main
canopy and emergent species Woop noted that flowering was general and heavy over the whole
crown, but in understorey species of e.g. Hopea and Vatica flowering was more sporadic and often
restricted to a few branches. Young trees of canopy species do not flower at all until their sympodial
crown is developed in direct sunlight. Woop found that some species failed altogether to flower,
several of which in Parashorea and Vatica were known to have flowered the previous year. It was of
particular interest that, of the species that did not flower, the majority were either montane or in the
peat swamps. In the latter habitat, mostly confined to the west coast, no dipterocarp flowered. From
forestry records over many years and personal observations of flowering periodicity in Shorea
curtisii and related species BURGESS concluded that local flowering is more frequent than generally
supposed, and this has been supported by others (e.g. ASHTON et a/., 1979), though fruit set rarely

246 FLORA MALESIANA [ser. I, vol. 97

occurs following a minor flowering. Closely related species differ in flowering periodicity.
Dryobalanops aromatica and Shorea leprosula populations, for instance, flower on average once
every two years, while S. parvifolia flowers only half as frequently and the important hill forest
species S. curtisii and S. platyclados at intervals not exceeding five years. In Dryobalanops aromatica
and Shorea curtisii local flowering can be found somewhere in Malaya almost every year and
sometimes occurs in two successive years in one area; this has also been observed in Shorea sect.
Pachycarpae in West Sarawak (e.g. SMYTHIES, 1958). BURGESS observed that individual trees of
Shorea curtisii do not, as a rule however, flower in two successive flowerings. He also observed that
in S. curtisii the flowering in 1968 could be subdivided into two almost distinct periods during which
two separate sets of individuals within a single gregarious population flowered. Planted dipterocarp
trees, and trees in forests previously selectively felled, flower more frequently, and generalisations
about dipterocarp flowering in natural conditions which are derived from observations on such
trees can be misleading (e.g. SASAH et al., 1979). Vatica rassak and Shorea macrophylla flower almost
annually in the arboretum of the Forest Research Institute at Kepong (Malaya) as apparently do
several species at Bogor and a Shorea stenoptera provenance in a remarkable trial plantation nearby
in W. Java, in which all plants are said to have been derived from a single tree near Pontianak that
was well known for its annual flowering.

Emergent species probably take many years to reach flowering age under forest conditions. At
Kepong cultivated saplings of Dipterocarpus oblongifolius have flowered after only 7 months
(KOCHUMMEN, 1961), but fruit was not set and cultivated trees there normally started flowering after
15-30 years (NG, 1966). TANG (1978) has recorded young trees of Shorea leprosula, growing
following selective felling, to set viable fruit after 7 years.

The cause of flowering remains obscure. It can occur between March and July, with a peak in May,
in Malaya and East Borneo; but a month or two later on average in N.W. Borneo. Both Woop and
BurGeEss have shown that the commonly held belief that general flowerings follow a period of
abnormal water stress is not consistently supported by data from rainfall stations; nevertheless the
fact that many other families (e.g. Burseraceae and Sapotaceae) also flower unusually heavily in a
good dipterocarp year (e.g. MEDWAY, 1972), indicates that an undefined climatic factor must be
involved. PALMER (1979) rightly pointed out that the climatic trigger must be easily observable
outside the forest as its effects are regional. NG (1978) has suggested that flowering may be initiated
by a period of high irradiation. WyCHERLEY (1973) demonstrated highly significant statistical
correlation between gregarious flowering of dipterocarps and preceding periods with both large
diurnal temperature ranges and high maximum temperature indicating high insolation; he deduced
that the latter is probably the main inductive factor, and this view is also supported by NG (1978).
Heavy flowerings never occur in successive years. It appears possible therefore that accumulation of
assimilates, including carbohydrates within the trees, takes place gradually following a flowering, so
that in each subsequent year the threshold, over which the climatically induced trigger is effective,
becomes lower until the combination of an adequate assimilate level and adequate climatic
stimulation induces formation of inflorescence primordia; the intensity of flowering may thus be
related to the degree to which the threshold is exceeded. Differential rates of inflorescence and
flower development among the species leads to spacing in the periods of anthesis of individual species.

Pollination. No reference occurs in the literature. Meliponid bees (Trigona spp.) are, according to
BurGEss, abundant in the crowns of flowering Shorea curtisii and other Malayan dipterocarps;
CHAN & APPANAH (1980) have found them to be principal pollinators of Dryobalanops and
Neobalanocarpus; as they are short distance foragers that tend to keep to a single crown, and as they
are glabrous or sparsely hairy and very efficient at cleaning themselves, they are unlikely to be
effective pollinators of trees that present many flowers at a time (D. H. JANZEN, comm.). SMITINAND
reported to me that honey bees are abundant round Dipterocarpus crowns, but they have not been
observed on the flowers of other genera. Thrips (Thysanoptera) frequent the flowers of many Shorea
and Hopea (personal obs.; CHAN & APPANAH, 1980) and are undoubtedly pollinators. Geometrid
moths and beetles visit some dipterocarp flowers at night, but have not been demonstrated to effect
pollination. The very large numbers of flowers on each tree, and the spatial isolation of the
understorey species, may be expected to lead to rarity of outcrossing; the infrequency of flowering
and its intensity would tend to prevent vector numbers from reaching adequate levels for effective
pollination even were they polylectic (promiscuous).

1982] DIPTEROCARPACEAE (Ashton) 247

Fruiting. Perhaps as a result of these problems of pollination, few fruit develop on each
many-flowered inflorescence, and in some years none, though heavy rain following anthesis is also
generally regarded as disastrous (SMYTHIES) and the main determinant of a successful crop. The fruit
are heavily parasitised by weevils of the genera Al/cydodes and Nonophyes, also the scolitid Poecileps
and some Lepidoptera (DALJEET SINGH, 1974), especially following minor flowerings when seedlings
rarely become established. These beetles are apparently not host specific, and increase greatly and
suddenly in numbers in flowering years though their life cycle is known to take 18 months. BURGESS
(1969) estimated that c. 80°, of Shorea curtisii seed on average is destroyed by these predators. The
genus Dipterocarpus is particularly susceptable.

Fruit development. As a consequence of the above fruiting years are less frequent than flowering
years. CHAN (1980) found in the 1976 flowering in West Malaysia that, though there was an interval
of 61 days between the onset of anthesis in the first and last of six Shoreas in sect. Mutica growing
together in Pasoh forest, fruiting started within 10 days of one another and the fruiting periods
otherwise completely overlapped. This can be accounted to the fact that the period between anthesis
and first fruit set was 82 days for the first to flower, 126 for the last. This phenomenon appears
general in the family; differentials in rates of development vary between related species rather than
between genera therefore. Together, these factors lead to occasional heavy fruitings in which
enormous quantities of seed from a majority of species in an area are set+concurrently and
germinate immediately on the forest floor.

Dispersal. Though the prolonged fruit sepals allow the fruit to gyrate and fall obliquely, once
within the main canopy they fall in random directions and on average almost vertically; many get
anchored in branches and die, and many main canopy and understorey species have re-evolved
short fruit sepals. WEBBER (1934) observed fruit being dispersed up to 4 mile in local high winds:
MULLER (personal comm.) has once seen the coastal cantonment of the Brunei Shell Petroleum
Company, a flat strip of cleared land behind the sea beach, inundated by the fruits of the Shorea
albida trees that fringe it on the inland side, observing a dispersal distance of c. 2 km. Such events
must be unusual and as RIDLEY (1930) noted, the vast majority of the fruit fall within 100 m of the
parent tree under forest conditions. BURGESS (1969) estimated that over one half of the seed of the
ridge-top species Shorea curtisii landed within 20 m of the parent tree. Even during a dry spell at
Bogor and with strong southern wind VAN STEENIS observed dispersal of Shorea grown in the
Botanic Garden over the large lawn in front of the palace not to exceed c. 500 m.

The fruit with their resinous pericarps are not favoured by many terrestrial animals, though wild
pigs devour them voraciously. The occasional heavy fruiting years undoubtedly minimise predation
and reduce seed mortality (BURGESS, 1969); he stated that damage by foraging ants is so severe in
Shorea curtisii that successful germination only occurs in heavy seed concentrations.

Some 20 dipterocarp species in West Malesia alone characteristically grow on periodically
swamped riverain alluvium and river banks. Most of these species, including Dipterocarpus apterus,
D. tempehes, Dryobalanops oblongifolius, Shorea macrophylla, S. palembanica, S. seminis, S.
splendida, S. sumatrana, Vatica pauciflora, V. rassak, and V. umbonata have large fruit with short
sepals and thick pericarps, but with these exceptions the fruits do not present any obvious
adaptation for dissemination by water. BURKILL (1922) briefly discussed the adaptation for water
dispersal in Vatica pauciflora (wallichii) and Shorea seminis (Isoptera borneensis) and found that the
fruit of the former floats for c. 22 days while that of the latter, when deprived of its corky sepals,
sinks within 24 days. There is no evidence that dipterocarp seed can withstand sea water.

Conditions for germination appear to be crucial but have been little studied. Fruit of Malesian
dipterocarps lacks dormancy, and attempts to induce dormancy by reducing water content and
temperature have only had limited success (TANG, 1971; TANG & TAMARI, 1973). In general seed of
Dipterocarpaceae is noted for its brief period of vitality; they do not stand drying out and seedling
stages are clearly adapted to temperature, moisture, and light conditions of the primary forest.
Dipterocarp seeds have no dormancy in nature and attempts at storage beyond a few weeks have
failed in Malesian species. Indian foresters (e.g. GUPTA, 1936-38) have noted that Dipterocarpus
seeds rarely survive where litter is thick on the forest floor, the radicle failing to penetrate it and
drying up. The gregarious riparian Indo-Chinese Dipterocarpus alatus appears to regenerate only at
prolonged intervals, when fruiting follows a flood in which the litter has floated away, leading to
even-aged stands. BURGESS (1969 personal comm.) has germinated species in Shorea sect. Mutica

248 FLORA MALESIANA [ser. I, vole9=

ben , a

Fig. 5. Ground-carpet of seedlings of Shorea multiflora (BURCK) SYM., 9 months after flowering. Note the
drip-tips. Brunei, Andalau For. Res., lowland dipterocarp forest, 100 m alt. (Photogr. ASHTON, Aug. 1959).

under controlled humidity conditions and found that species differ in their tolerance of dry
conditions at germination: the coastal hill and ridge-top species Shorea curtisii is paradoxically
highly sensitive to dry conditions at germination, though apparently more tolerant of water stress
than related species at maturity. He estimated that 40% of S. curtisii seedlings fail to survive the first
month following germination, and another 10°, succumb in the following dry season c. 3 months
later. Mortality in the life cycle is consequently greatest during fruit development and the 2—3 years
following seedling establishment.

General ecology. Following successful establishment carpets of seedlings of a single species are
seen around the boles of the parent trees. Fig. 5, 81. At this stage competition is therefore mainly
intraspecific, and it is interesting that, though this is the period of maximal mortality,
morphological and anatomical differences between allied species are at their least. Knowledge of the
ecology of dipterocarp seedlings is almost entirely empirical, derived from silvicultural experience.
SUNDERLAND (unpubl.) has demonstrated however that though seedlings of the slow-growing heavy
hardwood Shorea maxwelliana and the fast growing light hardwood S. leprosula share compensa-
tion points, total daily net photosynthesis and rates of dark respiration are much higher in the latter.
Though the former can survive under lower light intensities, neither can survive without the
occurrence of sunflecks.

If, as seems probable, dipterocarps form complex ectotrophic mycorrhizal associations (HONG,
1979), then the clumped distribution imposed by their reproductive biology will facilitate and be
enhanced by the events leading to the establishment of their rhizosphere associations. Many
basidiomycetes are known to produce carpophores in response to drought; under lowland rain
forest conditions carpophore formation appears to be, perhaps as a consequence, less frequent. In
wetter areas it may be that reproduction 1s largely vegetative therefore, the mycelia persisting and

1982] DIPTEROCARPACEAE (Ashton) 249

gradually spreading with the ever dispersing and coalescing clumps of the dipterocarp trees
themselves.

Given all these conditions I conclude with MERRILL (1923) that “it becomes perfectly evident that,
in order to explain the present distribution of Dipterocarpaceae, it is absolutely necessary to
postulate previous land connections from India to New Guinea over which, at some time(s) in
geologic history, it has been possible for certain species to march unimpeded.”

The morphological differences by which the dipterocarp species are mainly recognised, in bark
and twig as well as leaves, increase with age and reach their maxima once the crown has emerged
into direct sunlight, branching becomes sympodial and the mature tree leaf is attained. In the largely
understorey genus Vatica this differentiation does not occur and many species are difficult to
identify unless fruiting. KENWORTHY (1969) demonstrated the physiological significance of various
leaf characters of epidermal thickness, tomentum and wax deposits by which so many emergent
species are distinguished. He found that in Shorea curtisii the stomata of the glabrous seedling leaves
rapidly respond to water stress by closing, whereas in mature trees the stomata remained open at all
times of day and are clogged by the abundant filamentous wax secretions that give the leaf
undersurface of the species its characteristic appearance. The mature tree would act therefore as a
giant wick and would have little control of water loss other than by the development of epidermal
resistance. It is probable that the stomata are responsive in the freshly opened leaves, but rapidly
become clogged by detritus and wax. It appears that leaf change, which is probably frequent and
occurs at least once a year, is a crucial period which occurs only following times of rain when water
stress is minimal. The conspicuous changes in colour and albedo as the leaves expand, and
differences in these characters between individual trees, will create leaf temperature and hence
transpiration differentials (SMITH, 1909). KENWorTHy (1971) also pointed out that the net, which is
formed by the tertiary nerves and their associated sclerenchyma, effectively divides up the
dipterocarp leaf into compartments like an aircraft wing, so that rigidity is maintained after
prolonged drought or severe damage by predators.

Dipterocarp species vary greatly in growth rates; some, usually intolerant of low light intensities,
show rapid growth rates and reach mature habit within 60 years under forest conditions; others,
usually shade tolerant grow very slowly. The former probably have a life-span of c. 250, the latter
perhaps in excess of 1000 years, judged on the basis of girth growth data.

Range in non-seasonal zone. Climates with at least 2000 mm mean rainfall and no regular season
of pronounced water stress occur throughout Malaya, except the north-west, and in S.E. Peninsular
Thailand (Pattani); throughout Sumatra except the north-west (Atjeh) and south (Lampongs):
throughout Borneo except the extreme north-east (Kudat District) and south-east (particularly
near Pleihari in the rain shadow of the Meratus mountains); down the eastern side of the Philippine
archipelago from Cagayan Province of Luzon and including most of Mindanao; and throughout
lowland New Guinea and adjacent islands immediately to the west but excluding the region around
Port Moresby in the south-east and a belt in the south extending from the Fly River into West Irian
(Okaba to Merauke and Wassi Kussa). Within this region there 1s still considerable variation in
mean annual rainfall and its seasonality, and in some areas where seasonality is relatively
pronounced the dipterocarp flora becomes somewhat impoverished and species of the seasonal
evergreen forest become more abundant. Such is the case in eastern Luzon, much of eastern New
Guinea, eastern Borneo, much of Sumatra, Pattani in Thailand and northern Malaya including
Perak, Kedah, part of Kelantan and to a lesser extent Trengganu and northern Pahang and
Selangor. There is also evidence of recent changes in the boundaries of these forests: this may
explain the presence of Shorea leprosula and S. parvifolia in an outlier at Trang in Peninsular
Thailand, for instance.

Role in succession. It can be readily explained why dipterocarps are generally absent in secondary
forest in this zone, and why they take so very long to reinvade devastated land, for their very limited
means of seed dispersal combines with their sensitivity to water stress at germination and early
establishment to make them particularly unsuitable as colonisers. Colonisation is only known to
occur successfully therefore on clay soils with a high water retaining capacity in moist hilly districts
and periodically inundated alluvium (ASHTON, 1964). It is more difficult to understand why this
should not be true in the seasonal evergreen zone; here not only is Dipterocarpus alatus a rapid
coloniser of alluvium, but such species as Anisoptera costata and A. thurifera (R. JOHNS, unpubl.)

250 FLORA MALESIANA [ser. I, vol. 9?

Fig. 6. Hill dipterocarp forest of Shorea curtisii DyER ex KING, séraya, at c. 900 m alt., in Malaya (Photogr.
WYATT-SMITH).

1982] DIPTEROCARPACEAE (Ashton) 251

actually increase in density in secondary forest. Germination studies of these species as well as those
of the Dry Dipterocarp forests would be rewarding.

Edaphic conditions. As the role of the family differs entirely between the forests of the Sunda and
New Guinea non-seasonal zones the two must be considered separately. In western Malesia
(Sumatra, Malaya, Borneo and Philippines) Dipterocarpaceae dominate the forests on well-drained
yellow and red soils below 1300 m; these forests have hence been termed Mixed Dipterocarp forest
by me (1964). SYMINGTON (1943) had recognised that the coastal hills of Malaya bore a
characteristic forest flora, many elements of which, such as Shorea curtisii, S. ovata, S. glauca,
Vatica mangachapoi, Hopea beccariana, Dipterocarpus fagineus and Anisoptera curtisii also occur
along inland ridges. Fig. 6. BURGESS has demonstrated that the two habitats share a pronounced
tendency to prolonged periods of water stress differing from those of the semi-evergreen zone in
their unpredictability.

Increased water stress may explain why trees isolated by felling operations so frequently die;
death of mature trees in undisturbed forest on account of water stress has never been recorded. I
observed (1968) the sensitivity of Hopea enicosanthoides to prolonged submergence however. I had
earlier (1964) demonstrated that Mixed Dipterocarp forest on sedimentary rocks in Brunei varied
considerably in structure and floristics and correlated this variation largely to environmental
features that influence water status, including physiography and physical characteristics of the soil;
in my view nutrient status had little part to play in differentiating the vegetation. I demonstrated a
distinct floristic connection between the forests on very well drained yellow podsolic sands in Brunei
and those of the Malayan coastal hills, whereas the inland lowland Malayan element was confined
to clay soils, especially on broad low ridges.

My views were in conflict with those of SYMINGTON and WyYATT-SMITH who found no consistent
relationship between soils and forest variation in inland lowland Malaya. PoorE (1968) confirmed
their views in an intensive study of a limited area in Pahang, where he deduced that the commoner
species had wide edaphic amplitudes and were interchangeable so that, once seedlings had become
established, a process in which chance must play a large part, “possession is nine points of the law”’.

The conflict is largely resolved, and the lowland ecological range of the family more precisely
defined, in my recent work. I have shown that these lowland forests can be categorised into two
main groups: In one soil phosphorus levels in particular, and other elements to a lesser extent, are
relatively high and the forests are floristically rather uniform; in the other phosphorus levels are low
and the forests exhibit great variation which can be correlated with nutrient status (AUSTIN, ASHTON
& GREIG-SMITH, 1972). These observations are consistent with the theory that dipterocarps are
mycorrhizal. As soils with low nutrient levels are often those which are most freely drained, water
stress and limiting nutrient levels tend to go hand in hand but it is apparent, as this is not always the
case, that both factors have an important part to play in dipterocarp ecology. The low fertility soils
are principally those of the Tertiary and Quaternary sands at the margins of the Sundaland
continent, where the sediments have become impoverished by successive erosion and deposition
cycles since the Cretaceous; the Malayan Peninsula hardly possesses rocks younger than the
Cretaceous and this ancient land surface is covered by deep soils of moderate fertility, though even
here a subtle correlation does in fact exist between physiographic and floristic variation (ASHTON,
1976).

I have (1964) described the floristic and structural role of the family in the lowland forests. They
reach their zenith both in numbers of species and individuals on deep well drained yellow/red soils
of intermediate fertility, where nutrient levels are apparently limiting but not severely so. In Heath
forest on podsols dipterocarp diversity is much reduced and in peat swamp forests yet more so; in
both single dominant species are frequent. Examples are Shorea materialis in Heath forest and
Shorea albida, S. balangeran and Dryobalanops rappa in Bornean peat swamp forests. In the latter
forests diversity increases from the centres of the domed oligotrophic bogs outwards as soil fertility
increases, several species (Shorea inaequilateralis, S. pachyphylla, S. platycarpa, S. teysmanniana, S.
uliginosa, and Dipterocarpus coriaceus) being found exclusively in the Mixed Peat Swamp forests of
the margins (ANDERSON, 1963). Other species occur in both peat swamps and Heath forest, while
some 30 species are almost entirely confined to Heath forest; the ecotone between Heath and Mixed
Dipterocarp forest on yellow podsolic sandy soils is ill defined and many species are common to
both. More surprising is the marked decrease in species diversity on well drained soils of highest

N
Nn
i)

FLORA MALESIANA (ser. I, volRs2

Fig. 7. Even-aged stand of mixed dipterocarp forest, with e.g. Shorea almon Foxw. and S. polysperma (BLCO)
Me_rR. abundant in the canopy, recovering from total destruction due to a cyclone in 1944, in 1975. Note the
pyramidal, monopodial juvenile crowns. Mindanao, Suriago del Norte (Photogr. ASHTON).

1982] DIPTEROCARPACEAE (Ashton) 253

fertility; I have shown (1964) that there was a decrease in diversity on calcareous shales in
comparison to sandstones, most marked on the shallow skeletal soils on the narrow ridges and steep
slopes of the youthful N.E. Bornean physiography. Unpublished analyses from Sarawak show a
further decrease in both density and diversity on deep fertile soils; here gregariousness also
increases. The gregarious stands of Shorea retinodes on the slopes of the Barisan range, where soils
have been much affected by Quaternary volcanic activity and landslips remain common, may be
a comparable example. In some cases the explanation may partially lie in the isolation of these
small distinctive habitats of geologically recent origin, preventing the rapid build-up of diversity.
Fig. 7.

Low diversity and the presence of gregarious species also characterise excessively dry habitats in
the non-seasonal zone. Thus Shorea gratissima is confined to and gregarious on rocky headlands;
and Dryobalanops aromatica forms pure stands on coarse gravelly soils and yellow podsolic sands in
Atjeh and Malaya.

The /imestone dipterocarp flora is poor; though no species is known to be gregarious in the
non-seasonal zone Hopea ferrea is so on the rocky hills of the Langkawi Is. No species appears to be
confined to limestone, but species sometimes found on it include Cotylelobium malayanum, Hopea
aptera, H. billitonensis, H. cernua, H. dasyrrachis, Shorea guiso, S. havilandii. In the seasonal
north-west of the Malay Peninsula Shorea siamensis and S. roxburghii occur on it; and further north
outside Malesia, where mineral soils accumulate over the rock, a wide range of dipterocarps may
grow. Similarly, the dipterocarp flora on the soft coral limestones of eastern Mindanao does not
seem to differ from that on other substrates, and their roots penetrate the soft rock itself.

The intrinsically unstable and specialised habitat of river banks also carries a characteristic
dipterocarp flora of some 20 spp. in western Malesia. Most species are widespread but some (e.g.
Dipterocarpus oblongifolius, Hopea centipeda) occur only on the rocky banks of rapid inland rivers,
whereas others (e.g. Vatica venulosa, Shorea seminis) equally prefer alluvium along meandering
rivers of the plains.

The many similarities between the ecology of the dipterocarps and the undoubtedly ectotrophic
mycorrhizal Fagaceae, which assume dominance at high altitudes, is noteworthy.

Summarising: Comparatively few species are restricted to one vegetation type or one substratum
or type of bedrock. For example, many Vaticas occur in alluvial forest and near rivers, but seldom
exclusively so, and occur also on the hills, though they may be more common in the former habitat.
Hopea pentanervia occurs on mixed peat swamp over sand, podsols and cuestas, plateaus and
terraces near present or Pleistocene coastlines, and on ultrabasic rocks in N. Sabah; Shorea
polyandra is found on fertile, clay-rich soils on calcareous shales, igneous and volcanic rocks; Shorea
scabrida is found in freshwater swamp forest, on shallow peat overlying sand, and on skeletal soils
on ridges and plateaus, in Heath forest and in Mixed Dipterocarp forest. This apparent diversity of
habitats occupied nevertheless frequently conceals a common and sometimes rather specific
edaphic range.

Others are more clearly confined to certain edaphic habitats and some of these are cursorily listed
here:

Heath (kerangas) forest on podsols: Cotylelobium burckii, C. malayanum, Dipterocarpus
borneensis, Dryobalanops fusca, Hopea kerangasensis, H. micrantha, H. pterygota, H. vaccinifolia,
Shorea coriacea, S. induplicata, S. materialis, S. pallidifolia, S. retusa, S. revoluta, S. venulosa, Vatica
coriacea, V. parvifolia.

On sandy soil the following are frequently recorded: Anisoptera grossivenia, A. reticulata,
Cotylelobium melanoxylon, Dipterocarpus globosus, D. rigidus, D. sarawakensis, Dryobalanops
aromatica, Hopea beccariana, H. coriacea, H. treubii, H. vesquei, Shorea acuta, S. crassa, S.
dealbata, S. falcifera, S. flemmichii, S. geniculata, S. ladiana, S. laxa, S. rubella, S. rugosa, S.
scabrida, S. stenoptera, Upuna borneensis, Vatica borneensis, and V. oblongifolia sspp. crassilobata
and elliptifolia.

In peat swamps the following species are characteristic, some being gregarious there: Anisoptera
marginata, Dipterocarpus coriaceus, Dryobalanops rappa, Shorea albida (also in Heath forest), S.
balangeran (also in Heath forest), S. foraminifera, S. inaequilateralis, S. macrantha, S. pachyphylla,
S. platycarpa, S. teysmanniana, and S. uliginosa. Fig. 8, 96.

On ultrabasic soils the following are regularly recorded, but only the last is confined to them:

254 FLORA MALESIANA iser: I, voli92

Fig. 8. Stands of large trees in peat-swamp of Shorea albida SyM., alan; road tracé to oil well in Seria, Brunei, at 5
m alt. (Photogr. ASHTON, 1960).

1982] DIPTEROCARPACEAE (Ashton) 255

Dipterocarpus geniculatus ssp. grandis, D. ochraceus, Hopea pentanervia, Shorea coriacea, S.
tenuiramulosa.

It will be noted that all examples given occur in Borneo and many are endemic there; this reflects
the unusual habitat diversity as well as infertility of that island.

Altitude. The altitudinal limits of Dipterocarpaceae in the non-seasonal zone experience mean
minimum monthly temperatures comparable to those at the northern margin of the family range at
22° N, in Yunnan, K wangsi, S. Kwantung, and Hainan atc. 15° C. In areas of shallow soils, often of
low base status, such as those derived from the sedimentary rocks at the margin of the Sunda shelf
area in N.W. Borneo, altitudinal zonation of vegetation is obscured by variation in relation to soils
except at the extreme altitudinal limits and on isolated summits. In Malaya such zonation is clearer,
though the ecological relationship between the ridge and coastal hill flora has already been
described. The increase in raw humus accumulation above 1000 m is associated with the marked
affinity between the dipterocarp flora of the hill ridges and lowland podsolic soils. SYMINGTON
(1943) summarised the altitudinal zonation of dipterocarp species in Malaya: He recognised a
Lowland Dipterocarp forest zone below 300 m, with the coastal hills as a separate category, the
former with 108 spp., the latter with 27 of which 8 are shared; between 300-800 m are Hill
Dipterocarp forests with 49 spp., and between 800-1300 m (the altitudinal limit of dipterocarps in
Malaya) are Upper Dipterocarp forests with 15 spp. Within the two hill types there are 7 exclusively
hill species; of the rest 15 are exclusively coastal in the lowlands and of these 5 occur also in the
Upper Dipterocarp forests; the remaining 35 occur in Lowland Dipterocarp forests generally,
though of these but 4 are found in the upper zone. The existence of a small group of apparently
obligatory hill species is not confined to Malaya; some are widespread in western Malesia,
Dipterocarpus retusus extending eastwards to Sumbawa, while in Borneo there are a further 10
endemic montane taxa. A few ascend above 1000 m, for example to 1200 m: Shorea carapae, S.
revoluta, Vatica odorata ssp. mindanaensis; to 1300-1350 m: Shorea beccariana, S. flaviflora, S.
platyclados, S. rubra, Vatica heteroptera; to 1400 m: Shorea longisperma, Vatica dulitensis; to 1500
m: Shorea monticola, S. ovata, Vatica oblongifolia ssp. selakoensis; to 1600 m: Shorea venulosa; to
1650 m: Hopea cernua; to 1700 m: Vatica granulata ssp. sabaensis; to 1800 m: Shorea platyclados
(Atjeh).

At their upper limits the dipterocarp forests give place to oak-laurel forests in Malesia; it is
noteworthy that in Ceylon, where the Fagaceae, which share with Dipterocarpaceae gregariousness
and well developed ectotrophic mycorrhizal associations but which become dominant above 1300
m, are absent, 9 exclusively montane species of the endemic taxa Shorea sect. Doona and
Stemonoporus dominate the hill forests of the south-west up to 1700 m (AsHTON, 1972, 1977).

Synopsis: Role of dipterocarps in the frame of the forest. At their edaphic optimum in the lowlands
dipterocarps may comprise more than 80% of all emergent individuals in the forest, and up to 40%
of understorey trees. Under conditions of increasing proneness to aseasonal drought the emergent
stratum becomes less dense, the understorey more so, and dipterocarps decrease their represen-
tation in both except where gregarious species occur. Under the most mesic conditions the emergent
canopy becomes so dense as to be continuous, and the understorey consequently becomes diffuse in
the low light intensity; here total dipterocarp density as well as relative density is reduced.

In the non-seasonal evergreen forests of New Guinea dipterocarps play an altogether subsidiary
role. Only 14 spp. are so far known from these vast forests; 11 are endemic to New Guinea and
adjacent islands, and 9 are confined to local areas within the island. One species of Anisoptera, one
of Vatica and 12 of Hopea are represented. Only Anisoptera thurifera is truly emergent, and
gregarious stands often give a distinctive profile to the steep-sided crests. Several Hopeas, among
them H. forbesii and H. iriana, are canopy species, as is Vatica rassak in the more seasonal areas.
The family as a whole shows a distinct though not exclusive preference for ridge tops, noticeable
(though less markedly so) on the similarly youthful physiography of much of Borneo.

Single species as a rule individually merely play a minor part as contributors with many others to
the overall role of the family. Gregarious or semi-gregarious populations occasionally occur in
many, perhaps most, species including those of the understorey and are probably usually
attributable to chance favourable conditions at the time of establishment. In Malesia they are
habitual only in the species of the Dry Deciduous Dipterocarp (fire climax) forests, notably
Dipterocarpus obtusifolius, and to a lesser extent those of seasonal evergreen forests including

256 FLORA MALESIANA {ser. I, vol. 9?

ee 3 a

A
som =
ey Oo de %

. ‘a ‘a y ee
¥ . , ee \ ra Boe
pe el se) Yoo Ane
Fig. 9. Trunk-base of a colossal specimen of Shorea superba Sym. on basalt, half a mile south of Quoin Hill,

Balong area, Tawau, Sabah, forest guard K APIs BIN SISIRON at point of measurement: total height of tree 75 m,
clear bole 27 m, girth at 4 m above ground 95 m=3 m diam. (Photogr. G. H. S. Woop).

Anisoptera costata, A. thurifera, Dipterocarpus costatus, D. gracilis, D. grandiflorus, D. kerrii, Hopea
ferrea, H. forbesii, H. glabrifolia, H. gregaria, H. pierrei, Shorea guiso, S. hypochra, S. roxburghii, S.
selanica, Vatica cinerea and V. rassak on the one hand; — and on the other in species of limiting soil
conditions such as Dipterocarpus elongatus, D. oblongifolius, Hopea fluvialis, H. odorata, Shorea
seminis, S. sumatrana, Vatica pauciflora, V. rassak and V. umbonata on river banks and flood
plains, — Dryobalanops fusca, D. rappa, Shorea albida, S. balangeran and to a small extent
Anisoptera marginata, Shorea inaequilateralis, S. platycarpa, S. teysmanniana and S. uliginosa in
peat swamps, — and Anisoptera marginata, Cotylelobium burckii, C. malayanum, Dipterocarpus
borneensis, Dryobalanops fusca, Shorea materialis and sometimes others in Heath forest. Several
factors, including chance, may have played a part in the origin and maintenance of the extensive
gregarious stands of Dryobalanops aromatica (see VAN SLOOTEN, 1932; Lee, 1967) and Shorea
retinodes (VAN SLOOTEN, 1949).

1982] DIPTEROCARPACEAE (Ashton) 257

Literature: ANDERSON, Gard. Bull. Sing. 20 (1963) 131; ASHTON, Oxf. For. Mem. 25 (1964); Man.
Dipt. Brun. Suppl. (1968) 51; Biol. J. Linn. Soc. 1 (1969) 155; Proc. 2nd Aberdeen-Hull Symp. Mal.
Ecol. (1972) 35; Blumea 20 (1972) 357; Mal. For. 39 (1976) 56; Rev. Handb. Fl. Ceylon 1, 2 (1977)
166; ASHTON et al. (1979) in press; AUSTIN, ASHTON & GREIG-SMITH, J. Ecol. 60 (1972) 305; BURGEss,
Mal. For. 32 (1969) 438: Mal. For. 35 (1972) 103; BuRKILL, J. As. Soc. Str. Br. 86 (1922) 276, 281;
CHAMPION, Ind. For. Rec. Bot. 1 (1936); CHAN, Mal. For. 43 (1980) 438; CHAN & APPANAH, Mal.
For. 43 (1980) 132; CocKBURN, Mal. For. 38 (1975) 160; DaLsEET SINGH, Mal. For. 37 (1974) 24:
Gupta, Ind. For. 62 (1936) 525; ibid. 63 (1937) 734; ibid. 64 (1938) 15; HONG, Mal. For. 42 (1979)
280: KENWORTHY, Mal. Nat. J. 22 (1969) 129: Proc. Ist Aberdeen-Hull Symp. Mal. Ecol. (1971) 49:
KOCHUMMEN, Mal. For. 24 (1961) 236; Lee, Ph.D. Thesis, Univ. Malaya (1967); MCCLUuRE, Mal.
For. 29 (1966) 192: Mepway, Biol. J. Linn. Soc. 4 (1972) 117; MERRILL, Philip. J. Sc. 23 (1923) 8;
NG, Mal. For. 19 (1966) 290; Nature Malaysiana 3, n. 1 (1978) 12, col. phot.; PALMER, Mal. For. 42
(1979) 74; Poore, J. Ecol. 56 (1968) 143; RipLey, Disp. Pl. (1930) 51; SAsAH, TAN & ZULFATAH, Mal.
For. 42 (1979) 38; vAN SLOOTEN, Bull. Jard. Bot. Btzg III, 12 (1932) 7: ibid. III, 18 (1949) 263:
Reinwardtia 2 (1952) 1; Smirn, Ann. Perad. 4(1909) 229; SmyTuies, Sarawak Gaz. 84 (1958) n. 1206,
p. 146; STAMP, The Vegetation of Burma, Calcutta (1925); SyMiNGTON, Mal. For. Rec. 16 (1943):
TANG, Mal. For. 34 (1971) 84; ibid. 41 (1978) 294; TANG & TAMaRI, Mal. For. 36 (1973) 38; WEBBER,
Mal. For. 3 (1934) 18; Woop, Mal. For. 19 (1956) 193; Wyatt-SmitH, Mal. Nat. J. 7 (1952) 45. 91:
ibid. 8 (1953) 52; Mal. For. 17 (1954) 83; Mal. For. Rec. 23 (1963); WyCHERLEY. Micronesica 9
(1973) 75.

Morphology. Habit. See Fig. 9, 21, 25, 32, 51, 53, 55, 80, 111, 112, 117. The bole can be
cylindrical in cross-section, or fluted or ribbed, and is usually buttressed. See Fig. 29, 52, 59, 81. The
buttresses can be small or large, thin with a sharp edge as in Shorea sect. Shorea generally, or thick
with a rounded edge; the edge, from the soil surface to the apex where the buttresses merge into the
bole, is usually concave, but is sometimes straight or convex; the buttresses can terminate more or
less abruptly at apex and base, or continue as ribs up the bole as in Anisoptera sect. Anisoptera or
superficial roots over the ground. Flying buttresses are convex-edged buttresses originating from
the bole above the soil surface, and are differentiated from the stilt roots by being flat, therefore
elongate in cross-section, as opposed to being terete or spherical in cross-section; stilt roots can be
adventitious to the buttresses or arise from the bole. Flying buttresses and stilt roots are a particular
feature of Hopea, and are found at least occasionally in a majority of species with the exception of
those in sect. subsect. Hopea. Fig. 67, 71.

In the seasonal parts of S.E. Asia (Thailand, etc.) various species of Dipterocarpaceae have a
capacity for suckering and produce new stems from trunk-bases after cutting. However. the
Malesian rain-forest species generally lack this capacity and regenerate merely from seed.

The mode of branching. and the leaf arrangement, changes ontogenetically. The seedling
dipterocarp, after production of one or more pairs of opposite leaves. sends up a stem with spirally
arranged leaves. From the axils of these leaves arise the lateral branches: the leaves on the branches
are generally arranged distichously, consequently the branchlets are also alternately arranged and
the branches occupy one plane (plagiotropy), ascending, descending. or horizontal. The sapling
leader, or stem apex, grows continually at first, but soon adopts resting periods, growth proceeding
in flushes that are not necessarily seasonal at this stage though often synchronised among members
of a clump. The internodes are frequently longer towards the beginning of the growth flush than
towards the end, so that the stem leaves in some species, particularly Hopea sect. Dryobalanoides,
are largely bunched at the end of each growth stage of the leader; consequently the branches appear
to arise in whorls and the sapling and pole sized tree assumes a ‘pagoda shape’ (CORNER, 1940): each
flush of growth by the leader is coordinated with the sprouting of some of the axillary buds of the
previous flush to form new lateral branches.

Young dipterocarps therefore conform either to Massart’s model, that is, with rhythmic growth
and branching, or approach Roux’s, with continuous growth (HALLE, OLDEMAN & TOMLINSON,
1978).

The leaders have spiral leaf arrangement; if a leader dies or its growth is otherwise arrested at this
stage, new leaders emerge not from axillary but from tiny subsidiary buds (NG, 1976); whether a
single leader eventually achieves dominance over others varies with the species.

If the tree remains small, and never reaches the forest canopy, as in many Hopea species, it

258 FLORA MALESIANA [ser. I, vol 9?

frequently remains monopodial with this form of branching, and the crown remains lanceolate or
conical. Fig. 7, 36. If the leader reaches the canopy the subsequent lateral branching generally
become orthotropic, apical dominance Is lost, the earlier plagiotropic branches die and are lost and
a hemispherical or dome-shaped crown is formed. Fig. 1.

Alternatively, plagiotropic branches become orthotropic towards their tips (HALLE, in pass.).
Many Vatica species, however, which never become tall enough to reach the forest canopy, become
more or less sympodially branched and develop an irregular oblong crown; while Parashorea
macrophylla, and some members of the Shorea sections Mutica and Anthoshorea remain
monopodial for some time after they have emerged above the canopy and reached full height, and
may even send up further vertical leaders, apparently from axillary buds from the plagiotropic
side-branches. In most species with an emergent sympodial crown the leaves are spirally arranged
on the branchlets, but in species that tend to branch low, such as the riverside species, those species
in which some or all the branches become horizontal or pendant towards the apices, and the species
with compressed twigs, the leaf arrangement is frequently or always alternate. If the branches are
many and radiate from the bole apex, as in many emergent Hopea species, the crown of emergent
species takes on an evenly hemispherical appearance. If the main branches are few and large but
much branched towards the apices, the branchlets become bunched towards the ends of the large
branches into more or less hemispherical groups and the crown assumes the appearance of a
cauliflower head. If the twigs are stout and the leaves large the crown shape is uneven and the leaves
tend not to be confined to the perimeter of the crown. Rarely, the branches become pendent at the
apices and the crown becomes ‘weeping’ (e.g. Shorea inaequilateralis, S. quadrinervis). Fig. 111.

According to HALLE, OLDEMAN & TOMLINSON (1978) who have classified tree architecture into a
series of ‘models’, trees do not change from one model to another during life. They placed the
species of Dipterocarpus and Shorea they examined in Massart’s model in which the trunk is
orthotropic, the branches plagiotropic and in false whorls through rhythmic growth, and the
position of flowers various; and in Roux’s, which differs from Massart’s in that trunk growth is
continuous. From the foregoing description it would appear that the model may be mutable in
dipterocarps, for growth eventually always becomes rhythmic though it may not start thus, and
because in most emergent species successive branches become increasingly orthotropic with a
change from alternate to spiral leaf arrangement. This change in branching pattern would not imply
a change in model were such orthotropic branches arising through reiteration, that is from
supra-axillary buds following damage to the leader; but this does not seem to be the case.

Bark. Fig. 9, 22, 24, 31, 45, 52, 69, 102, 113, 114. THORENAAR (1926) first systematically examined
bark morphology in the family; SYMINGTON (1943) used bark characters extensively for field
diagnosis. WHITMORE (1962) described how weathering processes and tangential strain during
growth together act in conjunction with the growth of secondary phloem, expansion tissue
developed from phloem rays, phloem proliferation tissue developed from the phloem parenchyma,
and the periderms (each of which are laid down in several ways), to produce surface features
distinctive of both species and higher taxa; these features change ontogenetically from an initial
smooth surface. He thus rationalised bark description, which essentially comprises the surface
pattern (the visual summation of surface configurations) and the slash appearance (the visual
summation of the appearance of the inner and outer bark in oblique transverse section). Various
degrees and types of fissuring develop through tangential strain, and of flaking on account of the
disposition of the periderms. In a few taxa (e.g. most Vatica, and Shorea sect. Pachycarpae) the bark
remains smooth; in others (e.g. Anisoptera, Dipterocarpus, Shorea sect. Shorea and sect.
Richetioides) it almost always becomes distinctively flaky, while in yet others (e.g. Shorea subsect.
Mutica) it becomes distinctively fissured. In one known instance (fissured-bark Hopeas) a distinctive
bark configuration has no apparent taxonomic significance above the species level, and is not indeed
even consistently developed within the species. Seven distinct bark types, in three groups differing in
the amount of expansion tissue, were recognised; within each a number of categories exhibiting
lesser structural differences were defined as bark manifestations; it is these that characterise taxa at
generic and sectional level. In this account bark characters are summarised in generic and sectional
descriptions where appropriate, but have been omitted from species descriptions; reliable bark
descriptions of species can be found by reference to SYMINGTON and WHITMORE and also ASHTON
(1964, 1968). In particular they contribute important definitive characters for the sections of Shorea.

1982] DIPTEROCARPACEAE (Ashton) 259

Resin exudations from wounds in living bark and wood also differ in frequency, rate and mode of
crystallisation, translucency and colour and sometimes form useful subsidiary characters for
identification.

Roots. Fig. 10. Malesian dipterocarps lack taproots, though they are well known in Shorea
robusta GAERTN. f., and reported from Hopea parviflora BEDD., both of India. The laterals are of two
categories; an extensive branching system of superficial roots extending from the buttresses and
proliferating into a dense mat of fine roots, found by SINGH (1966) and HONG (1979) in many
Malayan species, and BAYLIS (with the author, unpubl.) in Hopea iriana, to be ensheathed in a dense
ectotrophic mycorrhizal mantle; and a diffuse coarse sparsely branching system, consisting of lines
of carrot-like ‘sinkers’ descending vertically from below the buttresses, sometimes also with
obliquely descending branches from the principal superficial roots. In acid peaty soils the former are
very well developed and the latter few; in fertile soils the converse; in deep fertile soils the latter can
penetrate at least 5 m beneath the surface. ANDERSON (1961) has described the extremely specialised
roots of the peat swamp species Shorea albida Sym. Here the buttresses give off a dense mat of roots
c. 30 cm above the permanent water table, on which litter falls and raw humus accumulates; beneath
there is a discontinuous cavity into which clumsy walkers easily fall; many stout roots descend from
the lower surface of the main superficial roots through this cavity into the waterlogged peat
substrate, anchoring the tree.

NG (1975) has described the occurrence in nature of root grafting between species in several
genera.

Buds. The dipterocarp resting bud is clad in appressed scales. Axillary buds are minute, except
prior to shoot expansion, in most species, but terminal buds are evident in Parashorea, Upuna and
Anisoptera, in most Shorea and in some species of other genera, while in Dipterocarpus they are
generally large, sometimes resinous, and their shape and indumentum provide valuable species
characters.

Parts of stem or leaf are frequently distorted in the young shoot, especially of saplings and young
trees, by certain wasps into distinct galls (CORNER, 1963; ANTHONY, 1973, 1974) which may be
globose and spinous, variously elongate and covered in umbricate fish-like scales, or merely simple
swellings. These galls appear to have no taxonomic value, and at least three forms are found in
Shorea ovalis.

Twigs. In most genera the twig surface, its diameter near the apex, its shape in cross-section, and
the shape and size of the stipule scars can be specifically diagnostic.

Leaf. Though the stipules are frequently fugaceous, in some species (especially in Dipterocarpus
and Hopea subsect. Pierrea) they are relatively persistent and the size and shape can be diagnostic.
The size and shape of blade and petiole, and the number of nerves, is usually specifically diagnostic.
The nerves are vertically transcurrent, bounded dorsiventrally by sclerenchyma to the epidermis.
The type of nervation is diagnostic for some genera and sections. If the nerves are very many,
slender, close, of equal length and straight to the margin with no discernable tertiaries, they are
termed ‘parallel’ (Dryobalanops); if as above, but unequal in length, and curving before the margin,
they are ‘dryobalanoid’ (some Hopeas); if the latter have distinct scalariform or reticulate tertiaries
and the nerves are very unequal and divisible into main nerves and shorter intermediate secondaries
they are termed ‘subdryobalanoid’ (some Hopeas). In Cotylelobium and Anisoptera the nerves curve
distally and unite forming a looped intramarginal nerve. Tertiary nerves of penninerved species are
generally scalariform but in most Vatica, Anisoptera, Cotylelobium and Shorea sect. Richetioides,
and in a few other species, they are reticulate.

Domatia occur in various species of the genera Vatica, Shorea and Hopea (Fig. 64a, 73a); they are
common in Hopea (cf. VAN SLOOTEN, 1941) and the saplings of Shorea in particular. They are often
diagnostic for a species, and are a product of the plant itself. Domatia are usually confined to the
axils of a few nerves towards the base of the midrib, but can extend the whole length, especially in
saplings; in Shorea platycarpa and some others they form a continuous series flanking the midrib,
while in young trees of Shorea leprosula they similarly flank the main nerves as well. Most frequently
they are pore-like, sometimes hairy, sometimes with the cavity enlarged, pale and stoutly rimmed
(e.g. Hopea nutans, Shorea parvifolia); sometimes they are scale-like as in Shorea platycarpa and S.
leprosula. Young leaves are usually suffused with magenta anthocyanin pigments; in Shorea sect.
Richetioides they are usually violet or dark crimson.

260 FLORA MALESIANA ser. I, volley?

er mgd

)
‘
a

Fig. 10. Structure of root-system of Shorea falcifera DYER ex BRANDIS with dug-out profile beneath it, Kuantan,
E. Malaya (Photogr. ASHTON, Sept. 1970).

1982] DIPTEROCARPACEAE (Ashton) 261

Extrafloral nectaries occur in many genera on the leaf upper-surface, often near the margins and
between vein divisions.

The petiole in mature leaves is thickened towards the apex into an umbo, often shrinking in the
drying process. As an exception the blade is peltate in Shorea peltata and leaves of young specimens
in Shorea chaiana, S. laxa, S. polyandra and some others. The umbo is obscure in Dryobalanops and
some Hopea species.

Epidermis and indumentum. GUERIN (1907) examined the leaf and twig epidermis anatomically in
many species and stated that from this alone most genera, and sometimes even species, can be
distinguished. I found that epidermal characters rarely provided evidence for species diagnosis
except where taxa were easily differentiated macroscopically; the size of epidermal cells, the
thickness of the walls and the position of domatia relative to the surface as well as the indumentum
varies more according to the part of the tree from which the leaf originates, and the age of the plant
than between closely related species. Thus I confirmed GUERIN’s conclusions that genera can
sometimes be identified anatomically, but the method was not valuable at the species level. GUERIN
also described long-stalked glandular hairs with unicellular heads on certain Stemonoporus species;
they appear identical to the glandular hairs that densely cover the young parts and inflorescence of
Upuna, and were also found, according to SYMINGTON (1941), among the indumental hairs of
Monotes; Rao (1953) found such hairs on the calyx of Shorea roxburghii G. Don. Short-stalked
glandular hairs with a multicellular head are a general feature common to most, if not all,
Dipterocarpaceae; the base of the stalk is typically sunken in an epidermal pit. A dense indumentum
of peltate emarginate scales occurs in Anisoptera, imparting a characteristic colour to the lamina
undersurface and twigs. The number of cells in the head often varies with the age of the plant and
therefore is of little value for diagnosis; thus in Anisoptera marginata they are at first 4-celled, later
8-celled.

Tomenta of evenly distributed unicellular hairs are present in nearly all species, though fugaceous
in many. Unicellular papillae occur on the ovary, stylopodium or connectival appendage of some
Shorea and Hopea species. In some Shorea species there is an indumentum of broadly lobed wax
secretions; in Hopea and some Shoreas this covering is more sparse and with slender acute lobes.
The macroscopic appearance is similar, except in colour, in all species; it is usually persistent, not
easily rubbing off.

Hairs arranged in fascicles are almost universal among the Dipterocarpaceae though often
fugaceous. The tufts may be short or long, sparsely or densely distributed: they may be of uniform
length, producing an ‘even’ tomentum, or of two or more different sizes, producing a ‘scabrid’
tomentum. In all species tomenta tend to become shorter and sparser towards the margins and
apices of organs; they are usually longest and most persistent on the leaf bud, and become
successively shorter, sparser, and less persistent on twigs, petiole, leaf midrib below, primary nerves
to tertiary nerves, and finally on the lamina surface itself. Thus many species possess persistently
tomentose midribs and glabrescent nerves, but never vice versa. In some Dipterocarpus this trend 1s
accompanied by a reduction in the bristles, while on the tertiary nerves the clusters are represented
by a single long bristle and a basal cushion representing the greatly shortened other members.

Hairs are most persistent on the inflorescence, secondly on the bud scales. Though the tomentum
tends to be longer, sparser, and more persistent in young trees, the colour and appearance furnishes
important diagnostic characters.

Inflorescence. The typical form is a semi-pendent bracteate, apical, axillary or occasionally
ramiflorous panicle once (if axillary) or twice (if terminal) branched in one plane with the flowers
secund, shortly pedicellate and nodding, developed in acropetal succession. Vateria, Upuna and
some Vatica possess cymes and the flowers are not secund; in Vatica the condition is clearly derived.
In Dipterocarpus generally, and occasionally in other genera, the inflorescence is few-flowered and
hardly or not branched. Most species bear apical as well as axillary inflorescences together on the
same twig; a few rarely or never develop terminal inflorescences, and of these several Hopea and
Vatica (e.g. V. sarawakensis) are strictly ramiflorous while some other Vatica (e.g. V. venulosa) and
Shorea (e.g. S. stenoptera, S. hemsleyana) approach this condition.

Flowers. The centrifugal stamens originate from a number of common bundles independent from
the gynoecium. Fig. 11. The gynoecium bundles break away from the common supply with the
stamens before the stamen supply begins to branch. In all genera but Upuna and some Stemonoporus

262 FLORA MALESIANA [ser. I, voleg2

Upuna borneensis Anisoptera grossivenia

Pie as Stamens.
Vf oe /---- Ovary.
FLORAL

S temonoporus SY WY Wren VASCULAR

lanceolata. SUPPLIES.
Sr é
ff A\S ~-Sepal.

Shorea havilandii

Shorea scaberrima

Fig. 11. Scheme of floral vascular supplies in five dipterocarp species.
®

1982] DIPTEROCARPACEAE (Ashton) 263

(Monoporandra) of Ceylon the number of primary stamen bundles is 10. In Upuna they are
independent of the perianth supply and continue between the perianth bundles in the pedicel; in
other genera the 10 stamen bundles are associated with the 10 perianth members, though in some
Shorea (e.g. S. geniculata) they appear not to unite within the length of the receptacle and pedicel. In
all species where the number of stamens is more than 10 the stamen fascicles originating from the
calyx supply are larger than those originating from the petals. In Shorea, Hopea, Vatica and
Cotylelobium the petal and sepal bundles, together with the staminal bundles originating from
them, are independent. Upuna, Anisoptera, Vateria, Stemonoporus and Dipterocarpus are alike in
that a single lateral branch arises constantly from the left or right of each sepal bundle which
supplies the petals; the staminal bundles from the sepal supply originate above this bifurcation.

In a recent paper C. Woon & H. KENG (1979) gave details with very numerous figures of the
stamens in Dipterocarpaceae.

Flowers are usually small, except in Dipterocarpus and Vateria. The shape and size of the bud and
the size and shape of the perianth members is diagnostic for some sections of Shorea. The number of
stamens in species with less than 20 is constant in normal flowers; in those with more than 20 the
number varies about a mean, but provides a useful specific diagnostic character. In Shorea the shape
of the filament is an important section character, and in many genera the shape of the connectival
appendage is diagnostic. The presence or absence of a stylopodium, the shape of the ovary and
stylopodium and the size of the style relative to that of the ovary are often diagnostic for sections
and sometimes species, particularly in Shorea and Hopea.

Fruit. The accrescent fruit calyx can usually be divided into two parts, the basal tube or cup,
which is adnate to the ovary in Anisoptera and some Vatica, and the distal lobes; in Shorea and
Parashorea, Dryobalanops, Neobalanocarpus and Hopea the sepals themselves are divided into the
distal wing-like lobes, and the usually somewhat broadened, thickened, saccate base adpressed to
the nut. In Cotylelobium and some Vatica the calyx cannot be so divided and only one measurement
is then given in the text. The number of long and short lobes is generically constant, though most
genera have representatives in which they are all short and equal; the latter condition is considered
to be an independent, secondary parallel reversion; see under evolutionary trends. The length of the
longer lobes, and rarely the length and size of the shorter lobes, can be specifically diagnostic.

The degree to which the calyx is united at the base into a cup or tube is generically constant,
Vatica excepted. In Dipterocarpus the tube may be spherical, or with five ribs, wings or tubercles
arising at the line of fusion of the calyx segments.

The size of the nut is often diagnostic. Its apex in Hopea bears the discernible remains and shape
of the stylopodium if present in the flower.

The structure of the seed has been described by CORNER (1976). In Upuna (and also
Stemonoporus, Vateriopsis and Pakaraimaea) there is a highly vascularised placental region which is
extended into a cup-shaped membrane in all but the last mentioned; this is the “cupule chalazique’ of
HEIM (1892).

Embryology and germination. RAO (1953, 1956) described early ovular development in the Indian
Shorea robusta GAERTN. f., S. roxburghii G. Don, Vateria indica L. and Hopea wightiana WALL. ex
W. & A. One ovule only was found to develop. Embryo-sac development as of the Polygonum type.
An initial oblique division of the zygote, followed by further oblique divisions of the daughter cells
leads to a 4- or 8-celled pro-embryo, following which further division is normal. The endosperm is at
first free nuclear. Fruit yielding more than one embryo were already known to BRANDIS; they are
unusual, but occasionally occur in many species and genera, while individual trees are known by
foresters to produce a high proportion. Foxwortny (1932) indicated that this is the general rule in
Shorea resinosa. MAuRY (1970a) found that each embryo developed from a separate ovule in two
species of the African Monotes (though it was not known how many ovules had been fertilised) but
in the gregarious riparian S.E. Asian Hopea odorata she was able (1970b) to confirm true
polyembryony, with up to 12 embryos developing from a single ovule. She reported possible
polyembryony also in the Malayan Shorea parvifolia; in several other Malesian species
false polyembryony of the Monotes type was demonstrated (1968, 1970b) and in some true
polyembryony suspected. Kaur et al. (1978) have now confirmed nucellar agamospermy in Shorea
ovalis and S. agami, and strongly inferred it in Hopea subalata and Shorea resinosa.

PIERRE (1889-91), Hem (1892), BRANDIS (1895), BURKILL (1917-25), SYMINGTON (1943) and

264 FLORA MALESIANA [ser. I, vol. 9?

:

Fig. 12. Sapling of Hopea dryobalanoides Mig. Note pendent leaves with drip-tips. Brunei (Photogr. ASHTON,
1958).

1982] DIPTEROCARPACEAE (Ashton) 265

Maury (1978) have described the diverse forms of mature embryo, the last three describing also
germination and seedlings from studies of selected species. Consistent differences exist between
some genera and sections, but many are obscured by bigger differences related to fruit size, which
can vary between closely allied species. As MAuRy (1978) showed, there can be much variation
within species, and unusual forms are associated with distinctive habitats, as in the Dry Dipterocarp
forest species Shorea roxburghii and S. siamensis, in which the germination is cryptocotylar. The
cotyledons are large, +unequal (subequal in many species of Shorea sect. Shorea, Richetioides,
Pentacme and some others, and in most Vatica and Cotylelobium); they are markedly unequal in
Dryobalanops, Neobalanocarpus, and Shorea sect. Anthoshorea, and also in sect. Doona where one,
large and ruminate, is retained within the pericarp at germination, while the other is small, laminar
and photo-synthetic. Often the larger cotyledon is folded round the smaller, which + encloses the
lignified placenta with the radicle lying outside. Both cotyledons are ruminate, much folded and
remain within the pericarp in Dipterocarpus and Stemonoporus, the embryo being freed by
elongation of the cotyledonary petioles; one is thus in Shorea sect. Doona; they are folded but
become freed in Anisoptera, Dryobalanops, Shorea sect. Anthoshorea and some large-fruited Vatica.
In some Vatica, and in Cotylelobium and Stemonoporus the cotyledons are laciniate. In Shorea sect.
Pachycarpae, many species of sect. Brachypterae and some other large-fruited Shorea they are
hastate, fleshy and prismatic, with the 2 inner faces flat and the outer convex; in many such species
they remain within the pericarp until the embryo has rooted as in Dipterocarpus, with the embryo
initially emerging by elongation of the cotyledonary petioles though unlike Dipterocarpus the
cotyledons are ultimately freed. In typical medium or small-fruited dipterocarps (including some
species of Dipterocarpus) germination is epigeal however, the cotyledons expand and become
laminar, remaining yellowish or reddish (violet or magenta in Shorea sect. Richetioides) but
photosynthesising to a limited extent.

The embryo usually remains small until immediately prior to germination, when the radicle
rapidly elongates and bursts through the fruit apex, splitting the pericarp irregularly or (in most
Vatica and in Vateria, Stemonoporus, many Hopea and Dryobalanops) +equally into 3 valves.

The first true leaves are opposite with interpetiolar stipules (except some Vatica), subsequent
leaves spiral except in Dryobalanops, Anisoptera, Vateria, and occasionally others with 2 or more
pairs of opposite seedling leaves. In Anisoptera the first 4 leaves sometimes appear in an estipulate
whorl.

Seedlings and saplings. The first series of spirally arranged leaves in dipterocarp seedlings are
characterised by their thinness, their long slender petioles, and their prominently acuminate apices
(Fig. 5, 12); the tomentum, if present, is longer and sparser than in mature trees; stipules are
narrower and more persistent. In Dipterocarpus the seedlings of all species are very similar in leaf
shape and tomentum, though the stipule and resting bud tomentum already have the diagnostic
characters that they possess in the mature tree. In most other genera the seedling is characteristic of
the section and frequently the species. The seedling leaves give way rather abruptly to leaves
resembling those of the mature tree but which are larger and more attenuate. This transitional stage
is often absent in Vatica and Hopea, but persists until the crown emerges in canopy species.

Taxonomic importance of the structure of embryo and seedling. The potential taxonomic
importance of the characters of the ripe embryo and the germinating seedling in the Dipterocarpa-
ceae was indicated by HEIM (1892) and BuRKILL (1920), and it has been in the Angiosperms as a
whole by STEBBINS (1974), who claimed that “seedlings tend to be constant within genera and
families, and to differ in a regular fashion between these categories, more than any other kind of
characters.”

In dipterocarps, these features in some cases confirm present concepts, and in others suggest some
new relationships (MAUuRY et al., 1975; MAuRY, 1978, 1979). The following characters have proved
useful:

Ripe embryo. The position of the hypocotyl which, on meridian section, is implanted in the
inferior half of the seed, in the superior half, in the middle, or in the apical quarter. When the free
cotyledonary lobes are located towards the right side and the folding axis towards the left one, the
hypocotyl is dexter; it can alternatively be vertical or sinister; the apical hypocotyl can be
horizontal-dexter or recurved-sinister.

The cotyledonary folds, in equatorial section, are covering (folded as the pages in a book), or

266 FLORA MALESIANA [ser. I, vol. 97

encircling (rolled pages), or juxtaposed (a lateral half for each cotyledon). In meridian sections
relative to the placental axis, the cotyledons are piled-up obliquely, or parallel, or superposed. The
apex of the lamina can be folded towards the tip of the hypocotyl.

The lamina of the cotyledons are entire; or lobate, in which case there are two thick lateral lobes
on both sides of the petiole, connected by a narrow, thin meridian zone (MAURY, 1978, vol. 2: p. 117)
or emarginate, that is of intermediary shape. The lamina shape in each of these types can be
transverse (type Anthoshorea: entire, type Shorea: bilobate), or elongate (type Richetioides: entire,
type Mutica: bilobate), or intermediate. In each of these types, the lamina can be either thin or fleshy.

Germinating seedlings. The pericarp either splits under pressure or from the unfolding cotyledons
and elongating hypocotyl, or dehisces independently prior to germination. The growth of the
cotyledons terminates either before the pericarp splits, or later on during the development of the
seedling. Cotyledonary nodes are characterized by the ratio of the number of gaps to the number of
resin ducts per cotyledon. Stomata, when fully developed, provide further useful characters
(Maury, 1978, p. 154). In Dipterocarpoideae the cotyledons reach their final size in the ripe embryo.
Germination leads only to unfolding of the cotyledons and the elongation of the hypocotyl. In this
case, therefore, the ripe seed contains all the systematic and phylogenetic information that
cotyledons can provide. The shape, structure, and organization of dipterocarpoid cotyledons are
constant but only at generic and lower levels. Nevertheless, the interpretation of these characters is
easier in seedlings when the cotyledons have fully unfolded, and when the phyllotaxy of the first
node and the epidermal characters of the first true leaves provide additional information.

Monotoideae and Pakaraimoideae therefore share independent dehiscence of the pericarp,
continued growth of the cotyledons following germination, apical folding of the cotyledons (not
confirmed in Pakaraimaea) and albumen in the ripe embryo, in all these differing from
Dipterocarpoideae. In the Asiatic subfamily, the group of genera Hopea, Neobalanocarpus, and
Shorea (with imbricate ripe fruit calyx) appear to be differentiated from the group of the genera
Dipterocarpus, Anisoptera, Upuna, Cotylelobium, Vatica, Vateria, and Stemonoporus (with valvate
ripe fruit calyx). Parashorea in the former group, and Dryobalanops, which is in the latter on the
basis of embryo characters, have subvalvate calyx in the ripe fruit.

Embryo and seedling characters provide novel insights in the presently recognized genera Shorea
and Parashorea on the one hand, and Vatica, Vateria, and Stemonoporus on the other hand
(Maury, /.c.). On this basis, Shorea sect. Doona, Pentacme, Anthoshorea, Shorea, Richetioides, the
combined sect. Pachycarpae, Mutica, Rubella, Brachypterae, and Ovalis (that 1s, subg. Rubroshorea
MEUER), and the genus Parashorea appear to be at the same taxonomic level. Within this grouping,
Shorea sect. Doona, Pentacme and Anthoshorea form a subunit, Shorea sect. Shorea, Richetioides,
and Rubroshorea another, while Parashorea stands alone but close to ‘Rubroshorea’. The
Rubroshorea grouping is indeed much more heterogeneous than the others.

The relations between and within the genera Upuna, Cotylelobium, Vatica, Vateria, and
Stemonoporus suggest that further discussion of Vatica, and also Stemonoporus might become
justified. The affinities are stronger between Upuna, Cotylelobium and Vatica sect. Sunaptea and
Vatica p.p. on the one hand, and between Vatica sect. Vatica p.p., Stemonoporus and Vateria on the
other hand. The complexity of the latter three groups (Maury, /.c.) requires additional study.

Literature: ANDERSON, Ph.D. Thesis, Edinburgh Univ. (1961); ANTHONY, Acad. Sc. 276 (1973)
193; Gard. Bull Sing. 29 (1974) 17; Marcellia 38 (1974) 99; AsHToN, Gard. Bull. Sing. 20 (1963)
229-284; Man. Dipt. Brun. (1964), Suppl. (1968); Gard. Bull. Sing. 22 (1967) 259-352; Blumea 20
(1972) 357-366; Rev. Flora of Ceylon 1 (2) (1977) 166-196; BRANDis, J. Linn. Soc. Bot. 31 (1895)
1—144; Burkill, J. Str. Br. R. As. Soc. 75 (1917) 43-48; ibid. 76 (1917) 161-167; ibid. 79 (1919) 39-44;
ibid. 81 (1920) 3-4, 49-78; ibid. 86 (1922) 271-291; ibid. 87 (1923) 218-222; Gard. Bull. Str. Settl. 3, 1
(1925) 4-9; CoRNER, Wayside Trees of Malaya (1940) 30; Ann. Bot. 27 (1963) 339; The seeds of
Dicotyledons | (1976) 33, 35, 119, & Atlas; Foxwortuy, Mal. For. Rec. 10 (1932); Gitc in E. & P.
Nat. Pfl. Fam. ed. 2, 21 (1925) 237-269; Gugrin, Bull. Soc. Bot. Fr., Mem. 11 (1907) 93; HALLE,
OLDEMAN & TOMLINSON, Trop. Trees & Forests. An architectural analysis (1978); HeIM, Bull. Soc.
Bot. Fr. 39 (1892) 149-154; Rech. Dipt. (1892) 1-186; HONG, Mal. For. 42 (1979) 280; Kaur et al.
Nature 271, 5644 (1978) 440; MaGurre et al. Taxon 26 (1977) 341-385; Maury, Bull. Soc. Hist. Nat.
Toulouse 104 (1968) 187-202; ibid. 106 (1970a) 277-281; ibid. 106 (1970b) 282-288; Thesis:

1982} DIPTEROCARPACEAE (Ashton) 267

Diptérocarpacees, du fruit a la plantule, Toulouse (1978) 2 vol.; MAURY-LECHON, Mem. Mus. Nat.
Hist. Nat. Paris, n.s. ser. B, 26 (1979) 81-106; Maury et al. Rev. Palaeobot. Palynol. 19 (1975)
241-289: METCALFE & CHALK, Anat. Dicot. | (1950); Mever, Acta Bot. Neerl. 12 (1963) 310-353:
MEUER & Woop, Sabah For. Rec. n. 5 (1964) 1-344; NG, Mal. For. 38 (1975) 153; ibid. 39 (1976) 91;
PigRRE, Fl. For. Coch. 3—4 (1889-91); Rao, Phytomorph. 3 (1953) 476-484; Proc. Nat. Inst. Sc.
India 21 (1955) 247-255; Curr. Sci. 25 (1956) 128-129; J. Mysore Univ. 15, 3 (1956) 7-15; SINGH,
Mal. For. 29 (1966) 13; VAN SLOOTEN, Bull. Jard. Bot. Btzg III, 17 (1941) 132; SYMINGTon, Mal. For.
Rec. 16 (1943); THORENAAR, Meded. Proefst. Boschw. 16 (1926); WHiTMoRE, New Phytol. 61 (1962)
191-220; Gard. Bull. Sing. 19 (1962) 321-371; C. Woon & H. KENG, Gard. Bull. Sing. 32 (dated
1979) 1-42, with numerous figures on 31 pl.

Anatomy. Stem. The whole family shares the combined characters of vestured pits and the
presence of tyloses. The Dipterocarpoideae are distinguished by two unique characteristics: The
lateral (and apical also in Stemonoporus) leaf traces separate from the central vascular cylinder well
below the node, passing up through the bark before entering the petiole; in Dryobalanops they arise
at or even before the previous node. BURCK, PIERRE and HEIM used such characters extensively for
taxonomic purposes, but BRANDIS was rightly cautious of such boldness before much more intensive
examinations have been undertaken; it is likely that such characters could prove useful, especially at
the species level. Vertical intercellular resin canals, scattered or in arcs, are universal also in the
subfamily, and occur characteristically in the pith of internodes and leaf traces; in Dryobalanops
there is a single central branching duct, in others varying numbers of peripheral ducts of varying or
constant diameter. These would also merit intensive investigations.

The wood anatomy has been treated exhaustively by GOTTWALD & PARAMESWARAN who
confirmed and extended the work of Descu, demonstrating that the great diversity of wood
anatomy provides valuable taxonomic characters at subfamilial, tribal, generic, infrageneric and
sometimes species level. The accompanying Table | is modified from their work and summarises the

Table |. Principal groupings of Malesian dipterocarps based on wood anatomy

1 2 Ben s4 5 6 i 8 9 10> DA et3
Vatica most spp. ar + 7 =7
Vatica § Vatica p.p. + + + +
Cotylelobium + *F + 3° a5
Upuna SS == at + =e
Anisoptera =F + + 55 +
Dipterocarpus =e ++ + +
Dryobalanops + _ ~ +
Hopea + +
Hopea ferrea ~ aE +
Neobalanocarpus heimii ~ - +
Shorea § Pentacme + +
§ Shorea _ 35 +
§ Anthoshorea + =e +
Shorea montigena + =F +
§ Brachypterae, Rubella,
Pachycarpae, Mutica + 55
Shorea leprosula, S. ovata,
S. teysmanniana + + +5
§ Richetioides =5 =F +
Parashorea + +
Parashorea smythiesii =F + ==
Key to wood characters:
1. Vessels solitary 6. SiOz frequent 10. Resin canals in medullary rays
2. Vessels grouped 7. SiO2 absent 11. Fibres thin-walled (<3 ym)
3. Uniseriate rays 8. Resin canals scattered 12. Fibres thick-walled (> 3 ym)
4. Rays storied 9. Resin canals in tangential bands 13. Fibres with bordered pits
5. SiO2 abundant, universal

268 FLORA MALESIANA [ser. I, vol. 97

principal grouping they distinguished in Malesian taxa. The diameter of vessels, the size of rays, and
the wood parenchyma distribution are also of more restricted diagnostic value.

Petiole. PIERRE, Heim, MAury, and other authors have used the petiolar anatomy, and
particularly the arrangement of the vascular bundles as seen in transverse section at the distal end
(the ‘caracteristique’), as a guide to classification and species determination. HEIM in particular
placed great weight on small differences observed in single specimens, as in his treatment of
Cotylelobium. | have found that, though some genera possess a characteristic basic arrangement,
variation within single species greatly exceeds variation between species except in some
Dipterocarpus, and sections Shorea and Hopea of those genera, where the anatomy is very complex
owing to the presence of several concentric arcs of vascular bundles. MAurRy (1978) has shown that
the sequential changes in petiolar anatomy in the first leaves of seedlings is distinctive at sectional
and generic level.

Literature: DEN BERGER & ENDERT, Meded. Proefst. Boschw. 11 (1925) 98: BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 1-144; Burck, Ann. Jard. Bot. Btzg 6 (1887) 145-193; DescH, Mal. For. Rec. 14
(1941); GoTTWALD & PARAMESWARAN, Bot. Jahrb. 85 (1966) 410-508; Heim, Rech. Dipt. (1891);
Maury, Thesis: Dipterocarpacees, du fruit a la plantule, Toulouse (1978); METCALFE & CHALK,
Anat. Dicot. | (1957) 212-220; MoLL & JANssonius, Mikrographie des Holzes der auf Java
vorkommenden Baumarten | (1906) 343; MULLER, Bot. Jahrb. 3 (1882) 446; PrERRE, Fl. For. Coch.
(1889-91) t. 212-259; SOLEREDER, Syst. Wert der Holzstruktur (1885) 81.

Palynology. The pollen grains in Dipterocarpaceae are mostly spherical and range in size from
17 um (Vatica havilandii) to 87 wm (Dipterocarpus humeratus). Pakaraimoideae and Monotoideae
are characterized by tricolporate pollen grains with a well developed endexine and a distinct foot
layer. Dipterocarpoideae have tricolpate pollen grains which lack an endexine and possess a thin,
often laminated basal layer which is probably homologous with a foot layer. Exine structure in
Pakaraimoideae is reticulate-columellate with straight columellae, in Monotoideae reticulate-colu-
mellate with columellae inclined towards the centre of the lumina in a tilioid pattern. In
Dipterocarpoideae exine structure varies between a) finely reticulate-columellate (Hopea, Shorea),
b) a structure in which columellae and tectum are intimately fused, forming urn-shaped structures
(Dryobalanops, Dipterocarpus) and c) a tilioid structure (Vatica). The surface of the muri may be
crenelated (Hopea, Shorea, Stemonoporus), crotonoid (Dryobalanops, Dipterocarpus) or smooth
(Dipterocarpus, Vatica). Stemonoporus is characterised by a fairly thick basal layer and operculate
colpi. Within Dipterocarpoideae a relation between flower size and pollen size has been established.
The large flowered genus Dipterocarpus has significantly larger pollen grains than genera with
smaller flowers such as Hopea, Shorea and Vatica, while Dryobalanops is intermediate. Within the
genus Dipterocarpus a similar correlation exists between size of flowers and of pollen. The pollen of
Pakaraimaea is of a generalised type, common in many Angiosperm families, but the pollen types of
Monotoideae and Dipterocarpoideae are more specialized. In Monotes, the thick wall with well
developed endexine and endoapertures may be related to the seasonal climate of dry evergreen
forest to which the genus is adapted. In most Dipterocarpoideae the endexine and consequently also
the endoaperture appears to have been lost, resulting in the development of a thin, very flexible wall,
with slit-like colpate apertures only.

The presence of a tilioid exine structure in Monotoideae and Dipterocarpoideae indicates affinities
with Tiliaceae (Brownlowioideae). The suggestion by KOSTERMANS that Pakaraimaea is close to
Schoutenia (Tiliaceae) is not supported by pollen morphology, since the latter genus has quite
different triporate-echinate pollen. The pollen morphology in Sarcolaenaceae (Chlaenaceae) is also
quite different from Dipterocarpaceae.

Literature: CARLQUIST, Brittonia 16 (1964) 231-254; KOSTERMANS, Taxon 27 (1978) 357-359:
MaAGumirE et al. Taxon 26 (1977) 341-385; Maury et al. Review Palaeob. Palynol. 19 (1975)
241-289. — J. MULLER.

Cytotaxonomy. Despite the ecological and economic importance of the Dipterocarpoideae, we
have at present still rather a limited knowledge of their cytology.

JONG & LETHBRIDGE (1967) and JONG (1976), using the squash technique, established the
occurrence of two main basic chromosome numbers, x = 7 and x = 11 among Indo-Malesian species
in nine genera studied. This is in accord with earlier chromosome counts by Roy & JHA (1965) on

1982] DIPTEROCARPACEAE (Ashton) 269

five Indian taxa representing four genera. Each basic number characterises a group of more or less
related genera: x= 11 the genera Anisoptera, Dipterocarpus, Upuna, Vatica, Vateria and Stemono-
porus, and x=7 Dryobalanops, Hopea, Neobalanocarpus, Parashorea, and Shorea.

These observations are, however, at variance with those of T1xter (1953, 1960) and PANCHO
(1971). Tix1eR published chromosome counts for some 13 Indo-Chinese taxa: 2n = 12 for Pentacme
(reduced here to Shorea sect. Pentacme) and Shorea, and 2n=20 for Anisoptera, Dipterocarpus,
Hopea and Shorea (see Table 2). PANCHO (/.c.) reported also 2n= 20 in two Philippine species of
Dipterocarpus. ROY AND JHA (1965) on the other hand, observed 2n = 22 in Dipterocarpus alatus, a
species also studied by Trx1ER (1953). That dysploidy or aneuploidy may exist within a genus or
species remains however to be firmly established, partly because differences in chromosome counts
appear to vary with observers rather than with taxa. No new basic numbers have yet been detected
in recent cytological studies on some dipterocarps from the Malay Peninsula, Borneo and Ceylon
(JONG & TAYLOR, unpubl.; SINGH, 1977).

The diploid numbers of one species of Stemonoporus, and two of Vateria, all 2n=22, and three
species of Shorea in the sections Doona and Pentacme, all 2n=14, conform to taxonomic
expectation (JONG, 1976; JoNG & Kaur, 1979), thus confirming on the one hand the position of
Stemonoporus and Vateria in the x=11 group of genera, and on the other ASHTON’s (1972)
reduction of Doona and Pentacme to Shorea.

Similarity in chromosome number, it should be emphasized, does not necessarily imply
resemblances in other features of the genome nor an unequivocal indication of close taxonomic
affinity. Dipterocarp chromosomes like those of most other tropical hardwood species are small in
size and rather uniform in morphology, and hence karyotypic variations are not readily discernible.
Although certain differences for example in the number of satellited chromosomes and
chromosome size have been noted by Roy & JHA (1965) among the five species they studied, the
taxonomic value of such variation in the family however, has yet to be properly assessed from a
wider and more representative sample (JONG & Kaur, 1979). Neobalanocarpus heimii (KING)
ASHTON illustrates another important point: this taxonomically problematic species is diploid
(2n= 14) and has a highly irregular pattern of meiosis, at least in the single individual studied (JONG
& LETHBRIDGE, 1967). Irrespective of the significance that might be attached to it (most probably a
reflection of hybridity), such a meiotic pattern could not have been anticipated from an examination
of somatic chromosomes. Thus meiotic information should be sought wherever possible in future
cytological surveys, especially in taxonomically interesting taxa.

Polyploidy. Bearing in mind the difficulties involved in cytological sampling of tropical rain-forest
trees, and the fact that much of the cytological information are based on cultivated specimens (often
on only a single representative of a species), the available data indicate that polyploids are rare in the
Dipterocarpoideae, although polyploid series are now known to occur in Hopea and Shorea
(2n=14, 21, 28). Hopea odorata (2n=14, 20) and Dipterocarpus tuberculatus (2n=20, 30) are
possibly examples of intraspecific polyploidy (see Table 2; also JONG, 1976; JONG & Kaur, 1979).
Hopea nutans and Shorea ovalis ssp. sericea are both tetraploid, and cytologically the latter behaves
as an autotetraploid (JONG & Kaur, 1979). A more extensive sampling might well disclose the
existence of related diploid cytotypes in these two species. Shorea ovalis is morphologically variable,
and it would be interesting to ascertain whether some of the distinct forms of this species are
associated with any cytological variation.

Of special interest are triploid or near-triploid taxa of Hopea and Shorea (JONG, 1976; KAuR et al.,
1978), namely, Hopea latifolia (as H. beccariana in JONG, 1976), H. subalata, and Shorea resinosa, all
2n =3x=21. Hopea odorata has 2n= 20-22.

In view of undecisive counts of 2n = 20, 21 and 22 encountered in root tips of Hopea beccariana,
H. subalata and H. odorata, SoMEGO (1978) proposed that Hopea may be dibasic, with x=7 and
x= 10 or 11, thus denying the possible occurrence of triploidy or near triploidy. Roy & JHA (1965)
earlier published a diploid number 2n = 14 for Indian plants of H. odorata making the interpretation
of 2n=20-22 for the same species (growing in Kepong Arboretum) as a near-triploid more
plausible.

The above species are able to produce viable seeds and, except for the first mentioned, also forma
varying proportion of seeds containing multiple embryos.

270 FLORA MALESIANA [ser. 1, vali

That chromosome sterility resulting from triploidy or near triploidy can be circumvented by
agamospermy has been inferred by KAUR et al. (1978) and JONG & KAuR (1979). Variations in the
somatic number could of course be due to the presence of accessory chromosomes although such
chromosomes have not previously been clearly identified in the Dipterocarpaceae.

The concurrence of triploidy and reproduction by seed points to the possible existence of
agamospermy in these plants. Indirect as well as some embryological evidence (in the case of Hopea
subalata) support such a deduction (KAuR ef al., 1978); conclusive evidence of nucellar
polyembryony has come from tetraploid Shorea ovalis ssp. sericea (SINGH, 1977; KAUR et al., 1978).
Thus there appears to be a close association between polyploidy, multiple seedlings, and
agamospermy and possibly with hybridity in the above mentioned cases; such a relationship is,
however, by no means a universal one for there are diploids such as Shorea agamii (2n= 14) in which
agamospermy Is also suspected to be in operation (KAUR et al., 1978; JONG & Kaur, 1979). This
species also produces multiple seedlings in varying proportions.

Three of the taxa studied by TIxtER (1953, 1960) might possibly be triploid, but their chromosome
numbers need confirmation: one form of Dipterocarpus tuberculatus, 2n = 30; Hopea odorata and
Shorea obtusa, both reported to have 2n = 20.

All the chromosome numbers published to date pertain to genera in the subfamily Dipterocarpoi-
deae and are listed in Table 2. Cytological information is not yet available for any member of
Cotylelobium, nor of the subfamilies Monotoideae and Pakaraimoideae.

Hybridisation. The triploid condition might have arisen in a variety of ways in the different taxa.
In the case of Hopea subalata, known only from Kanching Forest Reserve in Peninsular Malaysia
where a few small groups have been found (SYMINGTON, 1943) triploidy probably occurred as an
isolated event. By contrast, in H. odorata, a riverbank tree of widespread distribution occurring in
the evergreen forests of Indo-China, Burma, Thailand and northern Peninsular Malaysia
(SYMINGTON, 1943; ASHTON, pers. comm.), diploid and near triploid cytotypes are known, and
triploidy could have originated more than once in the different parts of its range, or alternatively it
could have spread with the help of agamospermy from a single origin. A more careful study of these
and other polyploids and their nearest relatives is required before greater understanding can be
attained. Similarly, it is uncertain whether the triploids are derived from occasional union of
haploid and unreduced gametes in a diploid cytotype, or from the habitual union of such gametes as
known only in Leucopogon juniperinum (see SMITH-WHITE, 1955), or from hybridisation between
diploid and tetraploid congeners. It is a well known fact that most triploids have low fertility
(although they are rarely completely sterile, LEwis, 1967), and are thus vulnerable to extinction
unless they possess some means of ensuring their perpetuation at least in the short term. It has been
inferred that agamospermy is the most likely mechanism (KAuR ef a/., 1978), and that a
combination of genes favouring apomixis could have been brought together through hybridisation.
The origin of the tetraploid status of Shorea ovalis ssp. sericea is still uncertain, but the occurrence of
chromosomal heterozygosity and agamospermy (JONG & KAurR, 1979) does strongly point to the
involvement of hybridisation in its origin.

The possible hybrid nature of Neobalanocarpus heimii has already been mentioned (see also JONG
& LETHBRIDGE, 1967; JONG, 1976) and further supportive evidence from meiotic examination of
additional individuals as well as from embryological and other studies of its reproductive biology
are required for a fuller understanding.

The best known dipterocarp interspecific hybrid in the aseasonal tropics is Shorea leprosula x S.
curtisii, both diploids of close affinity. The hybrid has been reported from several localities in
Peninsular Malaysia and from Singapore. Intermediate forms between Vatica rassak and V.
umbonata are found in unstable habitats where the ranges of the two species overlap in East Sabah.
Also some collections of Anisoptera costata and A. curtisii suggest local hybridisation in N.W.
Malaya. See further the notes under the species descriptions. Morphological hybrids are otherwise
rare in the Lowland Mixed Dipterocarp forests (ASHTON, 1969) although they appear to be more
common in the more seasonal regions, especially at zones of contact between allopatric taxa
(examples given in text). There is at present, however, no cytogenetic information on any of the
putative hybrids.

Literature: ASHTON, Bot. J. Linn. Soc. 1 (1969) 149-153; Blumea 20 (1972) 357-366; JONG in
Burley & Styles (eds.), Tropical Trees: variation, breeding and conservation; Acad. Press (1976)

1982]

DIPTEROCARPACEAE (Ashton)

Table 2. Published chromosome numbers of the Dipterocarpaceae

Name

Anisoptera KORTH.

costata KORTH.
(as A. cochinchinensis PIERRE ex
LANESS. )
? laevis RIDL.
laevis RIDL.
scaphula (ROXB.) KURZ
(as A. glabra KURZ)

Dipterocarpus GAERTN. /.

alatus ROXB.
alatus ROXB.
baudii KORTH.
cornutus DYER
costatus GAERTN. f.

(as D. artocarpifolius PIERRE ex

LANESS. )

elongatus KORTH.

(as D. warburgii BRANDIS)
intricatus DYER
kunstleri KING

(as D. speciosus BRANDIS)
oblongifolius BL.
obtusifolius TEYSM. ex. MIQ.
sarawakensis SLOOT.
tuberculatus ROXB.
tuberculatus ROXB. var.
turbinatus GAERTN. f.

Dryobalanops GAERTN. /.

aromatica GAERTN. f.

oblongifolia DYER

Hopea Roxs.

beccariana BURCK
glabraW. & A.

(as H. wightiana WALL. ex. W. & A.)
latifolia SYM.

(as H. beccariana BURCK)
nervosa KING
nutans RIDL.
nutans RIDL.
odorata ROXB.
odorata ROXB.
odorata ROXB.

pubescens RIDL.
sangal KORTH.
subalata SYM.
subalata SYM.

Neobalanocarpus ASHTON

heimii (KING) ASHTON

Parashorea KURZ

densiflora SYM.

Shorea Roxs. ex. GAERTN. f.

acuminata DYER

n

20,

2n

NNN bY WY
SoONNN ©

NN NY WY
NNN OS

Author

TIxiER, 1953

JONG & LETHBRIDGE, 1967
SoMmEGO, 1978

SOMEGO, 1978

TIXxIER, 1960

TIxiER, 1953

Roy & JHA, 1965

JONG & LETHBRIDGE, 1967
SOMEGO, 1978

TIXIER, 1960

PANCHO, 1971
Trxier, 1953

PANCHO, 1971

SOMEGO, 1978

TIxIER, 1953

JONG & LETHBRIDGE, 1967
TIXIER, 1960

TIXIER, 1960

TIXxiER, 1960

JONG & LETHBRIDGE, 1967
SomMEGO, 1978
JONG & LETHBRIDGE, 1967
SOMEGO, 1978

SOMEGO, 1978
JONG & LETHBRIDGE, 1967

JONG, 1976

SOMEGO, 1978

JONG & LETHBRIDGE, 1967
SoMEGO, 1978

TIXIER, 1953

Roy & JHA, 1965

Kaur et al., 1978
SOMEGO, 1978

JONG & LETHBRIDGE, 1967
SOMEGO, 1978

Kavr et al., 1978
SOMEGO, 1978

JONG & LETHBRIDGE, 1967
SoMEGO, 1978

SoMEGO, 1978

Kaur et al., 1978
SOMEGO, 1978

pbs

DD

FLORA MALESIANA

Table 2. (Continued)

Name n
(Shorea)
agamii ASHTON
argentifolia SYM.

assamica DYER
ssp. globifera (RIDL.) SYM.
bracteolata DYER
contorta VIDAL
curtisii DYER ex KING

gardneri (THW.) ASHTON
glauca KING

guiso (BLCO) BL.
hypochroa HANCE
leprosula Miq.

macrophylla (DE VR.) ASHTON
macroptera DYER
ssp. macroptera

maxwelliana KING
mecistopteryx RIDL.
multiflora (BURCK) SYM.
obtusa WALL.
ovalis (KORTH.) BL.
ssp. sericea (DYER) ASHTON 14
palembanica Mia.
parvifolia DYER

pauciflora KING

pinanga SCHEFF.
(as S. compressa BURCK)
pinanga SCHEFF.
pinanga SCHEFF.
platyclados SLOOT. ex. Foxw.
resinosa FOXw.
robusta GAERTN. f.
robusta GAERTN. f. 7
robusta GAERTN. f.
roxburghii G. DON
(as S. talura ROxB.) ij

siamensis Mia.
(as Pentacme siamensis (M1Q.) DC.
singkawang (MiqQ.) Mia.
smithiana SYM. a.
splendida (DE VR.) ASHTON
(as S. martiniana SCHEFF.)
stenoptera BURCK
sumatrana (SLOOT. ex THOR.) SYM. 7
trapezifolia (THW.) ASHTON
Upuna Sym.
borneensis SYM.
Vatica L.
cinerea DYER
odorata (GRIFF.) SYM. ssp. odorata
(as V. grandiflora DYER) 11

—

2n

[ser. I, vol. 9?

Author

KAUR et al., 1978
KAvrR et al., 1978
SOMEGO, 1978

SOMEGO, 1978

SOMEGO, 1978

JONG & Kaur, 1979

JONG & LETHBRIDGE, 1967
SOMEGO, 1978

JONG & Kaur, 1979
SOMEGO, 1978

SOMEGO, 1978

SOMEGO, 1978

JONG & LETHBRIDGE, 1967
SOMEGO, 1978

KAvr et al., 1978

KAvr et al., 1978
SOMEGO, 1978

SOMEGO, 1978

SOMEGO, 1978

SOMEGO, 1978

TIXIER, 1953

SOMEGO, 1978

JONG & LETHBRIDGE, 1967
SOMEGO, 1978

Kaur et al., 1978
SOMEGO, 1978

JONG & LETHBRIDGE, 1967
SOMEGO, 1978

TIXIER, 1960

TIXIER, 1960

JONG & Kaur, 1979
SOMEGO, 1978

KAvr et al., 1978

Rao, 1954

Roy & JHA, 1956, 1965
NANDA, 1962

Roy & JHA, 1965
SOMEGO, 1978

TIXIER, 1953

JONG & LETHBRIDGE, 1967
JONG & LETHBRIDGE, 1967
JONG & Kaur, 1979
SOMEGO, 1978

Kaur et al., 1978

JONG & LETHBRIDGE, 1967
JONG & Kaur, 1979

SOMEGO, 1978
SOMEGO, 1978

Roy & JHA, 1965
SOMEGO, 1978

1982} DIPTEROCARPACEAE (Ashton) 273

Table 2. (Continued)

Name n 2n Author
(Vatica)
pauciflora Mia.

(as V. wallichii DYER) 1] JONG & LETHBRIDGE, 1967
rassak KORTH.

(as V. papuana DYER) 22 JONG & LETHBRIDGE, 1967
stapfiana (KING) SLOOT. 22 JONG & LETHBRIDGE, 1967

79-84; JonG & Kaur, Mem. Nat. Hist. Mus. Paris B 26 (1979) 41-49; JONG & LETHBRIDGE, Notes
R. Bot. Gard. Edinb. 27 (1967) 175-184; Kaur, Ha, JONG, SANDS, CHAN, SOEPADMO & ASHTON,
Nature 271, n. 5644 (1978) 440-442: Lewis, Taxon 16 (1967) 267-271; NANDA, J. Ind. Bot. Soc. 41
(1962) 271-277; PANCHOo, Taxon 20 (1971) 794-795; Rao, Ind. For. 80 (1954) 551-552; Roy & Jua,
Sci. & Cult. 22 (1956) 236-238: J. Ind. Bot. Soc. 44 (1965) 387-397; SINGH (née AWTAR KAUR),
Unpublished Ph.D. Thesis, University of Aberdeen (1977); SMiTH-WHiTE, Heredity 9 (1955) 79-91;
SoMEGO, Mal. For. 41 (1978) 358; STEBBINS, Chromosomal Evolution in Higher Plants; Edward
Arnold, London (1971); SyMINGTON, Mal. For. Rec. 16 (1943); Trxter, Rev. Cytol. et Biol. Veg. 14
(1953) 1-2; ibid. 22 (1960) 65-70. — K. JONG.

Phytochemistry. General chemical properties were summarised by HEGNAUER (1966).
Production of oleoresins (balms, resins) is characteristic of most members of the family. Their
volatile portion consists mainly of sesquiterpenes such as humulenes, caryophyllenes, copaénes,
elemenes and guajenes (e.g. gurjunenes, apitonene); in some instances monoterpenoids predomi-
nate (i.e. borneol in the so-called “Borneo camphor from Dryobalanops aromatica. The
sesquiterpene alcohol spathulenol occurs in balms of species of three of the four subgenera of
Shorea (BISSET et al., 1971). The resin fractions of the oleoresins are composed of triterpenoids and
usually consist of neutral and acidic constituents. Dipterocarpol (= hydroxydammaradienone-II) is
a ketonic tetracyclic triterpene alcohol having the so-called dammarane skeleton; together with
similar compounds like dryobalanone it represents an outstanding feature of the subfamily
Dipterocarpoideae. The dammarane skeleton is also present in a number of acidic resin constituents
such as dipterocarpolic acid, dammarenolic acid (I) and shoreic acid (I]). Other dipterocarpaceous
resin triterpenoids possess the pentacyclic skeletons of ursolic acid (e.g. ursonic acid, asiatic acid,
the lactonic compound B [III] erc.), oleanolic acid (e.g. oleanolic acid, hederagenin efc.) and
betulinic acid (e.g. erythrodiol). Compounds I, II and III have an oxidatively cleaved A-ring: they
represent so-called A-ring seco-triterpenes, which seem to be rather characteristic of dipterocarps.
Many of the oleoresin constituents mentioned were described since 1966 (e.g. CHAN, 1969; CHEUNG,
1967, 1968; CHEUNG & FENG, 1968; CHEUNG & YAN, 1972; CHEUNG & WONG, 1972; GUPTA & SUKH
Dev, 1971: HARRISON et al., 1971; LANTZ & WOLFF, 1968). Some attention has been paid also to the
phenolic constituents of leaves, barks, woods and seeds. Dipterocarps tend to produce proantho-
cyanidins (i.e. oligomeric catechins formerly called leucoanthocyanidins) and gallic acid deriva-
tives. These polyphenolic compounds are building stones of condensed and hydrolysable tannins:
both types of tannins are present in taxon-characteristic ratios and amounts in many members of
the family. Two derivatives of gallic acid deserve mentioning. Ellagic acid, the dilactone formed on
hydrolysis of ellagitannins, was shown to occur in leaves and seeds of many species and bergenin, a
striking derivative of gallic acid, has been isolated up to this day from members of Dipterocarpus,
Shorea, Stemonoporus and Vateria (BHRARA & SESHADRI, 1966; Desai et al., 1967, 1971:
BANDARANAYAKE et al., 1977). Most probably both compounds will turn out in future to represent
good chemical characters of Dipterocarpaceae. The same may be true for Hopea-phenol, a phenolic
constituent of barks and heartwoods, which is presently known from species of Balanocarpus,
Hopea and Shorea (COGGON et al., 1965, 1966; MADHAV et al., 1967). Hopea-phenol was shown to
be a condensation product of four molecules of the trihydroxystilbene resveratrol. It is chemically
similar to the viniferin-type phytoalexins of Vitis vinifera. With regard to phenolic leaf constituents
BaTE-SMITH and WHITMORE (ex HEGNAUER, 1966) stressed the frequent occurrence of vicinal
trihydroxylation (ellagic acid, gallic acid; B-ring in the flavonoids myricetin and prodelphinidin) in
dipterocarps. As far as investigated, seed fats (oils) of Dipterocarpaceae are characterized by a

274 FLORA MALESIANA [ser. I, vol. 92

strong predominance of stearic and oleic acid. Sal fat (oil of seed kernels of Shorea robusta) was
shown recently to contain also small amounts (c. 4°%) of 9,10-epoxystearic acid (BRINGI, 1972).

BATE-SMITH & WHITMORE (1959) examined the phenols of fresh leaves in 28 species in 8 genera,
giving attention to those compounds known to be of chemotaxonomic interest elsewhere. A
grouping of genera was arrived at on the basis of the leucoanthocyanins present and their
abundance which little reflected grouping established by traditional means; no clear grouping of
Shorea species by their established sections was possible, though Neobalanocarpus was confirmed to
closely resemble Hopea.

Recently taxonomic potentialities of chemical characters at an intrafamiliar level were discussed
by several authors. Diaz et al. (1966) and Bisset et al. (1966, 1967, 1971) showed that the
composition of the oleoresins (sesquiterpene-fractions, triterpene-fractions) is rather characteristic
of the taxa Shorea sect. Doona, Anisoptera, Cotylelobium and Upuna and that in the genara
Dryobalanops, Dipterocarpus and Shorea the chemistry of oleoresins might be helpful to
classification beneath generic level. Subsequently BANDARANAYAKE et al. (1975, 1977) stressed the
systematic importance of resin composition in dipterocarps. According to these authors the
significant differences in resin composition between representatives of Shorea sect. Doona and other
sections do not agree with the proposition to merge Doona with Shorea. Very little work was
performed with representatives of the African subfamily Monotoideae (Monotes, Marquesia) which
lack resin ducts and seem to deviate chemically in several respects (D1Az et al., 1966) from the
Asiatic subfamily Dipterocarpoideae. At present a chemotaxonomic discussion of relationships
between these subfamilies seems to be premature. The same is true with regard to the recently
described New World dipterocarpaceous genus Pakaraimaea (MAGUIRE & ASHTON, 1977) which
according to KOSTERMANS (1978) would belong to Tiliaceae. The chemical evidence (GIANNASI &
NIKLAS, 1977) given for dipterocarpaceous affinity is inadequate. Confirmation of the preliminary
results reported by the authors mentioned as well as extension of phytochemical research are needed
before chemical characters can make a serious contribution to the classification of the taxa
concerned. Regarding relationships of Dipterocarpoideae with other plant families, the opinion held
in 1966 by the present author is still valid; we are not yet in a position to discuss relationships in
terms of chemical characters. On morphological arguments relationships with members of Malvales
are often postulated. Presently known chemical characters do not convincingly contradict such a
classification, but they form by no means strong evidence for such an affinity.

Literature. BANDARANAYAKE et al., Phytochemistry 14 (1975) 2043; ibid. 16 (1977) 699;
BATE-SMITH & WHITMORE, Nature 184 (1959) 795-796; BHRARA & SESHADRI, Curr. Sci. 35 (1966)
486; BisseT et al., Phytochemistry 5 (1966) 865; ibid. 6 (1967) 1395; ibid. 10 (1971) 2451; BRINGI,
Chemistry and Industry, London (1972) 805; K. C. CHAN, Phytochemistry 8 (1969) 1051; H. T.
CHEUNG, Tetrahedron Letters (1967) 2807; J. Chem. Soc. (1968) C, 2686; H. T CHEUNG & M. C.
FENG, J. Chem. Soc. (1968) C, 1047; H. T. CHEUNG & C. S. WONG, Phytochemistry 11 (1972) 1771;
H. T. CHEUNG & T. C. YAN, Austr. J. Chem. 25 (1972) 2003; CoGGON et al., J. Chem. Soc. (1965)
406; ibid. (1966) 439; Desat et al., Indian J. Chem. 5 (1967) 523; ibid 9 (1971) 612; D1az et al.,
Phytochemistry 5 (1966) 855; GIANNASI & NIKLAS, Taxon 26 (1977) 380; GupTA & SUKH Dev,
Tetrahedron 27 (1971) 635, 823; HARRISON et al., J. Chem. Soc. (1971) C, 2524; HEGNAUER,
Chemotaxonomie der Pflanzen 4 (1966) 31-44, 446-448, 487-488: KOsTERMANS, Taxon 27 (1978)
357; LANTZ & WOLFF, Bull. Soc. Chim. France (1968) 2131; MADHAV et al., Phytochemistry 6 (1967)
1155: MAGUIRE & ASHTON, Taxon 26 (1977) 341-385. — R. HEGNAUER.

Taxonomy. Historical review. The first mention of the family of great trees that dominate the
lowland and hill-forests of the Far East in European literature is in the diary of MARco POLo, who
recorded visiting Fansur in Sumatra, considered to be the present Baros on the west coast, where the
camphor tapped from the hinterland was at that time literally worth its weight in gold. Indeed, up
till the mid-eighteenth century it was only the Sumatran camphor which attracted the attention of
European travellers. It is mentioned in Dutch literature several times in the mid and late 17th and
early 18th century. RuMpHrus (1755) described the Arbor Camphorifera II occidentalis and also
mentioned an Arbor Koring (1741), which MERRILL (1917) regards as Dipterocarpus hasseltii BL.
(the name kéruing and variations of it being the Malay and Indonesian generic name); MERRILL’S
evidence is meagre however. The camphor tree receives further mention from CHARLES MILLER
(1778) who sent a specimen to BANKS from Sumatra.

1982] DIPTEROCARPACEAE (Ashton) 275

Meanwhile LINNaEus had received material from India of a plant that he described as early as
1737 as the genus Vateria, and in the Ist edition of Species Plantarum (1753) and 5th edition of
Genera Plantarum (1754) as Vateria indica. This he placed in his Class Polyandria Monogynia
between Mesua and Thea, and shortly after Microcos, Tilia, and Elaeocarpus. In 1771 he described
Vatica in Mantissa Plantarum, placing it in Dodecandria Monogynia with Befaria Mutis. DE
Jussieu (1789) placed Vatica and Vateria under “Genera alternifolia, hinc Guttiferis, inde Aurantiis
affinis’, with two other genera of undecided affinity — Allophyllus (now in Sapindaceae), and
Elaeocarpus. The two then known genera were thus brought together for the first time. In 1824 A. P.
DE CANDOLLE placed Vatica under Tiliaceae, but omitted mention of Vateria.

In 1825 SPRENGEL included Shorea GAERTN. and Dipterocarpus GAERTN. under Polyandria
Monogynia, citing Dipterocarpus in the Tiliaceae. Vatica and Hopea Roxs. he included in
Dodecandria Monogynia following LINNAEUS. In 1828 REICHENBACH placed the then known genera
in his Laurineae d. Pterigiae, adopting the latter name after CORREA’s genus Pferigium (1806), in
which had been included Dipterocarpus and Dryobalanops GAERTN. described the year previously.

In 1825 however BLuMe had created the family Dipterocarpeae, stating that it bore affinities to the
Tiliaceae in the contorted corolla, and to the Guttiferae in the resin ducts, superior ovary, many
stamens, and single exalbuminous seed.

LINDLEY (1836) put the ‘Dipterocarpeae’ with Sterculiaceae, Malvaceae, Elaeocarpaceae,
Tiliaceae and Lythraceae into his Alliance Malvales, commenting that BLUME had noticed affinities
with Guttiferae. So did MEISNER (1837) who placed Dipterocarpaceae next to Sterculiaceae,
Tiliaceae and the Madagascan Sarcolaenaceae (Chlaenaceae).

ENDLICHER (1840) on the contrary had put the family with the Class Guttiferae, with Chlaenaceae.
Ternstroemiaceae, Clusiaceae, Marcgraviaceae, Elatineae, Reaumuriaceae and Tamaricaceae, thus
far separated from Tiliaceae. He also erected the African genus Lophira BANKS ex GAERTN. (now
Ochnaceae), which GUILLEMIN (1830) had considered a dipterocarp, into an order of its own, and
reduced Shorea ROxB. ex. GAERTN. to Vatica L. This classification was subsequently followed by
LINDLEY (1846). BENTHAM & HOOKER (1862), besides accepting GUILLEMIN’s conclusions on
Lophira, included also Ancistrocladus WALL. PLANCHON (1849) had previously put both genera ina
group of their own allied to the Ochnaceae. BENTHAM & HOOKER resurrected Shorea, and
maintained the family in their Cohors Guttiferales, though stating its affinities to be with the
Tiliaceae as well as the Ternstroemiaceae.

The first complete monograph of the family appeared in A. DE CANDOLLE’s Prodromus (1868): he
enumerated 126 species in 13 genera including Monotes; 27 years earlier KORTHALS (1841) had
estimated the total known species at only 34. DE CANDOLLE again placed Ancistrocladus and
Lophira in separate families; he further described the first known African dipterocarp as Monotes
africanus, indicating by its name that the genus occupied an isolated place in the family. He claimed
the affinities of Dipterocarpaceae to be with Chlaenaceae and Ternstroemiaceae, with Lophiraceae
and Ancistrocladaceae as intermediate groups.

In 1874 Dyer monographed the genera Dipterocarpus and Dryobalanops. BURCK (1887) made a
study of East Indian dipterocarps based in large part on anatomical characters; he created no new
genera but united the genera Pentacme DC., Monoporandra THw. and Stemonoporus THw. with
Vateria and transferred part of the genus Hopea Roxs. to Doona THW.

Hetm’s ‘Recherches sur les Diptérocarpacées’ (1892) remains to this day the most detailed study
of the whole family. Though he frequently made anatomical studies of leaf, petiole, twig and fruit,
he gave particular weight to the characters of the stamens, the embryo and to the ‘caracteristique’
(the arrangement of the vascular bundles as seen in transverse section in the petiole at the umbo).
His system suffered because he was working at a time when herbarium collections were quite
inadequate in quality and in numbers for his task. The result was unfortunate; whereas BURCK five
years previously had recognised only 10 genera, HEIM maintained 30, in 8 series and 2 subseries; of
‘these 13 genera were new though based altogether on only 17 species, of which 11 were described
from single herbarium sheets. The genus Cotylelobiopsis HEI, for instance, was described from a
single sterile sheet in which the anatomy of the petiole was found to be unique in the family; the
specimen, which is lost, appears to represent fallen leaflets of Pseudosindora palustris Sym. in the
Leguminosae. Cotylelobium melanoxylon was represented under three binomials, each based on a
single specimen. Of the 4 species recognised by HEIM in his genus Richetia, SYMINGTON (1933: 153)

276 FLORA MALESIANA [ser. I, vol. 9?

later correctly reduced 3 to a single species already described by BURCK. The 4 genera united under
Vatica by BuRCK were redivided and placed in 2 series; Monotes DC. was removed to the Tiliaceae.

HeIM promised a more complete monograph at a later date, but in 1895 the Dipterocarpaceae
were treated by BRANDIS and GILG for the Pflanzenfamilien; later in the same year BRANDIS also
published an ‘Enumeration of the Dipterocarpaceae . . .’, based on the specimens at Kew and the
British Museum. Monotes was reunited with the Dipterocarpaceae, while none of HEIM’s genera
were accepted and few of his species. They maintained but 16 genera, being a return to DE
CANDOLLE’S generic concept, with the addition of Balanocarpus, Cotylelobium, Parashorea and
Isoptera, described subsequently to 1868; the reduction of Petalandra Hassk. to Hopea, and the
maintenance of Stemonoporus separate from Vatica, under which name it had been reduced by DE
CANDOLLE. They recognised 5 tribes in Dipterocarpaceae sens. str. following BRANDIS. Later GILG
(1899) placed the African dipterocarps in a separate subfamily. His account in the 2nd edition of the
Pflanzenfamilien (1925), which is the most recent of the whole family, is with this exception mainly a
reprint of the 1895 account. In 1941 SyMINGTON described the genus Upuna; this genus necessitates a
redefinition of BRANDIS’S tribes, and this has been discussed by me (1978). Two tribes are now
recognised in Dipterocarpaceae on the basis of the calyx, resin canal distribution and basic
chromosome number. One includes BRANDIS’s tribe Shoreae and Dryobalanops, while the remaining
genera are contained in the other.

Subdivision of the family. The family is subdivided into three subfamilies (MAGUIRE & ASHTON,
1977), which can be distinguished as follows:

1. Anthers basifixed; pollen tricolpate; exine 2—3-layered. Sepals, if ampliate, then generally
conspicuously unequal with 2-3 becoming alate in fruit. Ovary (2—)3-celled, each cell with 2
ovules. Wood, leaves and ovary with resin or secretory ducts; wood rays multiseriate. 13 genera:
tropical Asia and Malesia ._. . . . Dipterocarpoideae

1. Anthers basi-versatile; pollen tricolporate: exine 4 layered. Sepals equally accrescent, papyra-
ceous. Ovary (2-)3-4(5-)celled, each cell with 2-4 ovules. Wood, ovary and, commonly, leaves
without resin or secretory ducts.

2. Petals longer than sepals. Anthers little or deeply basi-versatile, connective little or
moderately projected as an apical appendage. Ovary 3—(4-)celled, each cell with 2 ovules.
Wood rays dominantly uniseriate. 2 genera: tropical Africa and Madagascar (1 species).

Monotoideae

. Petals shorter than sepals. Anthers deeply basi-versatile, connective conspicuously projected
as an apical appendage. Ovary 4~(5-)celled, each cell with 4 ovules. Wood rays dominantly
biseriate. 1 genus: tropical America . . . . . . . . +. . .Pakaraimoideae

i)

Recently KOSTERMANS (1978) suggested that Pakaraimaea would belong to Tiliaceae and might
even be congeneric with Schoutenia. This opinion rests on a superficial comparison and his
arguments are in part erroneous, e.g. on pollen and wood anatomy. It is now definitely proved that
Pakaraimaea belongs to Dipterocarpaceae (MAGUIRE & ASHTON, 1980).

Subdivision of the Dipterocarpoideae. Two tribes are recognised:

1. Fruit calyx lobes valvate at base, calyx cup vasculation not overlapping in flower. Vessels usually
solitary, resin canals scattered. Basic chromosome number (probably) x=11: Vateria,
Vateriopsis, Stemonoporus, Vatica, Cotylelobium, Upuna, Anisoptera, Dipterocarpus

Tribe Dipterocarpeae

2. Fruit sepals expanded and imbricate at the incrassate cupped base, calyx cup vasculation + over-
lapping at margin in flower. Resin canals in tangential bands. Basic chromosome number
(probably) x = 7: Dryobalanops, Parashorea, Hopea, Neobalanocarpus, Shorea. Tribe Shoreae

The two large genera Hopea and Shorea differ in a single character of the fruit calyx, indiscernible
in those species where the sepals are short and equal and which are assigned to their respective
genera according to the flora characteristics by which their sections are defined. Neobalanocarpus
heimii, with unique floral morphology and short subequal fruit sepals, is thus unassignable but in
several ways intermediate.

Generic and infrageneric delimitation (for a review see ASHTON, 1979a). A wide range of characters
of flower and fruit, leaf, bark and wood are available for distinguishing between dipterocarp genera.

1982] DIPTEROCARPACEAE (Ashton) 277

In tribe Dipterocarpeae the Malesian genera are marked by their internal uniformity and a unity
imposed by a large number of anatomical and morphological characteristics. This is particularly so
of the large genus Dipterocarpus. Vatica and Cotylelobium differ principally in flower and wood
structure, which are nonetheless rather uniform intragenerically; but it is remarkable that the two
distinctive forms by which the two sections of the large genus Vatica are distinguished are identically
repeated in Cotylelobium with a mere 5 species of which one, C. lewisianum (TRIM. ex HOOK. f-)
ASHTON of Ceylon, has short, equal, reflexed fruit sepals and a thick-walled, loculicidally sutured
pericarp. The monotypic Upuna is also very well distinguished, as is Anisoptera which nonetheless
contains two uniform sections differing only in floral structure.

The remaining genera pose more or less considerable problems of definition. Dryobalanops and
Parashorea are characterised by subequal typically long fruit sepals and are distinct anatomically
and in leaf venation. Parashorea shares alone with Dipterocarpus in the family its plicate vernation;
Dryobalanops has unique leaf venation; each are further distinguished by a constant and
characteristic floral and embryological structure. The remaining 253 Malesian species can be
subdivided into 18 groups, of varying size and distinctiveness but generally great internal
uniformity in diagnostic characters, on the basis of the androecium. In some, such as the large
sections Anthoshorea and Richetioides (also Doona, endemic to Ceylon) of Shorea, these characters
are correlated with equally constant and distinctive bark, wood, embryo and other characters. In
others, such as Hopea subsect. Hopea, Neobalanocarpus and Shorea sections Shorea, Pentacme and
Neohopea significant differences in floral, and in the case of Hopea fruit morphology, occur to
distinguish them from one another, yet all share an essentially uniform leaf, wood and bark
anatomy which often makes them impossible to assign in the field when sterile. The same is so of the
wood of Shorea sections Rubella, Brachypterae, Pachycarpae and Mutica, though here each
possesses distinctive bark manifestations, and indeed sections Brachypterae and Rubella more
closely resemble in floral structure sect. Anthoshorea though differing significantly in wood and bark
anatomy. We have here therefore a constellation of taxa, the most distinct of which approach the
status of genera equivalent to, for instance, Vatica and Cotylelobium, the least not meriting more
than subsectional status, yet all clearly part of a greater coherent group. Ideally all might be
included in a single common genus Shorea of many sections, yet as generic definitions were formerly
based on fruit calyx morphology the sections are presently included in two major genera, Shorea
and Hopea, differing only in a single character, the number of aliform fruit sepals. Species therefore
with short subequal fruit sepals — regarded in these taxa as a derived condition — are assigned to a
genus according to their sectional characteristics. Shorea isoptera, however, of the monotypic sect.
Neohopea, has 5 aliform subequal fruit sepals and unique floral morphology and is only assigned to
Shorea owing to its overall resemblance in bark, wood and habit, to members of the type section; it
would be undesirable to create a new genus for a single species in many respects intermediate
between such similar genera. Neobalanocarpus likewise has short subequal sepals and unique floral
morphology; it appears, on leaf characters, wood anatomy, biochemistry and habit to come closest
to Hopea, but is kept separate for historical reasons and as perhaps the most celebrated forest tree of
Malaya. Hopea and Shorea nevertheless are more or less recognisable as entities, the former being
generally smaller trees and frequently stilt-rooted; foresters in tropical Asia would damn the
botanist who undertook 86 nomenclatural changes in the quest of a spurious taxonomic ideal, and
so would I.

Specific and infraspecific delimitation (for a review see ASHTON, 1978). The subfamily is mainly
(though not exclusively) composed of morphologically well defined species in which with experience
delimitation should present no difficulty. A main source of misinterpretation has been owing to the
great difference in vegetative characters between the young and mature stages and hence to the lack
of field experience on the part of many monographers. The early collectors frequently did not have
the opportunity to collect leaf specimens from the crowns of the giant trees, and instead collected
fallen fruit and saplings. Later authors were unable to establish the true identity of species described
from such material, and redescribed material from the mature tree, when it became available, as a
new species. This confusion is accentuated by the uniformity of flower and fruit within some
sections, and the great value of leaf, tomentum and bark characters in diagnosis when the variability
during the life of each species is understood.

Size differences are not by themselves sufficient to distinguish species, neither are therefore

278 FLORA MALESIANA [ser. I, vol. 9?

differences of leaf size and shape together. Differences in fruit size are likewise unreliable and rarely
correlate with other characters; collections from one tree in different years often exhibit great
variation. A consistent discontinuity in leaf size, where correlated with differences in androecium or
gynoecium, in qualitative (but not quantitative) characters of indumentum, with qualitative
characters of the twig or stipule or with a discontinuity in the range 1n the number of leaf nerves does
constitute an adequate criterion for separating species however.

In the absence of any possibility of proving interfertility taxa are designated as subspecies in the
following circumstances:

(1) Where discontinuities occur in the range of quantitative characteristics of parts, or tomentum
distribution and density, but no qualitative difference exists.

(2) Ifa series of clearly defined, but closely similar, taxa exist which occupy distinct geographical
areas, even if intermediate forms are not known. The subspecies of Dipterocarpus conformis and
Shorea macroptera serve as examples.

(3) Ifa series of closely similar taxa occupy distinct, but greatly overlapping, areas and in at least
one area show intermediate forms. In Shorea parvifolia, for example, two subspecies are recognised
with greatly overlapping ranges; a third form (the ‘Perak form’ of SYMINGTON, 1943) occurs in
Malaya and in N.E. Borneo which possesses characters intermediate between the other two and
may represent a hybrid between the two subspecies.

By contrast, though several forms of Shorea pinanga occur, apparently with different ecological
ranges, the vast number of collections of this species show a bewildering range of more or less
continuous variation, made more confusing as the varying characters, leaf size, number of nerves,
and tomentum, also change much with the age of the plant; here subspecies are therefore not
recognised.

Polymorphism of the latter type is unusual, tending to occur within Malesia in species within
groups (such as Shorea sect. Pachycarpae and the New Guinea Hopeas) consisting of large numbers
of species confined to a single geographical area, possibly undergoing active diversification at
present. It is characteristic also of those species (e.g. Shorea siamensis, Dipterocarpus obtusifolius, D.
tuberculatus, Anisoptera costata, and A. thurifera) which occur in both seasonal and non-seasonal
climates and which become gregarious and regenerate freely in secondary forest or fire-savanna.
Here too the only well authenticated hybrid populations between markedly dissimilar species of
Dipterocarpus are known; examples are recorded by Foxwortny from Luzon, by KERR (1914) from
Thailand, and by PARKER (1927) from Burma.

Notes are included after the descriptive texts, which elaborate on complex patterns of
morphological variation in these taxa, including certain sections of Shorea, in which they prevail.

The apparently trivial yet remarkably constant differences which distinguish the multitude of
species of non-seasonal western Malesia, their morphological uniformity as a rule, their tendency
(see below, under evolutionary trends) in that region towards reduction of staminal size and
number, their occasional flowering and the existence of polyembryony and triploidy open the
possibility that apomixis may have become general under these climatic conditions. FEDOROV (1966,
1977), ASHTON (1969, 1977, 1978) and MEWER (1974) have discussed the problem of dipterocarp
speciation, but conclusive evidence is only now beginning to arrive (KAUR et al., 1978).

Evolutionary trends within the family (for recent reviews see MAGUIRE & ASHTON, 1977; ASHTON,
1979b). Fruit calyx. Without resorting to theories to decide which characters are primitive, it is safe
to assume that departure from regular actinomorphy in the dipterocarp flower must represent an
advanced condition. This, as the family name suggests, occurs in the calyx following anthesis. The
genera always possessing equal fruit sepals are the American subfamily Pakaraimoideae and the
entire African subfamily Monotoideae, the short-sepalled Vateria L. of Ceylon and S. India,
Vateriopsis of the Seychelles, Stemonoporus of Ceylon, and Dryobalanops with either short or
aliform fruit sepals. Vatica sect. Vatica is also characterised by short equal fruit sepals, whereas in
sect. Sunaptea they are unequal; species in the small genus Coty/elobium possess fruit identical to
those of the latter section but for C. /ewisianum (TRIM.) ASHTON of Ceylon in which they are as in the
type section. In Dipterocarpus, Parashorea, and all sections of Shorea and Hopea, however,
individual species occur also with short subequal fruit sepals: the evidence suggests that in these
latter genera this condition is a secondary reversion: of the 66 species among these genera with short
subequal sepals (3 in Parashorea and Dipterocarpus, Neobalanocarpus, 19 in Hopea, and 40 in

1982] DIPTEROCARPACEAE (Ashton) 279

Shorea) all but 8 are rare local endemics or confined to the island or country in which they occur. Of
the 7 relatively widespread species all but 2 are typically riverine or swamp species; this fact,
especially significant in a family usually characteristic of well drained land, suggests a secondary
adaptation to water dispersal. Further, the group among these genera where the short-sepalled
condition is most prevalent, and in which the non-riverine widespread species Shorea multiflora and
S. balanocarpoides occur, is sect. Richetioides of Shorea. In the unique leaf nervation, the great
restriction in most species of the tomentum so characteristic of the family, and in the characteristic
reduction of the number of pollen-sacs from the usual 4 to 2, they give every indication of being an
aberrant advanced group: this is supported by phytogeographic evidence, the section being large
but confined to Malaya, Sumatra, Borneo and the Philippines. Again. some of these local endemics
bear close resemblances to unequal-sepaled species of widespread distribution. As examples from
many, Shorea biawak and S. asahi, both known only from northern Borneo have close affinities with
S. maxwelliana and S. laevis respectively, both of which range through Borneo, Sarawak and
Malaya.

According to CoRNER (1949) an aril in an indehiscent fruit is a relic from a dehiscent state.
SYMINGTON (1941) described a vestigial aril-like structure in Upuna, and considered that the ‘cupule
chalazique’, which was observed by Heim (1892) in dried material of Stemonoporus spp. as a
cup-shaped process from the funicle, might also be an aril; this structure also occurs in Vateriopsis.
and the highly vascularised placenta of Pakaraimaea may be homologous. No known dipterocarp
has fruit dehiscent on the tree, but in Marquesia, Stemonoporus and Upuna the pericarp of the fallen
fruit splits along three loculicidal sutures at germination and in the first two there is sometimes
active recurving of the pericarp apex. The mature fruit of Pakaraimaea is unknown, but as the
pericarp is extremely thin along its five angles it would appear to be capsular. Such sutures occur
also in Vateria, Stemonoporus, Dryobalanops oblongifolius and in Vatica sect. Vatica. In other
Dryobalanops and some Hopea sutures are not discernible but the pericarp often splits into 3 equal
valves at germination; the pericarp of other dipterocarps is split more or less irregularly by the
expanding embryo. Of the taxa with loculicidal sutures only Upuna has unequal fruit sepals:
conversely all the natural groups within the Asian subfamily which have equal sepals as a constant
character are included among those with sutures, and of these all but Dryobalanops have fruit sepals
generally rotate or reflexed and shorter than the ripe seed.

Within Dipterocarpoideae there appears also to have been a reduction in the size of the seed and
thickness of the pericarp. In Vatica sect. Sunaptea the fruit is small, the pericarp thin-walled, and the
sutures obscure or absent. In the type section of Vatica most species have large fruit with
thick-walled pericarps. Some, such as V. vinosa, V. pallida, V. flavida and V. lobata have fruit as
small as those of sect. Sunaptea: these are further characterised by their comparatively thinner
pericarp and obscure sutures. All however are local endemics except for the riverine V. venulosa
which may well have extended its range rapidly and recently by water dispersal: hence they are
unlikely to be ancient species. In other genera the same correlations prevail with the exception of the
rather isolated genus Dipterocarpus, in which all the species have a large indehiscent nut enclosed in
a calyx tube, the clearly recent and still diversifying sect. Pachycarpae of Shorea in which the largest
fruited species are adapted to water dispersal, and a few other isolated Shorea species (e.g. S.
geniculata) where the large fruit appears to be recent rather than ancient, but whose adaptive
significance is not understood.

Stamens. The number of dipterocarp stamens may vary from 5 to c. 105. The most widespread
number is 15; 10 occur in some Hopea, some species of Shorea sect. Richetioides and sometimes in
abnormal flowers in sect. Mutica. The two Stemonoporus species formerly in Monoporandra, and a
single Bornean species of Vatica, possess 5 only. Genera with many stamens as a characteristic are
Pakaraimaea, Marquesia, Monotes, Vateria, Upuna, Dryobalanops and Dipterocarpus: it is usual
also in the type section of Anisoptera, and sect. Anthoshorea, Ovalis and Shorea of Shorea.
Phytogeographical evidence within Asia lends support to the suggestion that the primitive type had
numerous stamens, and that genera with 15, 10 or 5 stamens reached these numbers by reduction.
Vateria occurs in southern India, Ceylon, and the Seychelles whereas Stemonoporus, with 5 or 15, is
endemic to the wet zone of Ceylon, Anisoptera sect. Glabrae, with 15 stamens, ranges from Burma
and Indo-China through Malaya to Borneo, whereas sect. Anisoptera with many stamens occurs
from Burma and Indo-China through Malaya and Sumatra, Borneo, the Philippines. and across

280 FLORA MALESIANA [ser. I, voles?

Wallace’s line to New Guinea. Within Shorea, though the monotypic and probably advanced sect.
Ovalis, with many stamens, occurs in Malaya, Sumatra and Borneo, the large sect. Shorea and
Anthoshorea (with 15-~ stamens) are considerably more widespread than the equally large sect.
Mutica, Richetioides and Brachypterae (typically with 10-15 stamens with the exception of two
species). The former occur east of Borneo and the Philippines, and through India to Ceylon; the
latter are confined to the everwet region of western Malesia with the exception of one species in the
Moluccas. In this case a clear pattern emerges of a reduction of staminal number among local
endemic species of the everwet zone, especially in the geologically recent region of lowland Borneo.
The widespread species, often of the seasonal tropics and including the American and African
subfamilies, retain the primitive condition. This pattern is most clearly seen within those groups in
which staminal number is variable.

Wood anatomy. GOTTWALD & PARAMESWARAN (1966) independently regard a characteristic type
of multiple perforation, along with different stages of disintegration of the plate-membrane, in
vessels of Vateria and certain species of Vatica sect. Vatica, and of spiral thickenings in vessel and
parenchyma wells in the latter, as clearly indicating the primitive condition of the wood of these
groups in the Asian subfamily. However, the aberrant type of multiple perforation plate,
sporadically occurring in the woods of these taxa as well as the spiral thickenings to the vessel walls
can —in the opinion of P. Baas (personal comm.)— equally well be regarded as unusual
specialisations in the Dipterocarpaceae of no phylogenetic significance whatsoever (cf. the
occurrence of similar aberrant perforation plates in the wood of apple and pear trees, which cannot
be interpreted as an argument that Malus and Pyrus are primitive in the Rosaceae!). On the basis of
xylem anatomy Stemonoporus is closely allied with them, whereas Upuna is in many respects
intermediate between them and Anisoptera, Dipterocarpus and Cotylelobium. Shorea, Parashorea
and Hopea differ from other Dipterocarpoideae in their tangentially arranged resin canals (diffuse in
other genera). Their conclusions on phylogenetic affinities are closely in accord with those derived
from other evidence: “The evolutionary status within the (sub)family is characterised by a
concentration of the non-advanced characters in the taxa outside the tribe Shoreae; thereby the
genus Dryobalanops appears to form the connecting bridge. As a consequence, it is seen that in the
extreme small south-west area of distribution a larger number of genera are present with relatively
primitive characters than in the large Malesian region, usually regarded as the centre of origin of the
family.”

Petiole anatomy. The complex petiolar vascular supply, characteristic of many genera in the
family, reaches its greatest elaboration in Dipterocarpus, Vateria, and the type sections of Shorea
and Hopea. The supply is reduced to 3 peripheral bundles in many Hopea sect. Dryobalanoides and
is relatively simple in the other sections of Shorea, but also in the putatively primitive Vatica
however. It seems that the primitive condition was the complex one, but generalisations are difficult
as the complexity also varies much according to the size of the petiole; large-leaved Dipterocarpus
and Shorea (including species of sect. Pachycarpae) have very much more complex systems than
small-leaved, and genera with slender petioles such as Dryobalanops and Anisoptera have simpler
systems then Dipterocarpus, though more complex than most Hopea and Shorea (see e.g. MAURY,
1978). If the complex is more primitive, then Hopea sect. Hopea is nearest to the archetype in that
genus. This is supported by the leaf nervation, which most nearly approaches the type typical of
other genera, and by the bark morphology, which WHITMORE (1960) has shown to differ in no
important way from that of Shorea sect. Shorea; these two sections both appear to be primitive in
their genera, hence more nearly than others resemble the common ancestor and thus one another.
They further both share a wider distribution than other sections in their respective genera.

Family affinities. Up to the time of BRANpDIs’s Enumeration (1895) Dipterocarpaceae had been
variously associated with Aceraceae (GAERTNER), Lauraceae (REICHENBACH), Theaceae/Ternstroe-
miaceae (ENDLICHER, A. DE CANDOLLE), Malvaceae (LINDLEY), Tiliaceae (A. P. DE CANDOLLE,
BLUME, SPRENGEL, MEISNER, LINDLEY, HEIM, BRANDIS, and GILG), Sterculiaceae (HEIM), and
Guttiferae (DE JussIEU, BLUME, ENDLICHER, LINDLEY, BENTHAM & Hooker, and HEIM). Since
ENDLICHER the family has been more or less constantly put in the Guttiferales, this clearly allying it
with Guttiferae and Theaceae and separating it from Tiliaceae in the Order Malvales. It should be
remarked, however, that BENTHAM & HOOKER placed Chlaenaceae next to Dipterocarpaceae in
cohors Guttiferales which was immediately followed by cohors Malvales, the only difference being

1982] DIPTEROCARPACEAE (Ashton) 281

in the aestivation. This classification has been subsequently followed by Bessey (1915), HUTCHIN-
SON (1926), WETTSTEIN (1935), and PULLE (1950). LINDLEY (1836) at first placed Dipterocarpaceae in
his Alliance Malvales, but in ‘The Vegetable Kingdom’ (1846) followed ENDLICHER’s (1840) transfer
to Guttiferales. HEIM (1892) claimed the affinities to be mainly with the Ti/iaceae, but the only
subsequent author to adhere to this view was HALLIER f. (1912) who placed them in his Order
Columniferae, with marked affinities with Tiliaceae and Elaeocarpaceae. The Columniferae he
considered to be derived from ‘Protoberberidaceous’ ancestors, and far removed from the
Guttiferales, with Dilleniaceous ancestors.

The Ancistrocladaceae have been associated previously by many authors with the Dipterocarpa-
ceae owing to the superficial similarity of their fruit with its 5-merous perianth, persistent aliform
imbricate sepals, and the embryo with folded cotyledons enclosing the radicle and with the testa
intruding between the folds. The unilocular ovary with single ovule, the leaf-nervation, scandent
habit, presence of thorn-hooks, and anatomy indicate to me that the affinity is not close.

Dipterocarpaceae share with Guttiferae, Theaceae, and also Tiliaceae centrifugal stamens, so that
if CORNER’s (1946) contention is correct all these families share a common ancestry, though the
character gives no indication to which family the dipterocarps are most nearly associated.

With the Theaceae are shared the 5-merous perianth, imbricate persistent sepals, frequently
contorted corolla, numerous hypogynous stamens, 2-celled anthers generally dehiscing longitu-
dinally, superior generally 3-celled ovary with frequently 2 ovules per cell, axile placentation, and
seeds with scanty endosperm. The absence, in Theaceae, of stipules, stellate, tufted or glandular
hairs, connectival appendages, mucilage cells and usually resin canals and the frequently dentate
leaves, indistinct leaf nervation, and short-sepaled fruit calyx, readily distinguish the two families.

It is the presence of intercellular resin canals which has principally led systematists to associate
the Dipterocarpaceae with the Guttiferae; other characters in common are the vertically
transcurrent nervation, the usually many hypogynous stamens, usually contorted corolla, usually
racemose inflorescence, and persistent calyx with usually imbricate sepals. The absence of
endosperm, which has frequently been cited as a factor allying the two, but not Tiliaceae, is
erroneous; HEIM (1892) and others have already shown that some species in several groups retain
endosperm at maturity of the seed.

Of the characters shared with the Guttiferae, those of the androecium are shared also with
Theaceae and Tiliaceae, as also the contorted corolla and, in many, the nervation; the inflorescence
in the Tiliaceae is also frequently racemose while conversely Upuna, Monotes, some Vatica and one
Parashorea share the cymose inflorescence typical of most Tiliaceae; tribe Dipterocarpeae possesses
a subvalvate flower calyx; in Tiliaceae the calyx is always valvate in flower. The resin canals are
therefore the only character in common with the Guttiferae that are not also shared with some
members of the Tiliaceae. Canals are confined to the medulla in the Dipterocarpaceae, unlike the
Guttiferae.

Guttiferae differ notably in the absence of stellate or glandular hairs, the general presence of a
hypodermis, the papillose lower epidermis, absence of stipules, frequently unisexual flower, absence
of an appendage to the connective, absence of aliform fruit sepals, and opposite leaves (the
alternate-leaved South American genera Caraipa, Kielmayera, Haploclathra, Marila and Mahurea
are now considered to belong to the Theaceae or Bonnetiaceae). The leaf nervation in Guttiferae,
with many indistinct lateral nerves, is very dissimilar from the prominent pinnate nervation of
Dipterocarpaceae; Dryobalanops, and Hopea sect. Dryobalanoides, which are exceptional in sharing
a nervation superficially similar to Guttiferae, cannot be considered primitive types in the family.

With the Tiliaceae the Dipterocarpaceae share the same stamen characters; the closely related
Elaeocarpaceae bear connectival appendages similar to those of many dipterocarps; the genera
Grewia, Pentace, and Schoutenia, inter alia, possess persistent, expanded, and wing-like fruit sepals.
Characters shared with Dipterocarpaceae and Tiliaceae but not the previously discussed families are
mucilage canals in cortex and cells in the epidermis of many, the mixed uni- and multi-seriate rays
(except Monotes), the arrangement of bast fibres into outwardly tapering wedges, and the presence
of a complex indumentum which may include single or tufted uni-cellular hairs, short or long
stalked multicellular gandular hairs, and unicellular peltate or stellate hairs.

The floral vascular supply strikingly confirms the Malvalian allegiance (see VAN HEEL, 1966).
CORNER (1976) has further found that the seed coat conforms to that of Malvales. These facts argue

282 FLORA MALESIANA [ser. 1, velR9?2

against the family being included in the Guttiferales as opposed to the Malvales. The single
anomalous character of the resin canals is absent from the subfamily Monotoideae and
Pakaraimoideae. The Elaeocarpaceae are virtually always devoid of mucilage cavities; moreover the
suggestion that elongate mucilage cells have phylogenetically developed into resin canals lacks any
logical structural basis. The prominent pinnate nervation of dipterocarps, and frequently geniculate
petiole, is so similar to that of the Elaeocarpaceae that leaves on the forest floor are sometimes
almost unassignable. In the Tiliaceae the basal pair always send off lateral branches on the outer
side; this is also found in some dipterocarps, e.g. in Shorea robusta, Dipterocarpus nudus and D.
stellatus, and abnormally sometimes in other species. The dipterocarp leaf nervation, usually
entirely transcurrent owing to the presence of columns of sclerenchyma, as in Guttiferae and
Ochnaceae, is found sometimes also in Tiliaceae.

The genus Monotes, which with the genus Marquesia constitutes the African subfamily
Monotoideae, was transferred to Tiliaceae by HEIM (1892); indeed Marquesia excelsa R.E.FR. was
originally named (though not published) as a Schoutenia. The principal characters in which the
Monotoideae differ from Dipterocarpoideae are the uniseriate rays, absence of resin canals but
presence of elongate medullary mucilage cells, subversatile anthers with elongate slender filaments,
and the tricolporate pollen with 4-layered exine. Of these all but the first are common in the
Tiliaceae. The imbricate sepals, trilocular (in Marquesia incompletely septate as in Dryobalanops)
ovary with two ovules in each cell, and pinnate nervation with unbranched basal nerves, are typical
of Dipterocarpaceae. The cymose inflorescence is unusual in Dipterocarpoideae, but is widespread in
Tiliaceae. BANCROFT (1935) examined the wood anatomy of Monotoideae and found it to be quite
distinct from both families; this is confirmed by GOTTWALD & PARAMESWARAN (1966). The general
facies is more similar to Dipterocarpoideae, but differences were considered by them sufficient to
constitute a separate family. Resin was present in the wood as in Dipterocarpoideae, though not in
intercellular canals. Monotoideae are thus a clearly defined group.

The recent discovery of a dipterocarp subfamily in the Guyana Highlands (MaGutrE et al., 1977)
strengthens this link, for Pakaraimaea shares the stamen, pollen, calyx and many wood characters
of Monotoideae (though the wood rays are predominantly biseriate), yet has a generalised
Malvalian 4—5-celled ovary, each cell of which is 2—4-ovulate; in several respects therefore this South
American relict must be regarded as archaic within the family.

Another family, Sarcolaenaceae (Chlaenaceae), endemic in Madagascar, has often been claimed
to be a close relative of Dipterocarpaceae. MEISNER (1837) and BENTHAM & HOOKER (1862) had
already placed them close to that family; so did HUTCHINSON (1926) and METCALFE & CHALK (vol. 1,
1950, 22). According to the studies of CAvACO (1952) and CAPURON (1970) they exhibit a far greater
diversity than mostly admitted in handbooks, covering almost all characters of Dipterocarpaceae, in
particular those of Monotoideae.

Macutre et al. (1977) have pointed out that this family too has the indumentum and complex
petiolar anatomy characteristic of Malvales, but shares with Dipterocarpaceae alone in this order an
imbricate calyx, 3-celled ovary (here Pakaraimaea is the exception), and absence of paired basal leaf
nerves; they also indicate that a distinct leaf nervation with looped intramarginal nerve, and a
|—2-layered hypodermis are shared by genera in both families; DE ZEEUW (/.c.. 368) considered that
the wood anatomy more closely resembles that of Monotoideae and Pakaraimoideae than theirs
does even to Dipterocarpoideae; a resemblance to Tiliaceous wood exists too, but is more remote.

A preliminary anatomical study of Eremolaena boinensis also confirmed the presence of tufted
hairs with a rosette of glandular hairs at their bases and possibly of peltate scales on the twigs. The
anomocytic stomata, the bark anatomy, and the occurrence of mucilage cells in the mesophyll, pith
and primary cortex together with the indumentum points to both Tiliaceous as well as
Dipterocarpaceous affinities. In the wood anatomy, however, Sarcolaenaceae much more resemble
Monotoideae of the Dipterocarpaceae than Tiliaceae: ray and fibre type agree well and most
important of all Eremolaena shows vestured pits: a very important taxonomic character found in all
Dipterocarpaceae but not in Tiliaceae or other Malvales. METCALFE & CHALK say that the
Dipterocarpaceae and Sarcolaenaceae differ by the absence of resin ducts in the pith of
Sarcolaenaceae, but Monotes and Marquesia have not been examined in this respect, and could
equally well miss these. Though Sarcolaenaceae and Dipterocarpaceae could be considered to
represent different families, they appear to be distinctly allied at a higher level and could well belong

1982] _ DIPTEROCARPACEAE (Ashton) 283

or have been derived from the same ancient Tiliaceous matrix in Gondwanaland (MAGUIRE et al.,
1977).

History of the Dipterocarpaceae. In concluding the preceding chapters it is tempting to speculate
about the origin, dispersion, and development of this group in time.

The overriding contemporary concentration of dipterocarp species diversity of West Malesia
prompted authors (from MERRILL, 1923, to MEHER-HoMiI, 1979) to suggest a Far Eastern origin for
the family. The fossil evidence is ambivalent; PRAKASH (1972) and LAKHANPAL (1974) discussed the
fossil history of Dipterocarpaceae. PRAKASH assumed an Asian origin for Dipterocarpoideae, but
LAKHANPAL leaned toward a Gondwanic origin of the whole family, an idea originating with
CROIZAT (1952). AUBREVILLE (1976) argued for a biphyletic origin, with Dipterocarpoideae of
Laurasian, and Monotoideae of Gondwanic provenance.

Accepting the principle of continental drift one could rather easily imagine, on the basis of
comparative anatomy and morphology, a southern Gondwana origin with the development of a
‘Dipterocarpaceous ancestral stock’ in a Gondwana-continent, possibly in the Upper Cretaceous.
This implies of course a tropical to at least subtropical climate in a period long after the
Angiosperms originated and had already strongly diversified.

The subsequent Atlantic split then provided the South American continent with a section of the
dipterocarps of which Pakaraimaea, a generalised and in this sense archaic form within the family,
as the sole survivor; the reason why this West Gondwana offshoot did not lead to a separate diverse
and derived branch must remain obscure.

Also primitive in several respects was the other offshoot Monotoideae, which remained in a
central position in tropical Africa where it is now represented with 2 genera and c. 40 spp. (with one
which may have recently invaded Madagascar). Possibly from this same ancestral stock another
plant family developed in Madagascar, viz Sarcolaenaceae as it seems Monotoideae are their closest
allies.

The most successful 3rd offshoot was the Dipterocarpoideae of which the ancestors inhabited the
eastern tropical part of Gondwanaland. According to trustworthy fossils they were at least present
formerly in East Tropical Africa while Vateriopsis is a genus still found in the Seychelles, a relict ona
surviving peak of the submerged part of the Deccan plate. Whether the Deccan plate sailed from
S.E. Gondwanaland to the Laurasian shores of the Tethys with the dipterocarp pilgrim fathers to
S.E. Asia on board remains to be confirmed by fossil evidence from the late Cretaceous or early
Oligocene of India and East Africa, but the distribution and phylogeny of Vateriopsis, Vateria and
Stemonoporus strongly suggest that it did.

Probably not long after the Dipterocarpoideae reached S.E. Asia, which must have been by the
Oligocene according to the pollen records in Borneo, they rapidly diversified, as still shown by the
present wide representation and endemicity of supraspecific taxa in Ceylon. In all probability they
were already rain-forest constituents, as most are at present in Ceylon, as they would also have been
subject to oceanic (if more windy) conditions on a hypothetical rafted Indian subcontinent.

Migrating eastwards through S.E. Asia they finally invaded the Malesian area in the early
Tertiary, as testified by the Oligocene pollen record in Borneo. By the Miocene they had become
common, as shown by the abundance of fossil wood and their appearance as a regular constituent of
the pollen record, and they have presumably retained this position till the present day.

In continental S.E. Asia they have come to thrive in both the everwet and seasonal regions. By
their capacity for gregariousness and high stature they have overwhelmed the mixed lowland
rain-forest and created the present majestic forest profile of the large islands of West Malesia.

The Philippines seem to have been close enough, and I would assume at least once connected with
western Sundaland, to receive a generous supply. I do not believe that we can conclude from
dipterocarp geography alone, though, that these islands have been intermittently invaded by
dipterocarps as MERRILL (1923) discussed at length, though this would be compatible with the
geomorphological instability of the region bordering the east of Wallace’s line.

The land-connections between the Philippines and the Moluccas and New Guinea must have
been similarly inadequate and intermittent and apparently did not allow Parashorea, Dipterocarpus
etc., to enter the East Malesian Province. The 4 genera in this province (Celebes, Moluccas, New
Guinea) are: Anisoptera, Hopea, Shorea and Vatica. | accept DIELS’s contention (1922) that the few
dipterocarps of New Guinea represent a younger element in its flora, probably derived from the

284 FLORA MALESIANA [ser. 1, yoIN@e

northwest by way of the Philippines, Celebes and Moluccas. It is noteworthy that in New Guinea
Hopea shows fairly high and probably young speciation.

In South Malesia there are only 5 genera present in Java with few (10) species, and in the western
Lesser Sunda Is. only 2 genera with 3 spp. This paucity is ascribed to several causes; the volcanicity
and long period of human habitation in the moist humid lowlands of Java precluded intensive
collection there by the early Dutch botanists; further east poor land connections played a part,
though much of the present distribution can be explained by the dry climatic conditions in the
lowlands, for the petering out of the family eastwards is correlated with increasing drought; there
appears to have been no capacity for development of drought-resisting species such as happened in
continental S.E. Asia. It cannot therefore be safely concluded that South Malesia has not been a
source area or invasion track of dipterocarps to New Guinea, particularly if the fossil Shorea record
from Timor is correct.

Finally, there appears to have been a considerable extinction of dipterocarps in the Deccan
Peninsula as shown by the range map (Fig. 2); Anisoptera and Dryobalanops have become extinct
there, as has the latter also in Java. Whether Parashorea has ever occurred in the Deccan is
uncertain, let alone the Bornean endemic genus Upuna.

The monotypic genus Upuna, endemic in Borneo, deserves special attention. It is primitive in
several respects, notably it superficially resembles the primitive genus Monotes of a different
subfamily while it shares a putative aril with the Ceylonese genus Stemonoporus and Vateriopsis of
the Seychelles. It is most closely allied however to the Malesian genus Anisoptera, and to a lesser
extent Cotylelobium and Vatica, especially in its wood anatomy. The presence, in Borneo, of Upuna
emphasises that the story of dipterocarp development on its way east is undoubtedly more
complicated than the simple panorama given above would imply, but the fossil record is
unfortunately meagre.

All these characters correlate much more closely however with the aseasonal humid region of the
Mixed Dipterocarp forests and, if we accept the view that they are derived (see f.f.), we must also
accept the possibility that the subfamily originated in a seasonal tropical climate.

I firmly conclude from present knowledge of their reproductive biology, and in particular of the
fruit dispersal, lack of seed dormancy, ecology of establishment and seedling ecology that the spread
of dipterocarps must be overland. This is compatible with the hypothetical reconstruction above:
the big Sunda islands, which were one continuous land area for most of the Pleistocene period are
richest in dipterocarps, while the Celebes-Moluccan area could only intermittently have provided
land bridges to convey the dipterocarps to New Guinea; its archipelagic geomorphology with
frequently interrupted, partial land bridges seems to have impeded migration and prevented many
genera from completing the course.

Literature: ASHTON, Gard. Bull. Sing. 22 (1967) 250; A manual of the dipterocarp trees of Brunei
State and of Sarawak, Suppl. Kuching (1969); Biol. J. Linn. Soc. 1 (1969) 155; Ann. Mo. Bot. Gard.
64 (1977) 694; Gard. Bull. Sing. 31 (1978) 5; Mem. Mus. Hist. Nat. Paris sér. B, Bot. 26 (1979a) 128:
ibid. (1979b) 145; AUBREVILLE, Adansonia IT, 16 (1976) 205; BANCROFT, Amer. J. Bot. 22 (1935) 505;
BENTHAM & Hooker, Gen. PI. | (1862) 189; Bessey, Ann. Mo. Bot. Gard. 2 (1915) 1; BLUME, Bijdr.
(1825) 222; Mus. Bot. Lugd.-Bat. 2 (1852) 28; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 1; BRANDIS &
GiLG in E. & P. Nat. Pfl. Fam. 3, 6 (1895) 243; Burck, Meded. Lands PI. Tuin 3 (1886) 1; Ann. Jard.
Bot. Btzg 6 (1887) 145; CANDOLLE, A. DE, Prod. 16, 2 (1868) 604; CANDOLLE, A. P. DE, Tiliaceae, in
Prod. | (1824) 517; CapuRON, Adansonia 10 (1970) 247-265; Cavaco, Chlénacées, Fl. Madag. fam.
126 (1952) 37 pp.; CoRNER, Wayside Trees (1940); J. Arn. Arb. 27 (1946) 423; Ann. Bot. 13 (1949)
367; Gard. Bull. Sing. 17 (1960) 368; ibid. 18 (1960) 1; The Seeds of Dicotyledons (1976) 33, 35, 119;
CORREA DE SERRA, Ann. Mus. Paris 8 (1806) 397; ibid. 10 (1807) 159; CroizaT, Manual of
Phytogeography (1952) 423; DieLs, Bot. Jahrb. 57 (1922) 460; Dyer, in Hook. f. FI. Br. Ind. 1 (Jan.
1874) 294; J. Bot. 12 (Mar. 1874) 97; E-mer, Leafl. Philip. Bot. 4 (1912) 1475; ENDLICHER, Gen. PI.
(1840) 1012; Feporov, J. Ecol. 54 (1966) 1; Gard. Bull. Sing. 29 (1977) 127; GAERTNER, Fruct.
Suppl. 3 (1805); GiLG, Bot. Jahrb. 28 (1899) 127; in E & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 237;
GOTTWALD & PARAMESWARAN, Bot. Jahrb. 85 (1966) 410-508; GUERIN, Bull. Soc. Bot. Fr. Mem. 11
(1907) 93; GUILLEMIN, PERROTTET & RICHARD, Florae Senegambiae Tentamen, Paris (1830-33);
Hatter, Arch. Néerl. Sci. Exact. Nat. 3B, 1 (1912) 146; VAN HEEL, Morphology of the androecium
in Malvales. Leiden (1966); Hem, Bull. Mens. Soc. Linn. Paris 119 (1891) 954; ibid. 120 (1891) 958;

1982] DIPTEROCARPACEAE (Ashton) 285

ibid. 122 (1891) 970; Rech. Dipt. Paris (1892); HemMsLey, Rep. Challenger Bot. 1 (1884-85):
HUTCHINSON, Fam. FI. Pl. ed. 1, 1 (1926); pe Jussieu, Gen. Pl. (1789); Kaur et al. Nature 271. n.
5644 (1978) 440; Kerr, J. Siam Soc. 2 (1914) 9: KorTHALS, Kruidkunde (1841) 45-75: KOSTERMANS.
Taxon 27 (1978) 357; LAKHANPAL, Birbal Sahni Inst. Palaeobot. Spec. Publ. 1 (1974) 30-39:
LINDLEY, An introduction to Botany, ed. | (1832); Dipterocarpaceae, in A Natural System of
Botany (1836) 98; Dipterocarpaceae, in The Vegetable Kingdom (1846) 393; Linnaeus, Gen. Pl. ed.
1 (1737); Sp. Pl. ed. 1 (1753); Gen. Pl. ed. 5 (1754); Mantissa 1 (1767); ibid. 2(1771); Syst. Nat. ed. 12,
2 (1767); MaGutRE et al. Taxon 26 (1977) 343; MAGUIRE & ASHTON, Taxon 29 (1980) 225: MAURY.
Thesis: Dipterocarpacées, du fruit a la plantule, Toulouse (1978); MEHER-Homg1, Phytocoenologia 6
(1979) 85; MEISNER, Plantarum Vascularum Genera (1837) 35, Syn. 27; MERRILL, Interpretation of
Rumphius Herb. Amb. (1917) 366; Philip. J. Sc. 23 (1923) 14-17; Mever, Ann. Mo. Bot. Gard. 61
(1974) 806; MILLER, Phil. Trans. Roy. Soc. 68 (1778) 161; Parker, Ind. For. Rec. 13 (1927) 11, 24:
PLANCHON, Descr. Ancistrocladaceae in Ann. Sc. Nat. III, 13 (1849) 316; PoLo, The travels of
Marco Polo, tr. Aldo Ricci, London (1931); PRAKASH, Geophytologia 2 (1972) 198: PULLE,
Compendium van de Terminologie, Nomenclatuur en Systematiek der Zaadplanten, ed. 2, Utrecht
(1950); REICHENBACH, Nom. Bot. Hort. | (1846); Rrptey, FI. Mal. Pen. 1 (1922) 209: Disp. PI.
(1930); RoxBuRGH, FI. Ind. ed. Carey 2 (1832); RUMpuius, Herb. Amb. 2 (1741) 74; ibid. 7 (1755) 65,
68; SPRENGEL, Syst. Veg. ed. 16 (1825-28); STERN, Brittonia 17 (1965) 36; SyminGToN, Gard. Bull.
Str. Settl. 7 (1933) 129; J. Mal. Br. R. As. Soc. 19 (1941) 139; Bull. Bot. Gard. Btzg III. 17 (1941) 88:
Mal. For. Rec. 16 (1943); DE VRigsE. in Miquel, Pl. Jungh. (1850) 80; WALPERs, Ann. Bot. Syst. 1
(1848); WettsTEIN, Handbuch der systematischen Botanik ed. 4 (1935); WHITMORE, the systematic
morphology of bark in the Dipterocarpaceae, Ph.D. thesis, Cambridge (1960).

Silviculture. The advanced silvicultural systems developed in Malaya are generally applicable
in western Malesian Mixed Dipterocarp forests and are already described exhaustively by
WyatTT-SMITH (Mal. For. Rec. 23, 1963).

The principal silvicultural characteristics of dipterocarps in regions of Malesia where they are of
economic importance are their absence of coppicing when mature, irregular gregarious flowering,
poor fruit dispersal, lack of seed dormancy, high moisture requirements combined with a need for
little litter and friable soil conditions at germination, the ability of saplings once established to
persist for a number of years in the understorey in deep shade but the need for moderate to
abundant light to effect rapid growth. Though species of the seasonal tropics have been successfully
propagated by rooted cuttings (Momose, 1978), species of the aseasonal zone have only been
propagated with difficulty in this way. These and other characteristics vary among species and
timber groups. Silvicultural methods have evolved partially in response to increased ecological
understanding, partially owing to changes in forest economics and especially in timber values and
labour availability and costs. Plantations and other artificial regeneration are presently generally
too costly owing to the care required at and following germination and planting, and the necessity
for subsequent weeding. Though enrichment planting, i.e. the supplementing of natural regene-
ration by planting in specially opened gaps or lines in naturally regenerating forest, is sometimes
desirable owing to frequently patchy distribution of the preferred species it is doubtfully
economically justifiable until cheaper and simpler methods of propagation are developed. The
object of natural regeneration is to simplify forest composition so that trees are mostly restricted to
a few economic species and the crop is of approximately even age. Originally improvement fellings
were undertaken in unfelled forest. Later, in Malaya, the canopy was opened up prior to felling to
ensure a satisfactory seedling crop at the time of felling; periodic low intensity fellings ensured
retention of seed trees; eventually this proved uneconomic. In the meantime the development of
improved preservation techniques combined with increasing labour costs to make the heavy
hardwood species, which generally require at least 100 years to reach timber size, uneconomic to
regenerate, favouring in particular the light red merantis that can be cropped on a cycle as short as
50-70 years.

Young trees larger than saplings are, as might be expected. poorly represented within the canopy
in the forests of the Malay Peninsula and northern Borneo, and it is for this reason that the Malayan
Uniform System of silviculture, which advocates a single clear felling every 50-70 years followed by
regeneration of seedlings already established on the ground, is practised there. The Philippine
Selection System, which advocates selective felling of large trees at approximately twice the

286 FLORA MALESIANA [ser. I, vol 9?

Fig. 13. Cutting timber of Hopea dryobalanoides Miq. Tawau, Sabah, Kalabakan concession; tree 48 m tall, 80
cm diam. at 15 m (Photogr. G. H. S. Woop).

frequency of the fellings in the Malayan system, therefore relies on the existence of an adequate
stock of young trees to form each successive crop. There is little quantitative published evidence that
this always exists, but if it does it demonstrates a fundamental difference in structure and dynamics
between Philippine and other Mixed Dipterocarp forests; this, if correct, may well be due to the
more open canopy which might be associated with frequent wind damage in the typhoon zone.
Adequate regeneration persists on the ground between fruiting years in the better forests, and this
is why the modern Malayan Uniform System relies on a single felling followed by more or less
vigorous poisoning of undesirable species to create favourable light conditions for those preferred.
The merantis and other light demanders are generally well able to survive beneath the ensuing dense
growth of weed trees and woody climbers and eventually overtop them though this is reported not
to be the case, or for the young trees to become misshapen as a consequence, on the most fertile soils
where climbers are particularly dense, as in parts of E. Borneo. Under drier conditions, as on high
ridges and sandy soils, less rigorous poisoning is essential; in Sabah and elsewhere where labour

1982] DIPTEROCARPACEAE (Ashton) 287

Fig. 14. Forest ranger AMPON standing near logs of red and yellow séraya (Shorea spp.), Kalabakan concession,
Tawau, Sabah (Photogr. G. H. S. Woop).

shortages exist poisoning is often not undertaken yet the new crop is claimed still to be satisfactory.
The system has to be used intelligently and modified according to local circumstances to succeed; it
is ideal for the most productive lowland forests but for many others, especially many hill forests, a
satisfactory system is still awaited. In the meantime present overexploitation of the Mixed
Dipterocarp forests and abandonment of silvicultural operations is bound to lead to a future
hardwood shortage at a time when a glut in agricultural export crops is predictable; then drastic
price reversals will favour those who have maintained an adequate natural, or more especially
silviculturally improved, forest estate.

Uses. Timber. Fig. 13-14. Presently the medium or light weight pinkish or reddish hardwoods
known as red méranti (Mal.), red séraya (Sabah) or red luan (Philippines) are the chief export timber
in Malesia; they are produced by Shorea sect. Rubella, Brachypterae, Pachycarpae, Ovalis and
Mutica. They are used chiefly for light construction, furniture efc., but also for veneers. They are
liable to fungal attack in contact with the ground and do not take preservatives easily, though

288 FLORA MALESIANA [ser. I, vol. 92

Fig. 15. Slash and resin drippings of Hopea sangal Kortu. Brunei (Photogr. ASHTON).

Fig. 16. Tapping of resin (damar sibosa) in trunk of Shorea javanica K. & V. New tap-holes filled with white resin
in rows between older ones; with forest officer VERHOEF. Sumatra, distr. Baros, Kp. Sioidang (Photogr. J. BURER,
1937).

preserving their shape well on drying owing to their interlocked grain; they vary greatly in density
between species, and sometimes within (S. a/bida). A similar timber, yellow méranti from Shorea
sect. Richetioides, is a less attractive tawny yellow and less valuable. The whitish timber of white /uan
or white séraya (lighter Parashorea spp., especially P. malaanonan in Sabah and the Philippines) is a
major veneer wood; so also are the similar but siliceous white méranti or mélapi (Sabah) derived
from Shorea sect. Anthoshorea, and the equally siliceous mérsawa (Mal.), pénggiran (Sabah), or
palosapis (Philippines) derived from Anisoptera spp. Kapur, from Dryobalanops, is a popular pale
brown light hardwood. Previously the durable heavy hardwoods were valued, but growth rates are
too slow (see silviculture), extraction and milling too costly. Chéngal (Neobalanocarpus) of Malaya
was celebrated for its durability and favoured by the Malays for boat as well as house building.
Balau or résak (Mal.), guijo (Philip.) or selangan batu (Borneo), derived from Shorea sect. Shorea
yield valuable heavy hardwoods for heavy construction and decking. Other such heavy hardwoods
are derived from the larger Vatica and Cotylelobium knownas resak (Mal.) and narig (Philip.), some
Hopea known as chengal, mengerawan, giam (Mal.), yvakal (Philip.), and from Upuna, called upun
batu or penyau (Mal.).

Fat. The fruit of many Shoreas, and sometimes also Dryobalanops, have been boiled as a
vegetable by villagers in many countries. The fruit of Shorea sect. Pachycarpae, and also S. seminis,
S. sumatrana, S. palembanica, S. scaberrima, S. hemsleyana, S. macrantha, S. singkawang and
sometimes other species are collected and exported. The seed contains up to 70% fat; the fat is
similar to cocoa butter but has a higher melting point and is favoured in Europe for manufacturing
chocolates and cosmetics, and in the past for soap, candles and tallow. In Borneo the fat was used
for cooking, being stored as a stick owing to its high melting point. Soaking the seed in water

1982] DIPTEROCARPACEAE (Ashton) 289

increases the fat content as well as killing predators and those treated in this way (Black Pontianaks
of trade) are considered the best. The pericarp has a low nutritional value as animal food. The trees
are rarely grown in plantation owing to the irregularity of fruiting, but could prove a major
commodity if this problem were solved.

Resin. Fig. 15-16. The oleoresin of Dipterocarpus (gurjun oil of India; kanyin oil of Burma,
minyak kéruing of western Malesia), destructively tapped by wounding and firing the bark and
sapwood, is still used locally for caulking and varnish, being formerly an export commodity; it dries
very slowly. The clear crystalline resins of Neobalanocarpus and several Hopeas (damar mata
kuching) were formerly valued for varnish manufacture. The resin of Shorea is inferior, though used
locally for tallow.

Camphor. Dryobalanops aromatica, and to a lesser extent D. lanceolata and D. beccarii yield a
crystalline camphor which was being traded by Arabs in the 6th century and was already mentioned
by Marco POLo in 1299. The trade centred on the gregarious D. aromatica forest of North and East
Sumatra and Johore; it was sought from hollows within the trunks by drilling into the wood and
smelling; the trees were felled and chopped up to facilitate extraction. As few trees yielded any
quantity the search was conducted with great mystery and a cult developed involving rites of
exorcism and a camphor language to confuse malignant spirits. The other two species were tapped
simply by drilling and scraping the wounds. Besides being exported the camphor was used for
manufacturing perfume, incense and for embalming. Dryobalanops camphor ts distinguished from
others by the presence of d. borneol.

Other minor uses. The bark of many species is used locally for building the walls of farm huts, and
the leaves of large-leaved dipterocarps provide thatch. Charcoal, prepared from the resin of several
species, especially in Shorea sect. Richetioides was used for blackening teeth when this was
fashionable. The bark of some Vatica and Hopea spp. was once used as ‘/aru’, to prevent frothing
during the boiling of Arenga syrup for sugar manufacture, and to arrest fermentation of
toddy.

Selected literature: General: BURKILL, Dict. Pl. (1935) 162, 238, 284, 1187, 1254, 1663, 2222:
Heng, Nutt Pl. ed. 2 (1927) 1093; MEIER-DREES, Tectona 32 (1939) 954, 968, 986; Momose, Mal.
For. 4 (1978) 219. — Timber: DEN BERGER & ENDERT, Meded. Proefst. Boschw. 11 (1925) 98;
Descu, Mal. For. Rec. 14 (1941); PEARSON & BROWN, Commercial Timbers of India (1932); REYES,
Techn. Bull. 7, Philip. Dept. Agr. (1938); TAMesis & AGUILAR, Dept. Agr. Nat. Res. Pop. Bull. 44
(1935). — Fat: ANDERSON in Williams et a/., S.E. Asian plant genetic resources, Bogor (1975) 217;
Ba, Landb. 9 (1933) 211; Rowaan, Landb. 13 (1937) 314; Indische Mercuur 60, 27 (1937) 416:
SMYTHIES, Sarawak Gaz. 84, 1206 (1958) 146. — Resin: BUCKLY, Mal. For. Rec. 11 (1932); ENDERT,
Tectona 2, 28 (1935) 248; FoNTANOZA, Philip. J. Sc. 54 (1934) 77; Parus, Bidr. Kennis
Oost-Indische Damarhars, Leiden (1933); RAPPARD, Tectona 30 (1937) 897; SAsaKI, Mal. For. 43
(1980) 290 (seed storage and germination); SCHUITEMAKER, Tectona 26 (1933) 229; TAMARI, F.R.I.
Research Pamphlet 69 (1976); TANG & TAMARI, Mal. For. 36 (1973) 38.

- Notes. 1. The herbarium identification of dipterocarps must remain difficult, mainly for the
reason that besides leaves also fruit and flowers are required.

The generic key is largely based on sterile characters and those of fruit; I have added an additional
key to field groups which may be useful especially in the field.

Species of the genera Cotylelobium, Dipterocarpus, Dryobalanops, Parashorea and Vatica are also
largely keyed out by means of sterile characters together with fruit. So are the species of the large
genus Hopea, but in this key there are several leads, mostly for a few species, in which flowers are
required.

In the largest genus, Shorea, leaf characters alone are only diagnostic at sectional level for
Richetioides. Though bark and leaf characters together can provide a basis for keying out the vast
majority of Malesian species, sections —and hence species — are impossible to key reliably
without the valuable sectional characters provided by the former. This is because the combination
of ontogenetic and phenotypic variability in leaf characters of these emergent trees, combined with
the great number of species, makes it impossible to construct a key on this basis though the
experienced taxonomist can still identify most by careful matching with named material, based on
his knowledge of the intricate combination of subtle features by which the leaf of each species may
be diagnosed at least when mature. In this genus flowers provide invaluable sectional, and in some

290 FLORA MALESIANA [ser. I, vol. 9? ©

sections species characters, but fruit are only of limited value particularly for distinguishing the
species with short fruit sepals from the rest.

I do not believe, therefore, that the dichotomous key provides a practical means of identifying
sterile material in the larger genera.

In the forest it is a different matter, as SYMINGTON so excellently demonstrated. His, and all
subsequent, keys to sterile material include the field characters of bark and wood. Such keys are
practicable mainly on a provincial basis; they already exist for West and East Malaysia
(SYMINGTON, 1943; MEIJER & Woop, 1964; ASHTON, 1968); all but a handful of Bornean species are
included in the latter two. They are not appropriate to a regional monograph, and this account
therefore aims to provide the sound taxonomic base upon which forest botanists can build.

The species of Anisoptera and Shorea are here therefore mainly keyed out on sterile characters
and flowers, though in Shorea there are a number of leads, mostly for a few species, for which fruit is
required.

It would be an impossible task to frame two keys for each genus, one based on flower, the other on
fruit characters: in Vatica and Dipterocarpus, for instance, keys based on sterile characters and
flowers are impossible, while in Shorea reliance on either flower or fruit alone would be impossible
(unless bark characters were included) though the key based on flowers would approach
completeness. A reliable key based entirely on vegetative characters visible in the herbarium seems
out of the question.

It is a rather unfortunate situation which we have to accept. Also local keys for the various islands
would not bring much solution, since each of the three large Sunda islands harbours so many
species.

2. As far as the vernacular names are concerned, I have selected only a limited number, as there is
much overlapping.

3. BRANDIS, DYER, SYMINGTON, VAN SLOOTEN, I myself, and some other authors have entered in
their works quite a number of unpublished names in the synonymy; they were not rarely taken up
from herbarium labels where they were put from provisional identifications. They have here only
been taken up if they were also taken up in the Index Kewensis.

4. As to the spelling of sectional and subsectional epithets I have adhered to priority of the
original epithet given, which was either in the plural or singular.

KEY TO THE GENERA

Fruit calyx lobes valvate, not incrassate at base. Chromosome number x= 11. Tribe DIPTEROCARPEAE.
Eee of calyx united into a tube enclosing at least the basal half of the nut.
. Connectival appendages stout, tapering, at most 14 x length of anthers to long-filiform. Nut free.
Innovations not lepidote. Leaf without prominent inane nenel nerve; vernation plicate. Fig. 17-20
1. Dipterocarpus
3. Connectival appendages slender, at least 2 x length of anthers. Nut adnate to calyx tube. Innovations and
leaf beneath densely peltate lepidote. Leaf with prominent looped intramarginal nerve; vernation not
plcate FIG 27. 30s yo ee cal) de tye tue ah ee ss pos canyon ic RO
. Base of calyx not as above.
fi Branching of inflorescence truly cymose. Stamens at least 25. Fruit triangular in section. Fig. 33-34
3. Upuna
4. Branching of inflorescence racemose or subcymose. Stamens not exceeding 15. Fruit terete on section.
5. Anthers linear, setose along the lateral margins. Style at least 2 x length of ovary. Leaf with distinct looped

intramarginal nerve. Fig.37,40. . . . .4. Cotylelobium
5. Anthers broadly oblong to subglobose, glabrous. Style less than 2x length of ovary. Leaf without looped
intramarginal nerve. Fig. 41-42. . . . 20 ey Svauca

1. Fruit calyx lobes +imbricate and with a distinctly incrassate central swelling at base. Chromosome number
x= 7. Tribe SHOREAE.
6. Fruit calyx lobes all aliform, subequal, or all sort; ai into a shallow cup at base. Nervation densely
parallel. Fig.48,50 . . . . . 6. Dryobalanops
6. Fruit calyx sepals not as above, unequal if aliform or short and subequal. Nervation not as above.
7. Fruit sepals aliform, unequal, narrowly imbricate. Nut globose, verrucose, lenticellate. Vernation plicate.
Rig eSARS Tl iy Le . . . 7, Parashorea

7. Fruit sepals prominently imbricate. Nut ovoid, smooth. Vernation not plicate.

1982] DIPTEROCARPACEAE (Ashton) 291

8. Stamens 15: anthers linear-lorate, with very short terminal connectional appendages. Fruit sepals short,
Sebel ig GOs, we hee Ai Se Sitios kn pps aires an aw ope une & .8. Neobalanocarpus

8. Not as above.
9. Fruit calyx with 2 aliform and 3 short sepals; if they are all short and ast then identifiable only by

sectional characters of the flower. Fig. 62-76. . a1 «teen 9. copes
9. Fruit calyx with 3 aliform and 2 short sepals, rarely 5 aliform, unequal; if they a are all short and subequal
then identificable only by sectional characters of the flower. Fig. 77-116... . . ~~. 10. Shorea

FIELD KEY TO PRINCIPAL FIELD GROUPS

1. Nerves curving round towards the margin and anastomosing to form a distinct intramarginal nerve.

2. Bark surface at first smooth, becoming scaly and distinctly scroll-marked, inner bark homogeneous pale
brown. Leaf tertiary nerves indistinct; main nerves slender, hardly raised, aes not swollen at base of
lamina, not geniculate. Leaf undersurface lepidote . . . . .4. Cotylelobium

. Bark surface irregular section fissured, flaking but not scroll- marked: inner bark distinctly tangentially
laminated. Leaf tertiary nerves scalariform, well spaced, distinct; main nerves + prominent; petiole swollen
at base of lamina, geniculate. Leaf undersurface densely lepidote . . . . . . . 2. Anisoptera

1. Not as above.

3. Fresh leaves on crushing and cut wood + aromatic; leaf nervation parallel, nerves equal

i)

6. Dryobalanops
3. Fresh leaves on crushing and cut wood smelling resinous, but not aromatic; nervation not as above.

4. Bark with large pale warty lenticels more or less densely dotted over the surface, or arranged in groups,
more rarely in lines; leaf nerves straight, curving round only near the margin, usually distinctly undulate
between each nerve owing to the slight persistence of the plicate folding in bud.

5. Bark surface yellowish to orange-brown or rust, rarely pink-brown, more or less smooth or flaky;
terminal leaf buds prominent, stipule scars sg petiole aa swollen at base of leaf; leaf
coriaceous, not lepidote . . . . . . 1. Dipterocarpus

5. Bark surface mauve-brown to dark purplish brown or tawny brown; more or less closely shallowly
fissured, later flaking in small oblong pieces; petiole not or hardly swollen at base of leaf; terminal
buds usually small (excl. P. macrophylla); stipule scars short (excl. P. macrophylla, P. malaanonan), leaf
thin, white-lepidote beneath at least in saplings and seedlings, frequently so in mature trees

7. Parashorea

4. Lenticels small, usually inconspicuous; leaf nerves + curved from their bases; leaf not folded, vernation not
plicate.

6. Base cordate; leathery leaf undersurface with a dense felt of white hairs, with darker, pale brown,
nervation: bark surface dark chocolate-brown, closely fissured and flaking; buttress low, rounded

3. Upuna

6. Leaf not cordate at base, or, if cordate, without white tomentose undersurface.

7. Buttresses low, rounded; bark surface smooth, pale grey-brown; innner bark homogeneous cream to
pale brown; wood close textured, ray ends not glistening on tangential surface; leaf tertiary nerves
generally reticulate, not drying black. . . » 2s Os Vatical

7. Bark surface, if smooth, chocolate and grey dappled and with thin buttresses: or, if pale brown, wood
not dense, ray ends glistening on tangential surface; leaf tertiary nerves scalariform, or, if reticulate,

drying black.
8. Leaf nervation dryobalanoid or subdryobalanoid. Fig. 64b-c. . . . . many 9. Hopea
8. Leaf nervation notasabove . . . mrss Neobalanocarpus, : some 9. Hopea, 10. Shorea

(For distinction between these and their sections, see other keys.)

1. DIPTEROCARPUS

GAERTN. f. Fruct. 3 (1805) 50; BL. Bijdr. (1825) 223; DC. Prod. 16, 2 (1868) 610;
Dyer, FI. Br. Ind. 1 (1874) 294; J. Bot. 12 (1874a) 101, 152, t. 143-145, incl. sect.
Sphaerales, Angulati, Tuberculati, Alati et Plicati Dykr, /.c. 102, 103, 105, 107;
VESQUE, C. R. Ac. Sc. Paris 78 (1874) 625; J. Bot. 12 (1874) 149; BuRcK, Ann.
Jard. Bot. Btzg 6 (1887) 194; HEM, Rech. Dipt. (1892) 24; BRANpIs, J. Linn. Soc.
Bot. 31 (1895) 24; Gite in E. & P. Pfi. Fam. ed. 2, 21 (1925) 250; L..G. DEN
BERGER, Hand. Ned. Ind. Natuurwet. Congr. Batavia (1926) 400; SLoor. Bull.

292 FLORA MALESIANA [ser. I, vol. 9?

Jard. Bot. Btzg III, 8 (1927) 263; HEYNE, Nutt. Pl. ed. 2 (1927) 1093; PARKER, Ind.
For. Rec. 16, 1 (1931) 1; Burk. Dict. (1935) 838; Foxw. Mal. For. Rec: 10 (1932)
56; Philip. J. Sc. 67 (1938) 245; Sym. Mal. For. Rec. 16 (1943) 153; BROWNE, For.
Trees Sarawak & Brunei (1955) 102; SMITINAND, Thai For. Bull. 4 (1958) 1;
ASHTON, Gard. Bull. Sing. 20 (1963) 233; BACKER & BAKH. f. Fl. Java 1 (1963)
328; MEVER & Woop, Sabah For. Rec. 5 (1964) 230; ASHTON, Man. Dipt. Brunei
(1964) 16; ibid. Suppl. (1968) 6; Gard. Bull. Sing. 31 (1978) 5; SMITINAND, Thai
For. Bull. (Bot.) 12 (1980) 24. — Oleoxylon Roxs. Trans. Soc. Arts London 23
(1805) 413; WALL. Cat. (1828) n. 953, nomen. — Pterigium CorREA, Ann. Mus.
Paris 8 (1806) 397, p.p., quoad P. costatum (GAERTN. f.) CORREA; ENDL. Gen. PI.
(1840) 1013 (‘Pterygium’). — Mocanera BLANCO, FI. Filip. ed. 1 (1837) 446, p.p.,
quoad M. verniciflua BLANCO. — Duvaliella HEI, Bull. Mens. Soc. Linn. Paris 2
(1892) 1011. — Heimiatoma PrrrRE, Fl. For. Coch. 4 (1892) t. 259. — Fig. 17-26.

Medium-sized to large trees with thick, rounded, usually small and concave,
sometimes tall and straight buttresses. Crown usually relatively narrow, even or
irregular (not cauliflower-shaped), dome-shaped, frequently rather flat, open,
with a few large strongly ascending twisted branches. Bark surface pale or dark
grey to orange-brown, sometimes pink-brown; appearing smooth, shallowly
patchily flaked; or appearing square-section fissured, shaggy, with persistent
oblong flakes; + prominently densely warty lenticellate. Twigs variable, stout or
slender, terete or compressed, glabrous or tomentose; with distinct, usually
swollen and pale, amplexicaul stipule scars. Stipules large, hastate to lorate,
obtuse, +succulent, caducous, characteristically carpeting the forest floor in the
growing season. Leaves coriaceous, rarely thin, margin usually sinuate towards
the apex; nerves prominent beneath, straight, curved only towards the margin,
with traces of the plicate vernation remaining persistently between them, giving
the lamina a corrugated appearance (cf. Parashorea); tertiary nerves scalariform;
petiole distinctly geniculate, stout. Inflorescence racemose, short, stout, zig-zag,
few-flowered, somewhat irregularly sparingly branched; bracts as stipules but
smaller, fugaceous. Flowers large. Fig. 18. Buds ellipsoid. Calyx united round the
fruit into a tube, but not fused to it; lobes valvate: 2 long, oblong to spatulate,
+ distinctly 3-nerved, and 3 short, or all 5 short. Petals large, narrowly oblong,
strongly contorted, loosely cohering at base on falling, cream with a prominent
pink stripe down the centre. Stamens 15—40, persisting at first in a ring round the
ovary after the petals fall; filaments of variable length, broad, compressed,
connate at base, tapering apically, latrorse, with 4 pollen sacs, the inner 2
somewhat shorter than the outer 2; appendage to connective short, stout to long
filiform, slender, glabrous. Ovary enclosed in the calyx tube, the apex ovoid to
conical, shortly tomentose; stylopodium cylindrical to filiform, shortly tomen-
tose, narrowing gradually or abruptly into the glabrous filiform style. Fruit large.
Fig. 17. Calyx tube becoming +distinctly constricted into a distal neck as the nut
expands; lobes as in flower, but greatly expanded; nut ovoid, tomentose with a
short acute apical style remnant. Germination hypogeal, the intricately folded
subequal cotyledons remaining within the fruit and the plumule freeing itself by
elongation of the cotyledonary petioles; seed sometimes albuminous at germina-
tion.

1982] DIPTEROCARPACEAE (Ashton) 293

Distr. About 69 spp., in Ceylon, India, Burma, Thailand, Indochina, and Yunnan and 53 or 54 species in
Malaya, Sumatra, Java, Bali, Sumbawa, Borneo, Philippines and intervening islands. Fossil records from the
Tertiary in E. Africa. Fig. 19.

Ecol. Evergreen forests and savanna woodland below 1400 m. Some species become semi-gregarious on river
banks or alluvium (D. apterus, D. elongatus), podsols (D. borneensis), ridges (several species) and semi-evergreen
forest of seasonal climates (D. gracilis, D. costatus). Seedlings mostly require high light intensities for survival;
the genus is least common in dense valley forests.

Anat. One of the most clearly defined genera in the family; twigs with many resin canals, in |—2 concentric
rings, in the outer margin of the pith; leaf traces 3, arising in the distal 5 of each internode with 12 stipule traces;
distal end of petiole with 1—3 semicircles of vascular bundles, each with a resin canal, closed by an adaxial bar of
collateral vascular tissue.

Taxon. The genus has since Dyer (1874a) habitually been divided into five sections on the basis of the fruit
calyx tube; Sphaerales (tube round in cross-section); Tuberculati (with 5 distal tubercles); Angulati (S-angled);
Alati (5-winged) and Plicati (5-winged with the wings proliferated into folds thus obscuring the tube). These
characters are not only uncorrelated with others, and thus do not appear to define natural groupings, but are
inconsistent even within single species. Thus for example the winged fruit calyx tube of D. zeylanicus THw. is
frequently angled, even smooth; the tuberculate fruit of D. costulatus and D. kunstleri are sometimes distally
winged; the angled fruit of D. g/obosus is sometimes merely obscurely tuberculate and the narrowly winged fruit
of D. fagineus is sometimes hardly more than angled, while the wings of that of D. sublamellatus are partially
folded and place it in an intermediate position between Alati and Plicati. These sections are not therefore
adopted in this account (cf. Gard. Bull. Sing. 20, 1963, 234).

Uses. Light to medium timbers absorbing preservatives readily; used for railway sleepers and heavy
construction. The oleoresin is tapped in the semi-evergreen forests of Indochina and Burma, and sometimes
elsewhere, for varnishes and tallow, but cutting into the bole and wounding the tissues by burning.

Note. Following Rec. 75A of the Code I have treated the generic name as masculine, and not as feminine as
VAN SLOOTEN did.

KEY TO THE SPECIES
(sp. 54. D. orbicularis excepted)

1. Mature fruit calyx tube spherical, not angled, ribbed, tuberculate or winged.

2. Fruit calyx lobes vestigial, subequal... RAG SS Pee ee Oe ee veel teminenes
Fruit calyx lobes unequal, 2 expanded, aliform.
Fruit calyx tube verrucose-lenticellate . . . . . . =... =. =. ~. +. 2 D. verrucosus
: Fruit calyx tube smooth.
Pen ise | ee eet | A Pel o Ca Ra) Pe ae) CEMICuS

4. Stamens at least 20.
5. Shorter fruit calyx lobes less than 7 mm long, not recurved or revolute. Stamens 25. 4. D. caudiferus
5. Shorter fruit calyx lobes at least 8 mm long, prominently recurved and revolute. Stamens 30.
6. Twigs and buds densely ferruginous tomentose.

7. Leaf blade glabrescent. . . ne a ee ot ei she DEES
7. Leaf blade persistently tomentose beneath.
8. Leaves 8-15 by 4-10cm;tomentum short . . . . . . . . . . . 6. D. gracilis
8. Leaves 17-32 by 8-17 cm; tomentum long, tufted . . . . . . . . . +. 7. D. baudii
6. Twigs and buds buff or golden tomentose or glabrous.
9. Leaf undersurface shortly densely persistently buff pubescent. . . . . . 8. D. obtusifolius

9. Leaf undersurface sparsely caducously pubescent or glabrous.
10. Nerves at most 14 pairs.
11. Petiole at least 4 cm long. Fruit calyx tube ellipsoid... . . . . 9. D. rotundifolius
11. Petiole shorter than 4 cm long. Fruit calyx tube globose or turbinate.
12. Leaf bud persistently pubescent outside.
13. Leaf to 19 by 9 cm, ovate. Petiole 2.3-2.8 cm long. Twig persistently densely golden pubescent
10. D. chartaceus
13. Leaf to 13 by 5 cm, elliptic caudate. Petiole at most 2.5 cm long. Twig + glabrescent
11. D. caudatus
12. Leaf bud glabrous outside.

14. Inside of stipules densely pubescent. Nerves at most Il pairs. . . . . .12. D. kerrii
14. Inside of stipules glabrous. Nerves at least ll pairs . . . . . . . 13. D. hasseltii

10. Nerves at least 16 pairs.
(55 Fruit calyx tube Subglobase ets lodge OPP. TIA 2) es I reese

15. Fruit calyx ‘tube ‘obturbinates) <1 P08) © PU Ler 2 9. ee ee ISD littoralis

294 FLORA MALESIANA [ser. I, vol. 92

Fig. 17. Fruits in Dipterocarpus. All x 5. — A. D. cornutus Dyer. — B. D. grandiflorus (BLCO) BLco. — C. D.

acutangulus VesquE. — D. D. kunstleri KiNG. — E. D. pachyphyllus MEER. — F. D. elongatus KorTH. — G. D.

mundus SLoot. — H. D. retusus BL. (A KEP 77290, B anon. L sheet 955-159-403, C BNB 4890, D A 3705, E
MusIn 39208, FS 15131, GS 19046, H KosTERMANS 18730).

1982] DIPTEROCARPACEAE (Ashton) 295

1. Mature fruit calyx tube angled, ribbed, tuberculate or winged.

16. Fruit calyx tube narrowly elliptic-obovoid, with prominent, sharp or narrowly rounded ribs or wings
typically confined to the distal half and + terminating in tubercles. Leaf bud slender, falcate, densely
minutely greyish stellate puberulent. . . . . .. . . =. +. +. +. +. +. 16. D. kunstleri

16. Not as above.

17. Fruit calyx tube spherical, but with 5+prominent apical protuberances below the neck.
Renter ealyx tube ovold .f 2). RR ie SO oe Ue 17D ape
18. Fruit calyx not ovoid.
19. Leaves with 11—16 pairs of nerves.

20. Leaf beneath pale ochraceous pubescent . . . . . . . . . =. +. +. 418. D. rigidus
20. Leaf beneath glabrous.
21. Stipules pruinose outside, otherwise glabrous. . . . . . . . . . 19. D. costulatus
Zieotppies tomentose outside 2 EM SU . . . Diiglobosns

19. Leaves with more than 18 pairs of nerves.
22. Fruit calyx with 2 lobes aliform, more than twice as long as tube.

23. Leaf beneath glabrescent . . J See 2 8 ee ee D humeratns
23. Leaf beneath densely cream lepidote Fes) 28) eee es 940 De ys
22. Fruit calyx lobes subequal, vestigial 2 0 fo gee “ee Ejats 23.5Dclongatus

17. Fruit calyx tube + persistently angled, ribbed or r winged.
24. Fruit calyx tube with 5 wings continuing from base to apex, greatly proliferated and intricately folded,
obscuring the tube itself.
25. Leaf nervation beneath persistently yellow-brown hirsute. Twigs terete . . . 24. D. lamellatus
25. Leaf nervation beneath glabrous. Twigs compressed.
26. Leaves ovate-lanceolate, margin revolute. Nerves 15—20 pairs. Petiole stout, tomentose
25. D. lowii
26. Leaves broadly ovate, margin not revolute. Nerves 10-12 pairs. Petiole slender, glabrous
26. D. pachyphyllus
24. Fruit calyx tube with 5 angles, ribs or wings, straight or undulate but not intricately folded.
27. Fruit calyx tube at least 15 times as long as broad, distinctly ellipsoid.
28. Calyx tube angled rather than winged.
29. Leaves large, at least 15 cm long.

Seaves suborbicular. Nerves}9=12 pairs=.} 2: 2 55 9 794% 2 © 2 7.) 227. D_confertus

ja) Ceaves ovate or elliptic: Nerves 24-30 pairs. 2. Ss = a 2 Ae 8 Didyen
29. Leaves small, not exceeding 4 cm long.

Si stamens 15. Leaf undersurface glabrescent .© 2.0 2: Wrote Gee ~. &¢29°D:faginens

Sie Stamens 23-25. Leafiundersurface puberulent - = “2,9: 2° 3.6. 230! (Deinereus

28. Calyx tube distinctly winged.
32. Fruit calyx tube densely pale buff puberulent, with very narrow undulate wings.
SOmINeEVes:6—8 pairs!" es Nii fee Se A cae SR bP SEED cemivestitns
33. Nerves 16-20 pairs. . . bis. Wh alan tf v324 Dzoblonpifolins
32. Fruit calyx glabrous, wings if narrow straight.
34. Wings of calyx tube at least 8 mm wide, prominent.

Ssgstamenss30" = =: fF. 9. = 2: 2 Veg Si Se eS 2 Dy orandiflordus
35. Stamens 15.

36. Young twig, petiole and bud glabrous . . . . . . . +. + . 34. D. glabrigemmatus
36. Young twig, petiole and bud pubescent . . . . . . . +. +. 35. D. palembanicus

34. Wings of calyx tube at most 6 mm wide, narrow
37. Calyx tube wings most prominent in the distal half.
38. Calyx tube wings continuous from base to apex.
39. Leaf bud buff velutinate. Leaf undersurface sparsely so. Flower unknown 36. D. fusiformis
39. Leaf bud and leaf glabrous. Stamens 15... . . 37. D. mundus
38. Calyx tube wings + absent in basal 3 of tube. Stamens 25, Leaf undersurface and bud pubescent
38. D. borneensis
37. Calyx tube wings not as above. Stamens 15. . Seen. 39) Duds
27. Fruit calyx tube less than 15 times as long as broad, broadly, plkcasoil ie poh (sometimes with the
wings decurrent with the elec and thus appearing narrowly ellipsoid).
40. Fruit calyx tube persistently pubescent.

41. Leafand petiole glabrous .. Hse ob ba 05D Neentealatus
41. Leaves undersurface and petiole persistently tomentose.
42. (Nervéesiat most'14 pairs’) 0-21 044, oo elle 1 ) eee. Uh. (eee ei i enatatns

42 "Nerves atileast: 15 pairs pn 2 Se ae ee 2D Scontormis

296 FLORA MALESIANA [ser. I, weleg2

Te
ey

=i

ines

Will

Fig. 18. Flower details in Dipterocarpus. All x 5. — D. verrucosus Foxw. ex SLoot. A. Bud, B/. outer sepal, B2.
inner sepal, both from inside, C/. stamens from outside, C2. stamens from inside, C3. older stamen, D. pistil
(JAcoss 5461).

1982]

40. Fruit calyx tube’ glabrescent.

43. Fruit calyx tube angled rather than winged

43. Fruit calyx tube distinctly winged.

44. Wings of calyx tube exceeding 10 mm wide.
45. Leaves glabrous. Wings of calyx tube not decurrent with pedicel

DIPTEROCARPACEAE (Ashton)

297

. 43. D. acutangulus

44. D. sublamellatus

45. Leaves densely tomentose beneath. Wings of calyx tube decurrent with pedicel.

46. Leaves prominently cupped
46. Leaves applanate ;
44. Wings of calyx tube at most 9 mm 1 wide.

47. Leaves obtuse, broadly obovate, obtuse or retuse. . . . . . . 47.

47. Leaves not as above.

48. Wings of fruit calyx tube not continuous to base. Leaves large.

. 45. D. concavus
. 46. D. stellatus

D. sarawakensis

48. D. coriaceus

48. Wings of fruit calyx tube continuous to base. Leaves medium-sized.
49. Wings of fruit calyx tube broadest distally, to 9 mm broad. Leaves prominently cuspidate

49. D. cuspidatus

49. Wings of fruit calyx tube not broadest distally. Leaves not cuspidate in mature trees.
a Wings of calyx tube less than 3 mm wide, very narrow.

. Leaves at most 10 by 4.5 cm
a Leaves 10-19 by 4.5—9.5 cm

50. D. eurynchus
51. D. ochraceus

50. Wings of calyx tube to 8 mm wide, prominent.

52. Parts glabrous but for ovary

52. D. perakensis

52. Buds, twigs, petioles and nervation beneath densely persistent ochraceous pubescent

1. Dipterocarpus tempehes SLooT. Reinwardtia 5
(1961) 468, f. 4; MEER & Woop, Sabah For. Rec. 5
(1964) 270, f. 43B; ASHTON, Man. Dipt. Brun. Suppl.
(1968) 18, f.3.

Twigs, leaf bud, stipule outside, and petiole very
shortly densely evenly buff pubescent, leaf nervation
beneath and stipule within sparsely so; nervation
above caducously so. Twigs to 2 by 4mm @ towards
the compressed apices, much branched, slender,
becoming terete; stipule scars slender, dark. Bud to 12
by 3 mm, slender, lanceolate, acute. Stipule to 8 by 1.2
cm, lorate, acute. Leaves 6-12 by 3.5-8 cm, broadly
elliptic to obovate, thickly coriaceous, with broadly
cuneate base and acute to shortly abruptly acuminate
apex; prominently persistently folded between the

—— ~-
or) 10

Fig. 19. Density map of Dipterocarpus in Malesia.
Above the hyphen the number of endemics. below the
hyphen the number of non-endemics.

53. D. philippinensis

9-12 pairs of sharply ascending nerves; nerves at
35°—40°; tertiary nerves very slender, densely scalari-
form; petiole 1-2 cm long, short, drying densely buff
pubescent. Raceme to 2.5cm long, very short, axillary,
terete, densely shortly evenly buff pubescent, un-
branched, bearing to 3 distichous flowers. Flower bud
to 3 by 0.8 mm. Calyx and corolla typical, calyx
glabrous. Stamens c. 30, somewhat shorter than
anther; anther narrowly oblong, tapering into the
acicular appendage; appendage as long as anther,
prominent. Ovary ovoid, pubescent; style filiform,
pubescent except in the apical 1/3. Fruit pedicel
vestigial. Calyx glabrous; tube to 4 by 4 cm, turbinate,
with prominent but unraised pale lenticels; calyx lobes
vestigial.

Distr. Malesia: Borneo (Kapuas valley, Sarawak
west of the Niah R., E. Sabah south to Kutei).

Ecol. Locally abundant, fresh water swamps and
stream banks, clay rich alluvium.

Vern. Kéruing tépayan, k. asam, karup, bajan, bajan
uhit, tempéhes.

2. Dipterocarpus verrucosus Foxw. ex SLooT. Bull.
Jard. Bot. Btzg III, 8 (1927) 293; SLoor. in Merr. PI.
Elm. Born. (1929) 201; Foxw. Mal. For. Rec. 10
(1932) 71; SyM. Mal. For. Rec. 16 (1943) 189, f. 84B,
85; BROWNE, For. Trees Sarawak & Brunei (1955) 111;
ASHTON, Man. Dipt. Brun. (1964) 46, f. 6-7, pl. 6
(stem); ibid. Suppl. (1968) 19; MEER & Woop, Sabah
For. Rec. 5 (1964) 272, f. 3C, 43A. — Fig. 18, 20.
Twig, petiole, midrib and nerves beneath densely
caducous minute adpressed tufted gold-brown pubes-
cent; leaf bud and raceme densely, stipules sparsely,
persistently so. Twig 1.5-3 mm @ apically, terete or
slightly compressed; amplexicaul scars slightly
swollen. Bud to 12 by 2-3 mm, acute, narrowly falcate
to narrowly conical. Stiplule to 8 by 0.8 cm, narrowly

298 FLORA MALESIANA [ser. I, voles

Fig. 20. Dipterocarpus verrucosus SLOOT. a. Terminal bud and leaf of 75 m high sapling, b. young inflorescence
with stipules, c. stipules outside hairy, inside glabrous, d. older inflorescence without stipules, e. fruits, f. nut.
All x3 (a S 20286, b-c KEP 105026, d KEP 105157, e-f NT 429).

1982]

oblong, obtuse. Leaves 6-12 by 3.5-6 cm, ovate,
coriaceous; base cuneate; apex subacute or with to 5
mm long acumen; nerves 9-14 pairs, prominent, well
spaced, at c. 40°-SO°; tertiary nerves well spaced,
+sinuate; petiole |1.2—2 cm long, geniculate. Raceme
to 9cm long, axillary, unbranched or singly branched,
terete. Flower buds to 25 by 12 mm, fusiform. Calyx
shortly densely pubescent; corolla typical. Stamens 15,
exceeding style apex in bud; filaments as long as
anthers, narrowly deltoid, applanate; anthers nar-
rowly oblong, pubescent, tapering into the short
broad obtuse connectival appendage. Ovary conical,
tapering, pubescent; style glabrescent, as long as
ovary, swollen below apex. Fruit calyx tube to 1.5 by
1.3 cm, globose to slightly ovoid, with 5-7 mm @ neck,
glabrescent, dotted with verrucose pale brown lenti-
cels; 2 longer lobes to 9 by 2.2 cm, oblong to spatulate,
obtuse, c. 3 mm broad at the non-revolute base,
3-nerved, the 2 laterals continuing at least } of the
length; 3 shorter lobes to 3 by 2 mm, narrow, revolute
and recurved.

Distr. Malesia: Malaya (excepting more seasonal
areas), Sumatra (Asahan and Indragiri in east, Kam-
par-Siak on west coast), Singkep, Borneo (West
Borneo and Sarawak to S. W. Sabah, Tawau,
Nunukan).

Ecol. Mixed Dipterocarp forest on clay rich soil,
occasional on undulating land but locally common on
ridges below 650 m.

Vern. Kéruing merah, k. chaiér (Mal.), k. ladan, k.
daun halus, ariung (Sum.).

3. Dipterocarpus crinitus Dyer, Fl. Br. Ind. | (Jan.
1874) 296; VEsqugE, C. R. Ac. Sc. Paris 78 (March
1874) 627; Dyer, J. Bot. 12 (April 1874) 103, 154;
KoNGaaRewAS. Soc) Beng. Sc: 62) 25(1893)) 90:
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 28; RIDL. Agr.
Bull. Str. & F.M.S. 1 (1901) 55; Fl. Mal. Pen. 1 (1922)
214; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 273; ed. 2 (1927)
1095; Burk. J. Str. Br. R. As. Soc. 81 (1920) 58, fig.;
Dict. (1935) 843; Merr. En. Born. (1921) 398; SLOoT.
Bull. Jard. Bot. Btzg III, 8 (1927) 288; Foxw. Mal.
For. Rec. 10 (1932) 66; Sym. Mal. For. Rec. 16 (1943)
175, f. 84A, 85, 89; BROWNE, For. Trees Sarawak &
Brunei (1955) 108; ASHTON, Man. Dipt. Brun. (1964)
29, f. 6; ibid. Suppl. (1968) 13; MEWER & Woop, Sabah
For. Rec. 5 (1964) 246. — D. hirtus VESQUE, C. R. Ac.
Sc. Paris 78 (March 1874) 627; J. Bot. 12 (1874) 151;
Dyer, J. Bot. 12 (1874) 154.— D. tampurau (non
KortTH.) Burck, Ann. Jard. Bot. Btzg 6 (1887) 198,
p.p., BROWNE, For. Trees Sarawak & Brunei (1955)
108. — Fig. 21.

Young twig, leaf bud, stipule outside and petiole
persistently c. 3 mm long golden-brown tufted tomen-
tose, shorter on nervation beneath, midrib above and
margin; leaf fugaceous pubescent above. Twig c. 3 mm
@ apically, smooth. Bud 4-6 by 1.5—2 mm, small,
oblong, obtuse. Stipule c. 3 by 0.5 cm, lanceolate,
obtuse. Leaves 6-9 by 3—5 cm, elliptic, chartaceous,
obtuse or shortly acuminate, base obtuse, margin
revolute, persistently somewhat folded between the

DIPTEROCARPACEAE (Ashton)

299

13-15 pairs of nerves; petiole 1.5—2.5 cm long, slender.
Raceme to 12 cm long, terminal or axillary, terete,
golden long tomentose, unbranched or singly
branched, branchlets bearing c. 4 flowers; bracts to 30
by 8 mm, linear, acute, sparsely tomentose outside,
glabrous within. Flower bud to 3.5 by 0.8 cm. Calyx
and corolla typical, calyx glabrous. Stamens 15,
shorter than the style; filaments short, anthers fili-
form, tapering; appendage as long as anther, slender,
undulated towards the apex. Ovary ovoid-conical,
shortly pubescent; style and stylopodium filiform,
slender, 4-5 times as long as ovary, shortly pubescent
but for the distal 4. Fruit subsessile. Fruit calyx
glabrous, tube 1.5—1.8 by 0.6—0.8 cm, neck 0.4—0.6 cm
@; two longer lobes to 8 by 1.5 cm, oblong, lanceolate,
acute, prominently l-nerved; 2 shorter lobes to 3 mm
long, deltoid, acute.

Distr. E. Peninsular Thailand, and in Malesia:
Malaya (excluding seasonal areas), Sumatra (Asahan,
Indragiri, Bengkalis districts; Central Sumatra:
Sibolga, W. Indragiri), Borneo.

Ecol. Widespread on undulating land and low hills,
rarely to 850 m, on leached clay-rich soils in Mixed
Dipterocarp forest.

Vern. Kéruing (m)empélas, tampurau, k. bulu, k.
gombang, k. pékat, k. mérakluang, k. chaier (Mal.),
ariung, simarhalung (Sum.), resak empélas (Iban).

Note. Individuals, often in considerable numbers,
of this species are well known in Malaya to flower and
fruit outside general flowering years; the same occurs
in Borneo. Though seedlings are usually common in
the forest, as a rule very few seeds are viable.

4. Dipterocarpus caudiferus MerR. Philip. J. Sc. 29
(1926) 398; SLoor. Bull. Jard. Bot. Btzg III, 8 (1927)
302; BROWNE, For. Trees Sarawak & Brunei (1955)
107: SLoot. Reinwardtia 5 (1961) 459; ASHTON, Gard.
Bull. Sing. 20 (1963) 236; Man. Dipt. Brun. (1964) 25,
f. 6; ibid. Suppl. (1968) 11; MEER & Woop, Sabah
For. Rec. 5 (1964) 239, pl. 20 (habit), pl. 24, f.
35. — D. macrorrhinus SLoot. Bull. Jard. Bot. Btzg
III, 8 (1927) 300, f. 3.— D. kutaianus Stoort. Bull.
Jard. Bot. Btzg III, 16 (1940) 437, f. 2; BROWNE, For.
Trees Sarawak & Brunei (1955) 110.

Twig, leaf beneath and margin, petiole, raceme and
young leaf bud sparsely, 2-2.5 mm long, + fugaceous
or caducous silky long tomentose; subpersistent on
young trees, occasionally persistent on bud, twig, and
petiole. Twig c. 5 mm g, rather stout, terete; fre-
quently rather verrucose and sometimes (usually in
young trees) papery. Bud 10-25 by 3-5 mm, lanceo-
late, somewhat compressed. Stipules c. 7 cm long,
linear. Leaves 11-20 by 5-15 cm, elliptic, thinly
coriaceous, base obtuse or cuneate, apex prominently
to 8 mm long acuminate; margin frequently sinuate
towards the apex; midrib beneath often slightly
verrucose; nerves 12-20 pairs, dense, straight, at c.
40°—50°; tertiary nerves densely scalariform, slender;
petiole 3-4 cm long, rather long and slender, persis-
tently hispid on the knee. Raceme to 15 cm long,
terminal or axillary, terete, becoming angular on

300 FLORA MALESIANA ser. 1, vole:

Fig. 21. Habit of Dipterocarpus crinitus Dyer, kéruing ampélas. Brunei (Photogr. ASHTON).

1982]

DIPTEROCARPACEAE (Ashton)

301

drying, simple or singly branched; bracteoles to 30 by
2.5 mm, linear, glabrous, caducous. Flowers disti-
chous; buds to 5 by 0.8 cm. Calyx and corolla typical,
calyx glabrous. Stamens 25, shorter than the style;
filaments short; anthers narrowly oblong, tapering
apically; appendage to connective tapering, glabrous,
slightly shorter than the anther. Ovary ovoid-conical,
glabrescent; stylopodium twice as long as ovary,
narrowly cylindrical, densely tomentose; style half as
long as stylopodium, narrowly cylindrical, glabrous.
Fruit calyx entirely glabrous; tube to 3.5 cm @ and
long, obturbinate, tapering abruptly at the base
and gradually to the 0.8-1.3 cm @ neck; 3 longer
lobes 12-17 by 2-3 cm, oblong-lanceolate, obtuse,
tapering to c. 5 mm broad at the base, prominently
3-nerved; shorter lobes 4-6 by 5-7 mm, deltoid,
obtuse, undulate.

Distr. Malesia: Borneo (except S. and S.W.),
Banguey I.

Ecol. Clay soils in Mixed Dipterocarp forests, on
undulating land and hillsides below 800 m; sometimes
semigregarious on fertile soils.

Vern. Andri, damar laut, séndara (W. Borneo),
bajan, santiulit, témpehés (S.E. Borneo), kéruing
puteh.

Note. Geographically rather variable, especially in
S.E. Borneo where the closely related D. hasseltii also
occurs and can be difficult to distinguish from it when
not in fruit.

5. Dipterocarpus validus BL. Mus. Bot. Lugd.-Bat. 2
(1852) 36; WaALP. Ann. 4 (1857) 335; Mia. FI. Ind. Bat.
1, 2 (1859) 498; Ann. Mus. Bot. Lugd.-Bat. 3 (1867)
84, 85; DC. Prod. 16, 2 (1868) 614; Dyer, J. Bot. 12
(1874) 108, 153; Burck, Ann. Jard. Bot. Btzg 6 (1887)
202; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 37; MERR.
En. Born. (1921) 400; SLoor. Bull. Jard. Bot. Btzg III,
8 (1927) 272; AsHTON, Gard. Bull. Sing. 20 (1963)
237. — D. pilosus (non Roxs.) F.-Vill. Nov. App.
(1880) 20; VIDAL, Sinopsis (1883) pl. 14, f. d; Foxw.
Philip. J. Sc. 6 (1911) Bot. 244, pl. 34; ibid. 13 (1918)
Bot. 176; Merr. En. Philip. 3 (1923) 88. — D. warbur-
gil BRANDIS, J. Linn. Soc. Bot. 31 (1895) 32; Foxw. in
Merrill, Leafl. Philip. Bot. 6 (1913) 1952; Philip. J. Sc.
13 (1918) Bot. 178; Merr. En. Philip. 3 (1923) 91;
Heyneg, Nutt. Pl. ed. 2 (1927) 1098; SLoor. Bull. Jard.
Bot. Btzg III, 8 (1927) 305; Foxw. Philip. J. Sc. 67
(1938) 256; SLooT. Reinwardtia 5 (1961) 473; MEER
& Woop, Sabah For. Rec. 5 (1964) 273. — D. affinis
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 31; WHITFORD,
Bull. Bur. For. Philip. 10, 2 (1911) 70, pl. 72-73;
Foxw. Philip. J. Sc. 6 (1911) Bot. 246, pl. 35; ibid. 13
(1918) Bot. 176; Merr. En. Philip. 3 (1923) 88. — D.
lasiopodus PERK. Fragm. FI. Philip. (1904) 22; MERR.
Publ. Gov. Lab. Philip. 29 (1905) 30; En. Philip 3
(1923) 90; Reyes, Philip. J. Sc. 22 (1923) 322, pl.
13. — D. woodii Mrrr. Philip. J. Sc. 29 (1926) 399;
SLOOT. Bull. Jard. Bot. Btzg 8, III (1927) 303. — Fig. 22.

Large pale barked frequently buttressed trees.
Twigs, leaf buds, stipules outside, petioles and base of
inflorescence densely + persistently evenly long rufous

tomentose, becoming distinctly tufted as the twigs and
stipules expand; nerves and midrib beneath and
inflorescence sparsely evenly puberulent. Twigs c.
6-10 mm @, terete. Leaf buds to 9 by 1.3 cm,
lanceolate; stipules to 20 by 3 cm, lorate. Leaves 15—25
by 7.5-12 cm (to 40 by 20 cm in young trees),
elliptic-oblong to ovate, coriaceous; margin undulate,
prominently sinuate distally; base cuneate to obtuse
(narrowly peltate in young trees); apex acute to
prominently narrowly acuminate, to | cm long; nerves
22-28 pairs, straight, ascending at c. 50°, slender but
prominent beneath, +shallowly depressed above;
tertiary nerves densely scalariform, very slender,
evident but hardly elevated beneath; midrib promi-
nent beneath, +elevated above; petiole 3.5—S cm long,
prominently geniculate. Raceme to 14cm long, +un-
branched, axillary, bearing to 7 flowers. Flower buds
to 30 by 9mm; stamens c. 30, slightly shorter than style
at anthesis; filaments slender, c. + length of filiform
auriculate tapering anthers; appendage acicular,
slender, c. ? length of anthers; ovary small, ovoid,
hirsute as also the basal 4 of style; style stoutly
columnar, c. 4 times length of ovary. Fruit pedicel
expanding into the to 4 by 3.5 cm smooth turbinate
calyx tube; 2 /onger lobes to 25 by 3.5cm, lorate-spatu-
late, obtuse, c. 5mm broad at base; 3 shorter lobes to 6
by 6 mm, suborbicular, small, +recurved.

Distr. Malesia: Philippines (widespread), Borneo
(Tenom in north, and E. Coast of Sabah south-east-
wards to river Barito).

Ecol. Common and frequently gregarious both in
primary and secondary forest; on flat land, fresh water
swamp and on river banks; occasionally on low hills to
300 m.

Vern. Hagakhak (Philippines), kéruing kasugoi
(Sabah), Aambong, tampudau, kaladan (S.E. Borneo).

Note. There is some evidence of hybridisation with
D. kunstleri in the Philippines.

6. Dipterocarpus gracilis BL. Bijdr. (1825) 224; FI.
Jav. 2 (1829) 20, t. 5; WALP. Rep. 5 (1845) 123; Mia.
Fl. Ind. Bat. 1, 2 (1859) 497; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2 (1868) 609;
Dyer, J. Bot. 12 (1874) 102; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 196; BRANDIS & GILG in E. & P. Pf.
Fam. ed. 1, 3, 6 (1894) 256; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 30; K. & V. Biydr. 5 (1900) 117; MoLL &
JANSSONIUS, Mikrogr. Holz (1906) 358; Foxw. Philip.
J. Sc. 6 (1911) Bot. 248; ibid. 13 (1918) Bot. 177; ibid.
67 (1938) 249; Koorp. Exk. Fl. Java 2 (1912) 621;
HEyYNE, Nutt. Pl. ed. 1, 3 (1917) 273; ibid. ed. 2 (1927)
1094, 1095; Merr. En. Philip. 3 (1923) 89; GitG in E.
& P. Pfl. Fam. ed. 2, 21 (1925) 251; SLoor. Bull. Jard.
Bot. Btzg III, 8 (1927) 276; ibid. III, 16 (1940) 434;
Sym. Gard. Bull. S. S. 9 (1938) 321; Mal. For. Rec. 16
(1943) 177, f. 85; SMITINAND, Thai For. Bull. 1 (1954)
5; ibid. 4 (1958) 31; BACKER & BAKH. f. Fl. Java 1
(1963) 329; AsHTON, Gard. Bull. Sing. 20 (1963) 235;
Man. Dipt. Brun. (1964) 35, f. 6; ibid. Suppl. (1968) 15;
MEWVER & Woop, Sabah For. Rec. 5 (1964) 250, f.
37. — D. pilosus Roxs. [Hort. Beng. (1814) 93, nomen]

302

‘ : 3 , \ \
¥ y 2

ae a.

io LA ae > hy

Fig. 22. Close-up of bark, leaves and fruit of Dipterocarpus validus BL. Sabah (Photogr. G.H.S. Woop).

Fl. Ind. ed. Carey 2 (1832) 615; WALP. Rep. 5 (1845)
124; DC. Prod. 16, 2 (1868) 614; Dyer, Fl. Br. Ind. 1
(1874) 296; Kurz, Fl. Burma (1877) 115; BRANDIS &
GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1894) 244;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 27, p.p.; Ind.
Trees (1906) 65; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 274,
p.p., Troup, Silv. Ind. Trees 1 (1921) 39; GAMBLE,
Man. Ind. Timb. (1922) 71; Gitc in E. & P. Pfl. Fam.
ed. 2, 21 (1925) 251, p.p.; PARKER, Ind. For. Rec.
(Bot.) 13 (1927) 15; PARKINSON, Burma For. Bull. 27
(1922) 25. — Mocanera verniciflua BLCo, F1. Filip. ed.
1 (1837) 450. — D. marginatus KorTH. Kruidk. (1841)
64; WALP. Rep. 5 (1845) 124; BL. Mus. Bot. Lugd.-
Bat. 2 (1852) 37; Mia. Fl. Ind. Bat. 1, 2 (1859) 499;
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC. Prod. 16,
2 (1868) 613; Dyrr, J. Bot. 12 (1874) 105; BuRcK,

FLORA MALESIANA

[ser. I, vol. 9?

Ann. Jard. Bot. Btzg 6 (1887) 212; BRANpis, J. Linn.
Soc. Bot. 31 (1895) 38; Merr. En. Born. (1921) 399:
SLooT. Bull. Jard. Bot. Btzg III, 8 (1927) 328, f. 10,
p.p.; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 274; ibid. ed. 2
(1927) 1097. — D. vernicifluus BLco, Fl. Filip. ed. 2
(1845) 314; ibid. ed. 3 (1878) 217, t. 183; BL. Mus. Bot.
Lugd.-Bat. 2 (1852) 35; WaALp. Ann. 4 (1857) 335;
Mia. FI. Ind. Bat. 1, 2 (1858) 499; DC. Prod. 16, 2
(1868) 610; VIDAL, Sinopsis (1883) 15, t. 14 b; Rev. Pl.
Vasc. Filip. (1886) 59; Dyer, J. Bot. 12 (1874) 104;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 31; MerR.
Philip. J. Sc. 1 (1906) Suppl. 97; ibid. 3 (1908) Bot. 114;
Sp. Blanc. (1918) 268; En. Philip (1923) 91; Foxw.
Philip. J. Sc. 6 (1911) Bot. 248; ibid. 13 (1918) Bot. 177;
REYES, Philip J. Sc. 22 (1923) 321; HEYNE, Nutt. Pl. ed.
2 (1927) 1095. — D. fulvus BL. Mus. Bot. Lugd.-Bat.

1982]

2 (1852) 37; WacP. Ann. 4 (1857) 335; Mia. FI. Ind.
Bat. 1, 2 (1859) 499; DC. Prod. 16, 2 (1868) 613; Dyer,
J. Bot. 12 (1874) 108; BRANbis, J. Linn. Soc. Bot. 31
(1895) 40.— Anisoptera palembanica Mig. Sum.
(1862) 191, 485; Ann. Mus. Bot. Lugd.-Bat. (1867) 85;
DC. Prod. 16, 2 (1868) 616. — D. hispidus (non THwW.)
F.-VILL. Nov. App. (1880) 20. — D. velutina VIDAL,
Rev. Pl. Vasc. Filip. (1886) 59; PERK. Fragm. FI.
Philip. (1904) 22; Merr. Philip. J. Sc. 3 (1908) Bot.
114. — D. bancanus Burck, Ann. Jard. Bot. Btzg 6
(1887) 196; BRANDis J. Linn. Soc. Bot. 31 (1895) 31;
Heyne, Nutt. Pl. ed. 2 (1927) 1095. — D. skinneri
KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 91;
BRANDIS, J. Linn. Soc. Bot. 3 (1895) 26; RIDL. Fl. Mal.
Pen. | (1922) 214; Crais, Fl. Siam. Enum. | (1925)
137; HEYNE, Nutt. Pl. ed. 2 (1927) 1098: SLoot. Bull.
Jard. Bot. Btzg III, 8 (1927) 294; Foxw. Mal. For.
Rec. 10 (1932) 70, excl. var. hirtus RiDL.; BURK. Dict.
(1935) 845. — D. vanderhoevenii K. & V. Bull. Inst.
Bot. Btzg 2 (1899) 3; Bijdr. 5 (1900) 118; Mott &
JANSSONIUS, Mikrogr. Holz (1906) 356; Koorpb. Exk.
Fl. Java 2 (1912) 621. — Shorea mollis BOERL. Cat.
Hort. Bog. 2 (1901) 110. — D. angustialatus HEI,
Bot. Tidsskr. 25 (1903) 43; GUERIN, Fl. Gen. I.-C. 1
(1910) 365; Crars, Fl. Siam. Enum. 1 (1925) 133;
FISCHER, Kew Bull (1926) 457. — D. schmidtii HEm™,
Bot. Tidsskr. 25 (1903) 43.

Twig, leaf bud, stipule outside, leaf beneath, midrib
above and petiole densely persistently scabrid rufous
tomentose. 7wig c. 3 mm @ apically, terete or slightly
compressed, with minute cracks initiating from elon-
gated lenticels. Bud 10-14 by 3-S mm, narrowly
conical, obtuse. Stipule c. 5 cm long, narrowly
lanceolate. Leaves 8-15 by 4-10 cm, elliptic to ovate,
base obtuse; apex shortly acuminate; nerves 12-20
pairs, usually dense, at 40°—S0°; petiole 2-2.5 cm long.
Raceme to 9 cm long, terminal or axillary, terete,
shortly pale brown tomentose or glabrescent, singly
branched. Flower bud to 25 by 8 mm. Calyx and
corolla typical, calyx shortly tomentose. Stamens c.
30, exceeding the style; filaments short; anthers linear:
appendage to connective c. 2 times length of anther,
filiform, tapering, sparsely setose. Ovary ovoid-coni-
cal, tapering into the stylopodium; stylopodium and
style slender, filiform, tomentose in the basal half.
Fruit calyx glabrous; tube to 2cm @, smooth, globose;
2 longer lobes to 14 by 2.5 cm, narrowly spatulate,
obtuse, to | cm broad at base; 3 shorter lobes to 2.2 by
1 cm, ovate, constricted at base, the 2 sides becoming
revolute back to back.

Distr. Andamans, Chittagong, Burma, S.E. and
Peninsular Thailand, and in Malesia: Malaya, Suma-
tra, W. Java, Borneo (Sampit and the Rejang valley
eastwards), Philippines.

Ecol. Very widespread, often gregarious in sea-
sonal semi-evergreen dipterocarp forest on red soils;
becoming scattered, rare, and confined to fertile red
soils, in everwet areas (cf. Anisoptera costata), below
800 m.

Vern. Kéruing kesat, k. bungoh, k. daun halus
(Mal.), wuluk bulan (Jav.), bembang (Sum.). Philip-

DIPTEROCARPACEAE (Ashton)

303

pines: agan-an (Bicol), duha (Ibn.), kurimau (Ibn.),
lalian (TYag.), lanutan (Neg.), lauan (Zamb., Sul.),
pagsa hingau (Tag.), palamopang (Tag.), palohap
(Sbl.), pamalalian (Cag., Ibn.), pamantulen (IIk.),
pamantuling (Pang.), Pamarnisen (Ibn.), panao (Tag.,
Pang., Sbl., Ibn., Pamp.), putsa hingan (Tag.), sitam
(Ibn.).

Note. Specimens from the Indo-Burmese region
and northern Malaya, and also the Philippines are
usually more shortly sparsely evenly pubescent.

7. Dipterocarpus baudii KorTH. Kruidk. (1841) 59, t.
5; WALP. Rep. 5 (1845) 123; BL. Mus. Bot. Lugd.-Bat.
2 (1852) 36; Mia. Fl. Ind. Bat. 1, 2 (1859) 497: Sum.
(1862) 485; Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85;
DC. Prod. 16, 2 (1868) 609; SCHEFFER, Nat. Tijd. N. I.
31 (1870) 346; Burck, Ann. Jard. Bot. Btzg 6 (1887)
198: HEYNE, Nutt. Pl. ed. 2 (1927) 1095; Foxw. Mal.
For. Rec. 3 (1927) 42; SLoor. Bull. Jard. Bot. Btzg III,
8 (1927) 273; PARKER, Ind. For. Rec. 16 (1931) 3;
PARKINSON, Burma For. Bull. 27 (1932) 11; Foxw.
Mal. For. Rec. 10 (1932) 64; Burk. Dict. (1935) 842:
Sym. Mal. For. Rec. 16 (1943) 168, f. 85.—D.
duperreana PIERRE in Lanessan, PI. Util. Colon. Fr.
(1886) 297; Fl. For. Coch. 3 (1889) t. 219; BRANDIs, J.
Linn. Soc. Bot. 31 (1895) 7, 28; GUERIN, FI. Gen. I.-C.
1 (1910) 358; Rip. Fl. Mal. Pen. 1 (1922) 214; Crars,
Fl. Siam. Enum. | (1925) 134. — D. scortechinii KING,
J. R. As. Soc. Beng. Sc. 62, 2 (1893) 91; BRANDis, J.
Linn. Soc. Bot. 31 (1895) 28; Burk. J. Str. Br. R. As.
Soc. 81 (1920) 55, fig.; HEYNE, Nutt. Pl. ed. 2 (1927)
1095; Foxw. Mal. For. Rec. 10 (1932) 64.—D.
pilosus (non ROxB.) BRANDIS, J. Linn. Soc. Bot. 31
(1895) 27, p.p.; BRANDIS & GILG in E. & P. Pfl. Fam.
ed. 1, 3, 6 (1894) 244, p.p.; HEYNE, Nutt. Pl. ed. 1, 3
(1917) 224; Gita in E. & P. Pfl. Fam. ed. 2, 21 (1925)
251; Foxw. Mal. For. Rec. 3 (1927) 42.

Large tree. Leaf buds, stipules outside, twigs,
petiole and inflorescences densely persistently long
tufted rufous tomentose, leaf beneath and midrib
above, ovary apex, calyx and petals outside shortly so;
fruit calyx and leaves above glabrescent. Twig c. 4-6
mm @ apically, stout. Bud c. 15 by 8 mm, lanceolate;
stipule to 15 by 3 cm, lorate-lanceolate, becoming
sparsely tufted tomentose after expansion, tinged red.
Leaves 17—32 by 8-17 cm, +elliptic, thinly coriaceous,
drying rufous below, dark purplish above; base
narrowly obtuse; acumen to 2 cm long, tapering;
nerves 14-28 pairs, ascending, stoutly prominent
beneath; tertiary nerves dense; midrib prominent
beneath, applanate above; petiole 3-5 cm long, c. 3mm
@. Raceme to 6 cm long, axillary, generally un-
branched, bearing to 6distichous flowers. Flower budto
3 by 1 cm. Stamens 30, exceeding style at anthesis;
filaments slender, } length of the linear tapering
anthers; appendage acicular, c. } length of anthers;
ovary small, ovoid, pubescent; style stoutly columnar,
pubescent but for the apical 3. Fruit subsessile, pedicel
to 2 by 2 mm; calyx tube to 2 cm @, globose; 2 longer
lobes to 18 by 3 cm, lorate, obtuse, c. 6 mm wide at
base; 3 shorter lobes to 2 by | cm, elliptic.

304

FLORA MALESIANA

[ser. I, vol. 9?

Distr. Cochinchina, Cambodia, Burma, Thailand,
and in Malesia: Malaya, Sumatra (Atjeh south to
Padang highlands in W., and river Kampar in east,
southwards to Peranap).

Ecol. Lowland forest, undulating land.

Vern. Kéruing bulu, k.dadeh, k. sudoi, damar etoi, d.
minyak, néram bukit, stui, térak (Malaya); lagan,
kéruing, marakeloewang (Sumatra).

Note. Confused in the past with D. gracilis, from
which it differs in longer tomentum and larger size of
all parts. The related D. elongatus KoRTH., and also D.
hispidus THw. of Ceylon which differs in having a
cordate leaf with at most 16 pairs of nerves, would
appear to be geographical segregates.

8. Dipterocarpus obtusifolius TEYSM. ex Mig. Ann.
Mus. Bot. Lugd.-Bat. 1 (1863) 214; DC. Prod. 16, 2
(1868) 608; WALP. Ann. 7 (1869) 377; Dyer, FI. Br.
Ind. 1 (1874) 295; Kurz, Fl. Burma 1 (1877) 115;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 7, 27; Ind. Trees
(1906) 65, 701; GuERIN, FI. Gen. I.-C. 1 (1910) 358;
RYAN & Kerr, J. Siam Soc. 8 (1911) 3, incl. var.
subnudus RYAN & KERR; Troup, Silv. Ind. Trees
(1921) 39; Crais, Fl. Siam. Enum. (1925) 136;
PARKER, Ind. For. Rec. 16 (1941) 9; PARKINSON,
Burma For. Bull. 27 (1932) 23; TARDIEU, FI. Gen. I.-C.
Suppl. 1 (1943) 338; Sym. Mal. For. Rec. 16 (1943)
184, f. 85; SMITINAND, Thai For. Bull. 1 (1954) 51, incl.
var. glabricalyx et var. vestitus SMITINAND; ASHTON,
Gard. Bull. Sing. 31 (1978) 7.— D. vestitus WALL.
[Cat. (1828) 954; WALP. Rep. 5 (1845) 124; DC. Prod.
16, 2 (1868) 614, nomen] ex Dyer, FI. Br. Ind. | (1874)
295; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 270. — D.
punctulatus PIERRE, Fl. For. Coch. 3 (1889) t. 221;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 29; GUERIN, FI.
Gen. I.-C. 1 (1910) 357.

Small or medium-sized, usually misshapen and
often coppiced tree with open crown and thick, corky
deeply fissured bark. Twigs, leaf buds and petioles
densely pale fulvous to buff hirsute or glabrous; leaf
beneath and midrib above shortly densely evenly
persistently buff pubescent, with the nervation more,
or less (Malaya), hirsute; fruit glabrescent. Twig c. 7
mm @ apically, stout. Bud to 20 by 6 mm, lanceolate,
subacute; stipules to 7 by 1 cm, lorate, obtuse. Leaves
14-22 by 10-16 cm, broadly ovate, thickly coriaceous,
prominently plicate, crenate in the distal 4; base
obtuse or subcordate; apex obtuse or subacute; nerves
14-20 pairs, stout, prominent beneath, ascending;
petiole 2.5—4 cm long, c. 4mm @, stout. Flower bud to
2.5 by 1.2 cm; stamens c. 30, + equalling style at
anthesis; filaments lorate, tapering c. 4 as long as the
relatively stout tapering lorate anthers; appendage
acicular, c. 3 as long as anther; ovary ovoid, pubescent,
surmounted by a slender columnar style + thrice its
length and pubescent in the basal 5. Fruit pedicel to 2
by 3 mm, short; calyx tube to 2.5 cm @, subglobose,
smooth; 2 longer lobes to 15 by 3 cm, lorate-spatulate,
obtuse, abruptly tapering to c. 8 mm broad at the
subrevolute base; 3 shorter lobes to 15 by 10 mm,
ovate, obtuse, subrevolute.

Distr. Burma, Thailand, Indochina, and in Male-
sia: N. W. Malaya (Perlis).

Ecol. Rare, in Schima-bamboo forests in Malesia.
Elsewhere a characteristic and frequently gregarious
fire-resistant component of the Dry Dipterocarp
(savanna) forest.

Note. Very variable, especially in the amount and
density of tomentum, the more glabrous form (var.
subnudus RYAN & KERR) prevailing in the more humid
areas including N.W. Malaya; in this respect resem-
bling other dipterocarp species of the seasonal
regions, e.g. Dipterocarpus turbinatus, Anisoptera cos-
tata, Shorea siamensis.

9. Dipterocarpus rotundifolius Foxw. Mal. For. Rec.
10 (1932) 73, pl. 4; BuRK. Dict. (1935) 845; Sym. Mal.
For. Rec. 16 (1943) 188, f. 85.

Twigs, leaf buds, stipules outside, petiole and leaf
nervation beneath sparsely very long tufted golden
tomentose (especially saplings and young trees) or
glabrous; parts of petals exposed in bud and ovary
densely shortly evenly buff puberulent. Twig c. 5mm
@ apically, dark brown suffused with pale grey,
minutely verrucose-lenticellate; stipule scars promi-
nent. Buds to 4 by 2 cm, enormous, ellipsoid, obtuse;
stipules to 9 by 5 cm, elliptic, obtuse, concave. Leaves
8.5-16 by 8-15 cm, broadly ovate to suborbicular,
thickly coriaceous; base obtuse to cordate; apex
subacuminate or more typically obtuse; margin undu-
late, subrevolute; nerves 11(-13) pairs, spreading,
sharply prominent beneath, shallowly depressed
above; tertiary nerves remotely scalariform, distinctly
elevated beneath; petiole 4-10 cm long, 2-3 mm @,
very long. Flowers solitary on the 6 cm long axillary
peduncles, subtended and enclosed in a large, to 5 by 3
cm, elliptic obtuse concave subpersistent bract;
stamens 30, greatly exceeded by style at anthesis;
filaments slender, columnar, short; anthers long,
linear, tapering, prominently auriculate; appendage
acicular, slender, c. 4 length of anthers; gynoecium
puberulent but for the distal 4 of the style; ovary small,
ovoid; style slender, filiform, tapering, very long. Fruit
pedicel to 5 by 4 mm, expanding into the to 18 by 14
mm fusiform-ellipsoid smooth calyx tube; 2 longer
lobes to 9 by 2.5 cm, oblong, obtuse, c. 7 mm wide at
base, 3-nerved, the lateral 2 nerves short and the
central prominently laterally branched; 3 shorter
lobes to 14 by 7 mm, elliptic, obtuse, subrevolute.

Distr. Malesia: Malaya (Perak, and Trengganu
southwards on east coast).

Ecol. Local, in Mixed Dipterocarp forest on
coastal hills.

Vern. Kéruing mengkai.

Note. A curious and distinct form which is clearly
related to the widespread D. crinitus.

10. Dipterocarpus chartaceus Sym. Gard. Bull. S.S. 9
(1938) 322; Mal. For. Rec. 16 (1943) 169, f. 85,
86. — D. skinneri var. hirtus Ripv. Fl. Mal. Pen. |

1982]

DIPTEROCARPACEAE (Ashton)

305

(1922) 215; SLoor. Bull. Jard. Bot. Btzg III, 8 (1927)
295. — D. skinneri (non KING) RIDL. Fl. Mal. Pen. 1
(1922) 214, p.p.; Foxw. Mal. For. Rec. 10 (1932) 70,
P-p.

Large tree. Twig, leaf bud, stipule, inflorescence,
flower calyx, ovary apex and petals outside densely
persistently shortly golden pubescent; leaf undersur-
face sparsely caducously so; fruit glabrescent. Twig c.
4 by 3 mm @g, pale brown, somewhat compressed.
Buds to 7 by 3 mm, lanceolate-falcate; stipules to 8 by
9.8 cm, lorate-lanceolate. Leaf 8.5-19 by 4-9 cm,
elliptic-ovate or rarely obovate, crisply chartaceous
drying pale mauve and somewhat lustrous; base
cuneate; acumen to | cm long, tapering; nerves
10-12(-14) pairs, slender but distinctly elevated
beneath, ascending; tertiary nerves densely scalari-
form, hardly elevated; petiole 2.3-2.8 cm long, c. 2mm
@, slender. Raceme to 6 cm long, axillary, singly
branched or unbranched, bearing to 6 flowers. Flower
buds to 22 by 8 mm, fusiform-lanceolate. Stamens 30,
shorter than style at anthesis; filaments compressed at
base, tapering rapidly and then filiform, c. 5 length of
the relatively short linear tapering anthers; appendage
acicular, c. ? length of anther; ovary narrowly ovoid,
pubescent, tapering into the long slender filiform
style; style glabrous in apical 5. Fruit pedicel c. 1 by 2
mm, short; calyx tube to 18 mm @, subglobose; 2
longer lobes to 13 by 3.2 cm, lorate, somewhat
tapering distally, obtuse, c. 7 mm broad and revolute
at base; 3 shorter lobes to 4 by 4 mm, orbicular,
revolute.

Distr.
Malaya.

Ecol. Infrequent in lowland forest, sometimes in
areas periodically inundated, especially in seasonal
areas and near coast.

Vern. Kéruing kertas (Mal.).

Note. The fallen leaf is crisp and papery, drying
pale buff.

Peninsular Thailand, and in Malesia:

11. Dipterocarpus caudatus Foxw. Philip. J. Sc. 13
(1918) Bot. 177; ibid. 67 (1938) 256; MerR. En. Philip.
3 (1923) 89; SLoor. Bull. Jard. Bot. Btzg III, 8 (1927)
302, 303; ASHTON, Gard. Bull. Sing. 31 (1978) 8.

a. ssp. caudatus.

Leaf bud and stipule outside shortly densely pale
buff pubescent. Parts otherwise glabrous. Twig 1-3
mm @ apically, slender, + compressed, smooth, with
slightly swollen amplexicaul stipule scars. Bud 8-17 by
2-3 mm, narrowly falcate or linear; apex tufted with
slightly longer hairs arising from the outer surface.
Stipule c. 3.5 by 0.5 cm, linear, obtuse. Leaves 7—11 by
3.5-5 cm, broadly elliptic, + prominently plicate;
base broadly cuneate; acumen to 1.2 cm long, promi-
nent, narrow; nerves 9-12 pairs, prominent, at
35°—45°; petiole 1—-1.5 cm long, slender. Raceme to 12
cm long, slightly compressed; unbranched or singly
branched, the flowers secund, few; bracts unknown.
Flower bud to 3 by 0.8 cm. Calyx and corolla typical,
calyx puberulent. Stamens c. 30, shorter than the style;

filaments short; anthers short, linear, tapering; appen-
dage to connective slightly shorter than anther, stout
at base, tapering. Ovary conical, densely pubescent;
style and stylopodium narrowly cylindrical, some-
what stouter and more densely tomentose in the basal
half. Fruit calyx glabrous tube c. 2 cm long and g,
slightly obovoid, glabrous, minutely lenticellate,
tapering gradually to the pedicel, c. 1 cm @ at the neck;
2 longer calyx lobes to 14 by 3 cm, oblong, 3-nerved,
obtuse, rather abruptly narrowing to c. 5mm broad at
base; 3 shorter lobes 4-8 by 3-4 mm, variable,
strongly recurved and revolute.

Distr. Malesia: S. E. Philippines (Mindanao,
Luzon).

Ecol. Local in everwet zone.

b. ssp. penangianus (Foxw.) ASHTON, Gard. Bull. Sing.
Sing. 31 (1978) 8. — D. penangianus Foxw. Mal. For.
Rec. 10 (1932) 72, pl. 3 (germ. seeds); Sym. Mal. For.
Rec. 16 (1943) 185, f. 85; ASHTON, Man. Dipt. Brun.
(1964) 43, f. 6-7; ibid. Suppl. (1968) 16.

Twig, petiole, raceme, midrib and nerves shortly
sparsely fugaceous pubescent, twigs sometimes den-
sely persistently so. Leaves narrowly elliptic,
applanate, with narrowly cuneate base, hardly promi-
nent nerves beneath and 1.5—2.5 cm long petiole.

Distr. Malesia: Malaya (Perak, Penang, W. & E.
Johore, Kelantan, Pahang), Singapore, Sumatra
(Karimun, Musala), Borneo (Sarawak N.E. of river
Rejang, S.E. Sabah).

Ecol. Coastal hills, locally frequent.

Vern. Kéruing gasing, k. déran, songgi déran.

Note. Malayan collections are distinguished by
their denser more persistent indumentum on twigs,
and by their prominently lenticellate fruit calyx tube.

12. Dipterocarpus kerrii KiNG, J. R. As. Soc. Beng.
Sc. 62, 2 (1893) 93; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 28; Burk. J. Str. Br. R. As. Soc. 81 (1920) 55,
fig.; RIDL. Fl. Mal. Pen. 1 (1922) 215; Crat, Fl. Siam.
Enum. | (1925) 136; Soot. Bull. Jard. Bot. Btzg III, 8
(1927) 295; PARKER, Ind. For. Rec. 16 (1931) 15;
PARKINSON, Burma For. Bull. 27 (1932) 19; Foxw.
Mal. For. Rec. 10 (1932) 69; Burk. Dict. (1935) 844;
Philip. J. Sc. 67 (1938) 253; Sym. Mal. For. Rec. 16
(1943) 181, f. 85, 92; SMITINAND, Thai For. Bull. 1
(1954) 4; ibid. 4 (1958) 38; MeER & Woop, Sabah
For. Rec. 5 (1964) 259, f. 40. — D. obconicus Foxw. in
Elmer, Leafl. Philip. Bot. 6 (1913) 1951; Philip J. Sc. 13
(1918) Bot. 178; Merr. En. Philip. 3 (1923) 90; SLoor.
Bull. Jard. Bot. Btzg III, 8 (1927) 299. — D. pertur-
binatus Foxw. Philip. J. Sc. 13 (1918) Bot. 177. — D.
cuneatus Foxw. Philip. J. Sc. 13 (1918) Bot. 178;
Me_rr. En. Philip. 3 (1923) 89; SLoor. Bull. Jard. Bot.
Btzg III, 8 (1927) 300.

Large tree. Parts of petals exposed in bud, inside of
stipules and ovary apex densely silky cream pubes-
cent, parts otherwise glabrous. Twig c. 3 mm @,
subterete, blackish, minutely pale lenticellate. Bud to
12 by 3mm, lanceolate-falcate, drying black; stipule to

306

EE

pt Cs hl ,
vise ih ig st it,

Fig. 23. The sacred forest of Sangeh in Bali, bordering on rice-field, consisting of Dipterocarpus hasseltii BL.
(Photogr. DE VooGp).

8 by 0.5 cm, linear-lanceolate, subacute. Leaves 8-13
by 3.3-7 cm, + broadly elliptic, coriaceous, drying
dark chocolate-brown; base cuneate; acumen to 5mm
long, short, nerves (7—)9-11 pairs, slender but promi-
nent beneath, ascending; tertiary nerves densely sca-
lariform, very slender and barely elevated beneath;
petiole 2-2.8 cm long, slender. Inflorescence to 8 cm
long, singly branched or unbranched, bearing up to 5
flowers. Flower buds to 2.5 by | cm. Stamens c. 30,
exceeding style at anthesis; filaments filiform, slender,
c. 3 length of the short linear-lorate + tapering
subauriculate anthers; appendage acicular, slender, c.
+ length of anther; ovary small, ovoid, puberulent as
also the basal 4 of the stoutly columnar style. Fruit
pedicel to 3 by 3 mm, stout. Fruit calyx to 3.5 cm @,
globose to subturbinate, smooth; 2 longer lobes to 14
by 3 cm, lorate, tapering apically, obtuse, abruptly
constricted to c. 8 mm wide and subrevolute at base; 3
shorter lobes to | by | cm, prominent, suborbicular,
subrevolute.

Distr. Andamans, Burma, Peninsular Thailand
and in Malesia: Malaya (Malacca and Pahang north-
wards, coastal towards south of range), Sumatra
(Indragiri), N. Borneo (Sandakan area), Philippines.

Ecol. Semi-evergreen and evergreen coastal dipter-

FLORA MALESIANA

[ser. I, vol. 9?

ocarp forests in periodically or seasonally dry cli-
mates, on red lateritic soils, on undulating land and
hills below 400 m; frequently gregarious.

Vern. Kéruing gondol, k. chair, damar minyak
(Malaya), apitong, panao, malapanao, palsahingan
(Philippines).

13. Dipterocarpus hasseltii BL. Fl. Jav. 2 (1829) 22, t.
6; KorTH. Kruidk. (1841) 65; WALP. Rep. 5 (1845)
123; Mia. FI. Ind. Bat. 1, 2(1859) 497; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2 (1868) 609;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 196; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 30; K. & V. Bijdr. 5 (1900)
109; Rpt. J. Str. Br. R. As. Soc. 33 (1900) 42; ibid. 34
(1900) 94; MoLL & JANSSONIUS, Mikrogr. Holz (1906)
359; Koorp. Exk. Fl. Java 2 (1912) 620; BAKER f. J.
Bot. 62 (1924) 10; Cras, Fl. Siam. Enum. 1 (1925)
135; HEYNE, Nutt. Pl. ed. 2 (1927) 1096, 1097; SLoor.
Bull. Jard. Bot. Btzg III, 8 (1927) 280; ibid. III, 16
(1940) 436; Foxw. Mal. For. Rec. 10 (1932) 67; Philip.
J. Sc. 67 (1938) 251; BurK. Dict. (1935) 844; Mal. For.
Rec. 16 (1943) 180, f. 85, 91; SMITINAND, Thai For.
Bull. 4 (1958) 35; BACKER & BAKH. f. Fl. Java 1 (1963)
329; MEER & Woop, Sabah For. Rec. 5 (1964) 255;
ASHTON, Gard. Bull. Sing. 31 (1978) 8.— D. tam-

1982]

purau KorTu. Kruidk. (1841) 63; WALP. Rep. 5 (1845)
123; BL. Mus. Bot. Lugd.-Bat. 2 (1852) 36; Mia. FI.
Ind. Bat. 1, 2 (1859) 498; Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 85; DC. Prod. 16, 2 (1868) 609; Dyer, J. Bot. 12
(1814) 103; Burck, Ann. Jard. Bot. Btzg 6 (1887) 198,
p.p.; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 29; MERR.
En. Born. (1921) 400; SLoor. Bull. Jard. Bot. Btzg III,
8 (1927) 292, f. 1; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 274;
ibid. ed. 2 (1927) 1098; cf. ASHTON, Gard. Bull. Sing.
20 (1963) 234.— D. quinquegonus BL. Mus. Bot.
Lugd.-Bat. 2 (1852) 36; WALP. Ann. 4 (1857) 335;
Mia. Fl. Ind. Bat. 1, 2 (1859) 497; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; Merr. En. Born. (1921)
400. — D. pentagonus DC. Prod. 16, 2 (1868) 610;
Dyer J. Bot. 12 (1874) 104; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 199; BRANDis J. Linn. Soc. Bot. 31 (1895)
34; SLtoor. Bull. Jard. Bot. Btzg III, 8 (1927)
282. — D. lampongus SCHEFF. Nat. Tijd. N. I. 31
(1870) 146; Dyer, J. Bot. 12 (1874) 102; Burck, Ann.
Jard. Bot. Btzg 6 (1887) 197; BRANpis, J. Linn. Soc.
Bot. (1895) 31; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 272;
ASHTON, Gard. Bull. Sing. 20 (1963) 236, in obs. sub D.
gracilis. — D. balsamiferus BL. Mus. Bot. Lugd.-Bat.
2 (1852) 37; Mia. FI. Ind. Bat. 1, 2 (1859) 498; Ann.
Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2
(1868) 613; Dyer, J. Bot. 12 (1874) 108; Burck, Ann.
Jard. Bot. Btzg 6 (1887) 203; BRANpis, J. Linn. Soc.
Bot. 31 (1895) 40; K. & V. Bijdr. 5 (1900) 111; MoLL &
JANSSONIUS, Mikrogr. Holz (1906) 357; HEYNE, Nutt.
Pl. ed. 2 (1927) 1096. — D. trinervis (non BL.) Foxw.
Philip. J. Se. 6 (1911) Bot. 247; ibid. 13 (1918) Bot. 177;
Merk. En. Philip. 3 (1923) 91. — D. subalpinus Foxw.
in Elmer, Leafl. Philip. Bot. 6 (1913) 1950; Philip. J.
Sc. 13 (1918) Bot. 177; ibid. 67 (1938) 255; SLoor. Bull.
Jard. Bot. Btzg III, 8 (1927) 283. — Fig. 23, 24.

Parts of petals expanded in bud and ovary apex
densely persistently buff puberulent, calyx at first
frequently so, nerves beneath sparsely so or glabres-
cent, otherwise glabrous. Twig c. 4 by 2 mm g@
apically, somewhat compressed, black. Buds to 20 by 5
mm, falcate-lanceolate, drying black; stipules to 12 by
1 cm, lorate-lanceolate, subacute. Leaves 9-16 by
5-10 cm, elliptic, subcoriaceous, prominently plicate;
margin + prominently crenate; base cuneate; acumen
to 1 cm long, short; nerves 11—14 pairs, slender but
prominent beneath, ascending; petioles 2.54 cm long,
slender. Inflorescences to 10 cm long, axillary, bearing
to 4 + secund flowers. Flower buds to 3 by | cm,
fusiform. Stamens 30, shorter than style at anthesis;
filaments c. 5 length of anthers, long, broad and
compressed at base, rapidly tapering and filiform
above; anthers linear, somewhat tapering distally;
appendage acicular, very slender, c. 4 length of anther.
Ovary narrowly ovoid-lanceolate, tapering into the
very long slender filiform style; gynoecium puberulent
except for the distal 4. Fruit pedicel to 3 by 2mm; calyx
tube to 3 cm g, subglobose smooth; 2 longer lobes to
22 by 3 cm, lorate-spatulate, usually obtuse, 3-nerved,
c. 9mm wide at base; 3 shorter lobes to 15 by 13 mm,
suborbicular, subrevolute, prominent.

Distr. Peninsular Thailand, and in Malesia around

DIPTEROCARPACEAE (Ashton)

Fig. 24. Basal part of trunk of a relict tree of
Dipterocarpus hasseltii BL. near a coffee estate, Getas,
near Salatiga, Central Java (Photogr. ROEPKE, 1913).

the periodically dry borders of the everwet zone of the
Sunda shelf: Malaya (Central and North), Sumatra
(P. Simalur and Mentawei Is.), W. Java, Lesser Sunda
Is. (Bali) and S. and E. Borneo to S.E. Sabah,
Philippines including Palawan.

308

Ecol. Lowland dipterocarp forests on well-drained
but moist fertile red soils in valleys and on hillsides,
sometimes on calcareous soil, even limestone (Java);
sometimes gregarious; to 600 m.

Notes. I agree with MERRILL (En. Philip. 3, 1923,
89) in reducing D. subalpinus Foxw., based on ELMER
13521 from Cabadbaran, Mt Urdaneta, Agusan
Prov., Mindanao. Foxworthy (Philip. J. Sc. 67, 1938,
251) claimed that D. subalpinus differed from D.
hasseltii in having smaller leaves and fruit without the
distinctive large, suborbicular, shorter calyx lobes, in
this resembling D. gracilis BL. The problem is con-
founded by the frequently unusually sparsely tomen-
tose leaves of D. gracilis in the Philippines, especially
at higher altitudes where D. subalpinus appears to
prevail. However, the specimens presently available
convinced me that Philippine material is within the
range of variation of D. hasseltii from other parts ofits
range.

It is curious that, though this is clearly the species
that is venerated in the sacred forest of Sanggeh, Bali
collections from Lombok and Sumbawa Is., on a
similar but clearly natural site in the former, appear to
represent D. retusus (see there). I cannot distinguish
these two species in Java or the Lesser Sunda Islands
when young. Immature trees were seen to fruit
abundantly in Sanggeh. There is the possibility of
hybridization, in this part of the range, of these two
otherwise very distinct species.

14. Dipterocarpus retusus BL. Cat. (1823) 77, cum
tab.; Verh. Bat. Gen. K. W. 9 (1823) 178; Bijdr. (1825)
223; Fl. Jav. 2 (1829) 14, t. 2; WALP. Rep. 5 (1845) 122;
Mia. Fl. Ind. Bat. 1, 2 (1859) 497; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2 (1868) 609;
Dyer, J. Bot. 12 (1874) 102; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 197; BRANDIS & GILG in E. & P. Pfl.
Fam. ed. 1, 3, 6 (1894) 256; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 30; K. & V. Bijdr. 5 (1900) 112; Koorp. Exk.
Fl. Java 2 (1912) 21; GitG in E. & P. Pfl. Fam. ed. 2, 21
(1925) 251, 256, fig.; HEYNE, Nutt. Pl. ed. 2 (1927)
1097; SLooT. Bull. Jard. Bot. Btzg III, 8 (1927) 286;
ibid. III, 16 (1940) 435; Foxw. Mal. For. Rec. 10
(1932) 69: Sym. Mal. For. Rec. 16 (1943) 187, f. 85;
BACKER & BAKH. f. Fl. Java 1 (1963) 329; ASHTON,
Gard. Bull. Sing. 31 (1978) 9. — D. trinervis BL. Cat.
(1823) 78; Verh. Bat. Gen. K. W. 9 (1823) 178; Bijdr.
(1825) 223; Fl. Jav. 2 (1829) 11, t. 1, incl. var. elegans
BL. et var. canescens BL.; KORTH. Kruidk. (1841) 61;
WALP. Rep. 5 (1845) 122; LINDLEY, Veg. King. (1846)
393; HAssK. Pl. Jav. Rar. (1848) 270; SCHNIZLEIN, Ic. 3
(1857) t. 213; Mia. Fl. Ind. Bat. 1, 2 (1859) 496; Ann.
Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2
(1868) 608; ScHEFF. Nat. Tijd. N. I. 31 (1870) 346;
BAILLON, Hist. Pl. 4 (1873) 204; Dict. Bot. 1 (1878)
562; Tr. Bot. Med. Pharm. 2 (1884) 816; Dyer, J. Bot.
12 (1875) 102; Burck, Ann. Jard. Bot. Btzg 6 (1887)
195; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6
(1894) 256; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 29;
K. & V. Bidr. 5 (1900) 105; MoLL & JANSSONIUS,
Mikrogr. Holz (1906) 354; Koorp. Exk. Fl. Java 2

FLORA MALESIANA

[ser. I, vol. 9?

(1912) 620; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 274; ibid.
ed. 2 (1927) 1098; Gitc in E. & P. Pfl. Fam. ed. 2, 21
(1925) 251, 256, fig.; SLoor. Bull. Jard. Bot. Btzg III, 8
(1927) 284; ibid. III, 16 (1940) 436; BACKER & BAKH. f.
Fl. Java 1 (1963) 329. — D. spanoghei BL. Fl. Jav. 2
(1829) 16, t. 3; WALP. Rep. 5 (1845) 122; Mia. FI. Ind.
Bat. 1, 2 (1859) 497; Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 85; DC. Prod. 16, 2 (1868) 609; BurcK, Ann.
Jard. Bot. Btzg 6 (1887) 198, incl. var. cordata BURCK;
HEYNE, Nutt. Pl. ed. 2 (1927) 1097. — D. macrocarpus
VESQUE, C. R. Ac. Sc. Paris 78 (1874) 627; J. Bot. 12
(1874) 151; Dyer, J. Bot. 12 (1874) 153; PARKER, Ind.
For. Rec. (Bot.) 20, 15 (1934) 3. — D. pubescens K. &
V. Bull. Inst. Bot. Btzg 2 (1899) 2; Bijdr. 5 (1900) 115;
MoLt & JANssONIUS, Mikrogr. Holz (1906) 359;
Koorp. Exk. Fl. Java 2 (1912) 621; HEYNE, Nutt. PI.
ed. 1, 3 (1917) 274; ibid. ed. 2 (1927) 1097. — D.
tonkinensis A. CHEV. Bull. Econ. Indochine 20 (1918)
798. — Fig. 17H.

Medium-sized tree. Twigs, panicles, flower calyx
and corolla outside shortly densely buff puberulent or
glabrous; petiole, leaf bud and ovary densely persis-
tently pale orange-rufous silky tomentose or glabrous,
stipule caducously so, becoming tufted glabrescent on
expanding; leaf glabrescent. Twigs c. 8 mm @ apically,
stout, becoming pale brown. Buds to 2.5 by | cm,
ovoid-lanceolate, acute; stipule to 4 cm long, lanceo-
late, acute. Leaves 16-28 by 7-17 cm (to 50 by 70 cm in
young trees), + broadly elliptic-oblong, coriaceous,
prominently plicate, drying dark chocolate-brown;
nerves 16-19 pairs, prominent beneath, + depressed
above, ascending; tertiary nerves densely scalariform,
very slender and hardly elevated beneath; petiole 2.5—7
cm long. Panicles to 10 cm long, unbranched, axillary.
Flower bud to 3 by 1 cm, fusiform. Stamens 30,
exceeding style at anthesis; filaments short, filiform;
anthers long, linear, tapering; appendage acicular,
slender, c. + length of anther. Ovary small, ovoid,
densely pubescent as also the basal 3 of the stoutly
columnar somewhat tapering style. Fruit pedicel to 3
by 4 mm, stout; calyx tube to 3.5 mm @, subglobose,
sparsely minutely pale lenticellate, smooth; 2 longer
lobes to 25 by 4.5 cm, obtuse, prominently 3-nerved,
tapering abruptly to c. 12 mm wide at base; 3 shorter
lobes to 2 by 1.5 cm, prominent, ovate-elliptic, obtuse,
subrevolute.

Distr. Assam, N.W. Burma, N., S.E. & Peninsular
Thailand, Tonkin, and in Malesia: Malaya, Sumatra
(Gajo Lands, Angkola), Java, Lesser Sunda Is. (Lom-
bok, Sumbawa).

Ecol. Moist evergreen mid-mountain forests, at
800-1300 m in Malaya and W. Java but at lower
altitudes in seasonal areas, and as low as 100 m at
Lakhimpur, Assam and Lombok; locally common or
semi-gregarious.

Vern. Kéruing gunong (Malaya), pelahlar, palaglar
(Java).

Notes. The distribution, persistence and density of
the tomentum is very variable, even within a single
population, in E. Java and Burma; in Assam, N.
Burma and N. Thailand even the leaves are typically

1982]

persistently pubescent beneath, while in Malaya it is
uniformly glabrescent but for the buds and corolla.

The species, as here understood, has an interesting
distribution: at the extremities of its range in mainland
Asia and in the Lesser Sunda Is. (Lombok and
Sumbawa) it occurs in a seasonal climate down almost
to sea level and is relatively tomentose. In the everwet
tropics of Malaya and W. Java it is confined to
altitudes above 800 m and in the former is uniformly
glabrescent but for the buds and corolla.

D. littoralis is clearly a segregate from the wide-
spread D. retusus, endemic to the lowland forest of
Nusa Kambangan I., adjacent to the coast of S.
Central Java. I maintain it as a species. It would
appear that the ecological and geographical distribu-
tion, and diversification, of the two must be explained
in terms of regional Pleistocene history.

15. Dipterocarpus littoralis BL. Bijdr. (1825) 224: Fl.
Jav. 2 (1829) 17, t. 4; KorTH. Kruidk. (1841) 62;
WALP. Rep. 5 (1845) 122; Mia. Fl. Ind. Bat. 1, 2 (1859)
496; Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC.
Prod. 16, 2 (1868) 609; Dyer, J. Bot. 12 (1874) 102;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 198; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 30; K. & V. Bijdr. 5 (1900)
114; MoLL & JANssonius, Mikrogr. Holz (1906) 348;
Koorp. Exk. Fl. Java 2 (1912) 621; SLoor. Bull. Jard.
Bot. Btzg III, 8 (1927) 296, f. 2; BACKER & BAKH. f. FI.
Java 1 (1963) 329.

Young parts + densely pale ferrugineous pubes-
cent, persistent only on leaf bud, stipule outside and
ovary, becoming sparse though subpersistent on
petiole and midrib above. Twigs 8-11 mm g, stout,
terete, rough, with prominent large petiole scars. Bud
c. 20 by 15 mm, ovoid-lanceolate. Stipule to 15 by 4
cm, lanceolate, acute, caducous. Leaves clustered
round the twig apices, 16—25(—S2 in young trees) by
10—18(—28) cm, broadly ovate, thinly coriaceous, +
prominently persistently plicate; base obtuse or cor-
date; apex shortly acuminate; nerves 19-24 pairs,
straight, at 60°-70°, prominent beneath; tertiary
nerves laxly scalariform, slender but distinctly ele-
vated beneath; midrib prominent beneath, applanate
or somewhat elevated above; petiole 3—9(—12) cm
long, prominently geniculate. /nflerescences to 20 cm
long, glabrous, axillary and crowded round the twig
apices, lax, compressed, bearing to 3 flowers. Flower
buds to 35 by 10 mm, prominently pedicellate.
Stamens c. 30; filaments tapering, anthers narrowly
tapering, auriculate, pubescent; appendages shorter
than anthers. Fruit pedicel to 5 by 4 mm, prominent;
calyx tube to 3.5 by 3 cm, obturbinate, smooth; 2
longer lobes to 24 by 4 cm, lorate, narrowly obtuse or
subacute, c. 7 mm broad and somewhat revolute at
base; 3 shorter lobes to 10 by 6 mm, obtuse, with
revolute margins.

Distr. Malesia: Central S. Java (Nusa Kambangan
I. in Banjumas Res.).

Ecol. Apparently common in the mixed lowland
rain forest.

Vern. Lalar, pélahlar, kalahlar (Jav.).

DIPTEROCARPACEAE (Ashton)

309

Note. Apparently a lowland segregate of the
widespread submontane D. retusus. A collection at
Leiden (DE VRIESE s.n. sub n. 902. 146-236) is presum-
ably wrongly annotated as from Sumatra.

16. Dipterocarpus kunstleri KING, J. R. As. Soc. Beng.
Sc. 62, 2 (1893) 96; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 37; BRUHL & KING, Ann. R. Bot. Gard. Cale. 5,
2 (1896) 148, t. 180; Rip. Fl. Mal. Pen. 1 (1922) 217:
HEYNE, Nutt. Pl. ed. 2 (1927) 1094, 1096; SLoor. Bull.
Jard. Bot. Btzg III, 8 (1927) 327; Foxw. Mal. For.
Rec. 10 (1932) 83; Burk. Dict. (1935) 844; Sym. Mal.
For. Rec. 16 (1943) 182, f. 85; ASHTON, Gard. Bull.
Sing. 31 (1978) 10. — D. speciosus BRANbIs, J. Linn.
Soc. Bot. 31 (1895) 38; PERK. Fragm. FI. Philip. (1904)
22; Foxw. Philip. J. Sc. 6 (1911) Bot. 250, pl. 37; ibid.
13 (1918) Bot. 178; ibid. 67 (1938) 261; MERR. En.
Philip. 3 (1923) 90; SLoor. Bull. Jard. Bot. Btzg III, 8
(1927) 265. — D. exalatus SLOOoT. ex Woop, Gard.
Bull. Sing. 17 (1960) 486; Reinwardtia 5 (1961) 462;
ASHTON, Man. Dipt. Brun. (1964) 31, f. 6; ibid. Suppl.
(1968) 14; MEER & Woop, Sabah For. Rec. 5 (1964)
246, pl. 25b. — Fig. 17D.

Twig, lamina and petiole glabrous; leaf nervation
puberulent or glabrous beneath; leaf bud and stipule
outside persistently densely minutely pale grey
adpressed puberulent, appearing grey lepidote. Twig
to 5 mm @ apically, terete or slightly ribbed and
compressed; stipule scars prominent. Bud 12-15 by
2-3.5 mm, narrowly falcate, acute. Stipule to 7 by 0.8
cm, linear. Leaves 13-22 by 7-10 cm, elliptic to
broadly lanceolate, base cuneate; apex shortly acu-
minate (more prominently acuminate in young trees);
nerves 16-18 pairs, prominent beneath, at c. 40° to
50°; petiole 2-3 cm long. Raceme to 22 cm long,
terminal or axillary, slender, glabrous, somewhat
compressed, singly branched. Flower bud to 3.5 by 1.5
cm. Calyx and corolla typical, calyx glabrous.
Stamens c. 30, somewhat longer than the style;
filaments short, anthers linear, tapering; appendage to
connective as long as anther, slender. Ovary ovoid-
conical, shortly densely pubescent; stylopodium and
style about 5 times as long as the ovary, tapering from
the base, shortly tomentose but for apical 4. Fruit
calyx glabrous; tube to 5 by 2.5 cm, ellipsoid, tapering
gradually to the base and to the strongly constricted c.
1 cm @ neck; 5-ribbed or almost winged, the ribs c. 7
mm thick and 4 mm broad apically, either confined to
the apical half or continuing to the base as shallow
rounded ridges, terminating + abruptly distally as
obtuse tubercles; 2 longer lobes to 11 by 1.5 cm, of
very variable length, sometimes no longer than
shorter lobes, lanceolate, coriaceous, obtuse, tapering
abruptly to the c. 5 mm broad base, 3-nerved, nerves
indistinct; shorter lobes to 6 by 5mm, small, recurved,
somewhat thickened.

Distr. Malesia: Malaya (excluding seasonal area),
Sumatra (Atjeh, Tapanuli, Palembang, in north, west
and south-east; Simalur, Marsala, Banka), Borneo(S.E.
Borneo, Sabah, Brunei, Sarawak N.E. of Rejang),
Philippines (Luzon, Polillo, Negros, Samar, Basilan).

310

Ecol. Widespread, locally common, on undulating
or flat land, especially near streams.

Vern. Kéruing gombang, k.g. merah (Mal.), k. batu,
k. jombor, k. minyak, lagan laweh, 1. daun lebar
(Sum.), k. rapak, k. simpor, k. kuntum puteh, karang,
binawan, kambalong, tempudau, isak, tabuloh, karup
(Borneo).

Note. In N.E. and E. Borneo the calyx tube is +
obtusely ribbed and tuberculate distally, and the
wings often (not always) short or even rudimentary.
In the Philippines the species becomes very variable; in
some collections the fruit calyx tube is continuously
ribbed from base to apex and even winged suggesting
hybridisation with D. grandiflorus (BLCO) BL.; in
others the twigs and petioles are densely fulvous pilose
as in D. elongatus KorTH., which grows with it there.

Individual trees observed in the field in Borneo are
rather constant in the length of the calyx lobes, but
geographical variation appears continuous and to
some extent clinal, the lobes becoming shorter to-
wards the south-eastern limits of its range. I cannot
therefore recognize distinct varieties meriting taxo-
nomic status.

D. speciosus BRANDIS, based on VIDAL 2160 (in
fruit, K) from Luzon, is a specimen with tomentum
typical of D. kunstleri and fruit similar though with
narrower, more continuous ribs. I see no satisfactory
solution other than to unite the two names.

17. Dipterocarpus applanatus SLooT. Bull. Jard. Bot.
Btzg III, 16 (1940) 443, f. 5; MEWER & Woop, Sabah
For. Rec. 5 (1964) 237, f. 34; ASHTON, Man. Dipt.
Brun. Suppl. (1968) 11, f. 2, pl. 1 (stem-base).

Twig, base of inflorescence and inside of stipule
shortly evenly pale buff pubescent, base of petiole
caducously so; leaf bud and stipule outside longer
tomentose, longest towards apices. Twig S—11 mm @,
stout, pale with prominent slightly depressed dark
stipule scars. Bud 4-15 by 10-27 mm, broadly ellip-
soid to falcate, acute or obtuse. Stipule to 5 by 2 cm,
oblong, obtuse, concave. Leaves 12—30 by 9-20 cm,
usually large, + broadly elliptic, coriaceous but
undulate; base obtuse to subcordate; acumen short,
abruptly acuminate; nerves | 1—15 pairs, prominent, at
45°—70°; tertiary nerves remote, subreticulate; petiole
3.5—6 cm long, to 4mm @. Flowers unknown. Raceme
to 2 cm long, terete but rugose on drying, glabrescent,
singly branched or bifurcating, bearing distichous
fruit at c. 3 cm intervals. Fruit pedicel to 3 mm long,
tapering from base of calyx tube; calyx tube to 5 by 4.5
cm, glabrous, ovoid, bearing 5, to 8 mm wide,
prominent sharp-edged ribs; ribs widest distally and
there frequently undulate, sometimes absent basally; 2
longer calyx lobes to 19 by 4.5 cm, glabrous, lorate,
obtuse, tapering abruptly at the somewhat revolute
base; 3 shorter calyx lobes to 1 by 1 cm, suborbicular,
completely revolute.

Distr. Malesia: Borneo (W. Sarawak, E. Sabah,
Tidung and Bulungan).

Ecol. Locally common on sandy soils, especially in
valleys and on flat land near coasts.

FLORA MALESIANA

(ser. I, vol. 9?

Vern. Kéruing arong, k. daun besar.

Note. Apparently a segregate from D. kunstleri; it
may have arisen more than once from the parental
species.

18. Dipterocarpus rigidus RID. J. Str. Br. R. As. Soc.
82 (1920) 171; Fl. Mal. Pen. 1 (1922) 217; SLoor. Bull.
Jard. Bot. Btzg III, 8 (1927) 347; Reinwardtia 5 (1961)
463; Foxw. Mal. For. Rec. 10 (1932) 76, pl. 5; BURK.
Dict. (1935) 845; Descu, Mal. For. Rec. 14 (1941) 64;
Sym. Mal. For. Rec. 16 (1943) 187, f. 85; BROWNE,
For. Trees Sarawak & Brunei (1955) 110; ASHTON,
Man. Dipt. Brun. Suppl. (1968) 16, f. 3, pl. 2 (habit &
bark).

Twig, petiole, midrib above and leaf beneath
shortly densely evenly pale ocherous pubescent, leaf
above fugaceously so; leaf bud and stipule outside
(glabrous within in mature tree) longer tomentose.
Twig to 8 mm @ apically, stout, terete, with slender
stipule scars. Bud to 15 by 8 mm, conical, subacute.
Stipule to 7 by 1 cm, lorate, subacute. Leaves 13—25 by
8-16 cm, ovate, thickly coriaceous, with broadly
cuneate base; acumen to | cm long; nerves 12-16
pairs, at 40°-S0°; nerves and midrib prominent
beneath, depressed above; tertiary nerves scalariform,
evident beneath; petiole 3-6 cm long, geniculate.
Raceme to 15 cm long, terete, axillary, at first shortly
evenly ocherous pubescent towards the base, glabres-
cent; unbranched or rarely singly branched, bearing
distichous flowers at up to 3 cm intervals. Flower bud
to 3.5 by 1 cm. Calyx and corolla typical; calyx
glabrous. Stamens 24, slightly shorter than the style;
filaments narrowly lorate, slightly tapering, exceeding
length of anthers; anthers linear, tapering; appendage
acicular, c. 5 length of the anther. Ovary ovoid,
pubescent; style columnar, pubescent except for the
apical 4. Fruit glabrous. Pedicel 1 mm long, stout.
Calyx tube to 5 by 4.5 cm, subglobose, with 5 tubercles
(obscure at maturity) in the distal half; 2 longer calyx
lobes to 18 by 5cm, lanceolate to lorate, subacute, to 6
mm broad above the somewhat revolute base; 3
shorter lobes to 8 by 8 mm, suborbicular, revolute.

Distr Malesia: E. Malaya (Trengganu south-
wards), Sumatra (Riouw Arch., Singkep, Lingga),
Anambas Is., Borneo (Sarawak N.E. to Bintulu).

Ecol. Locally abundant on dry sandy soils on
coastal hills.

Vern. Kéruing chogan, k. utap, k. sungkit, k. pakat,
k. kélawar, k. kéluang, k. mérah, k. lékit, k. daun lebar,
k. d. panjang (Mal.).

Note. Sometimes difficult to distinguish from D.
costulatus and D. globosus, both of which can be more
or less tomentose when immature.

19. Dipterocarpus costulatus SLOoT. Bull. Jard. Bot.
Btzg III, 8 (1927) 315, f. 7; ibid. III, 17 (1941) 105;
Foxw. Mal. For. Rec. 10 (1932) 79; Sym. Mal. For.
Rec. 16 (1943) 174, f. 85; BROWNE, For. Trees
Sarawak & Brunei (1955) 108; MEVER & Woop,
Sabah For. Rec. 5 (1964) 244, pl. 25a (seedlings);
ASHTON, Man. Dipt. Brun. Suppl. (1968) 12, f. 2.

1982]

DIPTEROCARPACEAE (Ashton)

3

Vegetative parts of mature tree glabrous but for the
densely shortly buff pubescent inner surface of bud-
scales and stipules. Twigs and buds frequently
pruinose. 7wig to 7 mm @ towards the apex, some-
what compressed at first, becoming terete, smooth;
stipule scars slender, prominent. Bud to 30 by 6 mm,
falcate, acute, slender. Stipule to 16 by 2 cm, lorate,
subacute. Leaves 12-20 by 7-14 cm, broadly elliptic to
ovate, thickly coriaceous, with prominently cuneate
base, obtuse to shortly acuminate, persistently folded
between the 11-14 pairs of prominent nerves; tertiary
nerves slender, scalariform; petiole 3-6 cm long.
Raceme to 20 cm long, terete, glabrous, bearing
distichous flowers at to 4 cm intervals. Flower buds to
3 by 1.5 cm. Calyx and corolla typical, calyx glabrous.
Stamens c. 24, slightly shorter than the style; filaments
about 5 length of anther, compressed, tapering; anther
linear, tapering apically into the acicular appendage;
appendage somewhat less than 5 length of anther.
Ovary ovoid, densely pubescent; style columnar,
pubescent except in the apical 4. Fruit pedicel to 6mm
long, prominent. Calyx glabrous; tube to 1.5 by 2 cm,
subglobose, bearing 5 prominent flange-like tubercles
medially; 2 larger calyx lobes to 20 by 4 cm, broadly
lorate, obtuse, with to 6 mm wide revolute constric-
tion at the base; 3 shorter lobes to 7 by 6 cm,
suborbicular, obtuse, revolute.

Distr. Malesia: Malaya (excepting seasonal areas),
E. Sumatra (E. Atjeh, Langkat, Indragiri, P. Lingga),
west and north-east Borneo (Lower Kapuas, W.
Sarawak; Tawau Distr., Nunukan and Tidung).

Ecol. Local, on poor soils on flat and undulating
land, rarely to 600 m.

Vern. Kéruing paya, k. kipas, k. pékat, k. marak-
luang, k. ladan.

20. Dipterocarpus globosus VEsQuE, C. R. Ac. Sc.
Paris 78 (March 1874) 627; J. Bot. 12 (1874) 151;
Dyer, J. Bot. 12 (1874) 153; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 200; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 33; Mere. En. Born. (1921) 398; SLoor. Bull.
Jard. Bot. Btzg III, 8 (1927) 304; ibid. III, 17 (1941) 98;
BROWNE, For. Trees Sarawak & Brunei (1955) 109;
ASHTON, Man. Dipt. Brun. (1964) 33, f. 6-7, pl. 5
(stem); ibid. Suppl. (1968) 15. — D. beccarianus VEs-
QUE, C. R. Ac. Sc. Paris 78 (1874) 627; J. Bot. 12 (1874)
151; Dyer, J. Bot. 12 (1874) 153; Burck, Ann. Jard.
Bot. Btzg 6 (1887) 200. — D. beccarii Dyer, J. Bot. 12
(April 1874) 103, t. 144, f. 16; ibid. (May 1874) 153,
incl. var. glabrata.

Midrib and nerves below fugaceous pubescent;
young twig, raceme and petiole + persistently shortly
chestnut pubescent; leaf bud and stipule outside
densely persistently long chestnut tomentose. Twig
5-8 mm @ apically, stout, terete. Bud 1-2 by 0.8-1.2
em, ovoid, acute. Stipule c. 7 by 0.7 mm, narrowly
lanceolate, acute. Leaves 10-14 by 7-9 cm, thickly
coriaceous, broadly ovate, with sinuate margin;
prominently plicately folded between the nerves; base
broadly cuneate; acumen c. 4 mm long, broad; nerves
12-14 pairs, well spaced, prominent beneath, some-

what oblique (20°—30° apically and 50°-60° basally);
petiole 2-2.5 cm long, stout. Flowers unknown.
Raceme to 8 cm long, stout, axillary, singly branched.
Fruit calyx tube 2.5—3.5 cm long and broad, + ovoid,
glabrous, sometimes unribbed, usually + 5-ribbed
from the obtuse and often impressed base to the
constricted c. 8 mm @ neck; ribs obtuse or acute, most
prominent, slightly tuberculate, distally; 2 longer
lobes 11-15 by 2.5-3.5 cm, coriaceous, slightly
twisted, oblong, + broader in the distal half, obtuse,
gradually tapering to the flat c. 6 mm broad base,
glabrous, with three subequal closely parallel nerves,
prominent on the inner surface; 3 shorter lobes 4—5
mm long and broad, small, deltoid, + recurved,
revolute.

Distr. Malesia: Borneo (Sarawak, Brunei, S.W.
Sabah).

Ecol. Locally abundant on leached sandy soils in

Mixed Dipterocarp forest on coastal hills below 400
m.
Vern. Kéruing buah bulat.
Note. Differing principally from D. rigidus in the
absence of tomentum at maturity (also minor fruit
differences); in this bearing the same relationship as D.
humeratus to D. cornutus.

21. Dipterocarpus humeratus SLooT. Bull. Jard. Bot.
Btzg III, 8 (1927) 308, 311, f. 4; THORENAAR, Med.
Proefst. Boschw. 16 (1926) 108; AsHTon, Gard. Bull.
Sing. 20 (1963) 237; Man. Dipt. Brun. (1964) 36, f. 6,
pl. 12-13 (stem, bark); ibid. Suppl. (1968) 15; MEUER
& Woop, Sabah For. Rec. 5 (1964) 257, f. 39. — D.
validus (non BL.) BRANDIS, J. Linn. Soc. Bot. 31 (1895)
37, p.p., quoad specim. FORBES. — D. retusus (non BL.)
RIDL. Kew Bull. (1926) 60, p.p. — D. gibbosus SLooT.
Bull. Jard. Bot. Btzg III, 8 (1927) 311, f. 5.—D.
ursinus SLOOT. Reinwardtia 5 (1961) 470, f. 5.
Young twig, petiole, midrib and nerves beneath
persistently shortly cream pubescent; leaf bud and
stipule outside long fulvous tomentose, the hair tufts
becoming separated widely and partially caducous on
the expanding stipule. Twig 1-1.2 cm @ apically,
becoming uneven and thinly flaky, terete. Bud to 5 by
1.5 cm, narrowly conical, obtuse. Stipule to 10 by 2
cm, narrowly lanceolate, obtuse. Leaves 20-38 by
12-23 cm, broadly ovate; base obtuse; apex obtuse to
shortly acuminate; nerves c. 20 pairs, prominent, well
spaced, at 45°—60°; petiole 4-6 cm long, stout, promi-
nently geniculate. Raceme to 21 cm long, terminal or
axillary, terete or slightly compressed, long golden-
brown caducous tomentose at base, more shortly so
distally; usually unbranched. Flowers remote, disti-
chous; bud to 5 by 2 cm, large. Calyx and corolla
typical, calyx glabrous. Stamens c. 40, longer than
style; filaments c. 4 as long as anthers; anthers linear,
tapering; appendage to connective as long as anther or
slightly longer, slender. Ovary ovoid-conical, shortly
pubescent; stylopodium 2-3 times as long as ovary,
shortly pubescent the apex excepted. Fruit subsessile.
Calyx tube to 4 by 3.5 cm, globose, glabrous, with 5
obtuse tubercles in the apical half, terminating

312

abruptly at the c. 2.5 cm @ neck; 2 longer calyx lobes
to 18 by Scm, oblong, obtuse, tapering abruptly at the
to 1.2 cm broad revolute base, untwisted; 3 shorter
lobes to 1.5 cm long and broad, broadly ovate, base
subcordate, apex obtuse, revolute with the 2 halves
back to back.

Distr. Malesia: North and East Sumatra (Langsa;
Singkel, Tapanuli, E. coast, Palembang, Mentawai
Is.), Borneo (S.E. Kalimantan, Sabah, N.E. Sarawak,
Brunel).

Ecol. Local, on undulating land and ridges below
700 m.

Vern. Kéruing kerukup, k. latek bukit (Mal.).

Note. Clearly closely allied to D. cornutus, from
which it differs principally in the tomentum and
smooth fruit calyx.

22. Dipterocarpus cornutus Dyer, FI. Br. Ind. 1 (1874)
296; J. Bot. 12 (1874) 103, t. 143, f. 15; KING, J. R. As.
Soc. Beng. Sc. 62, 2 (1893) 93; BRANDIS, J. Linn. Soc.
Bot. (1895) 32; Ript. Fl. Mal. Pen. 1 (1922) 215;
SLooT. Bull. Jard. Bot. Btzg III, 8 (1927) 306; HEYNE,
Nutt. Pl. ed. 2 (1927) 727; Foxw. Mal. For. Rec. 10
(1932) 75; BurK. Dict. (1935) 842; Sym. Mal. For.
Rec. 16 (1943) 172, f. 85, 87, 88. — Fig. 17A.

Large pale-barked tree. Twigs, leaf buds, stipule
outside, petiole, leaf beneath including nervation,
midrib above, calyx, corolla outside and ovary apex
densely persistently pale ocherous-cream puberulent,
sometimes fulvous on twigs; fruit calyx glabrescent;
leaf bud, stipule and base of inflorescence frequently
also densely + caducously long rufous tufted tomen-
tose, the short even indumentum persisting after the
tufts drop. Twig c. 10 by 8 mm, stout, somewhat
compressed, pale brown, sometimes becoming papery
flaky. Bud to 6 by | cm, lanceolate, acute; stipule to 20
by 4 cm, lorate, narrowly obtuse. Leaves 15-30 by
7.5-18 cm, broadly elliptic to oblong-ovate, thickly
coriaceous, cream beneath, prominently persistently
plicate; base obtuse; apex obtuse or subretuse; nerves
18-21 pairs, prominent beneath; tertiary nerves very
slender, hardly elevated, densely scalariform; petiole
5-8.5cm long, c.4mm g, long, stout. /nflorescences to
28 cm long, to 5 by 3mm gat base, axillary, somewhat
compressed, unbranched, bearing to 10 distichous
flowers. Flower bud to 3.5 by 1 cm, fusiform. Stamens
30, exceeding style at anthesis; filaments short, broad,
applanate, deltoid; anthers c. 5 times length of
filaments, broadly linear-lanceolate, tapering; appen-
dage stoutly acicular, tapering, c. 5 length of anther.
Ovary ovoid, small, densely hirsute as also the basal 5
of the stoutly columnar + capitate style. Fruit shortly
pedicellate to subsessile; calyx tube to 2.5 cm long, to
3.7 cm g including the 5 prominent distal tubercles,
subglobose, densely verrucose lenticellate; 2 longer
lobes to 21 by 5 cm, oblong, narrowly obtuse,
abruptly constricted to c. 2 cm wide and revolute at
base; 3 shorter lobes to 15 by 15 cm, suborbicular,
revolute.

Distr. Malesia: Malaya, N. Sumatra (Atjeh, Lang-
kat), Borneo (S.E., from P. Laut to W. and E. Kutei).

FLORA MALESIANA

[ser. I, vol. 9?

Ecol. Lowland forest to 1000 m, on well drained
flat or undulating land or hills; frequently common or
semigregarious.

Vern. Kéruing gombang, k. dadek, k. (mara)kéluw-
ang, k. chaier, k. babi (Malaya), kéruing, tampudau,
tampurau, kapenkaluang (Borneo).

23. Dipterocarpus elongatus KorTH. Kruidk. (1841)
62; WALP. Rep. 5 (1845) 123; BL. Mus. Bot. Lugd.-
Bat. 2 (1852) 36; Mig. FI. Ind. Bat. 1, 2 (1859) 498;
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 83, 85; DC. Prod.
16, 2 (1868) 613; Dyer, J. Bot. 12 (1874) 108; BuRcK,
Ann. Jard. Bot. Btzg 6 (1887) 203; BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 32, 40; MeRR. En Born. (1921) 398;
SLoot. Bull. Jard. Bot. Btzg III, 8 (1927) 272;
Reinwardtia 5 (1961) 473, 474; ASHTON, Gard. Bull.
Sing. 20 (1963) 237. — D. apterus Foxw. Mal. For.
Rec. 10 (1932) 77, pl. 6; Burk. Dict. (1935) 842;
SLooT. Bull. Jard. Bot. Btzg III, 16 (1940) 439, f. 3;
Sym. Mal. For. Rec. 16 (1943) 167, f. 85; BROWNE,
For. Trees Sarawak & Brunei (1955) 107; ASHTON,
Man. Dipt. Brun. (1964) 23, f. 6; ibid. Suppl. (1968)
11. — Fig. 17F.

Twigs, leaf bud, stipule outside and petiole with +
dense, + caducous, very long red-brown tufted
tomentum; leaf nervation below more sparsely so or
glabrous. Twigs to 1.5 cm @ apically, terete, red-
brown, becoming cracked and thinly flaked. Bud to 6
by 1.5 cm, falcate, acute, glabrescent or remaining
tufted as the stipule expands. Stipule to 15 by 2.5 cm,
hastate, acute. Leaves 28-50 by 13-20 cm, elliptic,
coriaceous; base obtuse; apex shortly abruptly acu-
minate; nerves 25-38 pairs, prominent beneath, the
lamina persistently plicately folded between; tertiary
nerves distant, scalariform; petiole S—7 cm long, 0.5
cm @, stout. Flowers unknown. Raceme to 12 cm long,
terminal or axillary, rigid, rarely branched, red-brown
long tufted tomentose. Fruit calyx tube to 5 by 5.5cm,
at first tomentose, glabrescent, at first obovoid,
becoming globose, with 5 obtuse distal tubercles,
constricted to c. 1.5 cm @ at neck; lobes equal,
vestigial, to 8 mm long, becoming recurved, obtuse.

Distr. Malesia: Malaya (E. coast: Kelantan to
Central and E. Johore and Singapore), eastern Suma-
tra (Serdang and Langkat Distr.), Lingga Arch.,
Anambas Is. (S. China Sea), Borneo (Sarawak N.E. of
Rejang, Kapuas valley, S.E. Kalimantan).

Ecol. Fresh water swamp, usually on sandy, perio-
dically well drained, soil; locally common in primary
and secondary forest.

Vern. Kéruing latek, latek, k. babi (Brun.), k.
gumbang (Mal.), k. pasir (Sum.).

Note. Closely allied to the previous two species and
differing from D. humeratus principally in the fruit.

24. Dipterocarpus lamellatus Hook. f Trans. Linn.
Soc. 23 (1860) 159; Mia. Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 85; DC. Prod. 16, 2 (1868) 611; WaLp. Ann. 7
(1869) 377; Dyer, J. Bot. 12 (1874) 107, t. 145, f. 22;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 202; BRANDIS &
Gitc in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 256;

1982]

BRANDIS, J. Linn. Soc. Bot. 31 (1895) 39; MErR. En.
Born. (1921) 399; Gite in E. & P. Pfl. Fam. ed. 2, 21
(1925) 251; SLoor. Bull. Jard. Bot. Btzg III, 8 (1927)
347; MEVER & Woop, Sabah For. Rec. 5 (1964) 261, f.
41.

Large tree. Twigs, leaf buds, stipule, petioles, leaf
nervation beneath, midrib above, petiole and inflores-
cences densely persistently pale yellow-brown hirsute;
leaf surfaces and fruit calyx sparsely so. Twig c.4mm
@ apically, terete, + rugose. Leaf bud to 9 by 7 mm,
ovoid, acute; stipules unknown. Leaves 13-16 by 6-9
cm, elliptic, thinly coriaceous, + boat shaped with the
lower surface concave; base narrowly obtuse; acumen
to 8 mm long, abrupt, slender; nerves 15—17 pairs,
straight, ascending at c. 45°, slender but prominent
beneath, shallowly depressed above as also the mid-
rib; tertiary nerves laxly scalariform, slender but
prominent beneath, evident above; petiole 34 cm
long, c. 2 mm @g, slender. Inflorescence to 6 cm long,
axillary, apparently unbranched, bearing to 3 fruit
(flower unknown). Fruit pedicel to 4mm long, slender;
calyx tube to 1.8 cm g including the 5 densely
convoluted wings, subglobose; 2 longer lobes to 14 by
2.5 cm, spatulate, subacute, c. 8 mm broad at base; 3
shorter lobes to 14 by 7 mm, ovate, + revolute.

Distr. Malesia: Borneo (Sabah: Labuan (extinct),
Beaufort).

Ecol. Rare in Mixed Dipterocarp forest on low hills
near coast.

Vern. Kéruing jarang.

25. Dipterocarpus lowii Hook. f Trans. Linn. Soc. 23
(1860) 160; Mia. Ann. Mus. Bot. Lugd.-Bat. 3 (1867)
58; DC. Prod. 16, 2 (1868) 613; WaALP. Ann. 7 (1869)
377; Dyer, J. Bot. 12 (1874) 107, t. 145, f. 23; BuRcK,
Ann. Jard. Bot. Btzg 6 (1887) 202; BRANpis, J. Linn.
Soc. Bot. 31 (1895) 40; Merr. En. Born. (1921) 399;
Rpt. Fl. Mal. Pen. Suppl. (1925) 291; SLoor. Bull.
Jard. Bot. Btzg III, 8 (1927) 344, f. 14-15; in Merr. PI.
Elm. Born. (1929) 201; Foxw. Mal. For. Rec. 3 (1927)
43; ibid. 10 (1932) 91; Sym. Mal. For. Rec. 16 (1943)
183, f. 85; BROWNE, For. Trees Sarawak & Brunei
(1955) 110; ASHTON, Man. Dipt. Brun. (1964) 37, f. 6,
pl. 14 (stem, habit); ibid. Suppl. (1968) 15; MEVER &
Woop, Sabah For. Rec. 5 (1964) 261, f. 3b. — D.
undulatus VESQUE, C. R. Ac. Sc. Paris 78 (1874) 626; J.
Bot. 12 (1874) 151; Dyer, J. Bot. 12 (1874)
153. — Fig. 25.

Twig, midrib and nerves below + densely +
caducously shortly pale golden-yellow pubescent.
Twig 2-3 by 5-10 mm g, compressed, with broad
swollen stipule scars. Leaf bud c. 1.5 by 0.9 cm,
broadly conical, frequently falcate, obtuse. Stipule to
4 by 3 cm, broadly deltoid, acute. Leaves 15-20 by
6-10 cm, ovate-lanceolate, thickly coriaceous,
strongly folded between the 15—20 pairs of prominent
nerves; margin revolute; base obtuse or cordate; apex
obtuse or with to 6 mm long narrow acumen; petiole
1.5-3 cm long, stout, often wrinkled on drying.
Raceme to 3cm long, short, stout, terminal or axillary,
terete, simple or singly branched, flowers distichous;

DIPTEROCARPACEAE (Ashton)

313

bracteoles to 15 by 10 mm, elliptic to ovate, subacute,
densely shortly pubescent outside, puberulent within,
fugaceous. Flower bud to 4 by 1.2 cm. Calyx densely
golden-yellow pubescent, wings prominently undu-
late. Corolla typical. Stamens c. 30, shorter than the
style; filaments short; anthers narrowly oblong;
appendage to connective somewhat shorter than
anther, stout at base, tapering. Ovary ovoid-conical,
tapering, densely pubescent; stylopodium narrowly
cylindrical, pubescent, tapering to the somewhat
shorter glabrous filiform style. Fruit calyx glabres-
cent, tube c. 4 cm long and g (including wings),
globose, completely hidden by the intricate folds of
the 5 wings; 2 longer calyx lobes to 14 by 3.5 cm,
broadly oblong, obtuse, somewhat revolute, hardly
constricted at the base, concurrent with the wings of
the tube, with one long median nerve and two shorter
laterals; 3 shorter lobes 1.5—2 by 2 cm, broadly ovate,
recurved and concurrent with the wings.

Distr. Malesia: Malaya (Perak, E. coast), P. Sing-
kep, eastern Sumatra (Kuantan Distr., Langsa, Lang-
kat, Batu Is.), Borneo (Kapuas valley, Sarawak to
Sabah; Tidung).

Ecol. Well-drained leached, usually sandy soils, in
lowlands, especially on islands and near present and
former coastlines on low hills to 400 m; locally
common on ultrabasics in Sabah.

Vern. Kéruing sol, k. hijau, k. daun panjang, k. batu,
k. sindor (Mal.), bajan (Sabah), kKawaan kaput (W.
Borneo).

Note. Specimens from Malaya north of Johore and
from Sumatra and Singkep bear smaller leaves and
more slender twigs and petioles.

26. Dipterocarpus pachyphyllus Meer, Acta Bot.
Neerl. 12 (1963) 351, pl. 15; MEER & Woop, Sabah
For. Rec. 5 (1964) 265; ASHTON, Man. Dipt. Brun.
(1964) 41, f. 6; ibid. Suppl. (1968) 16. — Fig. 17E.
Freshly opened young parts evenly shortly pale
tawny tomentose, fugaceous on all but bud, stipule
and raceme. Twig to 4 by 2.5 cm @ towards apex,
compressed, smooth, with prominent raised stipule
scars. Bud c. 18 by 4mm, linear, acute. Stipule to 4 by
1.2 cm, linear to deltoid, subacute. Leaves 9-17 by 5-9
cm, broadly ovate, thickly coriaceous; base obtuse to
subcordate; acumen to | cm long, broad; nerves 10—12
pairs, distant, prominent, straight but curving
abruptly at the margin and coalescing to form an
indistinct intramarginal nerve; petiole 2-3.4 cm long,
slender, prominently geniculate. Raceme to 5 cm long,
terminal or axillary, terete, simple, flowers distichous.
Flower bud to 15 by 5 mm. Calyx densely golden-
tawny pubescent; wings prominently undulate. Cor-
olla typical. Stamens 23-25, shorter than style; fila-
ments as long as anthers, applanate at base, tapering;
anthers narrowly oblong; appendage to connective
somewhat shorter than anther, stout at base, tapering.
Ovary narrowly ovoid, densely pubescent; stylopo-
dium indistinct, tapering into style; style filiform,
glabrous except at base, somewhat shorter than ovary
and stylopodium. Fruit calyx entirely glabrous,

314 FLORA MALESIANA [ser. I, voleo?

Fig. 25. Dipterocarpus lowii Hook. f. (2 trees) as emergent in undisturbed profile of lowland dipterocarp forest,
near Bt. Puan, Belait, Brunei, c. 20 m alt.; pineapple cultivation in foreground (Photogr. ASHTON, Jan. 1960).

1982]

pruinose; tube to 2.5 by 2 cm including wings,
subglobose, completely hidden by the intricate folds
of the 5S wings; 2 longer lobes to 13 by 3 cm, spatulate,
subacute, with one long median nerve and two short
laterals; 3 shorter lobes to 1 by 1.5 cm, broadly
deltoid, recurved.

Distr. Malesia: Northern Borneo (Sabah, Brunei,
Sarawak E. of the Lupar).

Ecol. Scattered on leached clay soils in Mixed
Dipterocarp forest, undulating land and hillsides to
400 m.

Vern. Kéruing daun tebal, kéruing sol padi.

27. Dipterocarpus confertus SLoor. Bull. Jard. Bot.
Btzg III, 8 (1927) 322, f. 9; in Merr. Pl. Elm. Born.
(1929) 201; Bull. Bot. Gard. Btzg III, 17 (1941) 104; f.
14; Reinwardtia 5 (1961) 62, f. 1; Foxw. Mal. For.
Rec. 10 (1932) 62; BRowne, For. Trees Sarawak &
Brunei (1955) 108; ASHTON, Man. Dipt. Brun. (1964)
26, f. 6; ibid. Suppl. (1968) 12; MEWER & Woop, Sabah
For. Rec. 5 (1964) 242, f. 3a, pl. 26a (stem).

Young twig, leaf bud, stipule outside, petiole,
nerves and midrib above, nervation beneath, and
inflorescence persistently 1.5—3.5 mm long pale ful-
vous-brown tufted hispid; leaf surface caducously so.
Twig 0.8—1 cm g apically, stout, uneven, cracked and
marked by the thin amplexicaul stipule scars. Bud
1.2—2 by 1—1.7 cm, broadly ovoid, obtuse or subacute.
Stipule to 5cm long and broad, broadly ovate, obtuse,
caducous. Leaves (18—)22-35 by (14-)16-22 cm,
broadly obovate to orbicular, chartaceous, concave;
base obtuse or subpeltate, the lamina continuing as a
slight ridge above the base of the midrib; apex obtuse
or shortly acuminate; nerves 9-12 pairs, well spaced,
at 45° to 60°; petiole S—6 cm long. Inflorescence to 7 cm
long, subcymose, singly branched or unbranched,
short, bearing few distichous flowers; bracts and
bracteoles to 25 by 5 mm, linear, obtuse, sparsely
tomentose, caducous. Bud to 4 by 1 cm. Calyx and
corolla typical, calyx shortly tomentose. Stamens c.
25, as long as style; filaments as long as anther,
applanate at base, tapering and filiform below anther;
anther narrowly oblong, stout, tapering; appendage
almost as long as anther, stout at base, tapering and
filiform at apex. Ovary small, ovoid-conical, densely
pubescent; stylopodium indistinct; style 3 times as
long as ovary, columnar, slender, tomentose except at
apex. Fruit subsessile; calyx tube densely pale fulvous
long tomentose, c. 3 by 1.7 cm, narrowly obovoid,
with 5 indistinct ridges when mature, tapering basally
to the pedicel, only slightly constricted apically; 2 long
lobes to 14 by 3 cm, glabrous within, sparsely
tomentose outside, oblong-lanceolate, obtuse, taper-
ing to the constricted but non-revolute base, promi-
nently 3-nerved; 3 shorter lobes to 1.7 by 0.7 cm,
oblong, recurved distally.

Distr. Malesia: Borneo (S.E. Borneo, Sabah, Sar-
awak and Brunei).

Ecol. Mixed Dipterocarp forests below 800 m,
sometimes common on low hills and undulating
land.

DIPTEROCARPACEAE (Ashton)

315

Vern. Tampudau (S.E. Borneo), kéruing kobis
(Sabah, Sarawak).

Note. Sterile collections from Singkep could be
either this species or D. concavus.

28. Dipterocarpus dyeri PieERRE in Lanessan, PI. Util.
Colon. Fr. (1886) 297; Fl. For. Coch. 3 (1889) t. 216,
217; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 33; PARKER,
Ind. For. Rec. 16 (1931) 13; PARKINSON, Burma For.
Bull. 27 (1932) 15; Foxw. Mal. For. Rec. 10 (1932) 79,
p-p.; BuRK. Dict. (1935) 843; Sym. Mal. For. Rec. 16
(1943) 176, f. 85.

Large pale-barked semi-deciduous tree. Leaf buds,
stipules outside and ovary apex densely persistently
long rufous silky velutinate, twigs, leaf nervation
beneath and midrib above, inflorescence and calyx
caducously + sparsely so. Twig 10-15 mm g apically,
stout, pale brown, rugose, lenticellate; internodes
short, the leaves densely clustered at the ends of the
twigs. Buds to 25 by 15 mm, large, ovoid-lanceolate,
acute; stipule to 15 by 2.5 cm, lanceolate, subacute.
Leaves 16-40 by 7.5—14 cm, large, narrowly ovate to
elliptic, coriaceous; base broadly cuneate to subcor-
date; acumen to 5 mm long, short, broad, tapering;
nerves 24-30 pairs, ascending, slender but prominent
beneath as also the midrib; petiole 4-6 cm long, to 4
mm @g, relatively slender. /nflorescence to 16 cm long,
axillary, straight, borne in dense clusters behind the
leaves, each bearing to 6 distichous flowers. Flower
buds to 5 by 1.3 cm long, fusiform. Stamens c. 30,
exceeding style at anthesis; filaments slender, taper-
ing, + equal to the shortly lorate tapering distally
fimbriate anthers; appendage acicular, somewhat
shorter than anthers; ovary narrowly ovoid, sur-
mounted by a slender columnar style c. 15 its length.
Fruit pedicel to 3 mm long; calyx tube to 4 by 3 cm,
ellipsoid, with 5 narrow ribs running from the apex for
2 its length; 2 longer lobes to 20 by 5.5 cm, oblong-
oblanceolate, obtuse, c. 8 mm wide and somewhat
revolute at base; 3 shorter lobes to 15 by 6 mm, ovate,
subacute, revolute.

Distr. Cochinchina, Cambodia, S.E. and peninsu-
lar Thailand, Mergui, and in Malesia: Malaya (Perlis
and N. Kedah).

Ecol. Semi-evergreen Dipterocarp forest and Schi-
ma-bamboo forests at low elevations in the seasonal
zone, in moist sandy soil in valleys by streams but not
normally alluvium.

Vern. Kéruing daun besar, k. etoi (Malaya).

Note. PARKER observed hybridisation between this
species and D. alatus RoxB. in peninsular Burma.

29. Dipterocarpus fagineus VESQUE, C. R. Ac. Sc.
Paris 78 (March 1874) 625; J. Bot. 12 (1874) 149;
Dyer, J. Bot. 12 (1874) 152; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 200; KiNG, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 94, p.p.; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
33; BurK. J. Str. Br. R. As. Soc. 81 (1920) 51, 53, fig.;
Merr. En. Born. (1921) 398; Rip. Fl. Mal. Pen. 1
(1922) 216, p.p.; SLoot. Bull. Jard. Bot. Btzg III, 8
(1927) 318; Foxw. Mal. For. Rec. 10 (1932) 80; Sym.

316

FLORA MALESIANA

[ser. I, vol. 97

Kew Bull. (1937) 318; Mal. For. Rec. 16 (1943) 177, f.
85; BROWNE, For. Trees Sarawak & Brunei (1955) 108:
ASHTON, Man. Dipt. Brun. Suppl. (1968) 14, f. 2:
Gard. Bull. Sing. 31 (1978) 10.— D. prismaticus
Dyer, J. Bot. 12 (Apr. 1874) 104, t. 144, f. 17; ibid.
(May 1874) 152.— Duvaliella problematica Hem,
Bull. Mens. Soc. Linn. Paris 2 (1891) 1009: Rech.
Dipt. (1892) 72; BRANDIS & GILG in E. & P. Pfl. Fam.
ed. 1, 3, 6 (1895) 263; GiLc in E. & P. Pfl. Fam. ed. 2,
21 (1925) 258. — D. pseudofagineus Foxw. Mal. For.
Rec. 10 (1932) 82; Sym. Mal. For. Rec. 16 (1943) 186,
f. 85.

Young twigs, leaf buds, stipule outside (glabrous
within) and petiole shortly evenly densely persistently
pale golden-brown pubescent, occasionally glabrous:
midrib on both surfaces sparsely so. Twigs c.2 mm @
near the apices, ribbed, smooth to rugulose. Bud to 10
by 2.5 mm, falcate, acute. Stipule to 20 by 3 mm,
linear, caducous, sparsely pubescent outside, glabres-
cent within. Leaf 4~-9(-12) by 1.5-4.0(—5.5) cm, nar-
rowly elliptic to lanceolate; frequently glaucous
beneath; base cuneate; acumen to 8 mm long, slender:
nerves 8-10(—16) pairs, slender but elevated beneath,
at 30°—40°; tertiary nerves slender, subscalariform:
midrib applanate above, prominently terete beneath;
petiole 11-17 mm long, slender, geniculate, drying
buff pubescent. Raceme to 5 cm long, caducously
shortly evenly golden-buff pubescent, unbranched or
sometimes singly branched, bearing to 4 flowers.
Flower bud to 20 by 6 mm, fusiform. Calyx glabres-
cent. Corolla densely golden-buff pubescent outside,
sparsely so within. Stamens 15, shorter than style at
anthesis; filaments slender, tapering, c. 4 length of
anther. Gynoecium cinereous but for the distal 4 of the
style; ovary ovoid; style broadly columnar, somewhat
tapering. Fruit calyx and pedicel glabrous. Pedicel to 6
mm long. Calyx tube to 10 by 8 mm, subglobose to
ellipsoid, decurrent with pedicel at base, with 5 slender
acute ribs continuous to base or confined to distal end:
2 longer lobes to c. 8(—20) by 1.5 cm, lorate, obtuse; 2
shorter lobes to 6 by 5 mm, deltoid, subacute. Nut
apex densely golden-buff pubescent; style remnant 10
mm long, slender, pubescent.

Distr. Malesia: Malaya (Perak, Penang, coastal
Pahang and Trengganu), N.E. Sumatra (Riouw-
Lingga), Borneo (Sarawak). :

Ecol. Hill forests, locally gregarious on ridges,
especially at 500-800 m but down to 100 m near coast.

Vern. Kéruing pipit (Mal.).

Note. Collections from low altitude, previously
described as D. pseudofagineus generally, have longer
leaves with more nerves, are generally glaucous
beneath, and have fruit calyx ribs confined to the
distal half. There is no discontinuity in the variation
between the lowland and hill forms however and these
species are therefore not regarded as distinct here.

30. Dipterocarpus cinereus SLooT. Bull. Jard. Bot.
Btzg III, 8 (1927) 319, f. 8.

Large tree. Twig, leaf bud, stipule, petiole, leaf
undersurface, panicle and ovary densely persistently

buff pubescent; hairs longer on buds and stipules;
calyx thus at first, becoming sparsely so in fruit; parts
of petals exposed in bud densely puberulent. Twigs c. 2
mm @ apically, terete, rugulose, pale brown, much
branched. Buds to 12 by 3 mm, lanceolate-falcate;
expanded stipule not seen. Leaves 6-8 by 1.7-2.5 cm,
lanceolate, thinly coriaceous, distinctly persistently
plicate; base cuneate; apex shortly narrowly tapering-
acuminate; nerves 8—9 pairs, steeply ascending at c.
40°, slender but distinctly elevated beneath; tertiary
nerves densely scalariform, slender and obscure;
midrib prominent beneath, shallowly depressed
above; petiole 1.7—2.5 cm long, very slender. nflores-
cence to 4cm long, axillary, simple or singly branched,
bearing 4-5 secund flowers. Flower buds to 22 by 8
mm, fusiform. Stamens c. 25, exceeding style at
anthesis; filaments slender, somewhat longer than the
narrow tapering anthers and appendage; style colum-
nar, pubescent in the basal 4. Fruit pedicel to 3 mm
long, broadening into the to 14 by 10 mm broadly
ellipsoid or obovoid, sharply though narrowly
5-ribbed, calyx tube; 2 longer lobes to 5 by 1.2 cm,
broadly spatulate, narrowly obtuse, constricted to c. 4
mm broad at base; 3 shorter lobes to 5 by 5mm, ovate,
obtuse, + revolute.

Distr. Malesia: Central W. Sumatra (P. Musala).

Ecol. Lowland forest on leached soils, rather
common.

Vern. Lagan bras.

Note. Clearly closely allied to D. fagineus, from
which it differs principally in the number of
stamens.

31. Dipterocarpus semivestitus SLOOT. Bull. Jard. Bot.
Btzg III, 8 (1927) 342, f. 13; Sym. Mal. For. Rec. 16
(1943) 188, f. 85.

Parts glabrous but for densely puberulent inflores-
cence, fruit calyx tube, sparsely puberulent fruit calyx
lobes, and densely pubescent ovary. Twig c. 3 by 2
mm, + compressed apically, slender, much branched,
blackish, minutely pale lenticellate; stipule scars obli-
que. Buds to 8 by 2 mm, lanceolate-falcate, slender,
drying black; stipules unknown. Leaves 5.6—11 by
2.5-6 cm, narrowly elliptic, coriaceous; margin some-
what undulate; base cuneate; acumen to | cm long,
slender; nerves 6-8 pairs, steeply ascending, slender
and hardly elevated beneath; tertiary nerves densely
scalariform, hardly elevated beneath, set vertically to
the midrib; petiole 1.4-2.2 cm long, slender. Inflores-
cence to 7 cm long, simple. Flower unknown. Fruit
subsessile; calyx tube to 15 by 10 mm, narrowly
obovoid, with 5 sharp undulating ridges or narrow, to
2 mm wide, wings; 2 longer lobes to 6.5 by 1.5 cm,
spatulate, obtuse, c. 5 mm broad at base; 3 shorter
lobes to 5 by 4 mm, elliptic, subrevolute.

Distr. Malesia: S. E. Borneo (Marabahan Dis-
trict), Malaya (Perak).

Ecol. Rare in lowland forest on low lying, perhaps
almost swampy, land.

Vern. Kéruing padi, k. dadeh (Malaya), murtulang
(Borneo).

7

‘y Ya:

Fig. 26. The narrow-leaved seedlings of Dipterocarpus
oblongifolius BL. Brunei (Photogr. ASHTON).

32. Dipterocarpus oblongifolius BL. Mus. Bot. Lugd.-
Bat. 2 (1852) 36; WaLP. Ann. 4 (1857) 335; Mia. FI.
Ind. Bat. 1, 2 (1859) 498; Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 85; DC. Prod. 16, 2 (1868) 614; Dyer, J. Bot. 12
(1874) 105; ibid. (1874) 152; BurcK, Ann. Jard. Bot.
Btzg 6 (1887) 201; KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 95; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 37:
Merr. En. Born. (1921) 399; Rpt. Fl. Mal. Pen. 1
(1922) 216; Cras, Fl. Siam. Enum. | (1925) 136:
ENDERT, Bot. Versl. M.O. Borneo Exp. (1927) 248:
Stoot. Bull. Jard. Bot. Btzg III, 8 (1927) 338; Trop.
Natuur 17 (1928) 146, f. 9-10; Foxw. Mal. For. Rec.
10 (1932) 86; BuRK. Dict. (1935) 844; CorRNER,
Wayside Trees (1940) 211; Sym. Mal. For. Rec. 16
(1943) 184, f. 85: BROWNE, For. Trees Sarawak &
Brunei (1955) 110; ASHTON, Man. Dipt. Brun. (1964)
39, f. 6, pl. 9 (seedlings); ibid. Suppl. (1968) 16: MEUER
& Woop, Sabah For. Rec. 5 (1964) 264; SRIVASTAVA,
Mal. For. 40 (1977) 251, f. 1; CoRNER, Gard. Bull.
Sing. Suppl. | (1978) 44. — D. stenopterus VESQUE, C.
R. Ac. Sc. Paris 78 (1844) 625; J. Bot. 12 (1874) 150:
Dyer, J. Bot. 12 (1874) 152. — D. pulcherrimus RD.
Trans. Linn. Soc. Bot. 3 (1893) 283. — Fig. 26.
Twig, leaf bud, midrib on both surfaces, nervation
beneath, petiole, and raceme densely + evenly cadu-
cous pale yellow tomentose; persistently so on stipule
outside, calyx and ovary. Twig 2-3 mm @ apically,
terete or + compressed; amplexicaul stipule scars
inconspicuous. Bud c. 20 by 3mm, linear, compressed,
acute. Stipule 10-15 by 1.5 cm, linear, obtuse. Leaves
14-18(—25) by 4~-7(-9) cm, narrowly elliptic to lanceo-
late; base cuneate; apex gradually tapering; acumen to
1 cm long, slender; nerves 16-20 pairs, slender, at
40°—50°; petiole 13-2 cm long. Racemes to 18 cm long,
terminal and axillary, simple or singly branched, with
distichous flowers; bracts to 20 by 2.5 mm, linear.

DIPTEROCARPACEAE (Ashton)

317

Calyx densely cream tomentose. Stamens 15, shorter
than style; anther as long as the filament, narrowly
oblong, tapering from the base into the glabrous stout
appendage; + shorter than anther; ovary conical,
densely tomentose, tapering into the stylopodium, the
latter twice as long as the ovary, narrowly-cylindrical,
tapering into and + twice as long as the filiform
glabrous style. Fruit pedicel 1-2 mm long. Fruit calyx
tube 25—30 by 7-9 mm, narrowly obovoid or fusiform,
only slightly constricted at the c. 6 mm @ neck, with 5
wavy. c. 1 mm wide, thin wings from neck to base; 2
longer lobes 10-12 by 15 cm, narrowly spatulate,
obtuse, 3-4 mm wide at base, I-nerved with 2 small
lateral nerves at the base: 3 shorter lobes c. 10 by 3
mm, narrowly deltoid to linear, obtuse, recurved.
Saplings stenophyllous.

Distr. S. Peninsular Thailand (Pattani) and in
Malesia: Malaya (all major east-flowing rivers; rare in
Perak). Borneo (excepting most of Sabah).

Ecol. Gregarious on the banks of fast-flowing
inland rivers, germinating and becoming established
below the flood-line.

Vern. Neram (Mal.), /Jaran (S.E. Borneo), ensurai,
gansurai (northern Borneo).

Note. Leaves of seedlings are linear-lanceolate; cf.
VAN SLOOTEN (1928) and SrIvAsTAVA (1977). This
species is so characteristic of swift-running streams in
Borneo and Malaya, that CORNER (1940, 1978) gave
the name neram rivers to this type.

33. Dipterocarpus grandiflorus (BLCO) BLco, FI. Filip.
ed. 2 (1845) 314: ibid. ed. 3, 2 (1878) 218, t. 263: DC.
Prod. 16, 2 (1868) 612; Dyer, J. Bot. 12 (1874) 106. t.
145, f. 19: VipAL, Sinopsis (1883) t. 14A; Pl. Vasc.
Filip. (1886) 59; Burck, Ann. Jard. Bot. Btzg 6 (1887)
201: KinG, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 95;
BRANDIS & GILG in E. & P. Pfi. Fam. ed. 1, 3, 6 (1894)
256; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 37; RIDL.
Agr. Bull. Str. & F. M.S. 1 (1901) 55; Fl. Mal. Pen. 1
(1922) 216; Foxw. Philip J. Sc. 6 (1911) Bot. 251, pl.
36; ibid. 13 (1918) Bot. 179; ibid. 67 (1938) 259; Mal.
For. Rec. 10 (1932) 87; HEYNE, Nutt. Pl. ed. 1, 3 (1917)
273; ibid. ed. 2 (1927) 1095, 1096; MerR. Sp. Blanc.
(1918) 268; En. Born. (1921) 398; En. Philip. 3 (1923)
89: Burk. J. Str. Br. R. As. Soc. 81 (1920) 55; REYEs,
Philip. J. Sc. 22 (1923) 321; Crai, Fl. Siam. Enum. |
(1925) 134; Gite in E. & P. Pfi. Fam. ed. 2, 21 (1925)
25; Soot. Bull. Jard. Bot. Btzg III, 8 (1927) 333;
PARKER, Ind. For. Rec. 16 (1931) 5; PARKINSON,
Burma For. Bull. 27 (1932) 17; BurK. Dict. (1935)
843: Sym. Mal. For. Rec. 16 (1943) 178, f. 85, 90:
SMITINAND, Thai For. Bull. 4 (1958) 33; MEER &
Woop, Sabah For. Rec. 5 (1964) 252, f. 38.—
Mocanera grandiflora Buco, FI. Filip. ed. 1 (1837)
451. — D. blancoi BL. Mus. Bot. Lugd.-Bat. 2 (1852)
35.— Vatica trigyna Grirr. Notul. 4 (1854)
514. — D. motleyanus Hook f. Trans. Linn. Soc. 23
(1860) 159: Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867)
85; DC. Prod. 16, 2 (1868) 611; WALP. Ann. 7 (1869)
377. — D. griffithii Mig. Ann. Mus. Bot. Lugd.-Bat. 1
(1864) 213: DC. Prod. 16, 2 (1868) 612; WaLp. Ann. 7

318

(1869) 377; Dyer, FI. Br. Ind. 1 (1874) 299; J. Bot. 12
(1874) 107; Kurz, Fl. Burma 1 (1877) 116; RIDL.
Trans. Linn. Soc. Bot. 3 (1893) 283; KING, J. R. As.
Soc. Beng. Sc. 62, 2 (1893) 96; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 37; Ind. Trees (1906) 701; GAMBLE,
Man. Ind. Timb. (1922) 70.—D. pterygocalyx
ScHEFF. Nat. Tijd. N. I. 31 (1870) 347; Dyer, FI. Br.
Ind. | (1874) 298; HeEYNeE, Nutt. Pl. ed. 2 (1927)
1096. — Fig. 17B.

Very large hardly buttressed tree. Leaf bud, outside
of stipule, parts of petals exposed in bud, ovary apex
and sometimes twig densely evenly pale buff pubes-
cent, parts otherwise glabrous. Twigs to 12 mm @,
stout, becoming grey-brown; internodes short, the
leaves clustered around the twig endings. Buds to 2 by
1 cm, ovoid, acute; stipule to 18 by 5 cm, oblong-lan-
ceolate, subacute. Leaves 10-18 by S—12 cm, ovate,
coriaceous, + applanate; base obtuse or subcordate;
acumen to | cm long, short, nerves 15-17 pairs,
prominent beneath, spreading; tertiary nerves re-
motely scalariform, barely elevated; petioles 3-9 cm
long, very long, slender. /nflorescences to 18 cm long,
very long, slender, axillary, borne in dense groups
behind the leaves, unbranched, bearing to 3 remote
distichous flowers. Flower buds to 35 by 13 mm,
fusiform. Stamens 30, exceeding style at anthesis;
filaments broad, compressed, tapering, c. 4 the length
of the relatively short broadly linear-lanceolate taper-
ing anthers; appendages acicular, c. ¢% length of
anthers. Ovary tapering into the short stoutly colum-
nar style; ovary and style puberulent in the basal half.
Fruit pedicel to 2 by 4mm, stout. Calyx tube to 7 by 3.5
cm, ellipsoid, with 5 pruinose, to 1.5 cm wide,
prominent coriaceous wings continuous from base to
apex; 2 longer lobes to 22 by 3 cm, oblanceolate-spa-
tulate, obtuse, tapering abruptly to c. 1.3 cm wide at
base; 3 shorter lobes to 2 by 1.5 cm, elliptic, subrevo-
lute.

Distr. Andamans, Mergui, Peninsular Thailand,
and in Malesia: Malaya, north-east Sumatra (Atjeh,
Langkat, Bengkalis, P. Simalur, P. Musala, Riau
Arch., Banka, Billiton), P. Karimata, Borneo (from
lower Dayak in S.W. to Sabah in E.; incorrectly
recorded by BROWNE, For. Trees Sarawak & Brunei
(1955) 106 from Sarawak), Philippines (N.W. Luzon).

Ecol. Often common, becoming semi-gregarious,
in more seasonal climates in Semi-evergreen Diptero-
carp forests; on small islands and on coastal hills in
less seasonal areas, and in Malaya occurring fre-
quently with Shorea curtisii on inland ridges to 700 m.

Vern. Kéruing bélimbing, k. pekat, meluit (Malaya),
kéruing, lagan bras (Sumatra), tampudau, bajan, kéru-
ing (Borneo), apitong (Tag.).

Note. A remarkable relic from the once vast
Semi-evergreen Dipterocarp forests of the Pleistocene
Sundaland.

34. Dipterocarpus glabrigemmatus AsHTON, Gard.
Bull. Sing. 31 (1978) 11. — D. sp. ASHTON, Man. Dipt.
Brun. Suppl. (1968) 19.

Midrib and nerves sparsely pubescent, other vege-

FLORA MALESIANA

[ser. I, vol. 9?

tative parts glabrous. Twig c. 3 by 2 mm @g apically,
somewhat compressed and ribbed, lustrous; stipule
scars slender, pale, hardly raised. Leaf bud to 6 by 4
mm, acute. Stipule unknown. Leaves 6-9 by 4.5—6 cm;
broadly ovate; margin sinuate distally; base obtuse;
acumen to 5 mm long, short, broad; nerves 10-11
pairs, prominent beneath, at 45°-60° near the base,
straight but curving near the margin; midrib
applanate above, prominently terete beneath; tertiary
nerves subscalariform; petiole 1.5-2.5 cm long,
slender, drying rugose. Raceme to 10cm long, to 1mm
@ at base, axillary, unbranched, glabrous. Flower bud
to 25 by 12 mm. Calyx glabrous, prominently
5-winged; corolla typical. Stamens 15, subequal; fila-
ments compressed at base, tapering; anthers linear,
2-3 times as long as filaments; appendage to connec-
tive almost as long as anther, filiform, tapering. Ovary
ovoid, small, pubescent; style c. 5 times length of
ovary, pubescent except at apex. Fruit unknown.
Distr. Malesia: Borneo (Central Sarawak).
Ecol. Local, Mixed Dipterocarp forest, clay soil.

35. Dipterocarpus palembanicus SLooT. Bull. Jard.
Bot. Btzg III, 8 (1927) 336, f. 12; Sym. Mal. For. Rec.
16 (1943) 185, f. 85; ASHTON, Man. Dipt. Brun. Suppl.
(1968) 16, f. 6-7, pl. 6 (stem), p.p.; Gard. Bull. Sing. 31
(1978) 11. — D. alatus (non Roxs.) Foxw. Mal. For.
Rec. 10 (1932) 82.

a. ssp. palembanicus.

Twig, leaf bud and stipule outside densely fulvous
hirsute; petiole midrib above and leaf beneath densely
shortly puberulent; caducous on twigs and petioles,
otherwise persistent. Twig 2-3 mm @, terete, with a
rather papery, finely cracked surface. Bud 4-12 by 3-4
mm, shortly oblong to conical, obtuse. Stipule 2-3 cm
long, narrowly hastate, acute. Leaves 10-14 by 5-9
cm, broadly elliptic to ovate, thinly coriaceous; base
obtuse or cuneate, acumen short or to 2 cm long,
narrow; margin undulate; nerves slender, dense,
12-14 pairs, at 35°-45°, tending to be persistently
plicately folded between; tertiary nerves densely sca-
lariform, slender; petiole 2-3 cm long, prominently
geniculate. Raceme to 6 cm long, terminal or axillary,
terete, tending to wrinkle on drying, unbranched or
singly branched, the branchlets bearing distichous
flowers. Bud to 3.5 by 1.5 cm. Calyx and corolla
typical, calyx glabrous. Stamens 15; filaments com-
pressed at base, slender, tapering; anthers linear, c. 15
length of filament, tapering apically; appendage to
connective c. + length of anther, filiform, tapering,
reaching to 4 length of style. Ovary ovoid, densely
pubescent, tapering into style; style 34 times length of
ovary, stoutly filiform, densely pubescent and ribbed
in the basal 4, otherwise glabrous. Fruit calyx gla-
brous; tube to 3.5 by 1.5 cm, narrowly ellipsoid,
tapering gradually to the base and the c. 1 cm @ neck;
broadly 5-winged, the wings thin, to 8 mm broad, +
obtuse or subcordate at base, with the margin folded
over frequently at maturity; 2 longer calyx lobes to 10
by 3.5 cm, oblong, obtuse, base revolute, subcordate,

1982]

with 3 parallel nerves running down the whole length
close to the centre; 3 shorter calyx lobes to 5 by 10 mm,
broadly orbicular, strongly recurved, with narrow
base and revolute undulate margin.

Distr. Malesia: Malaya (Trengganu, Central
Johore), Singapore, Sumatra (Palembang), Borneo
(Central Sarawak).

Ecol. Local, Mixed Dipterocarp forest on hills,
clay soils, to 650 m.

Vern. Kéruing ternek (Mal.), lagan daun halus
(Sum.).

b. ssp. borneensis ASHTON, Gard. Bull. Sing. 31 (1978)
11. — D. palembanicus: ASHTON, Man. Dipt. Brun.
(1964) 41; MEER & Woop, Sabah For. Rec. 5 (1964)
266, f. 42.

Differing from ssp. palembanicus as follows: Twig,
petiole, midrib (both surfaces) and nerves beneath
densely shortly chestnut pubescent, caducous on
twigs; persistent, long, on bud, stipule and petiole;
lamina fugaceous pubescent. Leaves 7-11 by 3-6 cm,
oblong to ovate, with 12-14 pairs of nerves; petiole c.
15 mm long. Fruit calyx tube to 5.5 by 2 cm, wings to
15 mm broad, undulate, auriculate at base and apex.

Distr. Malesia: northern Borneo (Central and
N.E. Sarawak to E. Sabah and Nunukan I.).

Ecol. As ssp. palembanicus, and sometimes grow-
ing together with it.

36. Dipterocarpus fusiformis ASHTON, Gard. Bull.
Sing. 31 (1978) 12.

Large tree. Young parts densely buff velutinate,
persistent on leaf bud and stipule outside, becoming
sparse yet + persistent on twigs, petiole and leaf
undersurface, elsewhere caducous. Twigs c. 2 mm @
apically, slender, terete, + verrucose. Buds to 15 by 3
mm, linear, lanceolate; stipule to 30 by 6 mm,
lanceolate, acute. Leaves 7-17 by 3—7 cm, elliptic or
narrowly ovate, coriaceous; base cuneate or obtuse;
acumen to 12 mm long, slender, prominent; nerves
13-17 pairs, slender but prominent beneath, ascend-
ing at 30°-40°; tertiary nerves densely scalariform,
evident but hardly elevated beneath; petiole 2—2.6 cm
long, slender. Complete inflorescences unknown; in-
florescences singly branched or unbranched, axillary.
Flowers unknown. Fruit pedicel c. 1 mm long, short;
calyx tube to 28 by 18 mm, ellipsoid, with 5, to 6mm
broad, straight incrassate wings or narrow ridges,
continuous from base to apex but generally broader in
the distal half; 2 longer lobes to 10 by 2.6 cm, broadly
spatulate, obtuse, c. 7mm wide at base; 3 shorter lobes
to 5 by 5 mm, suborbicular, subrevolute.

Distr. Malesia: N.E. Borneo (Tawau area).

Ecol. Undulating well drained fertile soils; Mixed
Dipterocarp forest; rare.

Note. Clearly closely allied to D. mundus SLOOT. of
the Central Bornean hills, a species which is conspi-
cuously glabrous but for its corolla, ovary, and the
inside of its stipules. The two species are therefore very
different in appearance; flowers are still awaited and
may provide further differences.

DIPTEROCARPACEAE (Ashton)

319

37. Dipterocarpus mundus SLOoT. Bull. Jard. Bot.
Btzg III, 16 (1940) 446, f. 7; AsHTon, Man. Dipt.
Brun. Suppl. (1968) 15, f. 3. — Fig. 17G.

Parts glabrous but for pubescent inner surface of
stipules. Twig c. 2-3 mm @ apically, terete, much
branched; stipule scars slender, elevated but not
prominent. Bud to 14 by 3 mm, slender, acute. Stipule
to 4.0 by 0.6 cm, linear, caducous. Leaves 5.5—16 by
2.5-7.5 cm, narrowly elliptic to obovate; base nar-
rowly obtuse; acumen to 8 mm long; nerves 8-10
pairs, straight, slender but prominent beneath, at 30
to 50°; tertiary nerves remotely subscalariform; mid-
rib + applanate above, stout, prominent, terete
beneath; petiole 1.6—-3 cm long, slender, geniculate.
Raceme to 6 cm long, terete to somewhat compressed,
smooth, unbranched, bearing to 4 flowers. Flower
calyx and corolla typical. Stamens 15; filaments short,
compressed, anthers narrowly oblong; appendage to
connective c. 4+ length of anthers, filiform, tapering.
Ovary small, ovoid; stylopodium columnar, tapering,
pubescent, ending ina short glabrous style. Fruit calyx
and pedicel glabrous. Pedicel to 3 mm long, c. 1mm @.
Calyx tube to 3 by 1.2 cm, fusiform, bearing 5 wings,
each to 7 mm wide, broadest in the distal half,
incrassate; 2 longer lobes to 11 by 2.7 cm, oblong,
tapering to an obtuse apex, c. 5 mm broad above the
tube; 3 shorter lobes to 8 by 8 mm, broadly ovoid,
obtuse, somewhat recurved. Nut narrowly ovoid,
densely buff pubescent; style remnant to 8 mm long,
filiform.

Distr. Malesia: Central Borneo (Kapuas, Rejang
hinterland).

Ecol. Locally frequent on ridges, 400-600 m.

Vern. Késurai bukit.

38. Dipterocarpus borneensis SLOoT. Bull. Jard. Bot.
Btzg III, 16 (1940) 445, f. 6; BROWNE, For. Trees
Sarawak & Brunei (1955) 107; ASHTON, Man. Dipt.
Brun. (1964) 24, f. 6, 7; ibid. Suppl. (1968) 11.
Young twig, petiole, raceme, midrib, nerves and
sometimes tertiary nerves beneath sparsely shortly
rust pubescent, mostly caducous; leaf bud and stipule
outside densely persistently so, the hairs longer,
pinkish buff. Twig 3-5 mm @ apically, glabrous,
smooth but for the paler amplexicaul stipule scars,
occasionally rugulose. Bud 5—7 by 2.5—3 mm, ovoid,
subacute. Stipule to 40 by 4-5 mm, linear, acute.
Leaves 7-12 by 3-7 cm, broadly ovate to elliptic; base
obtuse or broadly cuneate; acumen to 8 mm long;
nerves 9-12 pairs, distant, at 50° at the lamina base,
down to 20° at the apex, prominent; margin fre-
quently undulate towards the apex; petiole 1.5—2.5cm
long, slender. Raceme to 6 cm long, singly or rarely
doubly branched, terminal or axillary, terete or +
compressed; bracteoles unknown. Flower bud to 4 by |
cm. Calyx and corolla typical, calyx glabrous.
Stamens c. 25, shorter than the style; anthers short,
narrowly sagittate; appendage stout, tapering, c. 4
length of anther. Ovary conical, shortly densely
pubescent; stylopodium and style c. 3 times length of
ovary, densely tomentose except at apex, slightly

320

FLORA MALESIANA

[ser. I, vol. 9?

swollen medially. Fruit calyx tube to 1.5 by 1 cm,
narrowly ovoid to ellipsoid, tapering gradually into
the pedicel and to the c. 8 mm g neck, with 5 narrow c.
1 mm broad, c. 6 mm, long wings in the distal half
terminating at the insertion of the cayx lobes; 2 longer
lobes to 7.5 by 1.5—1.8 cm, chartaceous, oblong,
obtuse, + abruptly constricted at the undulate mar-
gined base; 3 shorter calyx lobes c. 5 by 4mm, revolute
and recurved outwards.

Distr. Malesia: Eastern coastal Sumatra (Indra-
giri; Lingga Arch.: Singkep); Borneo (Sampit to
Muara Tewe and Ulu Mahakam in S.E. Borneo;
Sambas and Mampawah in W. Borneo; Sarawak and
Brunei).

Ecol. Common in heath forest on podsols below
350 m; rare in mixed swamp forest.

Vern. Lagan, kéruing daun halus (Sum.), ompal,
pendérawan, awang buah, béngkajarap, kéruing sindor
(Borneo).

Note. Closely resembling D. fagineus but with the
ridges of the calyx tube distinctly expanded into
narrow wings distally, and with longer tomentum
persisting on the nervation beneath.

39. Dipterocarpus nudus VESQUE, C. R. Ac. Sc. Paris
78 (March 1874) 626; J. Bot. 12 (1874) 150; Dyer, J.
Bot. 12 (1874) 152; BuRck, Ann. Jard. Bot. Btzg 6
(1887) 201; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 38;
Me_rr. En. Born. (1921) 399; SLoot. Bull. Jard. Bot.
Btzg III, 8 (1927) 325; BROWNE, For. Trees Sarawak &
Brunei (1955) 110; ASHToN, Man. Dipt. Brun. (1964)
38, f. 6, 7; ibid. Suppl. (1968) 16. — D. pentapterus
Dyer, J. Bot. 12 (Apr. 1874) 106, t. 144, f. 18; ibid.
(1874) 152.

Glabrous but for the buff pubescent inside of
stipules. Twig 3-4 mm @g apically, slender, smooth,
often tending to be triangular; with inconspicuous
stipule scars. Bud 15-30 by 2-3 mm, linear, terete.
Stipule to 6 by 0.5 mm, linear. Leaves 11-14 by 4-6
cm, narrowly elliptic, thinly coriaceous, margin undu-
late and + sinuate distally; base cuneate, apex to 6
mm long, acuminate; nerves 11—14 pairs, close, obli-
que (30°—-40°); tertiary nerves widely spaced, scalari-
form but rather sinuate, at c. 90° to nerves; petiole
2-4.5 cm long, slender. Raceme to 18 cm long,
terminal or axillary, glabrous, terete, becoming ribbed
on drying, zigzag; simple or singly branched, with
distichous flowers at wide intervals; bracts unknown.
Flower bud to 4 by 0.9 cm, slender. Calyx and corolla
typical, calyx glabrous. Stamens c. 15, shorter than the
style; filaments short; anthers narrowly oblong, taper-
ing. Ovary ovoid-conical, densely tomentose; style
and stylopodium about 3 times as long as ovary,
filiform, slightly tapering, densely tomentose at base,
glabrous at apex. Fruit glabrous; calyx tube c. 2.5 by 1
cm, ellipsoid to fusiform, broadest distally, the base
constricted abruptly at the pedicel, slightly constricted
at the c. 7 mm @ neck; 5 winged, the wings incrassate
but acute, rather narrow and widest (c. 3 mm) distally,
continuing the whole length of the tube; 2 longer calyx
lobes to 9.5 by 3 cm, lanceolate, equally 3-nerved,

obtuse, tapering gradually to the 3-4 mm wide base; 3
shorter lobes c. 4 mm long and broad, small, obtuse,
slightly revolute.

Distr. Malesia:
Brunei).

Ecol. Hillsides and ridges in Mixed Dipterocarp
forest to 650 m.

Vern. Kéruing lichin.

N.W. Borneo (Sarawak and

40. Dipterocarpus geniculatus VEsQuE, C. R. Ac. Sc.
Paris 78 (March 1874) 626; J. Bot. 12 (1874) 150;
Dyer, J. Bot. 12 (1874) 152; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 199; BRranpis, J. Linn. Soc. Bot. 31
(1895) 33; Merr. En. Born. (1921) 398; SLoor. Bull.
Jard. Bot. Btzg III, 9 (1927) 317; ibid. III, 17 (1941) 99,
f. 11; BROWNE, For. Trees Sarawak & Brunei (1955)
109; AsHTON, Man. Dipt. Brun. Suppl. (1968)
14. — D. angulatus Dyer, J. Bot. 12 (April 1874) 104;
ibid. (May 1874) 152.

a. ssp. geniculatus.

Young twig, leaf bud, stipule outside and raceme
densely shortly pale cream tomentose; petiole and
nervation beneath sparsely pubescent to glabrescent.
Twigs to 7 mm @, with rows of large longitudinally
elongated flat lenticels and conspicuous sinuate
amplexicaul stipule scars. Bud c. 2.5 by 2 cm, broadly
ovoid, acute. Stipule c. 6 by 2 cm, pink with a cream
tomentum when fresh, broadly lanceolate, acute.
Leaves 7-12 by 5-7 cm, elliptic to + obovate,
coriaceous, lustrous above; base obtuse, apex obtuse
or shortly acuminate; nerves 10-12 pairs, prominent
beneath, well spaced, at 45°-5S5°; petiole stout,
straight, 3-5 cm long. Raceme to 24 cm long, axillary,
terete, straight, singly, rarely doubly branched, the
flowers distant, distichous; bracts to 20 by 5 mm,
narrowly lanceolate, acute, shortly pubescent outside,
puberulent within. Flower bud to 3.5 by 1 cm. Calyx
and corolla typical, calyx shortly densely cream
tomentose. Stamens 30, shorter than the style; fila-
ments as long as the anthers; anther rather short,
narrowly oblong, tapering apically; appendage to
connective somewhat shorter than anther, stout at
base, tapering. Ovary ovoid-conical, shortly tomen-
tose; style and stylopodium + twice as long as ovary,
narrowly cylindrical, shortly tomentose basally, gla-
brous in the distal 4. Fruit calyx tube shortly densely
cream powdery tomentose, lobes sparsely so; tube c.
1.5 cm long and broad, obovoid, tapering to pedicel
and constricted to 1.2 cm g at the neck; 5-ridged, the
ridges | mm thick, stout, to 3 mm wide at the undulate
distal ends, continuing from the neck to the pedicel; 2
longer calyx lobes to 12 by 2.5 cm, oblong, obtuse,
tapering to the 6 mm broad strongly revolute base
with one branching central nerve running to the apex
and 2 shorter laterals; 3 shorter lobes c. 1 by 1.5 cm,
cordate, the sides recurved.

Distr. Malesia: Borneo (Sarawak from the
Kemena valley westwards, Lower Kapuas).

Ecol. Lowland dipterocarp forests on leached clay
soils, to 400 m.

1982]

DIPTEROCARPACEAE (Ashton)

EY

Vern. Kéruing kérubong, k. guntang, k. bélimbing
(Mal.).

b. ssp. grandis ASHTON, Gard. Bull. Sing. 20 (1963)
240; Man. Dipt. Brun. Suppl. (1968) 14. — D. genicu-
latus sensu ASHTON, Man. Dipt. Brun. (1964) 11, f. 6,
pl. 11 (habit, bark); MEER & Woop, Sabah For. Rec.
5 (1964) 248, f. 36.

Differing from ssp. geniculatus as follows: Twigs to
13 mm g; leaves 20-35 by 12-16 cm, petiole 8-10 cm
long; 2 longer fruit calyx lobes to 15 by 4 cm.

Distr. Malesia: N.E. Borneo (Sarawak from Sibuti
north-eastwards, Brunei, S.W. Sabah and Sandakan
Distr.).

Ecol. As ssp. geniculatus, but confined to ultrabasic
rocks in E. Sabah.

Vern. Kéruing kérubong, k. tangkai panjang.

41. Dipterocarpus costatus GAERTN. f. Fruct. 3 (1805)
50, t. 187; Roxs. Hort. Beng. (1814) 42; Fl. Ind. ed.
Carey 2 (1832) 613; BucH.-HAM. Mem. Wern. Nat.
Hist. Soc. 6 (1832) 299; WALP. Rep. 5 (1945) 124; DC.
Prod. 16, 2 (1868) 610; Kurz, J. As. Soc. Beng. 43, 2
(1874) 98; Fl. Burma (1877) 117; Kina, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 98; BRANDis, J. Linn. Soc. Bot.
31 (1895) 35; Ind. Trees (1906) 66; GAMBLE, Man. Ind.
Timb. (1922) 70; Crats, Fl. Siam. Enum. | (1925) 133;
PARKER, Ind. For. Rec. 13 (1827) 4; Foxw. Mal. For.
Rec. 10 (1932) 85; Burk. Dict. (1935) 842; Sym. Mal.
For. Rec. 16 (1943) 173, f. 85. — Pterigium costatum
Correa, Ann. Mus. Hist. Nat. Paris 8 (1806) 397, t.
65 — Shorea costata PRESL, Rostl. 2 (1825) 66. — D.
angustifolius W. & A. Prod. 1 (1834) 84; DC. Prod. 16,
2 (1868) 610. — D. lemeslei VEsquE, C. R. As. Sc.
Paris 78 (1874) 626. — D. insularis HANCE, J. Bot. 14
(1876) 241; Prerre, Fl. For. Coch. 3 (1889) t. 214;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 37.—D.
artocarpifolius PIERRE [ex LANESSAN, Pl. Util. Colon.
Fr. (1886) 297, nomen] Fl. For. Coch. 3 (1889) t. 213;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 37.—D.
parvifolius Hem, Bot. Tidsskr. 25 (1903) 43; GUERIN,
Fl. Gén. I.-C. 1 (1910) 365; Crarp, Fl. Siam. Enum. 1
(1925) 135.

Large unbuttressed tree. Twigs, leaf buds, midrib
above, petioles, inflorescences, flower calyx and petals
outside densely persistently evenly pale golden-brown
pubescent, leaf beneath and fruit calyx sparsely
somewhat caducously so. Twig c. 3 mm @g apically,
terete, pale brown, much branched. Buds to 10 by 4
mm, ovoid to lanceolate, acute; stipule to 5 by 1 cm,
lorate, tapering and subacute apically. Leaves 5.5-17
by 2.8-7 cm, ovate or elliptic, usually small, coria-
ceous; base obtuse or broadly cuneate; acumen to 6
mm long, short; nerves 11—13(—15) pairs, ascending;
tertiary nerves slender, elevated beneath: petiole
1.5—2.8 cm long, slender. /nflorescence to 7 cm long,
singly branched or unbranched, bearing to 7 disti-
chous flowers. Flower buds to 25 by 8 mm, fusiform.
Stamens 18-20, shorter than style at anthesis; fila-
ments short, lorate; anthers short, linear-lorate, taper-
ing; appendage acicular, very slender. c. 4 length of

anther; ovary ovoid, tapering into the columnar style;
ovary and basal 5 of style densely pubescent. Fruit
pedicel to 3 by 2 mm; calyx tube to 2 cm g, +
subglobose, with 5 continuous, to 2 mm wide, narrow
coriaceous wings; 2 longer lobes to 11.5 by 2.5 cm,
lorate-oblong, narrowly obtuse, c. 8 mm wide at the
subrevolute base; 3 shorter lobes to 15 by 15 mm,
suborbicular, revolute, prominent.

Distr. Andamans, Chittagong, Burma, Thailand,
Cambodia, Cochinchina, and in Malesia: Malaya
(from Negri Sembilan northwards).

Ecol. Scattered in Hill and Upper Dipterocarp
forests, 600-1000 m, and down to sea level in seasonal
areas in N.W. Malaya. Gregarious in lowland and hill
Semi-evergreen Dipterocarp forest of S.E. Asia.

Vern. Kéruing marakluang, k. bukit (Malaya).

Note. Allied to D. glandulosus THw. of Ceylon.

42. Dipterocarpus conformis SLooT. Bull. Bot. Gard.
Btzg III, 17 (1941) 102, f. 13.

a. ssp. conformis. — D. confertus (non SLOOT.) SLOOT.
Bull. Jard. Bot. Btzg III, 8 (1927) 324 (Sumatran
coll.).

Young twig, leaf bud, stipule outside, petiole,
raceme and leaf beneath persistently densely pale
pink-brown velutinate; leaf above caducously so
except on nerves and midrib. Twigs verrucose. Bud c. 8
by 4mm, ovoid, obtuse. Stipules c. 15 by 6mm, ovoid,
acute, cupped, fugaceous. Leaves 20-24 by 12-15 cm,
obovate, chartaceous; base obtuse or subcordate,
narrowly subpeltate; acumen 4-8 mm long; nerves
13-15 pairs, at c. 60°—70°; petiole S—6 cm long, slender,
rugose. Raceme to 6 cm long, short, terminal or
axillary, slender, ribbed on drying, unbranched or
singly branched, bearing few distichous flowers;
bracts unknown. Flower bud to 3 by 0.8 cm. Calyx and
corolla typical, calyx shortly tomentose. Stamens c.
30, shorter than style; filaments almost as long as
anther, slender; anther linear, short, tapering at apex;
appendage to connective slender, tapering, almost
equal in length to anther. Ovary ovoid to conical,
long, glabrescent at base, long tomentose medially,
shortly tomentose apically; style and stylopodium c.
4x length of ovary, slender, filiform, sparsely pubes-
cent at base, otherwise glabrous. Fruit calyx +
uniformly pubescent; tube c. 2.5 by 2 cm, ellipsoid,
neck narrowed to 1—1.2 cm g; with 5, to 10 cm wide,
incrassate wings continuing from pedicel to neck,
sometimes bent to one side but not undulate; 2 longer
calyx lobes to 10 by 2 cm, oblong, obtuse, constricted
abruptly to 7 mm broad at the base; 3 shorter lobes to
8 mm long and broad, ovate, not constricted at the
base, revolute.

Distr. Malesia: northern half of Sumatra (Langsa,
Atjeh, Ophir and Ayerbangis, W. Coast).

Ecol. Rare, lowland forests on hills.

Vern. Kéruing buah, lagan sanduk (Sum.).

b. ssp. borneensis ASHTON, Gard. Bull. Sing. 20 (1963)
28; Man. Dipt. Brun. Suppl. (1968) 12. — D. con-

522

formis: ASHTON, Man. Dipt. Brun. (1964) 28, f. 6;
MeEveER & Woop, Sabah For. Rec. 5 (1964) 244.

Differs from ssp. conformis as follows: Leaves 9-12
by 5-7 cm, nerves 15—18 pairs, petiole 17-25 mm long;
wings of calyx tube to 3 mm wide.

Distr. Malesia: Borneo (N.E. Sarawak, Brunei,
S.W. Sabah).

Ecol. Rare, Hill Dipterocarp forests, clay rich soils,
below 800 m.

Vern. Kéruing beludu kuning.

43. Dipterocarpus acutangulus VesQuE, C. R. Ac. Sc.
Paris 78 (1874) 626; J. Bot. 12 (1874) 150; Dyer, J.
Bot. 12 (1874) 152; SLoor. Bull. Jard. Bot. Btzg III, 8
(1927) 321; Reinwardtia 5 (1961) 457; Sym. Mal. For.
Rec. 16 (1943) 166, f. 85; BROWNE, For. Trees
Sarawak & Brunei (1955) 107; ASHTON, Gard. Bull.
Sing. 20 (1963) 240; Man. Dipt. Brun. (1964) 22, f. 6, 7:
ibid. Suppl. (1968) 11; MEER & Woop, Sabah For.
Rec. 5 (1964) 235, pl. 27 (habit). — D. appendiculatus
(non SCHEFF.) DYER, J. Bot. 12 (1874) 152; BuRCK,
Ann. Jard. Bot. Btzg 6 (1887) 200, p.p.; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 34, p.p.; MERR. En. Born.
(1921) 397, p.p. — D. tawaensis SLoot. Bull. Jard.
Bot. Btzg III, 8 (1927) 313, f. 6.— D. helicopteryx
SLooT. Bull. Jard. Bot. Btzg III, 16 (1940) 441, f. 4:
BROWNE, For. Trees Sarawak & Brunei (1955)
109. — Fig. 17C.

Twig, bud, stipule (outside only), petiole and leaf
beneath at first densely buff pubescent; persistent on
leaf bud and stipule, + caducous elsewhere. Twig c. 3
mm @ apically, becoming glabrous, terete, with
swollen amplexicaul stipule scars. Bud 3-12 by 2.5—S
mm, ovoid, broad and short, subacute; occasionally
glabrous. Stipule c. 5 by 0.8 cm, linear, obtuse. Leaves
7-10 by 3-6 cm, elliptic to ovate; base obtuse or
cuneate, apex with narrow c. 1 cm long acumen,
margin sinuate; nerves 7-12 pairs, at c. 30°-45°;
tertiary nerves dense, scalariform; petiole 1.5—2.5 cm
long, to 1 mm @. Flowers unknown. Raceme to 4cm
long, axillary, slender, glabrous. Fruit calyx glabrous;
tube to 2.5 cm broad and long, globose or ellipsoid,
becoming slightly impressed at the pedicel, constricted
to c. | cm @ neck; with 5 rounded or angular woody
ribs, 3-4 mm thick and c. 3 mm wide (often less
pronounced), initiating from the impressed base and
terminating abruptly at the insertion of the calyx
lobes; surface warty, glabrous, pruinose; 2 longer
lobes c. 10 by 2.5 cm, oblong-spatulate, slightly
twisted, to 5mm broad at base, 3-nerved, the 2 laterals
of variable length; 3 shorter wings c. 5 mm long and
broad, deltoid, subacute, with revolute margins.

Distr. Malesia: Malaya (rare: Negri Sembilan),
Borneo (Ulu Kapuas, W. Kalimantan; Sarawak,
Brunei, S.W. and E. Sabah).

Ecol. Mixed Dipterocarp forest, sandy soils on
coastal hills and inland ridges to 1000 m.

Vern. Kéruing merah, k. béludu.

Note. A polymorphic species (in this resembling D.
eurynchus q.v.); collections from N.W. Borneo and
Malaya bear distinctly larger more thickly coriaceous

FLORA MALESIANA

[ser. I, vol. 92

leaves and are in this respect intermediate between this
species and D. globosus VESQUE.

44. Dipterocarpus sublamellatus Foxw. Mal. For.
Rec. 10 (1932) 92, pl. 8; SLoor. Bull. Bot. Gard. Btzg
III, 17 (1941) 108, f. 15; Sym. Mal. For. Rec. 16 (1943)
92, f. 85; MEUER & Woop, Sabah For. Rec. 5 (1964)
269: ASHTON, Man. Dipt. Brun. Suppl. (1968) 17, f. 3.

Leaf bud and stipule outside (glabrous within)
densely long pale fulvous hirsute, rarely glabrous, leaf
nervation beneath sparsely caducously so, otherwise
glabrous. Twig to 3 mm @ towards the apex, much
branched, slender, dark, lustrous, smooth; stipule
scars pale, prominent. Bud to 12 by 8 mm, ellipsoid-
ovoid, subacute. Stipule to 40 by 8 mm, lorate, acute.
Leaves 5—15 by 3.5—8 cm, ovate to elliptic (immature),
coriaceous persistently prominently plicate; base
broadly cuneate; acumen short, slender, to 1 cm long
in young trees; nerves 8-12 pairs, prominent beneath,
ascending at 35°—5S°; tertiary nerves very slender,
densely scalariform; petiole c. 1.5(—3) cm long,
slender, geniculate, drying black or pruinose. Jnflores-
cence unknown. Flower bud to 3.5 by 1 cm. Calyx and
corolla typical; calyx glabrous. Stamens c. 24, as long
as style at anthesis; filaments lorate, slender, com-
pressed, somewhat longer than anther; anther linear,
tapering into the acicular appendage; appendage c. 4
length of anther. Ovary ovoid, pubescent; style colum-
nar, pubescent except in the apical 5. Fruit calyx tube
to 3 by 3 cm, globose, bearing 5, to 1.5 cm wide,
incrassate undulate wings; wings tapering distally,
auriculate at base: 2 longer calyx lobes to 12 by 3 cm,
lorate, obtuse, c. 6mm wide at the somewhat revolute
base; 3 shorter calyx lobes to 5 by 7 mm, suborbicular,
revolute.

Distr. Malesia: Malaya, Sumatra (Sibolga,
Tapanuli, Indragiri), Borneo (Sarawak, west of the R.
Suai in W.; Sebatik island and Nunukan in N.E.).

Ecol. Undulating land and low hills to 500 m,
locally common, especially in moist places.

Vern. Kéruing kérut, k. padi, k. sugi (Malaya),
lagan batu, 1. boeih, masibuk (Sumatra).

Note. Collections from N.E. Borneo, and one from
Central Sarawak (S 18428) have somewhat larger
parts, especially fruit, and less undulate calyx tube
wings.

Closely related to D. insignis THw. of Ceylon.

45. Dipterocarpus concavus Foxw. Mal. For. Rec. 10
(1932) 90, pl. 7; SLoor. Bull. Bot. Gard. Btzg III, 17
(1941) 101, f. 12; Sym. Mal. For. Rec. 16 (1943) 171, f.
85. — D. confertus (non SLooT.) SLooT. Bull. Jard.
Bot. Btzg III, 7 (1927) 324, quoad spec. Lingga.
Large buttressed tree. Twig, leaf bud, stipule out-
side, petiole, midrib above and inflorescence densely
shortly yellow-brown tufted tomentose, leaf under-
surface and nervation above more sparsely so, flower
calyx and ovary densely yellow-brown puberulent,
fruit calyx glabrescent. Twig c. 3 mm @ apically,
terete. Buds to 11 by 6 mm, narrowly ovoid, acute;
expanded stipule unknown. Leaves 10—23 by 6-17 cm,

1982]

DIPTEROCARPACEAE (Ashton)

323

broadly elliptic, + chartaceous, + prominently con-
cave beneath; base obtuse, apex + shortly promi-
nently cuspidate, obtuse; nerves c. 12 pairs, promi-
nently raised beneath as also the midrib; tertiary
nerves remotely scalariform; petiole 3.5—6 cm long, c.
3 mm @, long. /nflorescence to 6 cm long, axillary,
unbranched, bearing to 3 flowers. Flower buds to 3.5
by 1 cm, lanceolate. Stamens 25; mature flowers
unknown. Fruit subsessile; calyx tube to 4.5 by 3.5 cm
including the 5, to 1.5 cm wide, prominent continuous
coriaceous wings, the tube itself c. 1.5 cm @, subglo-
bose, the wings decurrent basally with the pedicel axis
and apically with the base of the lobes; 2 longer lobes
to 16 by 4cm, broadly lorate, obtuse, hardly tapering
but undulate at base; 3 shorter lobes to 1.5 by 1.2 cm,
suborbicular, with narrowly revolute margin.

Distr. Malesia: Malaya (Pahang and Perak north-
wards), Sumatra (P. Singkep).

Ecol. Local, on well-drained flat land.

Vern. Kéruing sendok, k. dadah, damar liat
(Malaya), k. lakis, k. jantong (Singkep).

Note. The Singkep specimens are sterile and might
be D. confertus.

46. Dipterocarpus stellatus VEsQuE, C. R. Ac. Sc.
Paris 78 (March 1874) 626; J. Bot. 12 (1874) 150;
Dyer, J. Bot. 12 (1874) 153; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 202; BRANDis, J. Linn. Soc. Bot. 31
(1895) 38; BoeRL. Cat. Hort. Bog. 2 (1901) 99; MERR.
En. Born. (1921) 400; SLooT. Bull. Jard. Bot. Btzg III,
8 (1927) 335; Reinwardtia 5 (1961) 465, f. 3; BROWNE,
For. Trees Sarawak & Brunei (1955) 111; ASHTON,
Gard. Bull. Sing. 20 (1963) 239; Man. Dipt. Brun.
Suppl. (1968) 17. — D. nobilis Dyer, J. Bot. 12 (Apr.
1874) 106; ibid. (May 1874) 153.

a. ssp. Stellatus.

Young twig, raceme, leaf bud, stipule outside,
midrib on both surfaces and petiole + densely
persistently long rust-brown tomentose, sparsely so
on leaf nervation beneath. Twig to 7 mm @, +
angular, with an uneven cracked surface and large
round petiole scars. Leaf bud 5-8 by 3-5 mm,
spherical, obtuse. Stipule c. 2 by 0.8 cm, deltoid, acute.
Leaves 20-25 by 12-16 cm, ovate, thinly coriaceous;
base cordate; acumen to 14 mm long, nerves 12-16
pairs, well spaced, at c. 55°-60°; petiole 4-S cm long.
Raceme to 20 cm long, axillary, angular on drying,
unbranched or singly branched. Flower bud to 6 by 2
cm, large. Calyx and corolla typical, calyx glabrous.
Stamens c. 30, subequal; filaments slender, com-
pressed at base, tapering; anthers linear, + twice as
long as filaments, expanded into small lateral knobs at
base, tapering; appendage to connective filiform, as
long as anther, reaching almost to style apex. Ovary
small, ovoid; style c. 5xlength of ovary, stoutly
filiform, densely pubescent at base, sparsely so dis-
tally, glabrous in the apical 4. Fruit sessile or on to 2
mm long stout pedicel. Fruit calyx tube to S by 2 cm,
obconical, the nut enclosed in the distal half, the basal
c. 1.4cm a narrow central axis on which the wings are

fused; glabrous, smooth, c. 2.5 cm @ at neck (includ-
ing wings); wings c. 1.2 cm wide, thin, undulate, +
bent over on one side at maturity, continuous from the
base to the calyx lobes; concurrent with the margins of
the lobes; 2 longer lobes c. 12 by 3 cm, oblong, obtuse,
with undulate margin at the c. 1.7 cm wide base; with 3
parallel nerves, close to the centre, continuing to the
apex; 3 shorter lobes c. 1 cm long and broad, deltoid,
obtuse, with an undulate revolute margin.

Distr. Malesia: Borneo (Sarawak west of the
Lupar).

Ecol. Hill forests, 500-800 m, local.

Note. Immature plants of the two subspecies can
be indistinguishable.

b. ssp. parvus ASHTON, Gard. Bull. Sing. 20 (1963) 239;
Man. Dipt. Brun. Suppl. (1968) 170. — D. conformis
sensu ASHTON, Man. Dipt. Brun. (1964) 45; MEUER &
Woop, Sabah For. Rec. 5 (1964) 268.

Differs from ssp. stellatus as follows: Twig c. 2mm
@; leaves 10-15 by 5-7 cm, base obtuse, acumen to 8
mm long, petiole c. 2 cm long; racemes to 10 cm long.

Distr. Malesia: Borneo (Rejang eastwards to E.
Sabah, Tidung, and Belajan R.).

Ecol. Undulating land and hills to 700 m on
leached clay rich soil, including acid volcanics.

Vern. Kéruing bulu, k. daun nipis.

Note. Closely allied to D. concavus of E. Malaya,
which differs principally in the concave leaf.

47. Dipterocarpus sarawakensis [BROWNE, For. Trees
Sarawak & Brunei (1955) 111, nomen] Soot. Rein-
wardtia 5 (1961) 465, f. 2; KOCHUMMEN, Mal. For. 25
(1962) 163; ASHTON, Man. Dipt. Brun. (1964) 43, f. 6;
ibid. Suppl. (1968) 17, pl. 3 (bark).

Twig, leaf bud, raceme, stipule outside, midrib on
both surfaces, and nerves below densely persistently
tawny tomentose; nerves above and tertiary nerves
beneath sparsely tomentose; leaf margin setose. Twig
3-4 mm g, terete. Bud 6-10 by 3 mm, ovoid, acute.
Stipule c. 1.5 cm long, small, narrowly ovate, obtuse.
Leaves 5.5-8 by 3.5-5 cm, broadly obovate,
applanate, margin straight and revolute towards the
cuneate base, becoming sinuate towards the obtuse or
retuse apex; nerves 7-8 pairs, distant, ascending
(30°-35°); petiole 0.7-1.0 cm long, stout. Raceme to 7
cm long, unbranched or singly branched, terete.
Flower bud to 2.5 by 1 cm. Calyx and corolla typical,
calyx glabrous. Stamens 15, shorter than the style:
filaments short; anthers narrowly oblong, relatively
short, stout; appendage to connective stout at base,
tapering gradually, short, 3 length of anther. Ovary
globose, tomentose as also the narrowly cylindrical
style and stylopodium. Fruit subsessile, glabrescent:
calyx tube to 2.5 by 1.3 cm, fusiform, broadest
towards the base, tapering to the pedicel and more
gradually to the 6 mm @ neck; 5-winged, wings thin
and papery, striated, strongly undulate, to 8 mm
broad towards the apex, tapering to the pedicel or
terminating somewhat above it, tapering more rapidly
apically and joining with the base of the lobes; 2 longer

324

calyx lobes to 9 by 2 cm, oblong, obtuse; bases
constricted to c. 6 mm broad, with slightly revolute
margins, with 2 lateral nerves running up half the
length of the lobe and one median nerve dividing into
three after the termination of the laterals; 3 short lobes
0.8-1.5 by 0.5 cm, ovate, completely revolute, nar-
rowly acute.

Distr. Malesia: Malaya (river Kemaman, Treng-
ganu, one record), S. Borneo (Barito), Sarawak and
Brunel.

Ecol. Locally frequent on leached sandy soils on
low coastal hills to 400 m.

Vern. Kéruing layang.

48. Dipterocarpus coriaceus SLooT. Bull. Jard. Bot.
Btzg III, 8 (1927) 331, f. 11; Sym. Mal. For. Rec. 16
(1943) 171, f. 85; BROWNE, For. Trees Sarawak &
Brunei (1955) 108; ANDERSON, Gard. Bull. Sing. 20
(1963) 157; ASHTON, Man. Dipt. Brun. Suppl. (1968)
12, f. 2. — D. dyeri (non P1ERRE) Foxw. Mal. For. Rec.
10 (1932) 80, p.p.

Leaf beneath, petiole and twig shortly densely
evenly pale pink-brown pubescent; leaf bud, stipule
outside and midrib beneath, twig and petiole of young
tree with to 2 mm long dense hairs; stipule glabrous
within. Twig 8-13 mm g, stout, terete, with subhori-
zontal stipule scars; internodes 3-10 mm long, short.
Bud 2.5—3 by 1.0-1.7 cm, ovoid-deltoid; stipule to 4 by
2 cm, narrowly ovate, subacute. Leaves 16-21 by
10-15 cm, broadly elliptic-ovate, thickly coriaceous,
with obtuse or broadly cuneate base and obtuse to
subacute apex; nerves 14-16 pairs, at 20°—60°, promi-
nent beneath, depressed above, the midrib likewise;
tertiary nerves obscure, scalariform; petiole 4.5—6 cm
long, prominently geniculate. Flower and inflores-
cence unknown. Fruit pedicel to 7 mm long, promi-
nent. Fruit calyx glabrous; tube to 3 by 2.2
cm,+broadly ovoid, with 5, to 5 mm wide, stout
wings or ribs, tapering abruptly at the apex and
gradually towards the base, absent in the basal j of the
tube; 2 longer lobes to 14 by 2.5 cm, spatulate,
narrowly obtuse, to 5 mm broad above the tube; 3
shorter lobes to 14 by 2.5 cm, spatulate, narrowly
obtuse, to 5mm broad above the tube; 3 shorter calyx
lobes to 12 by 6 mm, elliptic, obtuse, applanate to
slightly revolute.

Distr. Malesia: Malaya (Lower Perak, Pahang),
Sumatra (Indragiri), Borneo (Lower Kapuas, Suka-
dana, Lower Dayak, and Lundu to K. Balingian in
Sarawak).

Ecol. Local, in Mixed Peat swamp forest.

Vern. Kéruing paya (Sar.), kédau, tampurau (S.E.
Borneo).

Note. Malayan collections have smaller fruit with
narrower wings on the tube, and glabrescent leaf
beneath in mature trees.

49. Dipterocarpus cuspidatus ASHTON, Gard. Bull.
Sing. 23 (1967) 261, pl. 2; Man. Dipt. Brun. Suppl.
(1968) 13, f. 2.

Leaf bud densely persistently long buff tomentose;

FLORA MALESIANA

[ser. I, vol. 97

twig, petiole, midrib above and leaf nervation beneath
sparsely long pale grey-brown subpersistent tomen-
tose. Twig c. 1 mm @ apically, slender, terete, much
branched; stipule scars slender, obscure. Bud to 7 by 3
mm, oblong; stipule unknown, caducous. Leaves 6-11
by 2-4 cm, narrowly elliptic-ovate, applanate with
plicate folding almost disappearing when fully
expanded; base obtuse to rarely cuneate; acumen to 2
cm long, prominent, slender, cuspidate; nerves 8-9
pairs, slender but prominent beneath, at 40°—50°;
tertiary nerves densely scalariform; midrib slender,
prominent beneath; petiole 12-18 mm long, slender,
geniculate. Flower unknown. Raceme to 5 cm long,
terminal or axillary, terete, slender, sparsely persis-
tently long grey-brown tomentose, singly branched.
Fruit pedicel to 5 mm long, prominent. Calyx gla-
brous; 2 longer lobes to 8 by 1.8 cm, spatulate, obtuse
to subacute, c. 6 mm broad at the tube; 3 shorter lobes
to 9 by 4 mm, oblong, obtuse, somewhat revolute;
tube to 2 cm long, to 1.8 cm @, with 5 prominent
wings; wings thin, somewhat undulate, tapering into
the base but to 9 mm broad and subauriculate
apically. Nut to 3 by 1.6 cm, ovoid, greatly exceeding
the length of the calyx tube.

Distr. Malesia: Borneo (N.E. Sarawak).

Ecol. Rare: Mixed Dipterocarp forest on undulat-
ing land and low hills.

Vern. Kéruing runching.

50. Dipterocarpus eurynchus Mig. Sum. (1862) 485;
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC. Prod. 16,
2 (1868) 613; WaALP. Ann. 7 (1869) 377; BuRcK, Ann.
Jard. Bot. Btzg 6 (1887) 203; BRANobiS, J. Linn. Soc.
Bot. 31 (1895) 40; SLoor. Bull. Jard. Bot. Btzg III, 8
(1927) 273, 302; ASHTON, Gard. Bull. Sing. 20 (1963)
238; Man. Dipt. Brun. (1964) 30, f. 6; ibid. Suppl.
(1968) 14; Gard. Bull. Sing. 31 (1978) 12.—D.
eurynchoides SCHEFF. Nat. Tijd. N.I. 31 (1870) 346;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 203; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 40; SLooT. Bull. Jard. Bot.
Btzg III, 8 (1927) 273, 302.— D. appendiculatus
ScuerFF. Nat. Tijd. N.I. 31 (1870) 347; ibid. 32 (1873)
407; Dyer, J. Bot. 12 (1874) 104, p.p.; BURCK, Ann.
Jard. Bot. Btzg 6 (1887) 200, p.p.; BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 34, p.p.; MERR. En. Born. (1921)
397, p.p.; HEYNE, Nutt. Pl. ed. 2 (1927) 1094; SLoor.
Bull. Jard. Bot. Btzg III, 8 (1927) 326; Reinwardtia 5
(1961) 458; Foxw. Mal. For. Rec. 10 (1932) 83, in obs.;
Sym. Mal. For. Rec. 16 (1943) 167, f. 85.—D.
basilanicus Foxw. Philip. J. Sc. 13 (1918) Bot. 179;
ibid. 67 (1938) 259; MerRR. En. Philip. 3 (1923) 88.
Leaf bud densely persistently long buff tomentose;
twig, petiole and leaf nervation beneath sparsely long
pale grey-brown subpersistent pubescent; stipule per-
sistently so. Twigs to 2 mm g apically, terete, smooth
or verrucose; stipule scars slender. Bud to 10 by 3mm,
conical, obtuse or subacute. Stipule to 3 by 0.8 cm,
narrowly oblong, subacute. Leaf 4-6(-10) by
2-3.5(—4.5) cm, elliptic to obovate; base cuneate; apex
subacute to shortly acuminate (caudate in young
trees); nerves 8-9 pairs, at c. 40°; tertiary nerves

1982]

slender, dense; petiole 6-9 mm long, slender. Flower
unknown. Raceme to 6 cm long, axillary, terete or +
compressed, glabrous, simple or singly branched.
Fruit calyx glabrous; tube to 1.7 by 2 cm, 5 winged;
wings c. 2.5 mm wide, thin, continuous from base to
apex, straight, rigid, frequently becoming bent over to
one side; 2 longer lobes to 8 by 2 cm, oblong, obtuse,
tapering and revolute at base; 3 shorter lobes to 7 by 5
mm, ovate, obtuse, becoming revolute.

Distr. Malesia: Malaya (E. coast, Trengganu to
N.E. Johore), Sumatra (E. Atjeh, Langsa, P. Singkep,
Banka), Borneo (W. Borneo, Sarawak, Brunei), S.
Philippines (rare).

Ecol. Local, on undulating land in Mixed Diptero-
carp forest on leached clay soils, and on ridge tops to
700 m.

Vern. Kéruing baran, k. padi (Mal.), sémanto
minjak, kérukéh, ansang ansang, kéruing, kéruing
senium (Sum.).

Note. A polymorphic species. One distinctive and
well collected segregate is recognised here as a sepa-
rate species, D. ochraceus; others, presently ill under-
stood, may eventually deserve taxonomic definition:
The few Malayan and Sumatran collections have
markedly larger leaves with longer petioles, and
collections from Gunong Angsi, Negri Sembilan
(including KEP 23788, quoted by SYMINGTON (1943)
as a possible hybrid between D. gracilis and D.
costatus) come close to D. ochraceus. This species is
apparently derived from the widespread D. costatus of
the seasonal zone, as may be the closely similar D.
glandulosus THw. of Ceylon.

51. Dipterocarpus ochraceus MEER, Acta Bot. Neerl.
12 (1963) 351, pl. 14; MEER & Woop, Sabah For.
Rec. 5 (1964) 26.

Large tree. Young parts densely ocherous velu-
tinate, persistent on leaf buds, stipules outside, and
twigs, becoming sparse yet partially persisting on
petiole and nervation beneath, caducous on other
known parts. Twigs c. 4mm @ apically, rather stout,
terete, + minutely fissured. Buds to 20 by 8 mm, large,
ovoid-lanceolate, acute; mature stipules unknown.
Leaves 10-19 by 4.5-9.5 cm, broadly elliptic-ovate,
coriacous, + persistently plicate; base cuneate or
rarely obtuse; acumen to 15 mm long, broad; nerves
11-15 pairs, slender but prominent beneath, ascend-
ing at 30°—-60°; tertiary nerves densely subscalariform,
slender but distinctly elevated beneath; midrib promi-
nent beneath, + applanate above; petiole 1.5—2.5 cm
long, stout. Inflorescences and flowers unknown. Fruit
pedicel to 3 by 2mm, prominent. Calyx tube to 18 by
15 mm, broadly ellipsoid, with 5, to 2 mm broad,
continuous straight coriaceous wings; 2 longer lobes
to 8 by 1.8 cm, lorate-spatulate, narrowly obtuse, c. 6
mm broad at base; 3 shorter lobes to 6 by 6 mm, ovate,
subacute, subrevolute.

Distr. Malesia: N.E. Borneo (Kinabalu area).

Ecol. Ultrabasic and basaltic mountains, 600-700
m.
Vern. Kéruing ranau.

DIPTEROCARPACEAE (Ashton)

325

Note. A species which has presumably segregated
from the widespread D. eurynchus; see there.

52. Dipterocarpus perakensis ASHTON, Gard. Bull.
Sing. 31 (1978) 13. — D. alatus (non Roxs.) Foxw.
Mal. For. Rec. 10 (1932) 89, p.p.; SYM. Mal. For. Rec.
16 (1943) 166, f. 85.

Large tree. Parts glabrous but for the cream
puberulent ovary apex. Twigs c. 2 mm @ apically,
much branched, blackish. Buds to 9 by 2 mm,
lanceolate, drying blackish; stipules unknown. Leaves
4-7 by 8-13 cm, elliptic, coriaceous; base broadly
cuneate to obtuse; apex shortly acuminate; nerves
9-12 pairs, ascending, prominent beneath; tertiary
nerves densely scalariform, slightly elevated beneath;
petioles 1.5—3.2 cm long, slender. Flowers and inflores-
cences unknown. Fruit pedicel to 2 by 2 mm; calyx
tube to 2.2 cm @, subglobose, with 5 prominent
continuous to 8 mm wide coriaceous wings; 2 longer
lobes to 12 by 3 cm, oblong-lorate, obtuse, tapering
abruptly to c. 8mm wide at the revolute base; 3 shorter
lobes to 5 by 5 mm, suborbicular, subrevolute,
small.

Distr. Malesia: Malaya (Penang hill, Dindings,
Pangkor I.).

Ecol. Very local, in Lowland Dipterocarp forest on
coastal hills.

Note. This species is a segregate from D. eurynchus
Mia. and D. costatus GAERTN. f., clearly distinguished
by the relatively broad wings on the fruit calyx tube
and conspicuous lack of tomentum.

53. Dipterocarpus philippinensis Foxw. Philip. J. Sc. 6
(1911) Bot. 253; ibid. 13 (1918) Bot. 179; ibid. 67 (1938)
263; MERR. En. Philip. 3 (1923) 90.

Tall tree. Leaf buds, twigs, stipules, leaf nervation
beneath, midrib above, petiole, inflorescence and
ovary densely evenly shortly persistently pale
ocherous pubescent, leaf surface sparsely so; fruit
calyx glabrous. Twigs 3-4 mm @ apically, + terete,
pale brown. Leaf buds to 15 by 6 mm, lanceolate;
expanded stipules not seen. Leaves 9-17 by 3.5-8 cm,
narrowly ovate, thinly coriaceous; base cuneate; apex
acute or shortly indistinctly acuminate; nerves 11—18
pairs, slender but prominent beneath, ascending at
45°; tertiary nerves densely scalariform, sinuate, very
slender but somewhat elevated beneath; petioles
2.54.5 cm long, slender. Jnflorescence to 8 cm long,
unbranched, bearing to 4 flowers. Flowers unknown.
Fruit pedicel c. 1 mm long, short; calyx tube to 2.3 cm
@, subglobose, with 5, to 8 mm broad, prominent
continuous coriaceous wings; 2 longer lobes to 14 by
2.8 cm, lorate, obtuse, c. 1 cm broad at the subrevolute
base; 3 shorter lobes to 12 by 14 mm, ovate, suborbi-
cular, revolute.

Distr. Malesia: Philippines (Luzon; Bataan and
Abra Prov.).

Ecol. Rare, in Mixed Dipterocarp forest in sea-
sonal areas.

Vern. Ayamban (IIk.).

326

FLORA MALESIANA

(ser. 1, velRg2

7 @d
> : A4

Fig. 27. Anisoptera costata Kortu. A. Bud, A/. petal, A2. stamens and pistil, 43. pistil in section, all x 5, A4.

stamens from inside (left) and outside, x 10. — A. thurifera(BLCo) BL. B. bud, B/. petal, B2. stamens and pistil,

B3. pistil in section, all x 5, B4. stamens from inside (left) and outside, x 10.— A. laevis RiDL. C. bud, C/.

petal, C2. stamens and pistil, C3. pistil in section, all x 5, C4. stamen from inside, x 10 (A PoILANE 830, B
KOSTERMANS & SOEGENG 286, C S 24810).

Dubious

54. Dipterocarpus orbicularis Foxw. Philip. J. Sc. 13
(1918) Bot. 180; ibid. 67 (1938) 262; MeRR. En. Philip.
3 (1923) 907.

Leaf buds, twigs, leaf nervation beneath and pedun-
cle + sparsely fulvous hirsute, lamina beneath, midrib
above and ovary sparsely puberulent; fruit calyx
glabrous. Twigs c. 5 mm @ apically, terete, pink-
brown. Leaf buds to 12 by 6 mm, ovoid-lanceolate;
stipules not seen. Leaves 9-13 by 6.5—10 cm, broadly
elliptic, thinly coriaceous; base cuneate; apex mu-
cronate; nerves 10-12 pairs, slender but prominent
beneath, shallowly depressed above, ascending at 45°;
tertiary nerves laxly subscalariform, elevated beneath;
petioles 2.5—4.5 cm long, slender. /nflorescences to 12
cm long, unbranched, bearing to three flowers.
Flowers and mature fruit unknown. Young fruit
pedicel to 2 mm long; calyx tube with 5 prominent

chartaceous broad wings from base to apex; calyx
lobes unequal, 2 aliform.

Distr. Malesia: Philippines (Camarines Prov.,
Luzon).

Ecol. Mixed Dipterocarp forest at low elevations.

Notes. The type (FB 21719 from Camarines,
Luzon) consists of a leafy twig and an immature fruit.

This little known species, which in the absence of
ripe fruit cannot be placed in the key, resembles D.
grandiflorus BLCo in its fruit and D. gracilis BL. in its
indumentum. The leaf, almost orbicular and with few
nerves, is distinctive. Sterile specimens from Sabah
compared by FoxwortTny with this species belong to
the subsequently described D. confertus SLOOT., in
which the leaves are much larger, the fruit calyx
densely pubescent and with narrower wings. D.
orbicularis would appear to be vicarious with D.
stellatus VESQUE of Borneo and D. concavus Foxw. of
the Malay Peninsula.

1982] DIPTEROCARPACEAE (Ashton) 327

2. ANISOPTERA

KortTHu. Kruidk. (1841) 65; WALP. Rep. 5 (1845) 125; Ann. 4 (1857) 335; BL. Ann.
Mus. Lugd.-Bat. 2 (1852) 6; Mia. FI. Ind. Bat. 1, 2 (1859) 500; B. & H. Gen. Pl. 1
(1862) 500; DC. Prod. 16, 2 (1868) 615; Dyer, FI. Br. Ind. 1 (1874) 300; Burck,
Ann. Jard. Bot. Btzg 6 (1887) 220; HEIm, Rech. Dipt. (1892) 30; BRANDis, J. Linn.
Soc. Bot. 31 (1895) 40; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 258;
GILG in E. & P. Pfl. Fam. ed. 2, 21 (1925) 359; SLoor. Bull. Jard. Bot. Btzg III, 8
(1926) 3; HEYNE, Nutt. Pl. ed. 2 (1927) 1098; Foxw. Philip. J. Sc. 67 (1938) 263;
Sym. Gard. Bull. S. S. 8 (1934) 1; Mal. For. Rec. 16 (1943) 199, f. 99 (map);
WYATT-SMITH, Mal. For. 18 (1955) 70; ASHTON, Gard. Bull. Sing. 20 (1963) 230;
Man. Dipt. Brun. (1964) 9; MEER & Woop, Sabah For. Rec. 5 (1965) 292;
SMITINAND, Thai For. Bull. (Bot.) 12 (1980) 18. — Hopea Roxs. FI. Ind. ed.
Carey 2 (1832) 611, p.p., quoad H. scaphula RoxsB. — Mocanera BLco, FI. Filip.
ed. 1 (1837) 446, p.p., quoad M. thurifera BLCo. — Vatica (non L.) Dyer, FI. Br.
Ind. 1 (1874) 301, p.p., quoad V. scaphula. — Antherotriche TuRcZz. Bull. Soc.
Nat. Mosc. 2 (1846) 505.— Scaphula PARKER in Fedde, Rep. 30 (1932)
326. — Hopeoides CretTzoiu, J. Jap. Bot. 17 (1941) 408. — Fig. 27-31.

Medium-sized to very large trees, often with prominent, thick, rounded, tall
straight buttresses. Crown comparatively small, appearing irregularly hemi-
spherical and oblong, rather diffuse, with a few large twisted branches ascending
from the bole apex; branchlets not crowded towards the apices. Bark surface
rather shallowly irregular-section fissured, the fissures separating + flat flaking
ridges; dotted with warty lenticels. Young parts at first + densely lepidote with
emarginate peltate hairs; lamina undersurface persistently so. Twigs ribbed.
Stipules relatively large, narrow, fugaceous. Leaves oblong to obovate, base
usually obtuse, apex shortly acuminate; nerves curving round and anastomosing
at the apices, thus forming a looped intramarginal nerve (cf. Cotylelobium);
tertiary nerves densely or remotely scalariform; petiole distinctly geniculate.
Inflorescence long, lax, pendent, densely tomentose; bracteoles small, linear,
caducous. Flower bud distinctly pedicellate, distichous. Ca/yx + imbricate, rarely
valvate (A. /aevis); with 2 obtuse outer lobes slightly more thickened than 3 inner
acute lobes, united at base in an indistinct (in flower) tube round the partially
inferior ovary. Petals oblong-linear, falling separately. Stamens 15-65 in 3
verticils or irregular, the outer somewhat shorter than the inner, glabrous;
filaments rather short, slender, filiform, tapering, connate at base; anthers with 4
pollen sacs, the inner 2 shorter than the outer 2, latrorse; appendage to connective
short or long. Ovary semi-inferior, with a distinct stylopodium; style long or
short, obscurely trifid; stigma indistinct. Fruit calyx valvate with an ellipsoid
basal tube almost entirely enclosing and adnate to the nut, with 2 long narrowly
spatulate obtuse untwisted 3-nerved lobes, and 3 acuminate short lobes. Nut +
globose, enclosed but for the apex in the valvate tube, with distinct stylopodium.
Germination epigeal, pericarp splitting open irregularly apically; cotyledons
unequal; first leaves paired, with interpetiolar stipules, or in a whorl of 4, without
stipules; followed by spiral leaves as other genera.

Distr. 11 spp. from Chittagong and Indochina to New Guinea, in Malesia: 10 spp. Holocene fossils have been
found in the Siwalik beds of N.W. India. Fig. 28.

328 FLORA MALESIANA [ser. I, vol. 9?

Ecol. Lowland forests, rarely above 1000 m. Scattered in Mixed Dipterocarp, Mixed Swamp and Heath
forests of the humid zone but becoming gregarious in N. Malaya, Indochina and parts of New Guinea.

Uses. The pale siliceous wood is not durable without preservatives and blunts saws but is an important source
for veneers.

KEY TO THE SPECIES

1. Flower buds lanceolate. Stamens 25-60. Stylopodium in flower as broad as ovary, ovoid to oblong,

tomentose. Style short, 3- or 6-furcate, stigma minute. Spp. 1-8 1. Sect. Anisoptera
2. Stamens c. 25.
3. Nerves at most 14 pairs; leaves fugaceous pubescent orepilose . . . . . . .41. A. marginata
3. Nerves at least 15 pairs, petiole and nervation beneath + persistently pubescent.
4. Leaf undersurface brilliant gold lepidote, nerves depressed above. . . .2. A. curtisii
4. Leaf undersurface dull yellowish or greenish lepidote; nerves hardly or not depressed above
SSeS costata

is Stamens at Teast 30.
5. Twigs and nerves and midrib beneath dark rufous scabrid tomentose; lamina golden yellow lepidote
beneath; nerves and tertiary nerves + prominently depressed above.

6. Nerves 18—28 pairs: stamens.¢.36 = 2 coe: 2. ie «e502 eee ee Rn ta
ae Nerves 23=33 pairs; stamensc.60 .-. =...) ... ss SS Amegistocanpa
5. Twigs and nervation not as above.
iG Nerves 9-14 pairs; twigs and leaves glabrescent . . . . . . 6. A. reticulata
7. Nerves (10—)14-18(—20) pairs; twigs, petioles and nervation beneath pubescent.
8. Stamens 37-57; leaves greyish to brown lepidote beneath . . . . . . . . 7. A. thurifera
8. Stamens 35-38; leaves densely golden peltate beneath . . . .. . . 8. A. aurea
. Flower buds subglobose. Stamens 15. Stylopodium broad and disc-like, glabrous. Style slender, columnar;
stigma obscurely 3-lobed. Spp. 9—10. 2. Sect. Glabrae
9. Leaf 8-16 by 3.5-8 cm; undersurface pale grey-green lepidote; nerves 13-18 pairs . . 9. A. scaphula
9. Leaf 7-11 by 34 cm; undersurface rust to golden lepidote; nerves 10-14 pairs . . . .10. A. laevis

1. Section Anisoptera

ASHTON, Gard. Bull. Sing. 20 (1963) 231; Man. Dipt. Brun. (1964) 10. — Anthero-
triche TuRCZ. l.c. — Anisoptera sect. Pilosae HEIM, Rech. Dipt. (1892) 33; Sym.
Mal. For. Rec. 16 (1943) 199. — Anisoptera sect. Antherotriche HEM, Rech. Dipt.
(1892) 34.

Young parts and leaf below generally tomentose. Flower buds lanceolate;
anther ellipsoid; appendage to connective many times longer than anthers,
slender; stylopodium cylindrical to ovoid, conical, narrow; style short, stigma
minute.

1. Anisoptera marginata KorTH. Kruidk. (1841) 66, t.
6, la—13a; WALP. Rep. 5 (1845) 124; BL. Mus. Bot.
Lugd.-Bat. 2 (1852) 42; Mia. Fl. Ind. Bat. 1, 2 (1859)
501; Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC.
Prod. 16, 2 (1868) 615; Burck, Ann. Jard. Bot. Btzg 6
(1887) 220; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 43;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1894)
258; BoERL. Cat. Hort. Bog. 2 (1901) 100; HEYNE,
Nutt. Pl. ed. 1, 3 (1917) 275; ed. 2 (1927) 1099; MErRR.
En. Born. (1921) 401; Ripv. Fl. Mal. Pen. 1 (1922) 21;
SLooT. Bull. Jard. Bot. Btzg III, 8 (1926) 5; Foxw.
Mal. For. Rec. 10 (1932) 96; Sym. Gard. Bull. S. S. 8
(1934) 13, pl. 3C; Mal. For. Rec. 16 (1943) 206, f. 101,
103; Burk. Dict. (1935) 163; WyaTT-SMiTH, Mal. For.
18 (1955) 76; ANDERSON, Gard. Bull. Sing. 20 (1963)
157; ASHTON, Gard. Bull. Sing. 20 (1963) 232; Man.
Dipt. Brun. (1964) 14, f. 5; ibid. Suppl. (1968) 5;
MEER & Woop, Sabah For. Rec. 5 (1964) 298, f.
Fig. 28. Density map of Anisoptera KorTH. in Male- 49B.— A. grandiflora BRANDIS, J. Linn. Soc. Bot. 31
sia; number of endemics above the hyphen, number of (1895) 43; Merr. En. Born. (1921) 401; SLoor. Bull.

non-endemics below it. Jard. Bot. Btzg III, 8 (1926) 10. — A. mindanensis (non

1982]

DIPTEROCARPACEAE (Ashton)

329

Foxw.) WYATT-SMITH, Mal. For. 18 (1955) 77,
p.p. — Fig. 29.

Young twig, leaf bud, stipule outside (glabrous
within), midrib beneath and petiole shortly caducous
pubescent; panicle persistently so. Twig terete, becom-
ing angular on drying, smooth or with very fine
striation. Bud 2.3—-3.5 by 2 mm, ovoid, acute, com-
pressed. Stipule c. 8 mm long, linear, obtuse. Leaf
7-10 by 3.5-4.5 mm, oblong to + obovate, densely
golden-brown lepidote beneath, bullate in young
trees, otherwise applanate, with obtuse or broadly
cuneate base; acumen to 6 mm long; nerves 10-14
pairs, slender, at c. 65—70° to the midrib; petiole 1.5—2
cm long, slender. Panicle to 14 cm long, terminal or
axillary, lax, pendent, terete; irregularly doubly
branched. Flower bud to 8 by 3.5 mm, lanceolate,
acute. Calyx cupped at base; lobes subequal, narrowly
deltoid, the 3 outer slightly narrower, acute, the 2
inner obtuse. Corolla pale yellow; petals broadly
elliptic, fimbriate. Stamens c. 25; filaments short,
slender; anther oblong; appendage to connective c. 2
times length of anther, reaching apex of stylopodium.
Stylopodium cylindrical, shortly densely tomentose;
style short, tapering, stout at base, the basal half
tomentose, otherwise glabrous, trifurcate. Fruit calyx
shortly pubescent; tube c. 1.3 cm @, globose, to 8 mm
@ at the neck; 2 longer calyx lobes to 12 by 2 cm,
spatulate, obtuse; 3 shorter lobes 1.5—2 cm long, c. 3
mm broad at base, linear, acute; apex of nut almost
flat, tapering abruptly to the c. 3mm long, c. 1.5mm @
oblong stylopodium.

Distr. Malesia: Malaya (E. and W. coast, Perak
and Pahang southwards), Banka, E. Sumatra (Asa-
han southwards to Lampong), Borneo.

Ecol. Widespread but rarely common, Mixed Peat
Swamp forest; local, Heath forest on podsols, to 1200
m.

Vern. Mérsawa paya (Mal.), pangiran kérangas
(Sabah), kétimpun, resak gunong (S. Borneo), ténam,
mentanam, sésawah, résak pantai, maségar (Sum.).

2. Anisoptera curtisii DYER ex KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 100, incl. var. latifolia KING;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895)
263, t. 122, f. F; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
42; Burk. J. Str. Br. R. As. Soc. 81 (1920) 51, 63, fig.;
Merr. En. Born. (1921) 400; RipL. Fl. Mal. Pen. 1
(1922) 218; Gite in E. & P. Pfl. Fam. ed. 2, 21 (1925)
259; SLooT. Bull. Jard. Bot. Btzg II, 8 (1926) 11, p.p.:
Foxw. Mal. For. Rec. 3 (1927) 79; ibid. 10 (1932) 99;
Philip. J. Sc. 67 (1938) 271; Sym. Gard. Bull. S.S. 8
(1934) 15, pl. 4B; Mal. For. Rec. 16 (1943) 204, f.
100B, 101.

Large buttressed tree. Twig apices, stipules, leaf
buds, nervation beneath, petiole, panicles and calyx
+ densely gold-brown pubescent, caducous on twigs
and calyx; leaf beneath brilliant gold lepidote. Twig c.
2 mm @ apically, slender, becoming pale brown;
stipule scars obscure. Bud to 6 by 4 mm, ovoid, acute;
stipules to 7 by 2 mm, linear-lanceolate, acute,
fugaceous. Leaves (4-)7—14 by (1.5—)2.5—6 cm, nar-

Fig. 29. Habit of Anisoptera marginata KorTH. Note

the colossal size by the person standing in front of the

flying buttresses. Palembang (Photogr. THORENAAR,
1925).

rowly elliptic to oblanceolate, coriaceous; base
obtuse; acumen to | cm long, broad; nerves 15-25
pairs, spreading, with short secondary nerves, slender
but prominent beneath, narrowly depressed above as
also the midrib; tertiary nerves laxly subreticulate, +
elevated beneath; petiole 13-23 mm long, slender.
Panicle to 18 cm long, terminal or axillary, lax,
pendent, irregularly branched; branchlets to 2.5 cm
long, bearing to 7 flowers. Flower white; buds to 5 by 2
mm, lanceolate; 2 longer sepals lanceolate, subacute; 3
shorter sepals broad at base, prominently slender
acuminate; stamens 25, subequal; filaments somewhat
shorter than anthers, filiform; anthers broadly
oblong, somewhat tapering; appendage very slender,
c. 3 times as long as anthers; stylopodium broadly

330

FLORA MALESIANA

[ser. I, vol. 97

>
Bai
|
Y
K

FEES CIALLE

Fig. 30. Anisoptera grossivenia SLOOT. a. End of twig, b. single leaf, c. fruit, d. nut, all x 5 (aS 5819, b-dS 6514).

ellipsoid-cylindrical, densely gold puberulent, sur-
mounted by the short but prominent trifid style. Fruit
pedicel to 3 by 2mm, expanding into calyx tube; calyx
tube to 14mm g, subglobose; 2 longer lobes to 10.5 by
1.8 cm, spatulate, obtuse, c. 5 mm broad at base; 3
shorter lobes to 20 by 3 mm, linear-lanceolate, acute.

Distr. Malesia: Malaya (commonest in north),
Sumatra (P. Singkep).

Ecol. Mixed Dipterocarp forest on well drained
periodically dry soils, especially coastal hills and
inland ridges to 700 m.

Vern. Mérsawa kuning, rengkong (Malaya); kéru-
ing kutjing, mérsawa (Singkep).

Note. Closely related to A. costata; some collec-
tions from N.W. Malaya suggest occasional hybridi-
sation.

3. Anisoptera costata KorTH. Kruidk. (1841) 67, t. 6,
f. 1-9; BL. Mus. Bot. Lugd.-Bat. 2 (1852) 42; Mia. FI.
Ind. Bat. 1, 2 (1859) 501; Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 85; DC. Prod. 16, 2 (1868) 615; PreRRE in
Lanessan, Pl. Util. Colon. Fr. (1886) 298: Burck,
Ann. Jard. Bot. Btzg 6 (1887) 220; BraNopis, J. Linn.
Soc. Bot. 31 (1895) 43; Ripv. Agr. Bull. Str. & F.M.S.
1 (1901) 60; J. Str. Br. R. As. Soc. 54 (1910) 25, p.p.; FI.
Mal. Pen. 1 (1922) 218, p.p.; MeRR. En. Born. (1921)

400: SLoot. Bull. Jard. Bot. Btzg III, 8 (1926) 7, f. 1;
Reinwardtia 2 (1952) 8, f. 1, p.p.; THORENAAR, Med.
Proefst. Boschw. 16 (1926) 106; HEYNE, Nutt. Pl. ed. 2
(1927) 1098; Foxw. Mal. For. Rec. 3 (1927) 79; ibid.
10 (1932) 97; Sym. Gard. Bull. S. S. 8 (1934) 9, pl. 3A;
Mal. For Rec. 16 (1943) 204, f. 100A, 101; WyatTrT-
SMITH, Mal. For. 18 (1955) 74; BACKER & BAKH. f. Fl.
Java 1 (1963) 330; ASHTON, Man. Dipt. Brun. (1964)
11, f. 5, pl. 4 (stem); Gard. Bull. Sing. 31 (1978)
13. — Dryobalanops hallii KORTH. msc. in BURCK,
Ann. Jard. Bot. Btzg 6 (1887) 220, 243, nomen in
syn. — Dipterocarpus parallelus KORTH. ex BURCK,
Ann. Jard. Bot. Btzg 6 (1887) 220, 243. — A. oblonga
Dyer, FI. Br. Ind. 1 (1874) 301; Prerre, Fl. For. Coch.
3 (1889) t. 235, 236; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 42; Ind. Trees (1906) 67; GUERIN, FI. Gén. I.-C.
1 (1910) 369; GAMBLE, Man. Ind. Timb. (1922) 75;
Sym. Gard. Bull. S. S. 8 (1934) 10, pl. 3B; Mal. For.
Rec. 16 (1943) 208, f. 101; PARKINSON, Ind. For. Rec.
20 (1935) 7. — Shorea nervosa KuRz [Rep. Pegu App.
A (1875) XVIII, nomen] Fl. Burma 1 (1877) 119. — A.
cochinchinensis PIERRE in Lanessan, Pl. Util. Colon.
Fr. (1886) 268; Fl. For. Coch. 3 (1889) t. 235A, 253;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 42; Herm, Bot.
Tidsskr. 25 (1902) 44; Guerin, Fl. Gen. I.-C. 1 (1910)
367; CrarB, Fl. Siam. Enum. | (1925) 139; LECOMTE,

1982]

DIPTEROCARPACEAE (Ashton)

331

Bois Indochine (1926) 115. — A. robusta Pierre, FI.
For. Coch. 3 (1889) t. 236; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 42; GugrIN, Fl. Gén. I.-C. 1 (1910) 368;
CRAIB, Fl]. Siam. Enum. | (1925) 140; Sym. Gard. Bull.
S.S. 8 (1934) 11. — A. glabra (non KURZ) PIERRE, FI.
For. Coch. 3 (1889) t. 235B; GuERIN, FI. Gén. I.-C. 1
(1910) 361 (fig.), 369. — A. marginatoides Hem, Bot.
Tidsskr. 25 (1902) 44; GuERIN, FI. Gén. I.-C. 1 (1910)
369; CRAIB, Fl. Siam. Enum. 1 (1925) 139.— A.
mindanensis Foxw. Philip. J. Sc. 13 (1918) Bot. 181:
ibid. 67 (1938) 266; Merr. En. Philip. 3 (1923)
92. — Fig. 27A—-A4.

Twig of variable thickness, at first frequently
angular, becoming minutely striated or smooth,
terete. Bud 3-5 by 1.5-3 mm, ovoid, somewhat
compressed, acute. Stipule c. 8 by 3 mm, hastate,
acute, fugaceous. Leaves 6-18 by 7-11 cm, thinly
coriaceous, frequently slightly bullate, oblong to
obovate, undersurface grey-green lepidote to golden
or chocolate; base obtuse or broadly cuneate; acumen
to 5mm long; margin not revolute or only slightly so;
nerves 8-22 pairs, at 60°-70°; petiole 24 cm long.
Panicle to 20 cm long, terminal or axillary, angular,
pendent, doubly or trebly branched, branchlets bear-
ing up to 5 flowers; bracteoles to 4 by 2 mm, hastate,
acute, shortly densely pubescent. Flower bud to 12 by 6
mm, ovoid, acute. Calyx densely tomentose outside,
shortly pubescent within; lobes deltoid, + acute,
acuminate, of varying lengths. Corolla cream; petals
broadly hastate, acute, shortly puberulent on parts
exposed in bud. Stamens c. 25 (to 35 outside Malesia),
subequal; filaments short, compressed; anther oblong,
tapering apically; appendage to connective about
twice as long as anther, filiform, tapering, reaching,
almost to apex of stylopodium. Stylopodium cylindri-
cal, somewhat tapering, densely pubescent; style
short, trifid, pubescent at base, otherwise glabrous.
Fruit calyx shortly pubescent, tube glabrescent; tube
to 1 by 1.2 cm, globose, tapering gradually to the
pedicel, narrowed to 8 mm @ at the neck; 2 longer
calyx lobes to 16 by 1.5—2 cm, spatulate, obtuse, c. 5
mm broad at base; 3 shorter lobes to 20 by 4 mm,
variable, hastate, base slightly constricted. Nut apex
broadly conical, crowned by ac. 2.5 by 1.5mm oblong
stylopodium, shortly pubescent.

Distr. Burma, Thailand, Cambodia, Cochinchina,
and in Malesia: Malaya, Sumatra, W. Java (one
record), Borneo (S.E. Kalimantan, Sabah, Brunei,
N.E. Sarawak), Philippines (Mindanao, one record).

Ecol. Common, often gregarious, in Semi-ever-
green Dipterocarp forest and evergreen forest in
seasonal areas; rare but widespread in lowland forest
in everwet areas, to 700 m.

Vern. Mérsawa, m. késat, m. térbak, térbak, mér-
anti térbak, pokok pahit (Malaya), maségar, ténan,
méluwang tikus (Sumatra), kétimpun, laripung, damar
tingkis (Kalimantan).

Notes. A very variable species; forms vary from
epilose (A. mindanensis Foxw.) to densely pilose (A.
robusta PIERRE), another has relatively small few-nerved
leaves (Anisoptera sp. A. of Malayan literature).

As now understood, A. costata is distinguished by
its 25 stamens and generally grey-pubescent leaf with
generally at least 15 pairs of nerves. It occurs from
Mindanao (one definite record) through Borneo,
Java, Sumatra, and Malaya to Chittagong, Burma,
Thailand and Indochina. In Mindanao it is epilose but
for petals and ovary; in Borneo it has relatively small,
chartaceous, sparsely grey-brown pubescent leaves
with 15-20 pairs of nerves; in Java, Sumatra and
sometimes Malaya the leaves are similar but some-
what larger, with 22 pairs of nerves. In Malaya the
species becomes more variable however, usually being
relatively large-leaved, with a rather dense, often
golden tomentum; in the northwest a small, golden-
tomentose leaved form prevails with as few as 8 pairs
of nerves. In the Indochinese region the species varies
much in leaf size and tomentum, and in more seasonal
areas becomes shortly deciduous. No clear geographi-
cal differentiation is discernible, though variation is
greatest in southern Indochina. In summary, varia-
tion appears on the whole as continuous as in a
panmictic population, with geographically localised
forms appearing in the less seasonal areas.

There is a possibility of hybridisation with A.
curtisii in N.W. Malaya, and with A. megistocarpa in
Johore and Singapore (large-leaved golden tomentose
forms).

VAN SLOOTEN (1952) recorded A. costata from
Celebes and the Moluccas on the basis of sterile
material. It is indistinguishable from A. thurifera
(BLco) BL. when sterile; for phytogeographical rea-
sons I would prefer to tentatively associate these
numbers with the latter species.

4. Anisoptera grossivenia SLOOT. Bull. Jard. Bot. Btzg
III, 16 (1940) 431, f. 1; Wyatr-SmitH, Mal. For. 18
(1955) 75; BROWNE, For. Trees Sarawak & Brunei
(1955) 93; ASHTON, Gard. Bull. Sing. 20 (1963) 232:
Man. Dipt. Brun. (1964) 12, f. 5, pl. 3 (habit); ibid.
Suppl. (1968) 5; MEER & Woop, Sabah For. Rec. 5
(1964) 296, f. 50. — A. curtisii (non DYER ex KING)
Merr. En. Born. (1921) 400, p.p., quoad spec. Born.;
SLoot. Bull. Jard. Bot. Btzg III, 8 (1926) 11, p.p.,
quoad spec. Born. — Anisoptera sp. ‘B’ WYATT-SMITH,
Mal. For. 18 (1955) 79. — Fig. 30.

Leaf nervation beneath, petiole, stipule (outside,
shortly pubescent within), leaf bud, panicle and twigs
persistently rufous scabrid tomentose; lamina beneath
densely rich golden-yellow lepidote. Twig to 2mm g
apically, terete or + compressed, rugose when dry,
becoming minutely striated. Bud 3-5 by 2.5 mm,
ovoid, + compressed. Stipule c. 8 by 2.5 mm, hastate,
acute, fugaceous. Leaves 9-12 by 3—5S cm, oblong to
narrowly obovate, slightly bullate; base cuneate;
margin slightly recurved; acumen to | cm long,
narrow; nerves 18—28 pairs, dense, arched, at 60°—70°;
tertiary nerves conspicuous, semi-reticulate; looped
intramarginal nerve c. 1 mm from margin; petiole
2-2.5 cm long, geniculate. Panicle to 20 cm long,
terminal or axillary; slender, lax, pendent, angular;
regularly doubly branched, branchlets lax, to 5 cm

337)

FLORA MALESIANA

(ser. I, vol. 9?

long, bearing up to 8 flowers; bracteoles fugaceous.
Bud to 5 by 2.5 mm, lanceolate, acute. Calyx densely
rust-brown tomentose, cupped at the base; sepals
imbricate, acuminate; 2 inner sepals slightly longer,
broader, obtuse; 3 outer acute. Corolla magenta at
first, becoming rich yellow on opening; petals lanceo-
late, acute, shortly pubescent on parts exposed in bud.
Stamens c. 36; filaments slender, short; anthers sub-
globose; appendage to connective c. 3x length of
anther, reaching 3 height of stylopodium, slender.
Ovary half enclosed in calyx; stylopodium subcylin-
drical, tapering apically, densely shortly gold-brown
tomentose; style short, glabrous; stigma minute. Fruit
calyx sparsely puberulent; tube to 1.3 cm @, globose,
hardly constricted at the neck; two longer calyx lobes
to 20 by 2.5 cm, narrowly spatulate, obtuse, rarely
acute, to 5 mm broad at base; 3 shorter lobes of
varying length, occasionally to 6 by 0.4 cm, acute,
linear. Nut apex obtuse, with c. 8 mm long, c. 15 mm
@, acute oblong-ovoid stylopodium broadening into
the nut at the base; shortly densely rufous pubescent.

Distr. Malesia: Borneo (S.,S.E. & W. Kalimantan,
Sarawak, Brunei, S.W. Sabah).

Ecol. Lowland Dipterocarp forest on sandy soil,
on present or Pleistocene coastal hills.

Vern. Mérsawa, m. durian (Mal.), bénchaloi
(Brun.), péngiran késar (Sabah), bérbakau, mérbani,
pényau kérabak, p. rébong, p. batu, résak témbaga,
kélassih, képitun, ampereng, marlangat, chéngal padi.

5. Anisoptera megistocarpa SLooT. Bull. Jard. Bot.
Btzg III, 8 (1926) 12, f. 2; HoLtrum, Gard. Bull. S.S.5
(1931) 184; Foxw. Mal. For. Rec. 10 (1932) 98; Sym.
Gard. Bull. S. S. 8 (1934) 14, pl. 4A; Mal. For. Rec. 16
(1943) 207, f. 101; Burk. Dict. (1935) 164.— A.
costata (non KorTH.) RIDL. J. Str. Br. R. As. Soc. 54
(1910) 25, p.p.; Fl. Mal. Pen. | (1922) 218, p.p.; BURK.
J. Str. Br. R.. As. Soc. 75 (1917) 43.

Tall buttressed tree. Twigs, leaf bud, stipule out-
side, petiole, nervation beneath, panicle, flower calyx
and ovary densely persistently dark ferruginous sca-
brid pubescent; lamina beneath dark golden brown
lepidote. Twig c. 5 by 3 mm @ apically, stout, +
compressed, rugose, becoming dark brown verrucose.
Buds to 8 by 5 mm, ovoid; stipules to 12 by 6 mm,
fugaceous. Leaves (5.5—)9-20 by (2.5—)3.5—8 cm, nar-
rowly oblong-elliptic to oblanceolate, thickly coria-
ceous; margin subrevolute; base obtuse; acumen to |
cm long, tapering, downcurved; nerves 23—33 pairs,
dense, spreading, with short secondary nerves, slender
but prominent beneath, distinctly depressed above as
also the midrib and subreticulate tertiaries; petiole
2-2.8 cm long, to 3 mm @, stout. Panicle to 12 cm
long, terminal or axillary, pendent, irregularly later-
ally branched; branchlets to 3.5 cm long, bearing to 5
flowers. Flower white; buds to 8 by 3 mm, lanceolate; 2
longer sepals lanceolate, subacuminate; 3 shorter
sepals narrowly deltoid, subcaudate. Stamens c. 60,
subequal; filaments shorter than anther, filiform;
anthers oblong, attenuate; appendages filiform, twice
length of anthers. Stylopodium stoutly ovoid, pubes-

Fig. 31. Trunk-base of Anisoptera thurifera (BLCO) BL.
Sogeri, Papua (Photogr. ASHTON, Aug. 1970).

1982]

cent, surmounted by a short but distinctly trifid style.
Fruit pedicel to 3 by 2 mm. Calyx tube to 2 cm @,
globose; 2 longer lobes to 22 by 3.8 cm, spatulate,
obtuse, c. 6 mm wide at the base; 3 shorter lobes to 30
by 3 mm, linear.

Distr. Peninsular Thailand, and in Malesia:
Malaya (south from Perak and Pahang), Sumatra
(Langsa, Atjeh).

Ecol. Scattered in Mixed Dipterocarp forest on
undulating land and low hills.

Vern. Mérsawa merah, m. api,
(Malaya); beurmen (Sumatra, Atjeh).

sépah petri

6. Anisoptera reticulata ASHTON, Gard. Bull. Sing. 22
(1967) 260, pl. 1; Man. Dipt. Brun. Suppl. (1968) 5, f. 1.

Leaf, buds, and twig fugaceous golden puberulent.
Twig c. 2 mm @ apically, terete, rugulose, dark
chocolate-brown; internodes c. 1.5 cm long; stipule
scar short, horizontal. Leaf bud 2 by 2 mm, ovoid,
subacute. Stipule unknown. Leaf 4.5—13 by 2.2-5.5
cm, elliptic-obovate, coriaceous; base broadly
cuneate; apex tapering abruptly to a c. 5 mm long,
short acumen; nerves 9—14 pairs, prominent beneath,
arched, at 50°—55°; tertiary nerves subreticulate; mid-
rib applanate or somewhat depressed above, promi-
nent beneath. Petiole 15-35 mm long, c. 2 mm g,
terete, prominently geniculate and swollen distally,
drying black. Panicle to 6cm long, 2mm g at base (in
fruit); terminal or axillary, short, terete, + persis-
tently golden pubescent, singly branched. Bud to 8 by
3 mm, fusiform. Calyx lobes narrowly deltoid, un-
equal, valvate. Petals cream, lorate, pubescent on
parts exposed in bud. Stamens c. 35, subequal:
filaments short, compressed; anthers oblong,
attenuate; appendage to connective filiform. Stylopo-
dium oblong, obtuse, pubescent. Fruit pedicel 4 mm
long, c. 3 mm @g, stout, prominent. Calyx sparsely
caducous golden pubescent; tube to 2 cm @, subglo-
bose; 2 longer calyx lobes to 13 by 3 cm, lorate-spatu-
late, obtuse, tapering to c. 9mm broad above the tube;
3 shorter lobes to 20 by 3 mm, linear-lanceolate, acute.
Nut apex densely persistently yellow-brown pubes-
cent, crowned by a c. 4 mm long, 3 mm g, oblong
obtuse pubescent stylopodium.

Distr. Malesia: Borneo (N.E. Sarawak, Brunei,
S.W. Sabah).

Ecol. Rare, Mixed Dipterocarp forests on sandy
soils.

7. Anisoptera thurifera (BLCO) BL. Mus. Bot. Lugd.-
Bat. 2 (1852) 42; Wacp. Ann. 4 (1857) 336; DC. Prod.
16, 2 (1868) 615; VIDAL, Sinopsis (1883) t. 14, f. E;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 44, t. 2, f. 28:
BRANDIS & GiLG in E. & P. Pfi. Fam. ed. 1, 3, 6 (1895)
258; MERR. & ROLFE, Philip. J. Sc. 3 (1908) Bot. 115;
Merritt, For. Bur. Bull. Philip. 8 (1908) 48; Wuit-
FORD, Philip. J. Sc. 4 (1910) Bot. 703; For. Bur. Bull.
Philip. 10 (1911) 78; Foxw. Philip. J. Sc. 6 (1911) Bot.
256; ibid. 13 (1918) Bot. 181; ibid. 67 (1938) 267;
Merk. Sp. Blanc. (1918) 269; En. Philip. 3 (1923) 92;
Reyes, Philip. J. Sc. 22 (1923) 323; SLoor. Bull. Jard.

DIPTEROCARPACEAE (Ashton)

338

Bot. Btzg III, 8 (1926) 4; Sym. Gard. Bull. S. S. 8
(1934) 6, pl. 4C; AsHTton, Gard. Bull. Sing. 31 (1978)
15. — Mocanera thurifera Buco, FI. Filip. ed. 1 (1837)
446. — Dipterocarpus thurifer BLco, Fl. Filip. ed. 2
(1845) 310; ibid. ed. 3, 2 (1878) 212, t. 264; DC. Prod.
16, 2 (1868) 614. — Fig. 27B—B4, 31.

Notes. A. thurifera is now seen to be the eastern
analogue of A. costata, with a dstribution — if sterile
collections from Celebes and the Moluccas are inclu-
ded — from northern Luzon throughout the Philip-
pines to Celebes, the Moluccas and New Guinea. In
the Philippines geographically defined variation
occurs and a separate species has evolved within the
aseasonal parts of its range. In New Guinea local
variation is great and collections are presently inade-
quate to define geographical forms.

It is interesting that the species is known in both the
Philippines and New Guinea to be the only diptero-
carp which readily reinvades cultivated land.

I recognize the Philippine and New Guinea popula-
tions as geographical subspecies.

KEY TO THE SUBSPECIES

1. Leaves oblanceolate to lanceolate, prominently
acuminate. Stamens 35-47 . a. ssp. thurifera

1. Leaves obovate. Stamens 37-57
b. ssp. polyandra

a. ssp. thurifera. — Mocanera thurifera BLco, FI.

Filip. ed. 1 (1837) 446. — Mocanera mayapis BLco, /.c.

449. — Dipterocarpus thurifer BLco, FI. Filip. ed. 2

(1845) 310. — Dipterocarpus mayapis Buco, FI. Filip.

ed. 2 (1845) 313; DC. Prod. 16, 2 (1868) 610; Dyer, J.

Bot. 12 (1874) 108; BrRaANpis, J. Linn. Soc. Bot. 31

(1895) 40; Merr. Publ. Gov. Lab. Philip. 27 (1905) 21.

— Antherotriche lanceolata TuRCZ. Bull. Soc. Nat.

Mosc. 2 (1846) 515; WaALpP. Ann. | (1848) 113. — A.

lanceolata WALP. Ann. | (1848) 113; DC. Prod. 16, 2
(1868) 616; VIDAL, Phan. Cuming. (1885) 97; F.-VILL.

Nov. App. (1880) 20; BRANDIS & GILG in E. & P. Pfi.

Fam. ed. 1, 3, 6 (1895) 263, fig.; GiLG in E. & P. Pf.

Fam. ed. 2, 21 (1925) 259, t. 112. — Shorea mayapis
BL. Mus. Bot. Lugd.-Bat. 2 (1852) 33; WALP. Ann. 4
(1857) 338; DC. Prod. 16, 2 (1868) 632. — A. oblonga
(non DYER) F.-VILL. Nov. App. (1880) 20; VIDAL,
Rev. Fl. Vasc. Filip. (1886) 60. — Dipterocarpus
turbinatus (non GAERTN. f.) F.-VILL. Nov. App. (1880)
20. — A. vidaliana BRANDIS, J. Linn. Soc. Bot. 31

(1895) 44; PERK. Fragm. FI. Philip. (1904) 23; MrrR.
Philip. J. Sc. 1 (1906) Suppl. 97. — A. tomentosa
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 45. — A.
calophylla PERK. Fragm. Fl. Philip. (1904) 22. — A.
brunnea Foxw. Philip. J. Sc. 6 (1911) Bot. 254, pl. 40;-
ibid. 13 (1918) Bot. 181; ibid. 67 (1938) 270; MErRR. En.
Philip. 3 (1923) 92.

Tall or medium-sized, buttressed tree. Twigs, leaf
buds, stipules, petioles and leaves beneath + densely
persistently grey-green or pale to chocolate-brown
lepidote; panicles, flower calyx and ovary densely
grey-brown puberulent; panicle and calyx becoming

334

sparse or glabrescent in fruit. Twig c. 3mm @ apically,
terete, rugulose, pale brown. Leaf bud to 4 by 2 mm,
lanceolate; stipules to 8 mm long, linear. Leaves 6-15
by 2.5—6.5 cm, thinly coriaceous, elliptic to lanceolate
or obovate-oblanceolate, + coriaceous; base broadly
cuneate or obtuse; acumen to 1.3 cm long, slender,
down-curved and twisting over on pressing; nerves
(12-)14-18(—20) pairs, slender but distinctly elevated
beneath, less so above (as also the reticulate tertiary
nerves), arched, at 55°—80°, with or without short
secondary nerves; midrib prominent beneath, ob-
scure, depressed, above; petiole 1.7-3.5 cm long,
slender. Panicles to 20 cm long, terminal or subter-
minal axillary, lax, pendent; singly branched, branch-
lets bearing to 11 flowers. Flower buds to 9 by 3 mm,
lanceolate. Sepals narrowly deltoid; 2 longer sub-
acute, 3 shorter prominently acuminate. Stamens
45-47, subequal; filaments short, slender, filiform;
anthers narrowly oblong, somewhat tapering; appen-
dages very slender, + twice length of anthers. Stylo-
podium narrowly ellipsoid-cylindrical, puberulent dis-
tally, with prominent trifid style. Fruit pedicel to 3 mm
long, short. Calyx tube to 17 mm @, globose; 2 longer
lobes to 15 by 1.5 cm, spatulate, narrowly obtuse, c. 4
mm wide at base; 3 shorter lobes to 30 by 3 mm, linear.
Stylopodium short, conical.

Distr. Malesia: Philippines.

Ecol. Evergreen and Semi-evergreen Dipterocarp
forests below 750 m; common and often gregarious,
regenerating in secondary forest.

Vern. Mayapis, palosapis, palohapi, dagang, afu.

b. ssp. polyandra (BL.) ASHTON, Gard. Bull. Sing. 31
(1978) 16. — A. polyandra BL. Mus. Bot. Lugd.-Bat. 2
(1852) 42, t. 6; WALP. Ann. 4 (1857) 335; Mia. FI. Ind.
Bat. 1, 2 (1859) 501; Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 84; DC. Prod. 16, 2 (1868) 615; ScHEFF. Ann.
Jard. Bot. Btzg 1 (1876) 9; F.v.M. Descr. Not. App. 6
(1887) 97; Burck, Ann. Jard. Bot. Btzg 6 (1887) 220;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 45; Diets, Bot.
Jahrb. 57 (1922) 461; LANE-POOLE, For. Res. (1925)
22, 33, 119, 167; SLooT. Nova Guinea 14 (1926) 222;
Bull. Jard. Bot. Btzg III, 8 (1926) 15; Reinwardtia 2
(1952) 11, f. 2; WHITE & FRANCIS, Proc. R. Soc.
Queensl. 38 (1927) 247. — Dipterocarpus parviflora
Zipp. Flora 12 (1829) 281; Alg. Konst- & Letterbode
1, 19 (1829) 296; Bijdr. Natuurk. Wet. 5 (1830) 178;
Mia. FI. Ind. Bat. 1, 2 (1859) 501, nomen. — Diptero-
carpus microcarpus Zipp. ex Miq. Fl. Ind. Bat. 1, 2
(1859) 501, nomen in syn. — Anisoptera sp. Dyer, J.
Bot. 16 (1878) 99; BurckK, Ann. Jard. Bot. Btzg 6
(1887) 221; BRANDis, J. Linn. Soc. Bot. 3 (1895) 45;
Diets, Bot. Jahrb. 57 (1922) 461.— <A. forbesii
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 45; Dies, Bot.
Jahrb. 57 (1922) 491; Baker f. J. Bot. 61, Suppl. (1923)
5. — A. kostermansiana Ditmy, Reinwardtia 3 (1956)
347; ibid. 5 (1960) 267.

Differing as follows: Leaves 8.5—-18 by 3.5-8.5 cm,
elliptic to obovate, + coriaceous; base cuneate,
obtuse or cordate; apex shortly acuminate; nerves
(10—)12-14(-17) pairs, slender but distinctly elevated

FLORA MALESIANA

[ser. I, vole9?

on both surfaces. Stamens 37—S7. Fruit pedicel to 4 by
2mm, prominent; 2 longer calyx lobes to 7 by 0.6 cm; 2
shorter lobes to 7 by 0.6 cm.

Distr. Malesia: New Guinea, and possibly (sterile
material) from Celebes and Moluccas (Morotai, Hal-
mahera, Batjan, Obi and Aru Is.).

Ecol. Common, often gregarious, in lowland forest
on undulating land and hills to 600 m, especially on
sedimentary rocks and along ridges; regenerates pro-
fusely in secondary forest.

Vern. Baoti, tolu (Celebes), bolam, kora, hate besi
(Morotai), kara, hirus, kopodaka, owiru, kako (Hal-
mahera), damar utan, d. hiru, asamban, gawi, wewe
pérémpuan (Batjan), kora (Obi), jamar, doka (Aru),
aran marei, wuku, taire, damar papan, taai, baurai,
maniuri, armanuri, kansiopi, ansiopi, merait, kandau,
karalaka, garawa, karawa, warawa, balia, ordima,
barida (New Guinea).

Note. This species is extremely variable, paralleling
A. costata (q.v.). The range of variation in tomentum
is comparable in both species.

8. Anisoptera aurea Foxw. Philip. J. Sc. 67 (1938) 271,
pl. 1-2.— A. curtisii (non DYER ex KING) Foxw.
Philip. J. Sc. 6 (1911) Bot. 255, pl. 41; ibid. 13 (1918)
Bot. 181; WHITFORD, Bull. Bur. For. Philip. 10 (1911)
78; Merr. En. Philip. 3 (1923) 92; SLoort. Bull. Jard.
Bot. Btzg III, 8 (1926) 11; Sym. Gard. Bull. S. S. 8
(1934) 17.

Large tree with pale shaggily flaky bark. Leaf
undersurface densely golden lepidote; nerves and
midrib beneath, leaf buds, petiole, twigs and ovary
densely persistently pale brown puberulent; panicles
and fruit calyx sparsely so; parts of petals exposed in
bud densely greyish puberulent. 7Jwigs c. 3 mm @,
becoming terete, pale brown, rugulose. Leaf buds to 4
by 2 mm, lanceolate, acute; stipules not seen. Leaves
7-11 by 2.5-5.5 cm, oblong or oblanceolate, thinly
coriaceous; base cuneate or sometimes obtuse;
acumen to | cm long, slender, prominent, down-
curved and bending over on pressing; nerves
(15-)18-20 pairs, slender but distinctly elevated
beneath, less so but distinct above, at 65°—80°;
secondary nerves obscure or absent; tertiary nerves
densely reticulate, evident on both surfaces; midrib
slender but prominent beneath, obscure, deeply
depressed, above; petioles 15-27 mm long, slender,
prominently geniculate. Panicles to 12 cm long,
slender, lax, pendent, terminal or axillary, singly (if
axillary) or doubly branched; branchlets to 15 mm
long, bearing to 3 flowers. Flower buds to 6 by 3 mm.
Sepals narrowly deltoid, 2 longer subacute, 3 shorter
prominently acuminate; stamens 35-38, subequal;
filaments + equal to the oblong somewhat tapering
anthers, filiform; appendage very slender, c. 1? times
as long as anthers; stylopodium broadly ellipsoid-
cylindrical, densely golden stellate-puberulent,
crowned by the prominently trifid acute style. Fruit
pedicel to 4 mm long, slender; ca/yx tube to 7 mm @
subglobose; 2 longer lobes to 10 by 1.5 cm, narrowly
spatulate, narrowly obtuse, c. 3 mm wide at base; 3

1982] DIPTEROCARPACEAE (Ashton) 335

shorter lobes to 12 by 2 mm, linear; stylopodium
cylindric, tapering, subacute.

Distr. Malesia: Philippines (Luzon:
Camarines: Polillo).

Ecol. Mixed Dipterocarp forest, especially on
ridges to 200 m, in everwet zone.
Vern. Dugong, manapo,

(Laguna).

Quezon; malahapi, dagong

2. Section Glabrae

Hem, Rech. Dipt. (1892) 33; Sym. Gard. Bull. S. S. 8 (1934) 1; Mal. For. Rec. 16
(1943) 199; AsHTON, Gard. Bull. Sing. 20 (1963) 232; Man. Dipt. Brun. (1963)
10. — Scaphula PARKER in Fedde, Rep. 3 (1932) 326. — Hopeoides CrETZOWU, J.
Jap. Bot. 17 (1941) 408.

Young leaves and twigs epilose. Flower buds globose; anthers linear;
appendage to connective less than 5 length of anther, stout; stylopodium a
flattened discoid platform surmounting the ovary; style filiform, long, with

distinct trifid stigma.

9. Anisoptera scaphula (ROxB.) KURZ, Fl. Burma |
(1877) 547, 2 (1877) 586; PireRRE, Fl. For. Coch. 3
(1889) sub t. 235; Sym. Gard. Bull S.S. 8 (1934) 2, 4, pl.
1; Mal. For. Rec. 16 (1943) 209, f. 101, 104. — Hopea
scaphula Roxs. (Hort. Beng. (1814) 93, nomen] Fl.
Ind. ed. Carey 2 (1932) 611; WALP. Rep. 5 (1845) 128;
DC. Prod. 16, 2 (1868) 635; KURZ, Prelim. Rep. For.
Pegu (1875) App. A, 19, App. B, 29; Fl. Burma 1
(1877) 121, 547. — A. glabra Kurz, J. R. As. Soc.
Beng. Sc. 52, 2 (1873) 61; Prelim. Rep. For. Pegu
(1875) App. A, 16, App. B, 29; Fl. Burma 1 (1877) 112;
Dyer, FI. Br. Ind. 1 (1874) 301; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 41; Ind. Trees (1906) 67, t. 29; GUERIN,
Fl. Gén. I.-C. 1 (1910) 368. — Vatica scaphula Dyrr,
Fl. Br. Ind. 1 (1874) 301; KING, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 127; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
132; Ind. Trees (1906) 72; GAMBLE, Man. Ind. Timbers
(1922) 84. — A. thurifera (non BL.) RIDL. FI. Mal.
Pen. 1 (1922) 219; Foxw. J. Mal. Br. R. As. Soc. 5
(1927) 341, p.p.; Mal. For. Rec. 3 (1927) 78, p.p.; ibid.
10 (1932) 100. — Scaphula glabra PARKER in Fedde,
Rep. 30 (1932) 326; CreTzoru, Act. Faun. Flor. Univ.
Bot. 1, 9 (1933) 3; PARKINSON, Ind. For. Rec. 20, 15
(1935) 11. — Hopeoides scaphula Crerzow, J. Jap.
Bot. 17 (1941) 408.

Very tall buttressed tree. Parts glabrous but for the
greyish green lepidote leaf undersurface and young
parts. Twig c. 2 mm @ apically, slender, pale grey-
brown, terete. Buds to 2 by 2mm, small, acute; stipules
to 20 by 3 mm, linear, acute. Leaf 8-16 by 3.5—8 cm,
oblong-lanceolate to oblanceolate, somewhat charta-
ceous; base broadly cuneate to subcordate; acumen to
1 cm long, tapering; nerves 13-18 pairs, very slender,
arched, ascending, elevated on both surfaces as also
the very slender subreticulate tertiaries, with a few
short secondary nerves; midrib slender but promi-
nent, terete, beneath, channelled above; petiole 12—32
mm long, slender. Panicle to 9 cm long, terminal or
axillary. Flower white; buds to 6 by 5 mm, broadly
ovoid. 2 longer sepals lorate-lanceolate, subacute; 3
shorter sepals very short, deltoid, prominently slender
acuminate. Petals broadly ovate, glabrous. Stamens
15; filaments lorate, tapering, c. + length of the slightly

tapering lorate anthers; appendages very short, acicu-
lar. Stylopodium absent; style prominent, columnar,
glabrous, with obscurely trifid apex. Fruit pedicel to 3
by 2 mm, prominent. Calyx tube to 1.5 mm @,
subglobose; 2 longer lobes to 15 by 3 cm, spatulate,
obtuse, c. 4mm broad at base; 3 shorter lobes to 13 by
3 mm, linear-lanceolate.

Distr. Southern Indochina, Bangladesh (Chitta-
gong), Burma, Peninsular Thailand, and in Malesia:
Malaya (north of Negri Sembilan and Pahang).

Ecol. Semi-evergreen and Evergreen Dipterocarp
forests on undulating land and the lower parts of
valleys.

Vern. Mérsawa gajah, sanai, térbak, médang sawa,
kijal (Malaya).

10. Anisoptera laevis RIDL. Fl. Mal. Pen. | (1922) 219;
Foxw. Mal. For. Rec. 10 (1932) 101; Sym. Gard. Bull.
S. S. 8 (1934) 7, pl. 2; Mal. For. Rec. 16 (1943) 205, f.
100C, 101, 102; AsHToNn, Gard. Bull. Sing. 20 (1963)
233; Man. Dipt. Brun. (1964) 13, f. 5, pl. 5 (stem); ibid.
Suppl. (1968) 5; MEWER & Woop, Sabah For. Rec. 5
(1964) 298, f. 51. — Anisoptera sp. BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 41 (sub A. glabra). — A. glabra
(non KURZ) RipDL. Agr. Bull. Str. & F. M.S. 1 (1901)
60; J. Str. Br. R. As. Soc. 54 (1910) 25. — A. thurifera
(non BL.) Foxw. Philip. J. Sc. 6 (1911) Bot. 257, p.p.,
quoad sp. Mal.; J. Mal. Br. R. As. Soc. 5 (1927) 341,
p.p.; Mal. For. Rec. 3 (1927) 78, p.p. — A. mindanensis
(non Foxw.) WYATT-SMITH, Mal. For. 18 (1955) 77,
p.p. — Fig. 27C-C4.

Bud and twig apex + densely pale brown to rust
lepidote at first, becoming glabrous. Young twig c. 1.5
mm @, slender, terete, becoming rugose on drying;
later smooth or minutely striated. Bud 3-7 by 2 mm,
compressed, hastate to ovate, acute. Stipule c. 10 by 3
mm, glabrous, hastate to linear. Leaves 7-11 by 34
cm, oblong to + obovate, not bullate, epilose,
rust-brown lepidote beneath; base obtuse or broadly
cuneate; acumen to | cm long; nerves 10—14 pairs, at
50°—-60°, slender, curved; petiole 1.5—2 cm long. Pani-
cle to 12 cm long, terminal or axillary, lax, pendent,
angular, slender, densely shortly pale brown puberu-

336 FLORA MALESIANA [ser. I, vol. 9?

Fig. 32. Upuna borneensis SyM., upun batu, habit, about 40 m tall. Brunei, Labi Road (Photogr. ASHTON).

1982]

v.O$

DIPTEROCARPACEAE (Ashton)

337

W
Gomeiig”

aoe

O

Fig. 33. Upuna borneensis SyM. a. Apex of flowering twig, b. fruit, c. nut, all x} (SAN 15184).

lent to glabrous; alternately doubly branched, branch-
lets short, bearing to 11 flowers; bracteoles to 2.5 mm
long, lanceolate, shortly pubescent. Bud to 2 mm long,
subglobose. Calyx cupped at base; sepals subequal.
valvate, shortly grey-brown puberulent outside; 2
longer lobes slightly broader, obtuse, 3 shorter acute,
acuminate. Petals small, elliptic-oblong, glabrous,
pale yellow. Stamens 15, subequal, as long as or
slightly longer than the style; filaments short, tapering
to the anther; anther narrowly oblong, the inner cells
smaller than the outer; appendage to connective short,
erect. Ovary encased in the base of the calyx, covered
by a discoid stylopodium, crowned by a filiform

glabrous style; stigma 3-lobed. Fruit calyx glabrous at
maturity; tube 1.2-1.5cm g@, globose, to 8mm gat the
neck; 2 longer lobes to 15 by 1.5 cm, spatulate, obtuse,
c. 5mm broad at base; 3 short lobes to 1.5cm long, c. 3
mm broad at base, hastate to linear, margin slightly
revolute. Nut apex disc-like, shortly pubescent, flat:
style remnant 5-8 mm long, filiform, very slender.

Distr. Malesia: Malaya, Borneo (Sarawak, Brunei,
S.W. Sabah).

Ecol. Inland lowland and hill forests to 900 m;
widespread and often common.

Vern. Mérsawa durian, médang sawa, madang loh
(Mal.), péngiran durian (Sabah).

3. UPUNA

Sym. Bull. Jard. Bot. Bizg II, 17 (1941) 88, pl. I-I], f. I-Il. — Fig. 32-36.
Large, flaky-barked trees. Buttresses low, broad, rounded, single or grouped in
twos to fours round the base of the bole. Young parts caducous, inflorescence
persistently multicellular glandular tomentose. Stipu/e subulate, subpersistent.
Inflorescence cymose. Flower sepals subequal, imbricate, fused at the base to form
a shallow cup free from the ovary. Stamens 25-30; filaments compressed, dilated
at base, tapering and filiform below the anthers; anthers oblong to ovoid, latrorse:
appendage to connective filiform, many times length of anthers. Ovary ovoid,
without distinct stylopodium: style about twice as long as ovary, trifid towards

338 FLORA MALESIANA [ser. I, vol. 9?

v

TATA 7]
AG
sf >»

bo

/
Tit I
jz

if
RTs
X
&
x
x

Hi
py

Le

f

L
/
j
.e

y
Pl eee -
Me
A
+

ce /
[ee ese

ey

/

/
Uh

ue Je

/ f /

Fig. 34. Upuna borneensis SyM. — Flower details: A. bud, B/. outer sepal, B2. inner sepal, both from inside, C/.
stamens from outside, C2. Stamens from inside, D. pistil, all x10. — EF. Leaf from sterile twig, x3 (4A-D BRUN
3091, E SINCLAIR & KaApIM 10292).

apex; stigma minute. Fruit calyx with a distinct basal cup enclosing but not fused
with the nut; lobes valvate, chartaceous, 2 considerably longer than the other 3.
Nut ellipsoid, 3-angled, splitting into 3 valves at germination, with short acute
apical style remnant, tapering and narrow at base. Seed with distinct arillode.
Germination epigeal; cotyledons subequal, cordate.

Distr. Malesia: Borneo. Monotypic.

Note. The only dipterocarpoid genus not recorded from the present or past of S.E. Asia including Ceylon. An
isolated and in many ways primitive taxon, with leaves and twigs superficially resembling Monotes, dehiscent
pericarp and cymose inflorescence as in some Vatica, a rudimentary aril-like collar on the funicle resembling that
in Stemonoporus, androecium somewhat as in Anisoptera and gynoecium somewhat as in Cotylelobium, though

the gynoecium and androecium characters together are unique among those genera which share a valvate fruit

calyx. The wood anatomy, notably the diffuse distribution of intercellular canals, supports its affinity with the
last four genera.

Fig. 35. Trunk-base of Upuna borneensis SYM., same
place as fig. 32 (Photogr. ASHTON).

1. Upuna borneensis Sym. Bull. Bot. Gard. Btzg III, 17
(1941) 88, pl. JI-II, f. I-II; BRowne, For. Trees
Sarawak & Brunei (1955) 171; ASHTON, Man. Dipt.
Brun. (1964) 7, f. 4, pl. 2 (habit); ibid. Suppl. (1968) 3;
MeveR & Woop, Sabah For. Rec. 5 (1964)
327. — Fig. 32-36.

Young parts caducous multicellular glandular
tomentose; twig apices, cyme, leaf bud, stipule, petiole
and leaf nervation beneath densely persistently pale
chocolate-brown tomentose; lamina surface beneath
white tomentose. Twig to 3.5 mm @ apically, terete,
becoming glabrous, smooth to rugulose, uneven;
much branched; stipule scars small to obscure. Bud to

DIPTEROCARPACEAE (Ashton)

Fig. 36. Young, conical tree of Upuna borneensis SYM.
Forest Research Institute, Kepong, cultivated (Pho-
togr. HENDERSON).

12 by 6mm, ovoid to falcate, subacute. Stipules to 1.2
cm long, linear. Leaves 9-17 by 49.5 cm, oblong to
obovate; base cordate, acumen to 5 mm long, deltoid;
margin revolute; nerves 16-20 pairs, well spaced,
curved at the margin, raised beneath, at 60°—70
except at base, frequently with short secondary
nerves; tertiary nerves well spaced, scalariform, dis-
tinct; midrib prominent, terete beneath, depressed
above; petiole 1-2.5 cm long, geniculate. Cyme to 15
cm long, to 3-axillary, subterete, much branched;
bracts to 10 by 3.5 mm, lanceolate, acute, sparsely
tomentose, caducous. Flower bud to 5 by 2 mm,
narrowly ovoid, conical, acute. Calyx lobes densely
tomentose outside, glabrescent within, fused at base
forming a cup free from the ovary, imbricate; sub-
equal, 2 inner slightly more attenuate, obtuse, subacu-
minate; 3 outer acute, acuminate. Corolla deep purple,
dark yellow at margins, fading to pale red on falling;
petals broadly ovate, subacute, becoming reflexed
apically, sparsely tomentose on parts exposed in bud.
Stamens 25-30 in several verticils; filaments broad at
base, tapering abruptly and filiform below the subglo-
bose anthers; appendage to connective c. 3 times
length of anther, slender, curved. Ovary ovoid, den-
sely tomentose; style c. 3 times as long as ovary,

340 FLORA MALESIANA [ser. I, vol. 9?

Og
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LOY) 4!
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(OF

Fig. 37. Flower details in Cotylelobium melanoxylon (Hook. f.) PIERRE. A. Bud, B/. outer sepal, B2. inner sepal,
both from inside, C/. stamens from outside, C2. stamens from inside, D. pistil, all x10 (S 26853).

shortly pubescent in basal half, otherwise glabrous, densely fulvous tomentose, tapering to a to 5mm long
sometimes trifurcate at apex. Fruit calyx sparsely — slender style remnant, acute.

ashen tomentose; base narrowly cuneate, the lobes Distr. Malesia: Borneo (S. & W. Kalimantan, E.
united into a to 1 cm deep and wide cup; 2longerlobes _ Kutei F. R.; Sarawak, Brunei, S.W. Sabah).

to 13 by 2.7 cm, lanceolate, tapering gradually to the Ecol. Deep sandy soil in lowland Mixed Diptero-
subacute apex, constricted to 7 mm broad at base;3 carp forests, subcoastal hills.

shorter lobes to 7.5 by 1.7 cm, similarly shaped, Uses. Heavy construction timber.

subequal. Nut to 3.2 by 1.5 cm, narrowly ovoid, Vern. Pényau, upun batu.

4. COTYLELOBIUM

PrerRE, Fl. For. Coch. 3 (1889) sub t. 235; HEM, Rech. Dipt. (1892) 119; BRANDIS,
J. Linn. Soc. Bot. 31 (1895) 114; SLoor. Bull. Jard. Bot. Btzg III, 10 (1929) 393;
ibid. III, 12 (1932) 43; Sym. Mal. For. Rec. 16 (1943) 232, f. 111 (map); ASHTON,
Man. Dipt. Brun. (1964) 56; ibid. Suppl. (1968) 24; MevER & Woop, Sabah For.
Rec. 5 (1964) 323; ASHTON, Blumea 20 (1972) 358; SMITINAND, Thai For. Bull.
(Bot.) 12 (1980) 23. — Dyerella Hem, Rech. Dipt. (1892) 123. — Fig. 37-40.

1982] DIPTEROCARPACEAE (Ashton) 34]

Small, medium-sized or large trees; bole frequently twisted; buttresses low,
rounded, similar to those of Vatica. Crown hemispherical, rather small. Bark
surface greyish, at first smooth, hoop marked; becoming irregularly, frequently
shaggy, flaky, leaving a distinct scroll-marked surface below. Stipules fugaceous.
Leaves oblong to ovate-lanceolate, coriaceous, margin revolute, undersurface
lepidote; nerves curved, unraised above, hardly so beneath, bifurcating towards
the margin and anastomosing to form a looped intramarginal nerve, with shorter
indistinct intermediate nerves similarly bifurcating; tertiary nerves reticulate,
indistinct; midrib sunken above; petiole comparatively short, not geniculate.
Calyx lobes imbricate in flower; 2 obtuse outer lobes somewhat larger than 3 acute
inner lobes. Petals free, broad, elliptic-oblong, cream or pink. Stamens 15,
subequal, in 3 whorls, hence pairs alternating with single stamens; filaments
short, deltoid, connate at base; anthers narrowly oblong, latrorse, with 4 pollen
sacs, the inner 2 shorter than the outer 2, setose along the lateral margins;
appendage to connective short, slender. Ovary free from calyx, + globose;
stylopodium indistinct; style filiform, slender, many times longer than ovary,
shortly pubescent towards base; stigma small, trifid, slightly broader than style.
Fruit as in sect. Sunaptea of Vatica; calyx free from nut, with distinct filiform style

remnant.

Distr. 6 spp., Ceylon, Peninsular Thailand; in Malesia: 3 spp., in Malaya, E. Sumatra, Lingga, Anambas Is.
and Borneo. Fig. 38.

Ecol. Main canopy trees of dry acid soils, especially on coastal hills, but also on rentzinas over limestone and
mountain ridges inland to 1500 m; sometimes semi-gregarious.

Uses. A hard durable timber similar to that of Vatica is produced, but the trees are generally larger.

Note. The perianth and range of fruit variation strikingly resemble that of Vatica, and the wood anatomy
underlines this affinity; but the androecium, gynoecium and leaf nervation are quite distinct from that genus.
The Malesian species differ only in characters of the tomentum, and minor details of leaf shape and nervation;
they are variable and at times difficult to identify with certainty; the possibility of occasional hybridisation
cannot be ruled out.

KEY TO THE SPECIES

1. Leaf glabrescent, drying dull olive-brown . 1. C. melanoxylon
1. Leaf densely tomentose beneath.
2. Leaf 8-12 by 34.5 cm, oblong-lanceolate, drying dull olive-brown above, densely shortly evenly rich
yellow-ochre tomentose beneath, with darker midrib and petiole; intramarginal nerve close to margin
2. C. burckii
2. Leaf 6-8 by 2.2-3 cm, ovate-lanceolate, drying dark grey-brown above, densely dark grey scabrid
tomentose beneath with darker midrib and petiole; intramarginal nerve well within margin
3. C. lanceolatum

1. Cotylelobium melanoxylon (Hook. f.) PIERRE, FI.
For. Coch. 3 (1889) t. 235; Hem, Rech. Dipt. (1892)
119, 120; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3,
6 (1894) 268; BRANDis, J. Linn. Soc. Bot. 31 (1895)
115; Merr. En. Born. (1921) 408: SLoor. Bull. Jard.
Bot. Btzg III, 9 (1927) 78: ibid. 10 (1929) 403; ibid. 12
(1932) 44; Sym. Mal. For. Rec. 16 (1943) 236, f. 114:
BROWNE, For. Trees Sarawak & Brunei (1955) 96;
ASHTON, Man. Dipt. Brun. (1964) 59, f. 9; ibid. Suppl.
(1968) 24, pl. 5 (bark); MEER & Woop, Sabah For.
Rec. 5 (1964) 324, f. 59. — Anisoptera melanoxylon
Hook. f. Trans. Linn. Soc. 23 (1860) 160; DC. Prod.
16, 2 (1868) 616; WaLp. Ann. 7 (1869) 378; BURCK,
Ann. Jard. Bot. Btzg 6 (1887) 221. — Vatica melano-

xylon BENTH. & Hook. f. ex Mig. Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; Hem, Bull. Mens. Soc. Linn.
Paris 2 (1891) 956, incl. var. recta. — Vatica becca-
riana HEIM, /.c. 955. — Vatica harmandii Hem, I.c.
955. — C. beccarii PIERRE, Fl. For. Coch. 4 (1891) t.
258B; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 116; BEcc.
For. Born. (1902) 570, 591; MerRR. En. Born. (1921)
408; BROWNE, For. Trees Sarawak & Brunei (1955)
95.— C. harmandii Heim, Rech. Dipt. (1892) 122;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 115; Merr. En.
Born. (1921) 408; SLoort. Bull. Jard. Bot. Btzg III, 9
(1927) 78; ibid. 10 (1929) 405; BROwNeE, For. Trees
Sarawak & Brunei (1955) 95. — C. beccarianum HEI,
Rech. Dipt. (1892) 122; SLoort. Bull. Jard. Bot. Btzg

FLORA MALESIANA

[ser. I, vol. 9?

Slorn|o.

Fig. 38. Density map of Cotylelobium Pierre in
Malesia; number of endemics above the hyphen,
number of non-endemics below it.

III, 9 (1927) 78; ibid. 10 (1929) 405. — C. leucocarpum
SLooT. [Bull. Jard. Bot. Btzg III, 9 (1927) 78, nomen]
ibid. II, 10 (1929) 399, f. 2.— Vatica leucocarpa
Foxw. ex DEN BERGER & ENDERT, Med. Proefst.
Boschw. 11 (1925) 130; ex HEYNE, Nutt. Pl. ed. 2
(1927) 1129. — Fig. 37, 39.

Young leaves, twig, stipules outside, bud and
raceme densely shortly powdery grey tomentose,
fugaceous on leaf and midrib, + so on twigs,
persistent on racemes. Jwig to | mm @
apically, terete, slender, minutely striated with fine
cracks; stipule scars obscure. Bud c. 1.5 by 1 mm,
ovoid, acute. Stipules to 3 mm long, small, linear,
caducous. Leaves 5-10 by 2-6 cm, ovate-lanceolate,
oblong or obovate; base broadly cuneate or obtuse;
acumen to 8 mm; margin + revolute; nerves 10—13
pairs, slender, hardly raised beneath, with more
slender shorter parallel intermediates; intramarginal
nerve strongly looped, c. 2 mm within the margin;
petiole 9-12 mm long. Panicle to 8 mm long, axillary,
slender, singly branched; bracts and bracteoles to 10
by 3 mm, lanceolate, acute, shortly pubescent outside,
glabrescent within. Flower bud to 6 by 3 mm, ellipsoid,
obtuse. Calyx densely pubescent outside, sparsely so
within; lobes subequal, narrowly deltoid, acute. Petals
narrowly ovate, obtuse, sparsely pubescent on parts
exposed in bud. Stamens 15, subequal; filaments
linear, tapering, c. 3 times as long as the anther, half as
long as the style, sparsely ciliate along lateral margins;
appendage to connective c. 4 as long as the anther,
short, slender. Ovary ovoid, densely pubescent; style
filiform, slender, c. 3 times as long as the ovary,
pubescent towards base, otherwise glabrous. Fruit
calyx persistently pubescent towards base, glabres-
cent elsewhere; 2 longer lobes to 4.5 by 1.2 cm, oblong,
obtuse, c. 2mm broad at base; 3 shorter lobes 8-14 by
2-3 mm, hastate, acute, fimbriate; lobes united at base

melanoxylon (HooK. f.) PIERRE. Brunei (Photogr. G.
H. S. Woop, SAN 17547).

into a shallow cup c. 8 mm g. Nut c. 6 mm long and
broad, ovoid, with up to 5 mm long style remnant,
shortly pale buff tomentose.

Distr. Coastal Peninsular Thailand; in Malesia: S.
Malaya (N.E. Johore), Singapore, Sumatra (Singkep,
Lingga; Central Sumatra: Siak, Indragiri; P. Musala),
Borneo.

Ecol. Local, on dry, often sandy soils on coastal
hills and terraces, sometimes in secondary forest.

Vern. Resak, r. hitam (Brunei), r. témpurong
(Sabah), r. bukit, r. tembaga (Lingga), r. batu, r.
témbaga, r. padi, r. kéranji (W. Borneo), r. bunga (E.
Borneo).

Note. The species, especially C. malayanum, are
somewhat variable and the possibility of hybridisa-
tion between C. malayanum and C. melanoxylon
cannot be excluded.

2. Cotylelobium burckii (HEIM) Heim, Rech. Dipt.
(1892) 122; SLoor. Bull. Jard. Bot. Btzg III, 9 (1927)
78; ASHTON, Gard. Bull. Sing. 20 (1963) 243; Man.
Dipt. Brun. (1964) 57, f. 9, pl. 20-21 (habit, stem); ibid.
Suppl. (1968) 24. — Vatica burckii Hem, Bull. Mens.
Soc. Linn. Paris 2 (July 1891) 956.—C. flavum
Pierre, Fl. For. Coch. 4 (Oct. 1891) t. 258A; BRANDIS,
J. Linn. Soc. Bot. 31 (1895) 115, t. 3, f. 22-24; BoERL.
Cat. Hort. Bog. 2 (1901) 112; Becc. For. Born. (1902)
570; Merr. En. Born. (1921) 408; SLoort. Bull. Jard.
Bot. Btzg III, 12 (1932) 44; Sym. Gard. Bull. S. S. 8
(1934) 36; BRowneE, For. Trees Sarawak & Brunei
(1955) 171; ANDERSON, Gard. Bull. Sing. 20 (1963)

1982]

DIPTEROCARPACEAE (Ashton)

343

Fig. 40. Cotylelobium lanceolatum Crain. a. Flowering twig, x2/3. — C. burckii (HEIM) HEI. 6. Fruiting twig.
x2/3, c. fruit, d. nut, both x2, e. leaf from sterile twig, x2/3 (a S 28068, b S 12995, c-d KEP 32615, e bb. 15334).

157. — C. asperum SLootT. Bull. Jard. Bot. Btzg III, 10
(1929) 401, f. 3; ibid. 12 (1932) 1. — Fig. 40b-e.
Twig, bud, petiole, stipules outside, midrib beneath
and raceme densely shortly persistently buff tomen-
tose, slightly scabrid; leaf evenly densely ocherous
tomentose beneath. Twig c. 1.5 mm @ apically,
slender, smooth. Bud 1.5—2 by 1.5 mm, small. Stipules
to 8 mm long, narrowly deltoid, fugaceous. Leaves
8-12 by 3-4.5 cm, oblong-lanceolate; margin promi-
nently revolute; base broadly cuneate or obtuse;
acumen 2.5—7.5 mm long; nerves 10-12 pairs, indis-
tinct, the intramarginal nerve comparatively straight
and just within the margin; petiole 1.5—2 cm long.
Panicle to 15 cm long; to 3-axillary, rarely terminal,
terete or ribbed, irregularly doubly branched; brac-
teoles to 8 by 5 mm, elliptic, obtuse, shortly buff
pubescent. Flower bud to 10 by 4 mm, ovoid-lanceo-
late, acute. 2 calyx lobes long, linear, obtuse; 3 short,
lanceolate, acute; densely buff tomentose outside,
more sparsely within. Corolla cream; petals large,
narrowly oblong, obtuse, glabrous. Stamens 15, the
inner 5 slightly longer than the others; filaments short,
broad at base, strongly tapering; anthers narrowly

oblong, reaching half length of style, with sparsely
setose lateral margins; appendage to connective { as
long as anther, slender. Ovary subglobose, densely
tomentose; style slender, 3—4 times as long as ovary,
shortly pubescent but for the glabrous apical 4. Fruit
calyx as in C. melanoxylon but base more densely
tomentose, longer lobes c. 1.5 cm broad, persistently
shortly sparsely tomentose; 3 shorter lobes to 1.2 cm
long, linear, shortly sparsely tomentose on both
surfaces. Nut as in other species.

Distr. Malesia: Borneo (W. Kutei, Lower Dayak,
W. Borneo, Sarawak and Brunei N.E. to the Lim-
bang).

Ecol. Locally common on giant podsols, on raised
beaches, rare on sandstone cuestas, near present and
Pleistocene coastlines.

Vern. Résak durian, r. babalok, r. gunong, r. baru.

3. Cotylelobium lanceolatum CraiB, Kew Bull. (1913)
113; Fl. Siam. Enum. 1 (1925) 142. — C. malayanum
SLooT. Bull. Jard. Bot. Btzg III, 12 (1932) 43; Foxw.
Mal. For. Rec. 10 (1932) 247; BurK. Dict. (1935) 673;
Sym. Gard. Bull. S. S. 9 (1938) 349; Mal. For. Rec. 16

344 FLORA MALESIANA [ser. I, vole92

d

Fig. 41. Vatica umbonata (Hook. f.) BURCK. a. Habit, x5, b—c. young fruits, x5, d. ripe fruit, lateral view, e. ditto,
apical view, both nat. size (a SAN 68373, flowers from SAN 15367, b A 4743, c FRI 12496, d-e BRUN 933).

1982]

DIPTEROCARPACEAE (Ashton)

345

(1943) 235, f. 113, 114; BRowne, For. Trees Sarawak
& Brunei (1955) 95; ASHTON, Man. Dipt. Brun. (1964)
11, f. 9; ibid. Suppl. (1968) 24, pl. 4 (bark); MEVER,
Sabah For. Rec. 5 (1964) 324.— C. flavum (non
PiERRE) RiD-. J. Str. Br. R. As. Soc. 54 (1909) 25; FI.
Mal. Pen. 1 (1922) 239; SLoor. Bull. Jard. Bot. Btzg
III, 10 (1929) 396, f. 1, specim. BECCARI excl.; Foxw.
Mal. For. Rec. 3 (1928) 71; ibid. 10 (1932) 247. — Fig.
40a.

Twig, bud, petiole, stipule outside, lamina beneath,
and raceme persistently densely grey-brown to rufous
scabrid tomentose; tomentum on lamina beneath
dotted with scattered darker larger tufts. Twig c. | mm
@ apically, slender, terete. Bud c. 1.5 by | mm, small,
conical. Stipules c. 3 mm long, narrowly deltoid,
fugaceous. Leaves 6-8 by 2.5—3 cm, narrowly ovate-
lanceolate; margin revolute; base broadly cuneate;
acumen to 5 mm long; nerves 10-12 pairs, with a
prominently looped intra-marginal nerve c. 2 mm
from the margin; petiole 6-10 mm long, short, slender.
Panicle to 6 cm long, terminal or axillary, short, terete,
ribbed; irregularly singly or doubly branched; brac-
teoles to 4 by 3 mm, elliptic, obtuse, densely shortly

tomentose outside, sparsely so within. Flower bud to 8
by 3.5 mm, ovoid, obtuse. Ca/yx densely buff-tawny
tomentose outside, sparsely so within, lobes subequal,
deltoid, short; outer slightly narrower, more acute.
Corolla cream; petals short, narrowly oblong, obtuse,
glabrous. Stamens 15, the inner 5 slightly longer than
the rest; filaments short, triangular; anthers narrowly
oblong, with prominently setose margin; appendage
to connective short, slender. Ovary small, subglobose,
densely tomentose; style c. 3 times as long as ovary,
filiform, pubescent towards base. Fruit calyx as in C.
burckii but more scabrid tomentose; shorter lobes
1.2—2 by 0.4 cm, comparatively long, broadly hastate,
constricted at base. Nut as in other species.

Distr. Eastern Peninsular Thailand, and Malesia:
E. Malaya (from Trengganu southwards), Anambas
Is., Borneo (W. Borneo, Sarawak, Brunei, S.W. &
S.E. Sabah, Kutei, lower Mahakam), erroneously
recorded from Sumatra by SYMINGTON (1938).

Ecol. Locally abundant, podsolised soils in Heath
forest on terraces and sandstone ridges to 1500 m; on
peaty soils over limestone in W. Sarawak.

Vern. Resak, r. batu, r. bukit (Mal.).

5. VATICA

LINNE, Mant. 2 (1771) 152; B. & H. Gen. Pl. 1 (1862) 192, incl. sect. Isauxis (ARN.)
B. & H.; DC. Prod. 16, 2 (1868) 517; Dyer, FI. Br. Ind. 1 (1874) 301; BuRcK, Ann.
Jard. Bot. Btzg 6 (1887) 223, incl. sect. Sunaptea (GRIFF.) BURCK et sect.
Pachynocarpus (Hook. f.) BURCK, /.c. 225; HEIM, Rech. Dipt. (1892) 99; BRANDIS
& GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 268; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 116, incl. subg. Synaptea (GRIFF.) BRANDIS, /.c. 128, subg. Retinodendron
(KorTH.) BRANDIS, /.c. 119, subg. Isauxis (ARN.) BRANDIS, /.c. 127; SLooT. Bull.
Jard. Bot. Btzg III, 9 (1927) 67; Foxw. Philip. J. Sc. 67 (1938) 319; Sym. Mal. For.
Rec. 16 (1943) 211, f. 105 (map); ASHTON, Gard. Bull. Sing. 20 (1963) 243; Man.
Dipt. Brun. (1964) 61; ibid. Suppl. (1968) 25; MEER & Woop, Sabah For. Rec. 5
(1964) 301; SMITINAND, Thai For. Bull. (Bot.) 12 (1980) 81. — Seidlia KOSsTEL.
Allg. Med.-Pharm. FI. 5 (1836) 1945. — Vateria ARN. Ann. Nat. Hist. 1, 3 (1839)
155, pro sect. Isauxis ARN.— Retinodendron KortTH. Kruidk. (1840)
55. — Isauxis (ARN.) REICHB. Nom. (1841) 210. — Pteranthera BL. Mus. Bot.
Lugd.-Bat. 2 (1852) 30. — Sunaptea GrirF. Notul. 4 (1854) 516, corr. “Synaptea’
Kurz, J. R. As. Soc. Beng. Sc. 39, 2 (1870) 65. — Pachynocarpus Hook. f. Trans.
Linn. Soc. 23 (1860) 159; GiLc in E. & P. Pfl. Fam. ed. 2, 21 (1925) 266. — Elaeogene
Mia. Sum. (1862) 460. — Retinodendropsis HEM, C. R. Assoc. Fr. Pau 1892 (1893)
470. — Perissandra GAGN. Bull. Soc. Bot. Fr. 95 (1948) 27, cf. Jacoss, Blumea 15
(1967) 138. — Brachypodandra GAGN. l.c. 30. — Fig. 41-47.

Small to medium-sized, rarely large, trees; bole frequently sinuate, buttresses
thick, rounded, concave, usually small. Crown irregular, oblong, sympodial,
non-emergent. Bark surface usually grey mottled, smooth and hoop-marked, in
large trees becoming patchily flaked, occasionally scroll-marked. Young parts
usually + caducous powdery tomentose. Leaves variable; nerves curved, usually

346 FLORA MALESIANA [ser. I, vol. 9?

1979

Fig. 42. Vatica umbonata (HooK. f.) BURCK. A. Bud, A/. stamens from outside, A2. stamens from inside, 43.
pistil, all x10. — V. maingayi Dyer. B. Bud, B/. stamens from outside, B2. pistil, all x10 (A AMBULLAH 31457, B
Neth. Ind. For. Serv. F 922).

somewhat oblique to the midrib; tertiary nerves + reticulate, never truly
scalariform; petioles not geniculate. Inflorescences irregularly branched, race-
mose or sometimes partially cymose, short, rarely spreading. Flower buds ovoid
to lanceolate, of variable size. Calyx lobes + valvate, subequal. Peta/s narrowly
oblong, usually pale cream-white, not connate at base, falling separately. Stamens
15 in 3 verticils, single stamens alternating with pairs, short, the inner row slightly
longer than the outer row; filaments short, dilated at base, + tapering and
filiform below anthers; anthers broadly oblong, latrorse, the inner pollen sacs
smaller than the outer; appendage to connective short, rarely as long as anthers,
+ deltoid, stout. Ovary + broadly ovoid, conical, superior or semi-inferior,
shortly densely pubescent, without distinct stylopodium; style columnar, short,
stout, glabrous, somewhat expanded at apex and with a prominent conical
3-lobed stigma. Fruit calyx variable. Nut of variable size, broadly ovoid or
globose, with or without a distinct apical style remnant; pericarp splitting along 3
sutures at germination; germination epigeal (sect. Sunaptea; some sect. Vatica) or
hypogeal (some sect. Vatica) with the cotyledons remaining within the fruit;
cotyledons, if free from fruit, magenta to pale yellow, usually non-photosynthe-

1982] DIPTEROCARPACEAE (Ashton) ae

tic; first pair of leaves opposite with interpetiolar stipules, rarely alternate; the
succeeding leaves spiral.

Distr. About 65 spp. in Ceylon and southern and eastern India, Burma, Thailand, Indochina, S. China
(Hainan), and c. 55 spp. throughout Malesia excepting the Lesser Sunda Is. Fig. 43.

Ecol. Understorey trees, sometimes in main canopy, scattered in lowland forests and some species in hill
forests to 1600 m; several species semi-gregarious on river-banks.

Uses. A hard semi-durable timber is obtained, but the trees are small; used locally for house posts and other
minor construction.

Note. A genus clearly circumscribed by its extraordinary constancy of floral structure and also wood
anatomy; its closest affinity is with Cotylelobium (q.v.). The two main forms of fruit calyx appear to have evolved
only once and, judged by the wide geographical distribution of the two sections, would seem to mark an ancient
dichotomy in the genus.

KEY TO THE SPECIES!

1. Fruit calyx lobes equal. Spp. 1-24. 1. Sect. Vatica
2. Fruit calyx lobes corky, coalescing with each other and with the nut, forming a cup + enclosing it.
3. Nut ovoid or globose, not exceeding 2.5 cm long. Petioleto |1Smmlong . . . . 1. V. umbonata
3. Nut to 4 cm long, narrowly ovoid-ellipsoid. Petiole exceeding 18 mm long . . . . 2. V. stapfiana
2. Calyx lobes not as above.
4. Calyx lobes in fruit subcordate, somewhat revolute at base, not reflexed, + concealing nut.

5. Nerves 8-15 pairs, without distinct secondary nerves. . . . . . . . . +. 3. VY. venulosa
5. Nerves at least 16 pairs, with distinct secondary nerves.
6. Twigs, panicles and petiole persistently fulvous + flocculent pubescent . . . . 4. V. havilandii
6. Twigs and panicles caducous buff puberulent; petiole glabrous. . . . . . . 5. V. chartacea

4. Calyx lobes not cordate; nut completely exposed.
7. Nut ovoid, generally coming to exceed 2.3 cm long; pericarp thick, corky verrucose.
8. Fruit sepals to 5 mm long, broadly ovate to suborbicular, adpressed to the base of the nut.

Panto Sebys> cmyovoid iyi af “leg 1 (2 ewisd shtxcors vibee! ocet.6:V qpaucifiora
Se nto 5S) byel/8.cm, fusiform. sc) lott \h. 2 enempbe eam iin Sie iste 5 V Sridleyana
8. Fruit sepals linear-lanceolate.
10. Fruit sepals patent, becoming recurved and revolute resembling claws. . . . 8. V. soepadmoi
10. Fruit sepals not as above.
msg prominentiyabeaked)) i... | 4... Soe eee ee BOP Vo bella
11. Nut not prominently beaked.
12. Nut becoming asymmetrical; petiole exceeding 2cmlong. . . . . . . 410. V.rassak
12. Nut symmetrical; petiole shorter than 2 cm long.
13. Leaves to 20 by 7cm; nerves 10-12 pairs . . . . . . . =. +. +. #«.(211.V. granulata
13. Leaves at least 22 by 7 cm; nerves 15-28 pairs. . . . . . . . «412. V. sarawakensis
7. Nut smooth.
14. All parts entirely glabrous. .. 20s ae Re a) Sh) eee IBV albiramis

14. Young parts at least puberulent or pubescent.
15. Tomentum vinous.

h6@ Bwiesicompressed’ 2. 4 2 4 2 ow 4 4 WSU 9 eal oblongifolia
16. Twigs terete.
WE Calyx lobes thin, hardly revolute) "20, 27) =) ey. Culitensis
17. Calyx lobes, incrassate, a ae revolute.
LS sNue2 2 cmige- = RD OLDEN JM AT Oe e soanl GRVopedicellata
18. Nut not exceeding lcm @.
19 (Nerves O=19 pairs’ 0S 6 13. OT Re EOE een ee Nea ORAL
19: “(Nervés:12=20 pairs’. 2. 2° © 2 3) ne ee ee SV eavinosa

15. Tomentum not vinous.
20. Tomentum rufous.
21. Leaves large, oblong, with obtuse or cordate base. . . . . . . . 419. V. scortechinii
21. Leaves medium-sized, obovate, with narrowly cuneate base . . . . . . 20. V. globosa
20. Tomentum buff or pale brown.

(1) The following species are not inserted in the key: 52. V. elliptica, 53. V. pentandra, 54. V. cauliflora, 55. V.
glabrata, and 56. V. obtusa.

348 FLORA MALESIANA [ser. I, vol. 9?

Bes Fruit sepals incrassate, adnate tonut . . ober BS) 26. Se Evalobata
Fruit sepals not incrassate, becoming + revolute or r reflexed.
oe Stipulestlarge: elliptic, subpersisient =. 6 4) ee a Sec eet
23. Stipules minute, fugaceous.
24. Leaves and nut glabrescent. . . ee ere ere er 228) WV, ein
24. Leaves beneath and nut persistently pubescent rr meee rma oe 7 VY. ENTE

1. Fruit calyx lobes unequal. Spp. 25-5]. 2. Sect. Sunaptea
25. Fruit calyx lobes fused into a cup at base, adnate to the fruit.
26. Nut exceeding | cm @; fruit calyx lobes + patent, of varyinglengths . . . . 25. V. heteroptera
26. Nut at most 8 mm @; calyx with ascending lobes, 2 of which greatly exceed the others.
27. Tomentum cream to tawny; twigs + compressed at first.

28. Nerves 8-11 pairs «ty Wy ple) Melis OU xk ar aa Beg eee bo, SR oe sy ee OMB ema ERIE
28. Nerves 13-17 pairs. . . - oe. «oe ao a 4. RAVEiteysmanniana
27. Tomentum not as above; twigs terete.
29. Twigs and petioles sparsely palescurfy . . . . . . . .-. . +. +. +. 28. V. cinerea
29. Twigs and petioles densely persistently pubescent . . . . . . . +. +. +. 29. V. odorata
25. Fruit calyx lobes free to base.
30) Siwigsicompressed! > BIOL PN AL) TENE dite WG so eee ae AR Gs ee. AS? SRO AV COMIphessa

30. Twigs terete or ribbed.
31. Leaf obovate, thickly coriaceous, normally obtuse or retuse, margin revolute.

32. Inflorescence to 3 cm long, congested. Petiole exceeding 1.2cm . . . . . . 31. V. congesta

32. Inflorescence to 20 cm long, lax. Petiole lessthan| cmlong . . . . . . . 32. V. coriacea
31. Leaf not as above.

33. Nerves at least 22 pairs, Peraseny pubescent beneath” = «=. 47) 4) 229 S935 Gjavanica

33. Nerves less than 22 pairs, or, if 22, then glabrous.

34. Leaf nervation beneath pale Bown or ocherous pubescent.
35. Leaf undersurface persistently shortly buff scabrid pubescent; petiole 8-15 mm long
34. V. brunigii
35. Leaf beneath (but not nerves) sparsely sericeous, glabrescent; petiole 15-30 mm long.
36. Leaf base broadly cuneate. Nerves 10-11 pairs. . . . . . . . . 35. V. pachyphylla
36. Leaf base narrowly obtuse. Nerves 11-Il4 pairs. . . . . . . . . . 36. V. obovata
34. Leaf nervation glabrescent beneath.
37. Nerves hardly more prominent beneath than above.
38. Petiole at least 14 mm long.
39. Nerves 7-9 pairs; tomentum pink-brown . . . . . . . +. +. +. + 37. V. borneensis
39. Nerves 9-11 pairs; tomentum ocherous-buff . . . . . . . . ~~. 38. V. bantamensis
38. Petiole 1 cm long or shorter.
40. Ripe nut broadly ovoid, obtuse, densely tomentose.

41. Midrib raised above; base of leaf cuneate. . . . . . .39. V. mangachapoi
41. Midrib flat or slightly sunken above; base of leaf obtuse. » 6. 3 8) + eedO) Veipanvifolia
40. Ripe nut narrowly ovoid, tapering, acute, glabrous... . . . . 41. V. rynchocarpa

37. Nerves distinctly more prominently raised below than above...
42. Inflorescence, petiole, nerves and midrib beneath persistently shortly pale brown scabrid
tomentose. Leaf thin, obtuse.

AB wPetiolelexceedinealSammilong. %3)=2 ya //eas 2 Ve (22) 2 a eV reneran
43. Petiole at most 11 mm long.
44. Leaf elliptic to obovate, base narrowly Cuneate, 46 o5 4 epi) 2) ee ee ee vipes
44. Leaf elliptic-oblong to lanceolate, base obtuse .. .. 44, V. micrantha
42. Tomentum not as above, nervation beneath glabrescent. Leaf base generally cuneate.
45. Leaf drying yellowish olive, lustrous; tomentum ocherous buff. . . . 485. V. flavovirens

45. Leaf drying grey-brown or red-brown; tomentum pink-brown or rufous.
46. Tomentum very short, even.
47. Petiole at least 2 cm long; longer calyx lobes to 8 by2.5cm. . . . . 46. V. badiifolia
47. Petiole shorter than 2 cm; longer calyx lobes to6 by 1.8cm. . . . 47. V. perakensis
46. Tomentum scabrid or flocculent.
48. Major fruit calyx lobes coming to exceed 7 cm long.
49. Major fruit calyx lobes somewhat broadened at base; leaf nerves 12-22 pairs 48. V. nitens
49. Major fruit calyx lobes tapering at base; nerves at most 17 pairs.
50. Petiole scabrid pubescent; nerves 10-17 pairs. . . . . . . . .49. V. cuspidata
50. Petiole sparsely puberulent; nerves 9-11 pairs . . . . . . . +. 50. V. maingayi
AS. Majorfruit calyx lobes less thani6;cmlong’ = 5 5 5 =). 9 5 5) =) Se SSIVelow

1982] DIPTEROCARPACEAE (Ashton) 349

1. Section Vatica

ASHTON, Gard. Bull. Sing. 20 (1963) 250, incl. sect. Pachynocarpus (HOOK. f.)
Burck. — Seidlia KosTeL.— Vateria sect. Isauxis ARN. — Retinodendron
KorTH. — /sauxis (ARN.) REICHB.— Pachynocarpus Hook. f. — Elaeogene
Mia. — Vatica sect. Isauxis (ARN.) B. & H.— Vatica sect. Retinodendron
(KORTH.) BURCK et sect. Pachynocarpus (HooK. f.) BURCK, Ann. Jard. Bot. Btzg 6
(1887) 224. — Vatica subg. Retinodendron (KORTH.) BRANDIS et subg. Isauxis

(ARN.) BRANDIS, J. Linn. Soc. Bot. 31 (1895) 119, 127.

Calyx lobes equal.

1. Vatica umbonata (Hook. /.) BurcK, Ann. Jard.
Bot. Btzg 6 (1887) 232; SLoor. Bull. Jard. Bot. Btzg
III, 9 (1927) 132, 133; AsHTon, Gard. Bull. Sing. 20
(1963) 250; Man. Dipt. Brun. (1964) 78, f. 10; ibid.
Suppl. (1968) 36; MEIJER & Woop, Sabah For. Rec. 5
(1964) 320, f. 58, pl. 30A; ASHTON, Gard. Bull. Sing.
31 (1978) 17.

a. ssp. umbonata. — Pachynocarpus umbonatus HOOK.
Hook. f. Trans. Linn. Soc. 23 (1860) 159, t. 22; Mia.
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC. Prod. 16,
2 (1868) 617; WALP. Ann. 7 (1868) 378; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 135, t. 3, f. 25; BRANDIS &
GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 270, fig.;
Merr. En. Born. (1921) 409; RipL. Fl. Mal. Pen. |
(1922) 249, p.p.; GiLc in E. & P. Pfl. Fam. ed. 2, 21
(1925) 265. — V. verrucosa BURCK, Ann. Jard. Bot.
Btzg 6 (1887) 232, t. 29, f. 5; SLoor. Bull. Jard. Bot.
Btzg III, 9 (1927) 131; in Merr. Pl. Elm. Born. (1929)
205; BROWNE, For. Trees Sarawak & Brunei (1955)
102. — Pachynocarpus verrucosus HEIM, Rech. Dipt.
(1892) 107; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 136;
Merk. En. Born. (1921) 410; Ripi. Fl. Mal. Pen. 1
(1922) 249, p.p. — V. blancoana ELMer, Leafl. Philip.
Bot. 4 (1912) 1473; Foxw. Philip. J. Sc. 13 (1918) Bot.
196; ibid. 67 (1938) 326; Merr. En. Philip. 3 (1923)
102. — V. cupularis SLooT. Bull. Jard. Bot. Btzg III, 9
(1927) 132, t. 13; in Merr. Pl. Elm. Born. (1929) 205,
p.p.; HEYNE, Nutt. Pl. ed. 2 (1927) 1129; BROWNE, For.
Trees Sarawak & Brunei (1955) 100. — V. ramiflora
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 118,
p.p. — V. stapfiana (non SLOOT.) BROWNE, For. Trees
Sarawak & Brunei (1955) 102. — Fig. 41, 42A—A3.
Young twig, raceme, leaf bud, stipule and petiole
shortly sparsely pale grey-brown puberulent, persis-
tent only on leaf bud and stipules. Twig to 3 mm @
apically, stout, brittle, much branched, crooked,
covered with small linear striations, cracks and some-
times flakes. Bud to 2 by 1.5 mm, ovoid, subacute.
Stipules to 4 by 2 mm, hastate, subacute, early
caducous. Leaves 8-16 by 3—6.5 cm, somewhat coria-
ceous, elliptic; base + broadly cuneate; apex obtuse
or shortly acuminate; nerves 7-8 pairs, raised beneath,
slightly curved, at 45°-55°; no distinct secondaries;
midrib raised beneath, + applanate above. Petioles
7-15 mm long. Panicle to 12 cm long, singly, rarely
doubly branched, axillary or terminal, rigid, brittle,

falling apart before maturity. Flower bud to 1 cm long;
calyx shortly curved pubescent; flowers typical. Fruit
pedicel 2-5 mm long. Nut to 3 cm long and g,
pink-brown verrucose, compressed at base; calyx
united into a tube adpressed to and fused to nut;
differentiated into 5, to 2.6 by 1.8 cm, oblong-elliptic,
up to 5mm thick lobes by 5 deep longitudinal furrows;
nut 3-sulcate, exposed only at apex.

Distr. Malesia: Malaya (Pahang, Trengganu), W.
and N. Borneo, S. Philippines (Palawan).

Ecol. Locally abundant, alluvium river-banks;
scattered on hillsides, locally frequent on acid soils in
mountains at 900-1300 m.

Vern. Resak ayer, r. gunong, r. labuan.

Note. See also under 56. V. obtusa BURCK.

b. ssp. acrocarpa (SLOOT.) ASHTON, Gard. Bull. Sing.
31 (1978) 17. — V. acrocarpa SLooT. Bull. Bot. Gard.
Btzg III, 17 (1942) 241, f. 31; MEER & Woop, Sabah
For. Rec. 5 (1964) 320.

Nut ovoid, acute, the fruit sepals united to basal 4
only and reflexed.

Distr. Malesia: E. Borneo.

Ecol. Apparently always on or near river-banks.

Note. V. umbonata is frequently gregarious on
river-banks and the continuous variation found in this
habitat, especially in N.E. Borneo where the two
subspecies occur in the same area and in E. Malaya
where the closely related. V. stapfiana occurs in the
same habitat, suggests panmixis and some local
hybridisation. Some hybridisation appears to occur
with V. rassak in E. Sabah.

2. Vatica stapfiana (KING) SLooT. Bull. Jard. Bot.
Btzg III, 9 (1927) 129; ibid. III, 17 (1941) 137; Foxw.
Mal. For. Rec. 10 (1932) 274; Burk. Dict. (1935)
2225; SyM. Mal. For. Rec. 16 (1943) 228, f. 107,
109. — Pachynocarpus stapfianus KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 136; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 136; BRUHL & KING, Ann. R. Bot. Gard.
Calc. 5, 2 (1896) 161, t. 194; Burk. & Foxw. J. Str. Br.
R. As. Soc. 86 (1922) 279; RIDL. Fl. Mal. Pen. 1 (1922)
249; Craib, Fl. Siam. Enum. | (1925) 140. — Pa-
chynocarpus wallichii KiNG, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 135, p.p.; RIDL. Fl. Mal. Pen. 1 (1922) 250,
p.p. — Pachynocarpus grandiflorus Riv. J. Fed. Mal.
St. Mus. 10, 2 (1920) 127; Cras, Fl. Siam. Enum.

350 FLORA

MALESIANA

[ser. I, vol. 9?

[ j
nl )
eS

Z f
) 23

ley Jz
SO digm beg Seas
Ree Sey eon
MELT GG

aS —L. a Te
ae

Fig. 43. Density map of Vatica L. in Malesia; number
of endemics above the hyphen, number of non-ende-
mics below it.

(1925) 140. — Pachynocarpus umbonatus (non HOoK.
f.) Rw . FI. Mal. Pen. 1 (1922) 249, p.p. — Pachyno-
carpus verrucosus RIDL. Fl. Mal. Pen. | (1922) 249,
D-p.

Medium-sized tree. Twigs, petioles, stipules, pani-
cles, calyx outside and ovary + persistently densely
scabrid ferruginous puberulent, leaf undersurface
sparsely so or glabrescent, parts of petals exposed in
bud pale brown puberulent. Twigs 3-4 mm g, stout,
much branched, ribbed, becoming pale brown, rough.
Buds to 3 by 2mm, linear-lanceolate, acute, caducous.
Leaves 12-25 by 5-12 cm, typically elliptic-obovate,
+ thickly coriaceous; base cuneate; margins +
prominently revolute; acumen + abrupt, to 1.5 cm
long, prominent; nerves 7—9(—15) pairs, ascending,
arched, very stout and prominent beneath, evident
above as also the midrib; tertiary nerves subreticulate,
distinctly elevated beneath, hardly so above; petiole
1.7-3 cm long, 24mm g, stout. Panicle to 10cm long,
terminal or axillary, somewhat congested and irregu-
larly branched. Flower bud to 12 by 3 mm, fusiform;
petals pale yellow with a purplish patch at base;
staminal appendages c. 4 length of outer anther cells,
broadly deltoid; stigma short, hardly longer than
ovary, stout, expanding distally into the conical style;
flowers otherwise typical. Fruit subsessile or to 2mm
long pedicellate; calyx lobes equal, to 2.3 by 2.3 cm,
ovate, acute, united except at the apex and fused with
the to 4 by 2.5 cm ovoid apiculate protruding
verrucose corky pericarp.

Distr. Peninsular Thailand and
Malaya, Sumatra.

Ecol. Lowland dipterocarp forests, on hills, valleys
and stream banks, to 500 m.

Vern. Résak mempening, r. laru.

Note. A variable species closely related to V.

in Malesia:

umbonata, with which it appears to be undergoing
local hybridisation.

3. Vatica venulosa BL. Mus. Bot. Lugd.-Bat. 2 (1852)
32: WALP. Ann. 4 (1857) 337; Mia. FI. Ind. Bat. 1, 2
(1859) 502; DC. Prod. 16, 2 (1868) 623; BURCK, Ann.
Jard. Bot. Btzg 6 (1878) 232; Merr. En. Born. (1921)
409: SLtoor. Bull. Jard. Bot. Btzg III, 9 (1927) 78;
AsHTON, Man. Dipt. Brun. (1964) 79, f. 10; ibid.
Suppl. (1968) 36; Gard. Bull. Sing. 31 (1978) 17.

a. ssp. venulosa. — V. bancana SCHErFF. Nat. Tijd. N. I.
31 (1870) 348; ibid. 32 (1873) 407; Burck, Ann. Jard.
Bot. Btzg 6 (1887) 229, t. 27; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 128; BRANDIS & GILG in E. & P. Pfl.
Fam. ed. 1, 3, 6 (1895) 269; K. & V. Bijdr. 5 (1900) 127;
MoLL_ & JANssoNnius, Mikrogr. Holz (1906) 360;
Koorp. Exk. Fl. Java 2 (1912) 622; KoorD.-SCHUM.
Syst. Verz. (1913) Dipteroc. 4; HALL. f. Med. Rijks-
herb. 36 (1918) 4; Merr. En. Born. (1921) 408; GiLG in
in E. & P. Pfl. Fam. ed. 2, 21 (1925) 264; HEYNE, Nutt.
Pl. ed. 2(1927) 1128; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 96; Foxw. Mal. For. Rec. 10 (1932) 263; Sym.
Mal. For. Rec. 16 (1943) 217, f. 107; BROWNE, For.
Trees Sarawak & Brunei (1955) 99; BACKER & BAKH.

f. Fl. Java 1 (1963) 332; MEER & Woop, Sabah For.

Rec. 5 (1964) 305, f. 53. — V. schouteniana SCHEFF.
Nat. Tijd. N. I. 32 (1873) 408. — Dryobalanops
schefferi HANCE, J. Bot. 14 (1876) 307. — Retinoden-
dron bancanum KiNG, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 129. — Retinodendron kunstleri KING, J. R. As.
Soc. Beng. Sc. 62, 2 (1893) 129; BRUHL & KING, Ann.
R. Bot. Gard. Calc. 5, 2 (1896) 157, t. 189B. — V.
kunstleri BRANDIS, J. Linn. Soc. Bot. 31 (1895) 127, t.
3, f. 10; BRANDIS & GiLG in E. & P. Pfl. Fam. ed. 1, 3, 6
(1895) 270; Ripv. Fl. Mal. Pen. 1 (1922) 243. — V.
schefferi BRANDIS, J. Linn. Soc. Bot. 31 (1895)
128. — V. lutea RIDL. Kew Bull. (1926) 60.

Twig, raceme, leaf bud, stipule, midrib above and
petiole + persistently densely shortly pale pink-
brown puberulent; leaf beneath caducously so. Twigs
to 1.5 mm @g apically, thinly coriaceous, terete,
becoming smooth, glabrous. Bud to 2.5 mm long,
ovoid, acute. Stipules to 5 mm long, hastate, fuga-
ceous. Leaves 4-12 by 1.5—5 cm, elliptic to ovate-lan-
ceolate, thinly coriaceous; base cuneate, apex shortly
acuminate; nerves 7—12 pairs, slender, hardly raised
beneath, unraised above, curved at 60°—70°; without
distinct secondaries; midrib slender, prominent
beneath, slightly depressed above; petioles S-9 mm
long. Panicle to 3 cm long, terminal or 1-axillary,
ribbed on drying, singly branched. Flower buds to 1.3
cm long; calyx pink-brown pubescent; flowers other-
wise typical. Fruit pedicel to 2 mm long, hidden in the
bases of the calyx. Calyx glabrous; lobes subequal, to
3 by 1.3 cm, ovate, acute, revolute, bases cordate. Nut
to 1 cm g, globose, shortly sparsely pubescent,
completely hidden by, but free from, calyx; style
remnant to 1.5 cm long, acute.

Distr. Malesia: Malaya (Pahang, Perak), Sumatra
(Palembang), Banka, Billiton, W. Java (Bantam),
Borneo.

1982]

DIPTEROCARPACEAE (Ashton)

35]

Ecol. Very local, on alluvium river banks and fresh
water swamp.

Vern. Résak letop (Mal.), r. banka, r. puteh, r.
seluang, aboh.

b. ssp. simalurensis (SLOoT.) ASHTON, Gard. Bull.

Sing. 31 (1978) 18.— V. simalurensis SLoor. Bull.
Jard. Bot. Btzg III, 9 (1927) 100; HEYNE, Nutt. Pl. ed. 2
(1927) 1131.

Leaves 9-14(—20) by 3.5—5.5(-8) cm, glabrous;
nerves 12-15 pairs; petiole 8-14 mm long. Panicle to 7
cm long.

Distr. Malesia: W. Sumatra (P. Simalur, Mentawei
Is.).

Ecol. As the species.

4. Vatica havilandii BRANDis, J. Linn. Soc. Bot. 31
(1895) 133; Mere. En. Born. (1921) 409; SLoor. Bull.
Jard. Bot. Btzg ITI, 9 (1927) 95; Sym. J. Mal. Br. R. As.
Soc. 19 (1941) 155; Mal. For. Rec. 16 (1943) 220, f.
107; BROWNE, For. Trees Sarawak & Brunei (1955)
100; ASHTON, Man. Dipt. Brun. (1964) 71, f. 10; ibid.
Suppl. (1968) 33.

Young twig, raceme, leaf bud, stipule, and petiole
densely persistently deep fulvous-brown flocculent
tomentose; sparsely so, caducous, on leaf nervation
beneath. Twig to 2.5 mm @ apically, terete or slightly
compressed, smooth or slightly flaked. Bud to 4 by 3
mm, conical, subacute. Stipule to 12 by 2.5 mm, linear,
caducous. Leaves 8-17 by 2.5—5 cm, narrowly oblong
to obovate; base cuneate; acumen to | cm long,
narrow; nerves 15-20 pairs, slender, prominent
beneath, curved towards the apices, at 60°—70°, with
prominent short secondaries; midrib terete, promi-
nently raised beneath, slender, flat to slightly raised
above; petiole 1—-1.2 cm long. Panicle to 8 cm long,
terminal or axillary, terete, singly branched. Flower
bud to 5 mm long, subglobose. Calyx densely rust-
brown powdery tomentose; appendage to connective
as long as anther, stout; flowers otherwise typical.
Fruit pedicel 5 mm long, 2 mm @, pale rufous
pubescent, hidden in the base of the calyx. Calyx lobes
2.5 by 1.5 cm, ovate, acute, revolute, bases subcor-
date. Nut 12 mm @ globose, densely rufous pubescent,
completely hidden in, but free from, the calyx.

Distr. Malesia: Malaya (Perak, Pahang, Treng-
ganu), Borneo (Kapuas valley, W. and Central Sar-
awak, Brunei, Sandakan area).

Ecol. Rare, in Mixed Dipterocarp forests on hills
not far from the coast.

Vern. Resak degong.

Note. This and the following species share with V.
venulosa a very distinctive calyx and thus form a well
defined group within the genus.

5. Vatica chartacea ASHTON, Gard. Bull. Sing. 31
(1978) 18.

Medium-sized tree. Young twigs, stipules and pani-
cles caducous buff puberulent; pedicel, ovary and
parts of petals exposed in bud persistently so; parts
otherwise glabrous. Twig c. 2 mm g apically, much

branched, pale. Buds to 3 by 2 mm @, ovoid, acute;
stipule to 7 by 2 mm, lanceolate, caducous. Leaves
11-25 by 3-10 cm, oblong to obovate, thinly charta-
ceous and wrinkling on drying; base broadly cuneate
or obtuse; acumen to | cm long, prominent; nerves
16-20 pairs, slender but prominent beneath, distinctly
elevated above, arched, with short slender secon-
daries; tertiary nerves sinuately subscalariform,
slender and elevated on both surfaces; midrib stout,
terete and prominent beneath, distinctly elevated
above; petiole 10—22 mm long, slender. Panicle to 5cm
long, to 3-axillary, stout. Flower buds to 6 by 2 mm,
fusiform; sepals subequal, lanceolate, subacuminate;
anthers small, broadly oblong, tapering to the equally
long prominent stout appendages; style columnar,
somewhat longer than ovary, slightly tapering,
rimmed beneath the conical stigma; flowers otherwise
typical. Fruit pedicel to 6 mm long, very slender. Fruit
sepals to 6 by 1.5 cm, subequal, lanceolate, subacute,
cordate at base, 5—7-nerved, ascending and hiding the
to 13 by 11 mm ellipsoid nut.

Distr. Malesia: W. Borneo (Ulu Kapuas), N.E.
Borneo (Bintulu, Sarawak; Beluran and Sandakan to
Tawau).

Ecol. Moist low hillsides and banks of sluggish
rivers, very local.

Vern. Resak bunga, r. banka.

6. Vatica pauciflora (KORTH.) BL. Mus. Bot. Lugd.-
Bat. 2 (1852) 31, f. 7; DC. Prod. 16, 2 (1868) 618; Mia.
Sum. (1861) 191; BuRcK, Ann. Jard. Bot. Btzg (1887)
226; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 124;
ASHTON, Gard. Bull. Sing. 31 (1978) 19. — Retinoden-
dron pauciflorum KorTH. Kruidk. (1841) 58. — Va-
teria pauciflora WALP. Rep. 5 (1845) 126; DC. Prod.
16, 2 (1868) 626. — Elaeogene sumatrana Miq. Sum.
(1861) 460, 183; M. A. in DC. Prod. 15, 1 (1866) 1257;
HALL. f. Med. Rijksherb. 36 (1918) 14. — V. zollinger-
tana DC. Prod. 16, 2 (1868) 618; BuRcK, Ann. Jard.
Bot. Btzg 6 (1887) 227, t. 29-1; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 124; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 126. — V. wallichii Dyer, J. Bot. 12 (1874) 154;
Burk. & Foxw. J. Str. Br. R. As. Soc. 86 (1922) 273;
CrRaAIB, Fl. Siam. Enum. | (1925) 141; HEYNE, Nutt. Pl.
ed. 2 (1927) 1132; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 123, f. 12; Foxw. Mal. For. Rec. 10 (1932) 269;
Burk. Dict. (1935) 2225; Sym. Mal. For. Rec. 16
(1943) 230, f. 106, 107, 110. — V. lamponga Burck,
Ann. Jard. Bot. Btzg 6 (1887) 227, t. 29, f. 3a—d;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 123; BRANDIS &
GitG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 269; HEYNE,
Nutt. Pl. ed. 1, 3 (1917) 313; ibid. ed. 2 (1927) 1131;
GILG in E. & P. Pfl. Fam. ed. 2, 21 (1925) 264. — V.
forbesiana Burcxk, Ann. Jard. Bot. Btzg 6 (1887) 228:
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 124; HEYNE,
Nutt. Pl. ed. 2 (1927) 1131. — V. ruminata Burck,
Ann. Jard. Bot. Btzg 6 (1887) 227, t. 29, f. 4; HEYNE,
Nutt. Pl. ed. 2 (1927) 1132. — Pachynocarpus wallichii
KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 135, p.p.;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 136; Rip. FI.
Mal. Pen. 1 (1922) 250, p.p.; BURK. J. Str. Br. R. As.

352

Soc. 81 (1920) 75.— Pachynocarpus ruminatus
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 136.— V.
ovalifolia Ripv. J. Str. Br. R. As. Soc. 54 (1909)
26. — V. kelsalli Rov. J. Str. Br. R. As. Soc. 54 (1909)
27; Fl. Mal. Pen. 1 (1922) 244. — Pachynocarpus
umbonatus (non HOOK. f.) RIDL. Fl. Mal. Pen. 1 (1922)
249, p.p. — Pachynocarpus ridleyanus (non ANDERS.)
RIDL. Fl. Mal. Pen 1 (1922) 250, p.p. — V. sumatrana
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 120, f. 11;
HEyYNE, Nutt. Pl. ed. 2 (1927) 1131; BACKER & BAKH. f.
Fl. Java 1 (1963) 332.

Small or medium-sized tree. Outside of perianth +
persistently pale grey-brown cinereous; ovary, pani-
cles, and twig apices caducously so; elsewhere glabres-
cent. Twig c. 3 mm @ apically, much branched, pale
grey-brown, usually minutely rugulose. Buds small,
ovoid; stipules to 8 mm long. linear, becoming
reflexed. Leaves 6.5—20 by 2.2—8 cm, variable in size,
elliptic-lanceolate, thinly coriaceous; base cuneate;
acumen to 1.5 cm long, prominent; nerves 6—9 pairs,
arched, ascending, slender and hardly more elevated
beneath than above; petiole 10-18 mm long, smooth,
drying black. Panicle to 9 cm long, terminal or
subterminal axillary, many-flowered, irregularly
branching. Flower bud to 10 by 3 mm, fusiform;
appendages minute, hardly exceeding anthers; style
columnar, longer than ovary, expanding somewhat
distally below the small conical style; flowers other-
wise typical. Fruit pedicel to 7 by 3 mm, prominent;
calyx lobes to 5 mm @, hemispherical, incrassate, +
adnate round the impressed base of the nut; nut to 3 by
3 cm, ovoid, subacute, with 3 distinct loculicidal
furrows; pericarp thick, corky, verrucose.

Distr. Peninsular Thailand, and in Malesia:
Malaya, Sumatra (Palembang, Lampong), Banka.

Ecol. Banks of sluggish rivers, fresh water swamps;
common.

Vern. Resak laru, r. paya, r. ayer, r. pasir, damar
mata kuching (Malaya), r. padang, r. rawang (Suma-
tra).

Notes. Occupying the same habitat as V. umbonata
and indistinguishable from it when sterile; it is by the
fruit calyx nevertheless consistently distinguishable.
Though V. pauciflora is confirmed only as far north as
Songkhla, V. thorelii PIERRE of Cochinchina, based on
THOREL & HARMAND’S herb. Pierre 1586, is known
only in flower and is indistinguishable in that condi-
tion from it; moreover I have seen undoubted fruit of
V. pauciflora washed up on the beach at Kompong
Som, S.E. Cambodia.

See also 56. V. obtusa BURCK.

A specimen from Sikundur For. Res., W.N.W. of
Medan (N.E. Sumatra) (DE WILDE & DE WILDE-
Duyrses 19537) bears fruit which differs in having
greatly enlarged adnate sepals, in this resembling V.
umbonata, so far unknown from Sumatra. This may
represent a hybrid between the two species, neverthe-
less.

7. Vatica ridleyana BRANDIS in Hook. f. Ic. Pl. 25
(1895) t. 2401; J. Linn. Soc. Bot. 31 (1895) 122, t. 3, f.

FLORA MALESIANA

[ser. I, vol. 9?

5; Burk. & Foxw. J. Str. Br. R. As. Soc. 86 (1922) 277;
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 49, 73, f. 1d;
Foxw. Mal. For. Rec. 10 (1932) 268; Burk. Dict.
(1935) 224; Sym. Mal. For. Rec. 6 (1943) 227, f.
107. — Pachynocarpus ridleyanus ANDERSON, Index
Bot. Gard. Sing. (1912) 9; BURK. & Foxw. J. Str. Br.
R. As. Soc. 86 (1922) 272; RIDL. Fl. Mal. Pen 1 (1922)
250, p.p.-

Medium-sized tree. Twigs, petioles, panicles,
ovaries and parts of perianth exposed in bud +
persistently ocherous cinereous, parts elsewhere gla-
brescent. Twig c. 2-3 mm @g apically, much branched,
brown rugulose. Buds to 3 by 2 mm, ovoid. Leaves
6-14 by 3—7 cm, elliptic to narrowly ovate or obovate,
coriaceous, dull beneath; base cuneate; apex subacute
or shortly broadly acuminate; margin narrowly subre-
volute; nerves S—7 pairs, ascending, slender but promi-
nent beneath, evident above as also the midrib and
reticulate tertiary nerves; petiole 8-15 mm long, 2-3
mm @, stout. Panicle to 15 cm long but usually
shorter, terminal or axillary, irregularly branched.
Flower bud 7 by 2 cm, fusiform; appendage short,
deltoid; style somewhat longer than ovary, columnar,
expanding into the prominent conical stigma; flowers
otherwise typical. Fruit pedicel short, stout; calyx
lobes to 3 by 3 mm, short, deltoid, incrassate, relaxed,
adnate to the base of the 4.5 by 1.8 cm fusiform
beaked coarsely verrucose nut.

Distr. Malesia: Sumatra (Palembang), Singapore.

Ecol. Mixed Dipterocarp forest at low altitudes;
rare.

Vern. Resak buah cana.

8. Vatica soepadmoi ASHTON, Gard. Bull. Sing. 31
(1978) 19.

Small tree. Twigs, leaf buds, petioles, midrib above,
and panicles densely persistently pale brown scabrid
puberulent, nuts evenly so; nervation beneath sparsely
so. Twigs c. 2mm @, ribbed at first, becoming terete.
Leaf buds to 8 by 4 mm, lanceolate, acute. Leaves
7.5—12 by 3—5.5 cm, elliptic, oblong to narrowly ovate,
coriaceous, somewhat bullate between the nerves;
margin subrevolute; base obtuse; acumen to 1.5 cm
long, slender, prominent; nerves c. 11 pairs, arched,
tending to branch within the margin and form a +
indistinct looped intramarginal nerve, prominent
beneath, shallowly depressed above, with short
slender secondary nerves; tertiary nerves subreticu-
late, evident beneath, + obscure above; midrib
stoutly prominent beneath, evident but + channelled
above; petiole 10-15 mm long, slender. Panicle to 6.5
cm long, axillary, hardly branched. Flowers unknown.
Fruit pedicel to 6 mm long, prominent; calyx lobes
equal, to 18 by 10 mm, lanceolate, acute, recurved
inwards and + revolute thus resembling claws; nut
ovoid, not known at maturity.

Distr. Malesia: E. Sumatra (Upper Riouw: Pekan-
baru, Singkep).

Ecol. Low hills.

9. Vatica bella SLoor. Bull. Jard. Bot. Btzg III, 9

1982]

(1927) 102, f. 6; Foxw. Mal. For. Rec. 10 (1932) 265;
Burk. Dict. (1935) 2223; Sym. Mal. For. Rec. 16
(1943) 218, f. 107.

Medium-sized tree to 50 m tall. Twigs, leaf buds,
petioles, midrib above and panicle densely persis-
tently rufous cinereous, calyx and ovary caducously
so, leaf undersurface sparsely caducously so. Twig c. 2
mm @ apically, slender, terete, much branched, pale
brown. Bud to 3 by 2 mm, small, ovoid; stipules
fugaceous, not seen. Leaves S5—14 by 2.6 cm, elliptic-
obovate, coriaceous; base cuneate; acumen to | cm
long or short; margin frequently subrevolute; nerves
11 -12 pairs, arched, ascending, slender but distinctly
elevated beneath, evident above as also the midrib;
tertiary nerves densely subreticulate, hardly elevated
on either surface; petiole 5-15 mm long, short,
relatively slender. Panicle to 2 cm long, axillary,
congested, irregularly branched. Flower buds to 8 by 3
mm, lanceolate; appendage to connective as long as
anther cells, deltoid, prominent; style slender, twice as
long as ovary; stigma shortly conical; flowers other-
wise typical. Fruit pedicel to 3 by 2 mm; calyx lobes to
30 by 13 mm, lanceolate, acute, c. 6 mm wide at the
revolute base, becoming reflexed; nut to 2.5 by 2.5 cm,
turbinate, verrucose, with to 1 cm long persistent
tapering style remnant.

Distr. Malesia: Malaya (south from Perak and
Pahang).

Ecol. Locally common in Mixed Dipterocarp
forest, undulating land beneath 250 m.

Vern. Resak kéluang, r. laru, damar kéluang.

10. Vatica rassak (KORTH.) BL. Mus. Bot. Lugd.-Bat.
2 (1852) 31, incl. var. subcordata BL.; WALP. Ann. 4
(1857) 337; Mia. Fl. Ind. Bat. 1, 2 (1859) 502; Ann.
Mus. Bot. Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2
(1868) 619; BuRcK, Ann. Jard. Bot. Btzg 6 (1887) 225;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 125; BRANDIS &
GiLG in E. & P. Pfl. Fam. ed. 1, 3,6 (1895) 270; GRESH.
Ind. Mercuur 23 (1900) n. 37, tab.; Schets. Nutt. Ind.
Pl. (1900) t. 50; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 313;
ibid. ed. 2 (1927) 1130; Merr. En. Born. (1921) 409;
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 73, 104, f. 1c;
ibid. III, 17 (1942) 223, f. 22-24; ASHTON, Gard. Bull.
Sing. 31 (1978) 20. — Retinodendron rassak KORTH.
Kruidk. (1841) 56, t. 8. — Vateria rassak WALP. Rep.
5 (1845) 126. — V. papuana Dyer, J. Bot. 16 (1878)
100; Burck, Ann. Jard. Bot. Btzg 6 (1887) 229;
K.Scu. & HOLLR. FI. Kais.-Wilh. Land (1889) 52;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 127; BRANDIS &
GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 270; DIELs,
Bot. Jahrb. 57 (1922) 463; LANE-POOLE, For. Res.
Papua (1925) 120; SLoot. Nova Guinea 14 (1926) 226;
Bull. Jard. Bot. Btzg III, 9 (1927) 73, 112, f. 1b; MERR.
Philip. J. Sc. 30 (1926) 411; WHITE & FRANCIS, Proc.
R. Soc. Qsl. 38 (1927) 247; HEYNE, Nutt. Pl. (1927)
1129; Foxw. Philip. J. Sc. 67 (1938) 328; SLoor. Bull.
Jard. Bot. Btzg III, 17 (1942) 233, f. 27; apud Holth. &
Lam, Blumea 5 (1942) 214; Reinwardtia 2 (1952) 63, f.
21; MEWER & Woop. Sabah For. Rec. 5 (1964) 314, f.
56; ASHTON, Man. Dipt. Brun. Suppl. (1968) 35, f.

DIPTEROCARPACEAE (Ashton)

353

5. — Vateria papuana Dyer ex HEMSL. Bot. Chall. 1, 4
(1884-85) 123, 287, 296, t. 64B; K.Scu. & HOLtr. FI.
Kais. Wilh. Land (1889) 52; HeyNe, Nutt. Pl. ed. 2
(1927) 1129. — V. moluccana Burck, Ann. Jard. Bot.
Btzg 6 (1887) 226, t. 26; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 124; HeyNe, Nutt. Pl. ed. 1, 3 (1917) 313; ibid.
ed. 2 (1927) 1129.— Retinodendron moluccanum
Heim, Rech. Dipt. (1892) 104. — V. schumanniana
GILG, Bot. Jahrb. 18 (1894) Beibl. 45: 38; BRANDis, J.
Linn. Soc. Bot. 31 (1895) 127; BRANDIS & GILG in E. &
P. Pfl. Fam. ed. 1, 3, 6 (1895) 269, fig.; K.ScH. & LAuT.
Fl. Schutzgeb. (1901) 451; Diexs, Bot. Jahrb. 57 (1922)
463; (in Index Kewensis erron. under Shorea). — V.
celebensis BRANDIS, J. Linn. Soc. Bot. 31 (1895) 126, t.
3, f. 6; SLoor. Bull. Jard. Bot. Btzg III, 9 (1927) 77;
ibid. III, 17 (1942) 254; Reinwardtia 2 (1952) 65. — V.
subcordata (BL.) HALL. f. Med. Rijksherb. 36 (1918) 4;
SLoot. Bull. Bot. Gard. Btzg III, 17 (1942) 228, f.
25-26. — V. celebica SLooT. Bull. Bot. Gard. Btzg III,
17 (1942) 237, f. 28-29. — Fig. 44.

Twig, petioles, buds, and stipules outside (glabrous
within) very shortly evenly persistently pale buff
pubescent, leaf nervation beneath sparsely so. Twig c.
3 mm @g, stout, crooked, ribbed, becoming rugose,
flaky, pale grey-brown; stipule scars prominent, hori-
zontal; internodes 1-3 cm long. Bud to 4 by 4 mm,
ovoid-conical, subacute. Stipules to 14 by 4 mm,
lorate, subacute, subpersistent. Leaves 13-32 by 5-11
cm, oblong to narrowly elliptic, thickly coriaceous;
base broadly cuneate to subcordate; acumen to 1.5cm
long; nerves (10—)16—20 pairs, prominent beneath,
slightly elevated above, arched at 50°—-60°, with short
hardly elevated secondary nerves; tertiary nerves
reticulate; midrib prominent beneath, applanate
above; petiole 2-2.5 cm long, stout, not geniculate,
drying pale buff pubescent. Panicle to 14 cm long,
terminal or axillary, ribbed, at first shortly evenly buff
pubescent, becoming pale brown flaky; irregularly
branched, with many branches near base, appearing
fascicled. Flower bud to 14 by 3 mm, fusiform; calyx
densely shortly pale buff pubescent; flowers otherwise
typical. Fruit glabrous. Pedicel to 3 mm long, stout.
Calyx lobes to 12 by 7 mm, deltoid, acute, incrassate,
reflexed, recurved. Nut to 5 by 3.5 cm, oblong,
symmetrical and obtuse to ovoid, more or less
prominently attenuate-acute and asymmetrically
twisted, sometimes irregularly pitted, furrowed at the
sutures, minutely verruculose and rugulose; pericarp
thick, corky.

Distr. Malesia: Borneo (E. of Rejang valley and
Sampit, commonest on E. coast), S. Philippines (Sulu
Is.: Tawi Tawi), Celebes, Moluccas (Sula Is.: Mangoli;
Morotai, Halmaheira, Batjan, Obi Is., Aru Is.), New
Guinea, Sudest I.

Ecol. River-banks in Borneo, elsewhere also on
hills to 400 m, locally abundant.

Vern. Résak irian, r. damau, r. ayér, r. tébong
(Sabah), damar déréh, d. putéh, nunuh, singkodoh
putéh (Celebes), kokolaka, bou-ura, por, damar hiru,
manauri, laintoti, wakaju, imoimo, asuk, baia, guim-
bur, lagima, mutani, owi, simbiau (New Guinea),

354

Fig. 44. Fruiting twig of Vatica rassak (KORTH.) BL.
near Sarmi, c. 200 km west of Hollandia, Irian Jaya
(Photogr. KARSTEL, Dec. 1957).

doyong (Sula Is.), damar hiru, salo hiru, damar atung,
geru (Moluccas).

Notes. Another variable semi-gregarious chiefly
riparian species (see e.g. V. umbonata).

The above description defines my interpretation of
this variable species. The large, oblong-elliptic, coria-
ceous leaves with long petiole, and the large, corky
nut, are characteristic though the nut shape is very
variable (in part owing to the degree of maturity on
herbarium specimens). Its distribution into seasonal
areas and its semi-gregarious ecology parallel that of
other polymorphic species such as Anisoptera costata
Korth., A. thurifera(BLco) BL., and Vatica umbonata
(HooK. f.) BURCK. The variation is locally too great to
clearly distinguish geographical subspecies.

Some hybridisation appears to occur with V.
umbonata in East Sabah.

11. Vatica granulata SLooT. Bull. Jard. Bot. Btzg III,
95 (1927) Mel OZ rbd LIE <(1941) MSGsafes 20;
ASHTON, Man. Dipt. Brun. (1964) 70, f. 10; ibid.
Suppl. (1968) 32; Gard. Bull. Sing. 31 (1978) 21.

a. ssp. granulata.

Y oung twig, leaf bud, stipule, petiole and nervation
beneath densely shortly yellow scabrid tomentose at
first, glabrescent or sparsely pubescent on nervation.
Twig to 3 mm @g apically, stout, angular, persistently
papery flaky. Bud to 4 by 3 mm, conical, subacute.
Stipule to 6 by 4 mm, hastate, subacute, caducous.
Leaves 10-20 by 2.7—7 cm, coriaceous, narrowly
obovate; base narrowly obtuse; acumen to 6 mm long;
nerves 12-14 pairs, prominent beneath, well spaced,

FLORA MALESIANA

[ser. I, vol: 9?

slightly sunken above. Petiole 1.2-2 cm long, stout.
Flowers unknown. Inflorescence unknown. Fruit pedi-
cel to | mm long, fruit subsessile. Calyx lobes to 7 by 4
mm, chartaceous, deltoid, brittle, reflexed, glabrous.
Nut to 3.5 by 4cm, ovoid, obtuse or subacute, coarsely
granulate, dehiscing along 3 distinctly furrowed
sutures at germination.

Distr. Malesia: Borneo.

Ecol. Widespread, locally abundant, on high ridges
at 500-1200 m.

Vern. Resak ranting bersisek.

b. ssp. sabaensis ASHTON, Gard. Bull. Sing. 31 (1978)
Pail.

Differing as follows: Leaf by 6-10 cm, petiole 12-20
mm long. Stipules to 3 by 2 cm, oblong to lanceolate,
acute, obtuse to subcordate, subpersistent. Fruit
sepals to 20 by 12 mm, elliptic, revolute, reflexed.

Distr. Malesia: Borneo (Crocker range, S.W.
Sabah to Kelabit Highlands, N.E. Sarawak).

Ecol. As type subspecies, to 1700 m.

Vern. Reésak pengasoh.

12. Vatica sarawakensis Heim, Bull. Mens. Soc. Linn.
Paris 2 (1891) 970; Rech. Dipt. (1892) 109; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 124; SLoor. Bull. Jard. Bot.
Btzg III, 9 (1927) 106, f. 7; BRowNeE, For. Trees
Sarawak & Brunei (1955) 101; ASHTON, Gard. Bull.
Sing. 20 (1963) 252; Man. Dipt. Brun. (1964) 77, f. 10:
ibid. Suppl. (1968) 36; Gard. Bull. Sing. 22 (1967) 262;
MEER & Woop, Sabah For. Rec. 5 (1964) 319, f.
57. — Retinodendropsis aspera HEIM, C. R. Assoc. Fr.
Pau 1892 (1893) 470; cf. ASHTON, Gard. Bull. Sing. 22
(1967) 262.— V. ramiflora SLooT. Bull. Jard. Bot.
Btzg III, 9 (1927) 118, p.p.; in Merr. Pl. Elm. Born.
(1929) 205; Bull. Bot. Gard. Btzg III, 17 (1942) 240, f.
30; Reinwardtia 5 (1961) 479; BRowNnE, For. Trees
Sarawak & Brunei (1955) 101.

Twig, raceme, leaf bud, stipule and petiole densely
pale brown to fulvous scabrid tomentose, sparsely so
on leaf nervation beneath. Twig to 5 mm @ apically,
stout, becoming ribbed, smooth or finely cracked. Bud
to 8 by Smm, conical, subacute. Stipule to 15 by Smm,
narrowly hastate, acute. Leaves 22-35 by 7-15 cm,
thinly coriaceous, oblong to obovate; base obtuse;
acumen to 1.4 cm long, deltoid; nerves 15-28 pairs,
prominent beneath, flat or slightly raised above;
petiole \-2 cm long, to4mm @, stout. Panicle to 12cm
long, axillary to ramiflorous (rarely terminal), to
3-axillary, terete or angled; much branched, the
branchlets cymose; bracts to 8 by 4 mm, narrowly
deltoid, subacute, densely rufous tomentose, subper-
sistent. Flower bud to 8 mm long; calyx shortly
red-brown pubescent; appendage to connective over 4
length of anther, prominent; flower otherwise typical.
Fruit glabrous; pedicel to 2 mm long. Calyx lobes
subequal, to 16 by 7 mm, oblong-hastate, obtuse,
becoming reflexed, somewhat revolute. Nut to 2.5 by
2.5 cm, subglobose to ovoid, verrucose-lenticellate,
with 3 distinct sutures uniting at the apex; style
remnant short but prominent.

1982]

Distr. Malesia: Borneo (Sarawak to Sabah; S.E.
Borneo to P. Laut and Balikpapan).

Ecol. Scattered in Mixed Dipterocarp forest on
clay rich soils on undulating land and hills below 1000
m.

Vern. Résak daun bésar, r. sarawak, damar tingkis.

Note. The fruit sepals of E. Borneo specimens (V.
ramiflora SLooT.) are foliose and rather broad, these
on Sarawak specimens narrow and _incrassate.
Further collections may justify distinction of two
subspecies in V. oblongifolia.

13. Vatica albiramis SLoor. Bull. Jard. Bot. Btzg III,
9 (1927) 101, f. 5; in Merr. Pl. Elm. Born. (1929) 205;
AsHTON, Man. Dipt. Brun. (1964) 66, f. 10; ibid.
Suppl. (1968) 30; MevER & Woop, Sabah For. Rec. 5
(1964) 305, f. 52.

Parts glabrous but for the shortly sparsely caducous
pubescent calyx and shortly fulvous caducous pubes-
cent ovary. Twig to | mm g apically, slender, smooth
to rugulose. Bud to 2 by | mm, conical, acute. Stipule
to 16 by 3.5 mm, hastate, subacute, caducous. Leaves
8-20 by 3-7 cm, thinly coriaceous, elliptic to lanceo-
late; base + narrowly cuneate; acumen to 1.5cm long,
narrow; nerves 4-6 pairs, rather broad, slightly raised
on both surfaces, more prominently so beneath,
curved, continuing almost parallel to the margin
before terminating, at 45°—50°(—70°); midrib broad,
rounded, slightly raised on both surfaces; tertiary
nerves subscalariform; petiole 1-1.3 cm long. Panicle
to 28 cm long, terminal or axillary, lax, spreading,
glabrous, terete; doubly or trebly branched. Flower
bud to 1.5 cm long, slender; calyx shortly sparsely
pubescent; corolla rich lemon-yellow; appendage to
connective rudimentary; flowers otherwise typical.
Calyx lobes subequal, to 12 by 4 mm, ovate, obtuse,
tapering to 3.5 mm broad at base, coriaceous, gla-
brous, becoming reflexed. Nut to 1.2 by 1.2 cm,
globose, without style remnant, shortly fulvous
pubescent.

Distr. Malesia: Borneo (Rejang valley to Kina-
balu; E. Sabah).

Ecol. Clay rich soils, low hills and ridges; to 1400 m
in Kinabalu area.

Vern. Résak ranting puteh, r. puteh.

Note. Differing from the closely related V. hullettii
of Malaya in the narrow fugaceous hastate stipules
and totally glabrous young parts and petiole.

14. Vatica oblongifolia Hook. f. Trans. Linn. Soc. 23
(1862) 160; Mia. Ann. Mus. Bot. Lugd.-Bat. 3 (1867)
85; DC. Prod. 16, 2 (1868) 619; WaLP. Ann. 7 (1868)
378; Burck, Ann. Jard. Bot. Btzg 6 (1887) 229;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 126, t. 3, f. 11;
Merk. En. Born. (1921) 409; SLoor. Bull. Jard. Bot.
Btzg III, 9 (1927) 109, f. 8; ibid. II, 17 (1941) 135;
BROWNE, For. Trees Sarawak & Brunei (1955) 101;
ASHTON, Man. Dipt. Brun. (1964) 75, f. 10, p.p.; ibid.
Suppl. (1968) 34; Gard. Bull. Sing. 22 (1967) 264, pl.
3-7; MEER & Woop, Sabah For. Rec. 5 (1964) 314, f.
55, pl. 30b (stem). — Fig. 45.

DIPTEROCARPACEAE (Ashton)

Fig. 45. Stem-base of young tree of Vatica oblongifolia

Hook. f. ssp. oblongifolia, with flowering and fruiting

twigs. Sibulu R., Mengalang For. Res., Beaufort

Distr. (Photogr. G. H. S. Woop, July 1954, KEP
80255).

Note. The taxa I here recognise as subspecies are
rather constant and rarely difficult to identify.
Though the full range of variation they together
manifest is great, the distinctive characters of the
depressed twig and short dense vinose indumentum
unites them as a single entity.

KEY TO THE SUBSPECIES

1. Petiole at most 14 mm long; leaf elliptic
b. ssp. elliptifolia
1. Petiole at least 15 mm long.
2. Fruit calyx lobes more than 10 by 6 mm; nerves at
least 26 pairs.
3. Leaf broadly oblong, base + cordate
c. ssp. crassilobata
3. Leaf narrowly obovate, base + cuneate
d. ssp. multinervosa

356

2. Fruit calyx lobes at most 4 by 3 mm; nerves at
most 18 pairs.

4. Leaf narrowly elliptic, base cuneate, petiole

1.5—2.5 cm long e. ssp. Selakoensis

4. Leaf obovate to oblong, base obtuse, petiole

2.5—5 cm long a. ssp. oblongifolia

a. ssp. oblongifolia. — ASHTON, Gard. Bull. Sing. 22
(1967) 264, pl. 3. — V. furfuracea BuRCK, Ann. Jard.
Bot. Btzg 6 (1887) 228; Becc. For. Born. (1902)
570. — Fig. 45.

Young twig, raceme, calyx, leaf bud, stipule, petiole
and leaf nervation densely shortly evenly persistently
vinous cinereous. Twig c. 4 by 2 mm @ apically,
compressed; becoming terete, smooth to finely
cracked and flaked. Leaf bud to 3 by 5 mm, conical,
subacute, compressed. Stipule to 4.5 by 1.2 cm,
oblong, obtuse. Leaves 10-31 by 4.5—10 cm, coria-
ceous, oblong to obovate; base obtuse; apex +
abruptly tapering, with or without an up to | cm long
acumen; nerves 10-18 pairs, prominent beneath,
curved distally, at 50°-80°; with short + prominent
secondary nerves; midrib prominent, terete beneath,
depressed above; petiole 2.5—-S cm long, geniculate.
Panicle to 8 cm long, compressed, singly or doubly
branched, terminal or to 3-axillary. Flower bud to 1.5
cm long, slender; petals cream, purplish towards base
at first; flowers typical. Fruit pedicel to 8 mm long.
Calyx lobes to 3 by 2 cm, equal, thickly coriaceous,
deltoid, acute, + reflexed. Nut to 2 cm long and @,
globose, faintly 3-sulcate.

Distr. Malesia: Borneo (widespread).

Ecol. Leached shallow soils on low hills and on
ridges to 700 m inland.

Vern. Résak mambangan, r. daun panjang.

b. ssp. elliptifolia ASHTON, Gard. Bull. Sing. 22 (1967)
265, pl. 7; Man. Dipt. Brun. Suppl. (1968) 34.

Twigs c. 3 by 2mm @. Leaves 12-20 by 3.5—7 cm,
elliptic or obovate; base broadly cuneate; acumen to |
cm long, short, slender; nerves 14-16 pairs, slender,
prominent beneath; petiole 10-14 mm long, short.
Fruit calyx lobes to 8 by 4 mm, oblong, obtuse,
revolute.

Distr. Malesia: Borneo (Central Sarawak, Brunei).

Ecol. Rare, deep sandy soils on subcoastal hills.

c. ssp. crassilobata ASHTON, Gard. Bull. Sing. 22
(1967) 265, pl. 5; Man. Dipt. Brun. Suppl. (1968) 34.

Twigs c. 4 by 2 mm @, stout. Leaves 11.5—21 by
4.5-10.5 cm, broadly oblong; base obtuse or cordate;
acumen short, broad; nerves 16-23 pairs, prominent
beneath; petiole 1.8—2.8 cm long. Fruit calyx lobes to
15 by 12 mm, obovate, obtuse to emarginate, subro-
tate.

Distr. Malesia: Borneo (Sarawak, Brunei).

Ecol. Local, Mixed Dipterocarp forests on deep
leached sandy soils near Pleistocene coastlines.

d. ssp. multinervosa ASHTON, Gard. Bull. Sing. 22
(1967) 265, pl. 6; Man. Dipt. Brun. Suppl. (1968) 34.

FLORA MALESIANA

[ser. I, voleg2

Twig c.4 by 2mm, stout. Leaves 14-31 by 4-8.5 cm,
narrowly obovate; base narrowly obtuse or cuneate;
acumen to 1.5 cm, long, slender; nerves 18—27 pairs,
very prominently elevated beneath, with prominent
short secondaries and tertiaries; petiole 1.5—2.5 cm
long. Fruit calyx lobes to 13 by 8 mm, oblong, obtuse,
reflexed.

Distr. Malesia:
Nunukan I.).

Ecol. Frequent on deep fertile clay-rich soils;
especially on volcanic and basement rocks.

Borneo (Sarawak, Sabah to

e. ssp. Selakoensis ASHTON, Gard. Bull. Sing. 22 (1967)
264, pl. 4; Man. Dipt. Brun. Suppl. (1968) 34.

Twig c. 2 by 1 mm, slender. Leaves 6.5—22 by 2.5-6.5
cm, narrowly elliptic, base cuneate, acumen to 1.5 cm
long, slender; nerves 11-18 pairs, slender, hardly
raised beneath; petiole 1.5—2.5 cm long. Fruit calyx
lobes to 4 by 3 mm, oblong, obtuse.

Distr. Borneo (W. Sarawak).

Ecol. Abundant in summit forests of granodiorite
mountains, 600-1400 m.

15. Vatica dulitensis Sym. Gard. Bull. S.S. 8 (1934) 35,
pl. 10; BRowng, For. Trees Sarawak & Brunei (1955)
100; ASHTON, Man. Dipt. Brun. (1964) 69, f. 10, pl. 22
(bark); ibid. Suppl. (1968) 31; MEER & Woop, Sabah
For. Rec. 5 (1964) 308.

Young twig, raceme, leaf bud, stipule (both sur-
faces), petiole, and leaf nervation beneath shortly
densely dark vinous tomentose, caducous on ner-
vation, persistent elsewhere. Twig to | mm @ apically,
slender, much branched, becoming smooth glabrous.
Bud to 2 by 1.5mm, broadly ovoid, obtuse. Stipule to
3 by | mm, narrowly hastate, acute, caducous. Leaves
4-11 by 0.8-3.2 cm, thinly coriaceous, narrowly
obovate to elliptic-lanceolate, base cuneate; acumen
to 1.5 cm long, caudate; nerves 10-12 pairs, slender,
hardly raised beneath, curved, at 50°—60°, with short
slender secondaries; midrib slender, prominent
beneath, applanate above; petiole 6-10 mm long,
slender. Panicle to 2.5 cm long, singly or doubly
branched, terminal or axillary. Flower bud to 6 mm
long, small. Ca/yx densely vinous cinereous; corolla
cream; appendage to connective short, acute; flowers
otherwise typical. Fruit pedicel to 3 mm long, slender.
Calyx lobes equal, to 14 by 5mm, oblong, obtuse, base
to 5 mm broad, glabrous outside, vinous caducous
puberulent within, becoming rotate to reflexed. Nut to
8 mm @, globose; style remnant to | mm long, linear,
densely shortly vinous tomentose.

Distr. Malesia: Borneo (Sarawak to Sabah, Bulun-
gan).

Ecol. Locally abundant on shale ridges, to 1350 m
in Kinabalu area, occasional on undulating clay rich
soils in lowlands.

Vern. Resak tiong.

16. Vatica pedicellata BRANDIS, J. Linn. Soc. Bot. 31
(1895) 125, t. 3, f. 12-14; Merr. En. Born. (1921) 409;
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 111, f. 9;

1982]

DIPTEROCARPACEAE (Ashton)

357

BROWNE, For. Trees Sarawak & Brunei (1955) 101;
ASHTON, Man. Dipt. Brun. Suppl. (1968) 35, f. 5.

Twig, bud, petiole and stipule persistently densely
shortly vinous pubescent. Twig c. 2 mm @ apically,
slender, subterete to slightly compressed, lax; stipule
scars short, horizontal. Bud to 4 by 2 mm, ovoid,
compressed, acute. Stipule to 7 by 2 mm, lorate,
subacute, caducous. Leaves 9-23 by 3.5-7.5 cm,
medium-sized, elliptic to lanceolate, coriaceous, with
somewhat revolute margin; base obtuse, rarely
cuneate, acumen to 2 cm long, slender; nerves 9-15
pairs, slender, hardly raised on either surface but more
prominent beneath, at 60°—80°, arched, tending to
anastomose distally forming an indistinct looped
intramarginal nerve; with many short slender secon-
daries; tertiary nerves reticulate; midrib prominent
beneath, slender but elevated above yet set in a distinct
trough; petiole 1-2.5 cm long, not geniculate, terete,
drying rugulose, shortly evenly vinous pubescent.
Panicie to 8 cm long, axillary, nbbed, compressed,
slender, densely shortly evenly persistently vinous
pubescent; singly or slightly doubly branched,
branchlets bearing to 6 + secund flowers; bracteoles
fugaceous. Flower bud to 6 by 2 mm, fusiform; calyx
vinous puberulent; petals cream with a purplish
suffusion outside; flowers otherwise typical. Fruit
entirely shortly evenly vinous pubescent, caducous on
calyx, otherwise persistent. Pedicel to 8 mm long,
slender. Calyx lobes to 6 by 4 mm, oblong, obtuse,
revolute, reflexed. Nut to 18 by 22 mm, becoming
subglobose, subacute; sutures obscure.

Distr. Malesia: Borneo (Sarawak west of the Lupar).

Ecol. Locally frequent in Heath forest, usually on
shallow rocky soils near coast.

Note. Clearly close to V. oblongifolia but lacking its
distinctive compressed twig.

17. Vatica rotata ASHTON, Gard. Bull. Sing. 22 (1967)
270, pl. 13; Man. Dipt. Brun. Suppl. (1968) 36, f. 5.

Twigs, leaf bud, petiole, and midrib beneath persis-
tently densely vinous sericeous. Twig c. 1 mm g,
slender, much branched, terete, smooth. Leaf bud c. 2
by | mm, ovoid, subacute. Stipule unknown, fuga-
ceous. Leaves 5.5-10 by 3-5 cm, broadly elliptic,
ovate, coriaceous; base obtuse; acumen to 8 mm long;
nerves 9-11 pairs, arched, at 65°—80°, hardly raised on
either surface; tertiary nerves slender, densely reticu-
late; midrib prominent beneath, evident towards base
above, otherwise obscure, depressed; petiole 8-12 mm
long, slender, drying vinous to black. Flowers un-
known. Fruit and inflorescence entirely persistently
evenly vinous sericeous. Panicle to 1.5 cm long,
terminal or axillary, slender, terete, singly branched.
Fruit pedicel to 9 mm long, slender. Calyx lobes to 7 by
4 mm, equal, suborbicular, revolute, rotate. Nut
(immature) c. 4 mm g, subglobose, obtuse, with 3
indistinct sutures.

Distr. Malesia: Borneo (Central Sarawak, Kapuas
valley).

Ecol. Mixed Dipterocarp—Heath forest ecotone,
deep sandy soil.

18. Vatica vinosa ASHTON, Gard. Bull. Sing. 19 (1962)
318, pl. 32; Man. Dipt. Brun. (1964) 79, f. 10: ibid.
Suppl. (1968) 37.

Young twig, raceme, leaf bud, stipule, petiole and
nervation beneath + persistently shortly evenly rich
vinous pubescent, tending to rub off leaf undersur-
face. Twig to 1.5 mm @ apically, slender, much
branched, becoming pale grey and brown mottled,
glabrous, frequently thinly cracked and flaked. Bud to
1.5 mm long and g, ovoid, obtuse. Stipule to 5 by 2.5
mm, oblong, subacute, caducous. Leaves 6-15 by
1.54.5 cm, elliptic to lanceolate; base narrow, obtuse
to cuneate; acumen to 1.2 cm long, Narrow; nerves
12-20 pairs, slender, + prominent beneath, curved,
close set, at 50°—70° to the midrib, with short slender
secondary nerves; midrib slender, prominent, terete
below, flat above; petiole 6-13 mm long, slender.
Panicle to 7 cm long, terminal or axillary, terete, singly
or doubly branched; flower bud to 6 mm long, small;
calyx sparsely vinous cinereous; appendage to connec-
tive short, obtuse; flowers otherwise typical. Fruit
calyx densely shortly vinous puberulent on both
surfaces; lobes to 9 by 8 mm, equal, obovate, +
abruptly revolute, thickened, reflexed at base but
curving outwards and + rotate apically; nut to 8 mm
@. globose, densely vinous puberulent; style remnant
short.

Distr. Malesia: Borneo (N.E. of Rejang valley to
E. Sabah and Tarakan).

Ecol. Fertile clay-rich soils in Mixed Dipterocarp
forest below 300 m.

Vern. Resak tangkai unggu.

19. Vatica scortechinii (KING) BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 122, t. 3 f. 9, p.p.; RiDL. Fl. Mal. Pen. 1
(1922) 244; Crai, FI. Siam. Enum. | (1925) 141, p.p.:
SLOOT. Bull. Jard. Bot. Btzg III, 9 (1927) 73, 108. f. le,
p.p.; Foxw. Mal. For. Rec. 10 (1932) 267, p.p.; BURK.
Dict. (1935) 2224; Sym. J. Mal. Br. R. As. Soc. 19, 2
(1941) 155; Mal. For. Rec. 16 (1943) 227, f. 107;
MEWER & Woop, Sabah For. Rec. 5 (1964) 319. — Re-
tinodendron scortechinii K1NG, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 128; BRUHL & KING, Ann. R. Bot. Gard.
Calc. 5, 2 (1896) 157, t. 190.

Medium-sized tree. Young parts ferruginous ciner-
eous, caducous except on twig apices and panicles.
Twig 3-6 mm @ apically, straight, terete or ribbed
especially along the leaf traces, with conspicuous
slightly downpointing stipule scars. Buds to 5 by 3
mm, ovoid-lanceolate, acute, hidden within the to 35
by 12 mm lanceolate subpersistent stipules. Leaves
13-40 by 5-16 cm, oblong-lanceolate or oblanceolate,
thinly coriaceous; base obtuse or occasionally cor-
date; margin + narrowly revolute: apex subacu-
minate to acute; nerves 14-25 pairs, arched, slender
but prominent beneath, distinctly elevated above as
also the midrib, with many short indistinct secondary
nerves; tertiary nerves reticulate, distinctly elevated
on both surfaces; petiole 13—25(-40) mm long, 3—5 mm
@, stout, characteristically puckered on drying. Pani-
cle to 9cm long, axillary to ramiflorous; flower buds to

358

14 by 5 mm, fusiform; anthers oblong, tapering,
relatively long; appendages very short; style some-
what longer than ovary, columnar, capitate; stigma
conical, prominent; flowers otherwise typical. Fruit
pedicel to 5 by | mm, slender, long; calyx lobes to 7 by
4mm, lanceolate, subacute, reflexed at base, upcurved
distally; nut to 15 mm @, subglobose, with 3 indistinct
loculicidal grooves.

Distr. Malesia: Malaya (Perak, Selangor, Pahang,
Trengganu).

Ecol. Local, on undulating land near coast and to
1800 m on inland ridges.

Vern. Réesak langgong.

Note. Some sterile collections from the east coast
closely resemble V. sarawakensis, so far not confirmed
from Malaya.

20. Vatica globosa ASHTON, Gard. Bull. Sing. 22
(1967) 269, pl. 12; Man. Dipt. Brun. Suppl. (1968) 32,
iis Oe

Twig, leaf bud and petiole persistently rufous-
brown sericeous. Twig c. 2 mm @ apically, terete,
smooth, grey; stipule scar short, horizontal, obscure.
Bud c. 2 by 2 mm, ovoid, acute. Stipule unknown.
Leaves 7-18 by 3-6.5 cm, obovate, somewhat charta-
ceous; base narrowly cuneate; acumen to 2 cm long,
slender, caudate; nerves 12-16 pairs, slender but
prominent beneath with the lamina concave between
them, distinctly elevated above, at 35°—60°, with many
short secondary nerves; tertiary nerves reticulate;
midrib prominent on both surfaces, more so beneath
than above; petiole 7-15 mm long, terete with a
distinct furrow above, drying rugulose. Panicle to 3
cm, terminal or axillary, frequently several in an axil,
congested, ribbed or terete, persistently rufous seri-
ceous; irregularly + singly branched; bracteoles fuga-
ceous. Flower bud to 5 by 2 mm, lanceolate. Calyx
densely vinous pubescent outside, glabrous within;
corolla sparsely so, cream on opening; appendage to
connective conical, c. + length of anther; flowers
otherwise typical. Fruit entirely persistently rufous
sericeous. Calyx lobes to 4 by 3 mm, equal, oblong,
obtuse, reflexed. Nut to 20 mm @, globose, obtuse,
3-sutured.

Distr. Malesia: Borneo (Mukah to Niah valleys,
Central Sarawak), W. Borneo (Ulu Kapuas).

Ecol. Very local, Mixed Dipterocarp forest on low
hills.

21. Vatica lobata Foxw. Mal. For. Rec. 10 (1932) 276,
pl. 23; Burk. Dict. (1935) 2224; Sym. Mal. For. Rec.
16 (1943) 221, f. 107.

Small tree. Young parts, calyx and ovary +
caducous buff cinereous; leaf buds, panicles and petals
outside persistently so, otherwise glabrous. Twig c. 2
mm @, slender, pale brown, sometimes chartaceous
flaky, terete. Buds small, ovoid; stipules fugaceous.
Leaves 7-24 by 2-9.5 cm, elliptic-lanceolate, thinly
coriaceous; base + narrowly cuneate, subdecurrent;
acumen to 15 mm long, slender; nerves c. 5(—8) pairs,
ascending, slender but distinctly elevated beneath, less

FLORA MALESIANA

[ser. I, vol. 9?

so above, as also the midrib; tertiary nerves laxly
reticulate, hardly elevated on either surface; petiole
8-12 mm long, c. mm g, relatively short. Panicle to 5
cm long, terminal or axillary, shortly branched.
Flower buds to 6 by 2 mm, fusiform; appendages very
short, deltoid; style stoutly columnar, somewhat
longer than ovary, expanding at base and at the
prominent conical style; flowers otherwise typical.
Fruit pedicel to 2 by 1 mm; calyx lobes to 10 by 8 mm,
equal, elliptic, incrassate, smooth, lustrous, adnate to
the 17 by 15 cm ovoid smooth apiculate nut.

Distr. Malesia: Malaya (E. coast: Trengganu to
Johore).

Ecol. Local, near streams.

Vern. Resak paya, r. laru.

22. Vatica hullettii (RIDL.) ASHTON, Gard. Bull. Sing.
31 (1978) 21. — Capura hullettii Ripv. J. Str. Br. R.
As. Soc. 54 (1910) 36. — Otophora hullettii RIDL. FI.
Mal. Pen. 1 (1922) 494; RADLK. in E. & P. Pfl. R. Heft
98 (1932) 775, cf. LEENH. Blumea 17 (1969) 88. — V.
stipulata RID. J. Str. Br. R. As. Soc. 82 (1920) 172,
nom. illeg.; Fl. Mal. Pen. 1 (1922) 244; SLoor. Bull.
Jard. Bot. Btzg III, 9 (1927) 76; Foxw. Mal. For. Rec.
10 (1932) 253; SyM. Mal. For. Rec. 16 (1943) 229.
Small tree. Ovary, calyx and panicle caducous pale
brown puberulent, petioles persistently so. Twigs c. 2
mm @ apically, smooth, with prominent ribs decur-
rent from leaf traces; stipule scars obscure, horizontal.
Buds hidden between to 33 by 22 mm large ovate to
elliptic subacute coriaceous 3—5-nerved persistent
stipules. Leaves 9-30 by 2-9 cm, narrowly elliptic,
coriaceous, frequently somewhat bullate; base
cuneate; acumen to 2 cm long; nerves 8-11 pairs,
slender but distinctly raised beneath, elevated above as
also the midrib, arched, ascending, with short second-
ary nerves; tertiary nerves reticulate, slightly elevated
on both surfaces; petiole to 3 cm long, 1-3 mm @,
drying characteristically pale cream brown rugulose.
Panicle to 15 cm long, lax, irregularly branched,
slender. Flower buds to 12 by 2 mm; appendages small,
deltoid; style longer than ovary, slender, greatly
expanding into the prominent conical stigma; flowers
otherwise typical. Fruit pedicel to 4 by 1 mm, promi-
nent; calyx lobes to 20 by 14 mm, ovate, subacute,
loosely enclosing the base of the nut but the sides
becoming completely revolute; nut to 2 cm @, subglo-
bose, verruculose, with 3 indistinct loculicidal sutures.
Distr. Malesia: Malaya (Negri Sembilan, Malacca,
Johore).
Ecol. Rare, lowland dipterocarp forest on hills.
Note. Related to V. albiramis (q.v.).

23. Vatica pallida Dyer, Fl. Br. Ind. 1 (1874) 302;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 121; Rup. FI.
Mal. Pen. 1 (1922) 244; SLoor. Bull. Jard. Bot. Btzg
ITI, 9 (1927) 73, 103, f. la; Foxw. Mal. For. Rec. 10
(1932) 266; BurK. Dict. (1935) 2224; Sym. Mal. For.
Rec. 16 (1943) 225, f. 107. — Retinodendron pallidum
KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 128; BurK.
J. Str. Br. R. As. Soc. 81 (1920) 51, 62, 63, fig.

1982]

Small, low-branched tree. Young parts pale brown
caducous puberulent, persisting on into the young
fruit on calyx and ovary. Twig c. | mm @ apically,
slender, straight, terete, smooth, pale grey-brown.
Buds minute. Leaves 6-16 by 2-5.6 cm, narrowly
elliptic, thinly coriaceous; base narrowly cuneate;
acumen to 2cm long, slender, subcaudate; nerves 9-11
pairs, arched, ascending, slender, hardly elevated on
either surface as also the reticulate tertiary nerves and
somewhat more prominent midrib; petiole 5-8 mm
long, short, slender. Panicle to 4 cm long, terminal or
subterminal axillary, few branched. Fruit pedicel to 4
by | mm, slender, long; calyx lobes to 8 by 3 mm,
lanceolate, loosely clasping the c. 1 cm @ subglobose
smooth mucronate nut.

Distr. Malesia: Malaya (Penang).

Ecol. Common in the forests on the slopes of
Penang hill, to 350 m.

Vern. Resak kéchil.

Note. Closely allied to V. philastreana PIERRE of
southern Indo-China and also to V. /anceaefolia BL.
of Assam.

24. Vatica flavida Foxw. Mal. For. Rec. 10 (1932)
272, pl. 22; BurK. Dict. (1935) 2223; Sym. Mal. For.
Rec. 16 (1943) 220, f. 107.

Medium-sized tree. Twigs, petioles, leaf buds, mid-
rib above, panicles and nut densely persistently tawny
scabrid pubescent, leaf nervation beneath sparsely so,

DIPTEROCARPACEAE (Ashton)

359

calyx outside caducously so; outsides of petals densely
evenly cream puberulent. Twigs 2-3 mm @ apically,
prominently ribbed at first, becoming pale brown,
terete. Buds to 6 by 4 mm, ovoid-lanceolate, acute.
Leaves 6-11 by 2.5—S cm, elliptic to narrowly obovate,
coriaceous; base broadly cuneate to obtuse; acumen
to 1 cm long, slender, prominent; nerves 9-11 pairs,
prominent beneath, distinctly depressed above with
the leaf surface bullate between, arching within the
margin; tertiary nerves subreticulate, distinctly ele-
vated beneath, hardly so above; midrib stoutly promi-
nent beneath, shallowly depressed though evident
above; petiole 11-15 mm long. Panicles to 3.5 cm long,
short, axillary, twice branched from the bases, many-
flowered and appearing congested about the bases of
the leaves. Flower buds to 8 by 3 mm, fusiform; sepals
unequal, ovate-lanceolate; corolla pale yellow;
stamens 15; anthers subglobose, appendages promi-
nent, broadly deltoid; style columnar, slightly longer
than ovary; stigma conical; flowers otherwise typical.
Fruit pedicel to 4 mm long, prominent but hidden in
the reflexed sepals; calyx lobes to 8 by 6 mm, equal,
broadly lanceolate, becoming reflexed with the apices
recurved upwards; nut to 10 by 10 mm, broadly ovoid
to subglobose, with 3 distinct loculicidal furrows.

Distr. Malesia: Malaya (S. Perak).

Ecol. Rare, in forest on swampy land.

Vern. Resak padi.

2. Section Sunaptea

(GrirF.) BuRcK, Ann. Jard. Bot. Btzg 6 (1887) 223; AsHTon, Gard. Bull. Sing. 20
(1963) 250.— Sunaptea GriFF., corr. ‘Synaptea KuRz.— Pteranthera
BL. — Vatica sect. Euvatica B. & H. Gen. Pl. 1 (1862) 192; Dyer, FI. Br. Ind. 1
(1874) 301; Burck, Ann. Jard. Bot. Btzg 6 (1887) 224; KING, J. R. As. Soc. Beng.
Sc. 62, 2 (1893) 102. — Vatica subg. Synaptea (GRIFF.) BRANDIS, J. Linn. Soc. Bot.

31 (1895) 128.

Fruit calyx lobes unequal, chartaceous, with 2 lobes longer than the other three,

not becoming reflexed.

25. Vatica heteroptera Sym. J. Mal. Br. R. As. Soc. 19
(1941) 147, pl. 5B; Mal. For. Rec. 16 (1943) 221, f. 107.

Medium-sized tree. Young parts pale brown cadu-
cous cinereous, panicle and nut + persistently so.
Twig c. 2 mm @ apically, slender, rather straight,
terete, smooth, pale brown. Bud to 2 by 2 mm, small,
ovoid; stipule fugaceous. Leaf 9-20 by 3-8 cm,
narrowly oblong-elliptic, thinly coriaceous; base nar-
rowly obtuse to broadly cuneate; acumen to 2 cm
long; nerves 11-14 pairs, slender but prominent
beneath, evident above, arched, with or without short
secondary nerves; midrib slender but prominent
beneath, sharply elevated above; tertiary nerves den-
sely subreticulate; petiole 11-18 mm long, c. 2mm @,
relatively short. Panicles to 12 cm long, terminal or
axillary, rather straight; flowers unknown. Fruit pedi-

cel to 3 by 2mm; calyx lobes to 2.5 by 1 cm, lanceolate,
subacute, unequal but patent; nut to 15 mm @, to 12
mm long, depressed-globose; shortly mucronate.
Distr. Malesia: Malaya (Perak, Pahang).
Ecol. Locally frequent in Upper Dipterocarp forest
on ridges, 1000-1300 m in the Malayan Main Range.
Vern. Resak gunong.

26. Vatica maritima SLooT. Bull. Bot. Gard. Btzg III,
17 (1942) 245, f. 32; ASHTON, Man. Dipt. Brun. (1964)
72, f. 10; ibid. Suppl. (1968) 33; MEER & Woop,
Sabah For. Rec. 5 (1964) 310.

Young twig, panicle, leaf bud and petiole densely
shortly evenly + persistently cream pubescent. Twig
to 8 mm @ apically, ribbed or somewhat compressed,
becoming terete, glabrous, smooth. Bud to 4 by 3 mm,

360

ovoid, subacute. Stipule unknown. Leaves 8-16 by
3-8 cm, broadly or narrowly ovate, coriaceous; base
broadly cuneate to subcordate, acumen to | cm long,
short, broad; nerves 7-10 pairs, raised on both
surfaces, more prominently so beneath, with short,
slender secondary nerves; tertiary nerves reticulate;
petiole 2-2.7 cm long, not geniculate. Panicle to 11 cm
long, axillary, rarely terminal, lax, angular; branchlets
to 1.5 cm long, short, with up to 12 close flowers;
bracteoles to 3 by 2 mm, ovate, subacute, sparsely
pubescent. Flower bud to 1.4 cm long; calyx densely
pale brown, tomentose, flowers otherwise typical.
Fruit pedicel to 4 mm long; ca/yx puberulent at base,
otherwise glabrous; 2 longer lobes to 5 by | cm,
oblong-spatulate, obtuse, base to 3 mm broad, fusing
with shorter lobes forming an up to 5 mm @ shallow
cup adnate to the base of the fruit; 3 shorter lobes to 13
by 4 mm, lanceolate, acute. Nut to 7 by 7 mm,
subglobose, sparsely cream pubescent; style remnant
to 1 cm long, slender.

Distr. Malesia: Borneo (N.E. Kalimantan; Brunei;
W. Sabah), also S. Philippines (Palawan).

Ecol. Very local, dry coastal hills in non-seasonal
lowland forest.

Vern. Resak laut, r.l. timor (Borneo).

27. Vatica teysmanniana BuRCK, Ann. Jard. Bot.
Btzg 6 (1887) 230; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 133; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 314; ibid.
ed. 2 (1927) 1131; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 90, f. 3. — Sunaptea teysmannii HEIM, Rech.
Dipt. (1892) 100.

Medium-sized to large tree. Twigs, leaf buds and
petioles densely persistently pale fawn scabrid pubes-
cent, midrib above and panicles + caducously so;
parts of petals exposed in bud and ovary densely
evenly persistent-puberulent, calyx and pedicel cadu-
cously so. Twig c. 4 by 2mm @ apically, compressed at
first, becoming terete, + ribbed, blackish, with long
+ prominent pale horizontal somewhat descending
stipule scars. Buds to 6 by 5 mm, ovoid, acute. Leaves
9.5—34 by 3-11 cm, variable in size, narrowly elliptic to
lanceolate, lustrous, thickly coriaceous; margin nar-
rowly subrevolute; base broadly cuneate to obtuse;
apex shortly retuse, obtuse or to | cm long broadly
acuminate; nerves 13-17 pairs, arched, slender but
prominent beneath, less (but distinctly so) above as
also the midrib, sometimes with a few short + obscure
secondary nerves; tertiary nerves densely reticulate,
distinctly elevated on both surfaces; petiole 13-25 mm
long, c. 3 mm @g, stout, not geniculate. Panicles to 15
cm long, terminal or axillary, lax, spreading, rather
regularly singly or doubly branched; branchlets to 7
cm long, bearing to 7 flowers. Flower buds to 12 by 3
mm, large. Fruit pedicel to 6 mm long, slender,
prominent; calyx lobes unequal; 2 longer lobes to 11
by 1.8 cm, lorate-spatulate, obtuse, c. 4 mm wide at
base; 3 shorter lobes to 18 by 6 mm, lanceolate, acute;
nut to 6mm @, globose.

Distr. Malesia: Sumatra (E. Coast: Bengkalis),
Banka.

FLORA MALESIANA

[ser. I, vol. 97

Ecol. Mixed swamp forests.

Vern. Reésak ayér, r. paya, r. badouw, r. siantén.

Note. The vegetative parts evoke V. coriacea of
Borneo, but the fruit calyx, inflorescence and bracts at
once distinguish the species.

28. Vatica cinerea KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 104; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 131;
BRUHL & KING, Ann. R. Bot. Gard. Calc. 5, 2 (1896)
150, t. 183A; Rip. J. Str. Br. R. As. Soc. 61 (1912) 50;
Fl. Mal. Pen. 1 (1922) 243; HEYNE, Nutt. Pl. ed. 1, 3
(1917) 312; Crap, Fl. Siam. Enum. 1 (1925) 140;
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 80; Foxw.
Mal. For. Rec. 10 (1932) 256; BurK. Dict. (1935)
2223; Sym. J. Mal. Br. R. As. Soc. 19 (1941) 157; Mal.
For. Rec. 16 (1943) 218, f. 107. — V. lankaviensis
Rip. J. Str. Br. R. As. Soc. 54 (1909) 27; Crais, FI.
Siam. Enum. | (1925) 141. — Synaptea cinerea RIDL.
Fl. Mal. Pen. 1 (1922) 243. — Synaptea lancaviensis
Rpt. Fl. Mal. Pen. 1 (1922) 241.

Small or medium-sized tree. Twig, petiole and
panicle sparsely pale pink-brown puberulent; outside
of perianth and ovary densely so, caducous except on
petals and ovary. Twig c. 1 mm @ apically, slender,
much branched, pale brown, terete, smooth. Bud
small, ovoid; stipules fugaceous. Leaf 5.5—14 by 1.5—5
cm, elliptic-lanceolate, thinly coriaceous; base
cuneate, apex acute to subacuminate; nerves 9-11
pairs, arched, ascending, slender and hardly more
elevated beneath than above as also the midrib and
densely reticulate tertiary nerves; petiole 5-11 mm
long, slender. Panicle to 9 cm long, terminal or
axillary, irregularly branched. Flower bud to 8 by 3
mm, lanceolate; appendages very short; style colum-
nar, slender, somewhat longer than ovary, capitate;
stigma large, conical; flowers otherwise typical. Fruit
pedicel to 3 by | mm, slender, prominent; calyx united
into an up to 7 mm @ cup at base adnate to the basal 5
of the ovary; 2 longer lobes to 7 by 1.8 cm, spatulate,
obtuse, c. 3mm wide at base; 3 shorter lobes to 15 by 5
mm, lanceolate, acute; nut to 7 mm @, subglobose,
with filiform style remnant.

Distr. South Tenasserim, Peninsular Thailand,
and in Malesia: Malaya (Kedah, Perlis, Langkawi).

Ecol. Dry ridges, headlands, limestone and other
rocky places, in Schima-bamboo forests, to 600 m.

Vern. Reésak, r. laut.

29. Vatica odorata (GriFF.) SYM. J. Mal. Br. R. As.
Soc. 19 (1941) 156; Mal. For. Rec. 16 (1943) 224, f.
107; ASHTON, Gard. Bull. Sing. 22 (1967) 263; ibid. 31
(1978) 23.

a. ssp. odorata. — Hopea faginea WALL. Cat. (1828)
963, nomen. — Shorea pinangiana WALL. Cat. (1828)
157, nomen; KuRZ, J. R. As. Soc. Beng. Sc. 43, 2 (1874)
96, nomen pro syn.; KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 105, nomen pro syn. sub V. faginea. — Sunaptea
odorata GriFF. Notul. 4 (1854) 516; Ic. Pl. As. (1854) t.
585A, f. 5; Herm, Rech. Dipt. (1892) 113. — Hopea
grandiflora WALL. [Cat. (1828) 958, nomen] ex A. DC.

1982]

Prod. 16, 2 (1868) 634. — Synaptea grandiflora KURZ,
J. R. As. Soc. Beng. Sc. 39, 2 (1870) 65; PIERRE, For.
Fl. Coch. 4 (1891) t. 240-242. — Anisoptera odorata
Kurz, Flora 30 (1872) 191; J. R. As. Soc. Beng. Sc. 43,
2 (1874) 96; Fl. Burma | (1877) 112. V. grandiflora
Dyer, Fl. Br. Ind. 1 (1874) 301; KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 101; BRANDis, J. Linn. Soc. Bot.
31 (1895) 129: Ind. Trees (1906) 72; Foxw. Mal. For.
Rec. 10 (1932) 260. — V. faginea Dyer, FI. Br. Ind. 1
(1874) 301; Kurz, J. R. As. Soc. Beng. Sc. 43, 2 (1874)
96; KinG, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 105;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 129; Ind. Trees
(1906) 72; BRANDIS & GILG in E. & P. Pfl. Fam. 3, 6
(1895) 270; GukrIn, FI. Gen. I.-C. 1 (1910) 391; Ripv.
Fl. Mal. Pen. 1 (1922) 242, p.p.; GitG in E. & P. Pf.
Fam. ed. 2, 21 (1925) 265; HEYNgE, Nutt. Pl. ed. 2
(1927) 1129; SLoor. Bull. Jard. Bot. Btzg III, 9 (1927)
81; Foxw. Mal. For. Rec. 10 (1932) 260.— V.
astrotricha HANCE, J. Bot. 14 (1876) 241; LANgss. PI.
Util. Colon. Fr. 1 (1886) 299; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 130; BRANDIS & GILG in E. & P. Pf.
Fam. ed. 1, 3, 6 (1895) 270; FiscHER, Kew Bull. (1926)
457; SLooT. Bull. Jard. Bot. Btzg IIT, 9 (1927) 80. — V.
dyeri PIERRE ex LANESS. Pl. Util. Colon. Fr. 1 (1886)
299: KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 106;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 128, t. 3, f.
15-17; GugrIn, Fl. Gen. I.-C. 1 (1910) 387, 392, fig.;
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 82; Foxw.
Mal. For. Rec. 10 (1932) 258; Burk. Dict. (1935)
2223. — Sunaptea astrotricha PIERRE, For. Fl. Coch. 4
(1891) t. 240; Hem, Rech. Dipt. (1892) 118. — Sunap-
tea dyeri PIERRE, For. Fl. Coch. 4 (1891) t. 241; RIDL.
Fl. Mal. Pen. 1 (1922) 240.— Sunaptea faginea
PreRRE, For. Fl. Coch. 4 (1892) 242; Ripv. Fl. Mal.
Pen. 1 (1922) 242. — V. curtisii KiNG, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 105; BRANbIs, J. Linn. Soc. Bot.
31 (1895) 131; BRUHL & KING, Ann. R. Bot. Gard.
Calc. 5, 2 (1896) 151, t. 183B; SLoort. Bull. Jard. Bot.
Btzg III, 9 (1927) 81; Foxw. Mal. For. Rec. 10 (1932)
257; BurRK. Dict. (1935) 223. — V. fleuryana TARDIEU,

Not. Syst. 10 (1942) 137; Fl. Gen. I.-C. Suppl. 1 (1943)
~ 359, f. 39 (9-16). — V. tonkinensis [CHEVALIER, Bull.
Ec. Indochine 20 (1918) 799, nomen] TaRpDiEu, Not.
Syst. 10 (1942) 137; Fl. Gén. I.-C. Suppl. (1943) 357, f.
39 (1-8).

Young twig, raceme, leaf bud, stipule, and petiole
+ densely persistently pale yellowish brown to ful-
vous tomentose. 7wig 1.5 mm @ apically, terete,
glabrous, rugose, frequently finely flaked. Bud to 2 by
1.5mm, ovoid, obtuse. Stipule to 8 by 2 mm, oblong,
obtuse, caducous. Leaves 8-16 by 2.7-5.5 cm, nar-
rowly elliptic to ovate, thinly coriaceous, base obtuse
or cuneate; acumen to 8 mm long; nerves 11-15,
prominent beneath; midrib prominent beneath,
applanate above or slightly depressed; petiole 8-12
mm long. Panicle to 7 cm long; terminal or axillary
singly branched. Flower bud to 8 mm long; calyx
densely shortly pale grey-brown tomentose; appen-
dage to connective short, obtuse; flowers otherwise
typical. Fruit pedicel c. 3 mm long, slender. Fruit calyx
at first powdery tomentose, glabrescent, united in an

DIPTEROCARPACEAE (Ashton)

361

up to Smm deep, to 8 mm @, cup at base and fused to
nut; 2 longer lobes to 5.5 by 1.5 cm, thin, spatulate,
obtuse, base to 2.5 mm broad, 3 shorter lobes to 14 by
4 mm, hastate, acute. Nut to 7 mm long and g,
globose, shortly densely yellow-brown to fulvous
tomentose, the basal half adnate with the calyx cup;
style remnant to 2 mm long, linear.

Distr. Tenasserim, Thailand, Indochina, S. China
(Kwangsi), and in Malesia: Malaya (Negri Sembilan
& Pahang northwards), Borneo (Tawau, S.E. Borneo).

Ecol. Scattered on dry ridges in coastal forests in +
seasonal areas.

Vern. Resak runting késat.

b. ssp. mindanensis (FOxw.) ASHTON, Gard. Bull. Sing.
22 (1967) 263; Man. Dipt. Brun. (1964) 75 (as V.
odorata; f. 10); ibid. Suppl. (1968) 34 (ditto); Gard.
Bull. Sing. 31 (1978) 24. — V. mindanensis Foxw. in
Elmer, Leafl. Philip. Bot. 6 (1913) 1957; Philip. J. Sc.
13 (1918) Bot. 196; ibid. 67 (1938) 327; MerRR. En.
Philip. 3 (1923) 102; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 71; MEWER & Woop, Sabah For. Rec. 5 (1964)
313. — V. sorsogonensis Foxw. Philip J. Sc. 13 (1918)
Bot. 196. — V. aerea SLOOT. Bull. Bot. Gard. Btzg III,
17 (1941) 133, f. 19.

Differing from ssp. odorata in the 15-20 mm long
geniculate petiole and leaf drying greyish rather than
yellow-brown.

Distr. Malesia: Borneo (Kapuas hinterland; Sar-
awak to Crocker range; Tawau); Philippines (Luzon,
Leyte, Mindanao).

Ecol. Common on ridges between 1000-1200 m, in
non-seasonal areas.

Vern. Resak biabas (Borneo).

30. Vatica compressa ASHTON, Gard. Bull. Sing. 22
(1967) 267, pl. 10; Man. Dipt. Brun. Suppl. (1968) 30,
feos

Young parts (except stipule within) evenly pale
pink-brown pubescent, caducous except on leaf bud
and stipule outside. Twig c. 3 by | mm @, compressed,
smooth; stipule scars c. 2 mm long at first, pale,
horizontal, prominent. Bud c. 3 by 3 mm, ovoid, acute,
compressed. Leaves 11-19 by 5-8.5 cm, ovate-elliptic,
coriaceous; base obtuse; acumen to | cm long; nerves
10-15 pairs, slender, only slightly raised beneath, less
so above, arched, at c. 50°, with short indistinct
secondary nerves; tertiary nerves scalariform; midrib
prominent beneath, applanate or raised above; petiole
15-23 mm long. Panicle to 7 cm long, somewhat
compressed, sparsely pale pink-brown pubescent,
irregularly singly or doubly branched. Flower bud to 9
by 4 mm, lanceolate; calyx densely pale fulvous
pubescent; corolla densely pubescent on parts
exposed in bud; flowers otherwise typical. Fruit
pedicel to 8 mm long, slender, pubescent. Calyx
puberulent to glabrescent; 2 longer lobes to 6.5 by 1
cm, lorate, + obtuse, c. 3mm broad at base; 3 shorter
lobes to 20 by 6 mm, deltoid, acute, revolute. Nur to 5
mm @, shortly ovoid, densely shortly buff pubescent;
style remnant to 3 mm long glabrous.

362

FLORA MALESIANA

[ser. I, vol. 9?

Distr. Malesia: Borneo (Sarawak W. of Lupar
valley).

Ecol]. Local in Heath forest, often with impeded
drainage.

31. Vatica congesta ASHTON, Gard. Bull. Sing. 22
(1967) 268, pl. 11; Man. Dipt. Brun. Suppl. (1968) 31,
fess

Young twigs, buds, stipules and petioles densely
shortly pale ocherous-brown scabrid pubescent, leaf
beneath and midrib above towards base sparsely so.
Twig c. 3 mm g towards apex, stout, at first angular,
rugose, becoming striated and chartaceous flaky. Bud
to 4 by 3 mm, broadly ovoid-conical, acute. Stipule to
8 by 3 mm, narrowly deltoid, acute. Leaves 8-22 by
3.5-8 cm, oblong-elliptic to obovate, thickly coria-
ceous, subacuminate, obtuse or retuse; margin revo-
lute; base obtuse; nerves 7—11 pairs, slightly raised
above, stout and prominent beneath, at 40°—65°;
tertiary nerves prominent beneath, less so above,
reticulate; midrib hardly raised above, stout, promi-
nent, striated, beneath; petiole 1.2—2.5 cm long, stout,
rugulose. Panicle to 3 cm long, angular, densely
shortly ocherous-brown scabrid tomentose, axillary
to ramiflorous, fascicled, short, and congested; singly
branched, branchlets to 1.8 cm long. Flowers disti-
chous; bud to 8 by 3 mm: calyx lobes densely shortly
ocherous-brown tomentose, deltoid, acute, somewhat
spreading in bud; petals lorate, obtuse, densely pubes-
cent on both surfaces; appendage to connective short,
bifid: style short, sericeous; flowers otherwise typical.
Fruit pedicel sparsely shortly ocherous-brown scabrid
tomentose, fruit otherwise glabrous. Pedicel to 4mm
long, slender. 2 longer calyx lobes to 12 by 2.2 cm,
lorate, obtuse, prominently recurved. Nut to 12 by 8
mm, ellipsoid, glabrous; style remnant to 6 mm long,
slender.

Distr. Malesia: Borneo (W. and Central Sarawak;
W. Kalimantan: Lower Kapuas).

Ecol. Rare, low hills, Mixed Dipterocarp forest.

32. Vatica coriacea ASHTON, Gard. Bull. Sing. 19
(1962) 314, pl. 30; Man. Dipt. Brun. (1964) 68, f. 10:
ibid. Suppl. (1968) 31.

Young twig, panicle, leaf bud, stipule and petiole
densely shortly ocherous pubescent, caducous on all
but bud. Twig to 5 mm g apically, stout, ribbed,
glabrous, smooth. Bud to 7 by 5mm, ovoid, subacute.
Stipule to 13 by 5S mm, hastate, subacute. Leaves
6.5-15 by 2.2-6 cm, thickly coriaceous, obovate, +
strongly cupped towards margin; base narrow, obtuse
or cuneate; apex obtuse to retuse; nerves 10—11 pairs,
slightly raised on both surfaces, indistinct, curved, at
50°—60°, with short slender secondary nerves; midrib
stout, prominent, terete beneath, slightly raised
above; petiole 1-1.5 cm long. Panicle to 20 cm long,
axillary or terminal, singly branched; limits of inflor-
escence ill-defined, often bearing leaf-like bracts.
Flowers to 1.8 cm long; calyx densely shortly pale
grey-brown pubescent; flowers typical. Fruit pedicel to
7 mm long, slender; calyx shortly persistently pubes-

cent outside, glabrescent within, free to base; 2 longer
lobes to 7 by 2.3 cm, oblong, obtuse, to 3.5 mm broad
at base; 3 shorter lobes to 2 by 6 mm, hastate, acute.
Nut to 8 mm g, and globose, puberulent or glabres-
cent, rugulose, frequently crowned with an up to 2mm
long linear style remnant.

Distr. Malesia: Borneo (Sarawak, Brunei).

Ecol. Locally frequent, Heath forest on podsols on
coastal raised beaches, and on rentzinas over lime-
stone in the west.

Vern. Résak daun tébal.

33. Vatica javanica SLOoT. Bull. Jard. Bot. Btzg III,
16 (1940) 451, f. 9; BACKER & BAKH. f. FI. Java 1
(1963) 332; ASHTON, Gard. Bull. Sing. 31 (1978) 22.

a. ssp. javanica.

Medium-sized to large tree. Twigs, petioles, pani-
cles, pedicels and base of calyx outside densely
persistently tawny scabrid pubescent; leaf undersur-
face frequently more sparsely so; fruit calyx cadu-
cously so. Twig c. 3 mm @ apically, stout, +
compressed at first and ribbed, becoming terete,
smooth. Leaf bud to 8 by 5 mm, ovoid, acute. Leaves
13-24 by 6-10 cm, elliptic-oblong or obovate,
applanate, thinly coriaceous; base obtuse or subcor-
date: acumen to 1.5 cm long, + prominent; nerves
22-25 pairs, + straight, ascending, + distinctly
branching and arching within the margin forming a
looped intramarginal nerve, stoutly prominent
beneath, elevated above as also the midrib, without
distinct secondary nerves; tertiary nerves subscalari-
form, prominent beneath, evident above; petiole 2-3
cm long, c. 3 mm g, stout. Panicles to 12 cm long,
terminal or axillary, rather straight, irregularly dou-
bly branched. Flower buds to 11 by 4 mm, fusiform;
sepals narrowly lanceolate, unequal; anthers subglo-
bose: appendages very short; style c. 15 times as long
as ovary, expanding distally; stigma conical; flowers
otherwise typical. Fruit pedicel to 8 mm long, slender;
calyx lobes unequal, free to base; 2 longer lobes to 7.5
by 1.7 cm, spatulate, obtuse; 3 shorter lobes to 30 by 7
mm, lanceolate, slender; nut globose, unknown at
maturity.

Distr. Malesia: W. Java (Priangan Regencies, once
collected).

Ecol. Primary forest, 950 m.

b. ssp. scaphifolia (KOSTERM.) ASHTON, Gard. Bull.
Sing. 31 (1978) 22. — V. scaphifolia KOSTERM. New &
Crit. Mal. Pl. 3 (1955) 2, f. 1.

Leaves + prominently boat-shaped with the lower
surface concave, + distinctly bullate between the
nerves, the nerves and sometimes tertiary nerves
consequently + channelled above.

Distr. Malesia: S.E. Borneo (Samarinda, Balikpa-
pan).

Ecol. Locally frequent in Lowland Dipterocarp
forest on well drained undulating land.

34. Vatica brunigii ASHTON, Gard. Bull. Sing. 22

1982]

(1967) 267, pl. 9; Man. Dipt. Brun. Suppl. (1968) 30, f.
Si;

Young twigs, buds, stipules, petioles, and leaf
beneath persistently shortly yellowish buff scabrid
pubescent; leaf above fugaceous flocculent tomentose.
Twig c. 2mm @ towards apex, slender, terete. Bud to 3
by 2 mm, ovoid-conical, obtuse. Stipule to 5 by 2mm,
lorate, obtuse; acumen to | cm long; nerves 9-12 pairs,
unraised above, slender but prominent beneath, at
65°-80°; midrib hardly raised above, prominent
beneath; tertiary nerves scalariform, obscure above,
slightly raised beneath; petiole 8-15 mm long. Panicle
to 12 cm long, terete or angular, densely shortly
persistently ocherous scabrid tomentose, terminal or
axillary but confined to young twigs; singly (if axil-
lary) or doubly (if terminal) branched, branchlets to 6
cm long; bracteoles to 3 by | mm, elliptic, sparsely
pubescent. Flowers distichous; flower buds to 8 by 2
mm, ellipsoid; calyx densely shortly ocherous-grey
pubescent; peta/s narrowly oblong, obtuse, sparsely
pubescent outside, glabrescent within; appendage to
connective short, erect, exceeding apex of anther; style
columnar, longer than ovary; flowers otherwise typical.
Fruit pedicel and calyx sparsely buff pubescent, nut
densely scabridly so. Pedice/ to 3 mm long, slender; 2
longer calyx lobes to 6.5 by 1.5 cm, lorate to spatulate,
obtuse; 3 shorter lobes to 15 by 6 mm, ovate, acute,
slightly recurved. Nut to 9 by 7 mm, ovoid, subacute.

Distr. Malesia: E. Sumatra (P. Musala, Lower
Langkat, Riouw), Borneo (Pontianak, Sarawak,
Brunel).

Ecol. Heath forest on shallow podsols, terraces,
plateaux and cuestas, coastal areas and dry ridges to
700 m.

35. Vatica pachyphylla Merr. Philip. J. Sc. 13 (1918)
Bot. 311; En. Philip. 3 (1923) 102; Foxw. Philip. J. Sc.
67 (1938) 325.

Twigs, petioles, panicle, parts of petals exposed in
bud and ovary densely persistently pale tawny some-
what scurfy pubescent, leaf undersurface and midrib
' above + caducously so. Twig c. 3 mm @g, stout,
rugose, with prominent raised petiole scars. Leaves
7-15 by 2.8-7.5 cm, elliptic, coriaceous, dull beneath,
lustrous above; base + broadly cuneate, acumen to |
cm long, slender, prominent, nerves 10-11 pairs,
ascending at c. 45°, rather straight, prominent
beneath, somewhat less so above, as also the midrib;
tertiary nerves densely reticulate, distinctly + equally
elevated on both surfaces; petiole 17-23 mm long.
Panicle to 7 cm long, axillary, stout, irregularly
doubly branched. Flower bud to 9 by 3 mm; sepals
narrowly deltoid, unequal; anthers narrowly oblong,
tapering; appendages slender but prominent; style
slightly longer than ovary, expanding distally, capi-
tate; stigma conical; flower otherwise typical. Fruit
pedicel to 4mm long, slender; 2 longer calyx lobes to 7
by 1.3 cm, broadly spatulate, obtuse, c. 5mm broad at
base; 3 shorter lobes to 20 by 8 mm, narrowly
elliptic-lanceolate, acute; nut to 8 mm @, broadly
ovoid or subglobose, mucronate.

DIPTEROCARPACEAE (Ashton)

363

Distr. Malesia: Philippines (E. Luzon, Polillo).

Ecol. Scattered in non-seasonal evergreen diptero-
carp forests below 80 m.

Vern. Hagakhac na itim, dadiangao, tamahuan
(Camarines), manapo (Tayabas), yacal (Polillo), banic
(Cagayan).

36. Vatica obovata SLooT. [Med. Proefst. Boschw. 2
(1925) 132, nomen] Bull. Jard. Bot. Btzg III, 9 (1927)
89) f. 2.

Medium-sized tree. Twigs, petiole, leaf buds, sti-
pules, nut, fruit pedicel and midrib and nerves beneath
densely + persistently evenly pale brown puberulent,
leaf undersurface and fruit calyx sparsely so. Twig c. 2
mm @ apically, ribbed at first, becoming terete. Leaf
bud to 5 by 4mm, ovoid, acute; stipule to 8 by 3 mm,
lanceolate, fugaceous. Leaves 8-15 by 9.5—7 cm,
elliptic to obovate, thinly coriaceous; margin nar-
rowly subrevolute; base narrowly obtuse; acumen to
1.5 cm long, downcurved and twisting over on
pressing; nerves 11-14 pairs, arched, slender but
prominent beneath, evident and slightly elevated
above as also the midrib; tertiary nerves subscalari-
form, very slender but slightly elevated on both
surfaces; petiole 15-20 mm long, slender, prominently
geniculate. Flowers and inflorescences unknown.
Fruit pedicel to 5 mm long; calyx lobes unequal, 2
longer lobes to 6.5 by 1.5 cm, spatulate, subacute, c. 5
mm broad at base; 3 shorter lobes to 1.5 by 6 mm,
lanceolate-acuminate; nut to 6 mm @, subglobose.

Distr. Malesia: Sumatra (Palembang).

Vern. Resak lingga.

37. Vatica borneensis BurcK, Ann. Jard. Bot. Btzg 6
(1887) 230; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 133:
Merk. En. Born. (1921) 408; SLoor. Bull. Jard. Bot.
Btzg III, 9 (1927) 87; BROWNE, For. Trees Sarawak &
Brunei (1955) 100; AsHTON, Gard. Bull. Sing. 20
(1963) 252; Man. Dipt. Brun. (1964) 68, f. 10; ibid.
Suppl. (1968) 30. — V. urbanii Hem, Bull. Mens. Soc.
Linn. Paris 2 (1891) 956; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 133; MERR. En Born. (1921) 409; SLoorT.
Bull. Jard. Bot. Btzg III, 9 (1927) 77.— Sunaptea
borneensis HEIM, Rech. Dipt. (1892) 116. — Sunaptea
urbanii HEM, Rech. Dipt. (1892) 115. — V. beccarii
Dyer ex BRANDIS, J. Linn. Soc. Bot. 31 (1895) 133,
nomen in syn.

Young twig, panicle, leaf bud, midrib beneath and
petiole + persistently densely evenly pink-brown
puberulent. Twig to 1.5mm g apically, slender, ribbed
or terete, becoming smooth or rugulose. Bud to 2 by
1.5 mm, ovoid, acute. Stipule unknown. Leaves 6-10
by 2.5-5 cm, elliptic, coriaceous; base cuneate;
acumen to 6 mm long; nerves 7-9 pairs, hardly raised
on either surface, at 30°—35°, strongly curved, with
short secondary nerves; midrib slender, prominent
beneath, + applanate above; petiole 1.5—2.5 cm long,
slender, not swollen distally. Panicle to 5 cm long,
axillary, terete, densely pink-brown pubescent. Flower
bud to 1 cm long; densely pink-brown pubescent;
flowers otherwise typical. Fruit pedicel c. 5 mm long,

364

FLORA MALESIANA

[ser. 1, vokeag4

Fig. 46. Close-up of inflorescence of Vatica banta-
mensis (HASSK.) B. & H. ex Mia. Cult. Hort. Bog. July
1955:

slender; calyx caducous rufous powdery tomentose;
lobes free to base; 2 longer lobes to 5.5 by 1.5 cm,
oblong-spatulate, to 4mm broad at the non-revolute
base; 3 shorter lobes to 18 by 7 mm, unequal, narrowly
ovate, tapering, acute, slightly constricted at base, not
revolute. Nut to 1 cm @, globose, puberulent to
glabrous, crowned by a short acute style remnant.

Distr. Malesia: Borneo (Sarawak, Brunei).

Ecol. Leached sandy soils, coastal hills and inland
ridges to 900 m; pole forest.

Vern. Resak kémudi.

38. Vatica bantamensis (Hassk.) B. & H. ex Mia.
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 85, cf. B. & H.
Gen. Pl. 1 (1862) 192; Burck, Ann. Jard. Bot. Btzg 6
(1887) 231, t. 28; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
132; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6
(1895S) 270; K. & V. Bijdr. 5 (1900) 129; Koorp. Exk.
Fl. Java 2 (1912) 622; Gite in E. & P. Pfl. Fam. ed. 2,
21 (1925) 265; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 87; BacKER & Baku. f. Fl. Java 1 (1963)
332. — Anisoptera bantamensis Hassk. Retzia 1

(1855) 140; Wace. Ann. 4 (1857) 336; Mia. FI. Ind.
Bat. 1; 2, (1859); S01: (DE, Prody 162 a Gis6s)
615. — Synaptea bantamensis Kurz, J. R. As. Soc.
Beng. Sc. 39, 2 (1870) 65; PiERRE, For. Fl. Coch. 4
(1892) t. 258; Heim, Rech. Dipt. (1892) 113. — Fig. 46.

Medium-sized tree, to 30 m. Young parts densely
evenly pale ocherous buff puberulent, persistently so
on ovary, panicle, calyx, and parts of petals exposed in
buds; becoming sparse or glabrous on fruit calyx.
Twig c. 2 mm @ apically, terete, + rugulose. Buds
minute. Leaves (4.5—)7.5-18 by (1.8-)3.5-7.5 cm,
elliptic or lanceolate, coriaceous, + lustrous; base
cuneate; acumen to | cm long, slender; nerves 9-11
pairs, slender, arched, ascending, distinctly and
almost equally elevated on both surfaces as also the
short + obscure secondary and densely reticulate
tertiary nerves; midrib prominent beneath, distinctly
but less prominently elevated above; petiole
(10-)14-22 mm long, slender, distinctly geniculate.
Panicle to 7 cm long, terminal or axillary, irregularly
doubly branched. Flower bud to 9 by 3 mm, fusiform;
sepals unequal; anthers narrowly oblong, tapering;
appendage very short, conical; style longer than
ovary, columnar, expanding from base; stigma shortly
conical; flowers otherwise typical. Fruit pedicel to 5
mm long, slender; calyx lobes unequal, united into a
shallow, to 5 by 10 mm cup at base adnate to the nut; 2
longer lobes to 9 by 2.5 cm, broadly spatulate, obtuse,
c. 6mm broad at base; 3 shorter lobes to 25 by 9 mm,
lanceolate, narrowly subacuminate; nut to 10 mm @,
subglobose.

Distr. Malesia: W. Java (S.W. Bantam: Udjong
Kulon).

Ecol. Rare, evergreen forests.

39. Vatica mangachapoi BLco, FI. Filip. ed. 1 (1837)
401; DC. Prod. 16, 2 (1868) 623; VIDAL, Sinopsis
(1883) t. ISB, f. 1-6; Rev. Pl. Vasc. Philip. (1886) 61:
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 134; Merr.
Publ. Govt. Lab. Philip. 27 (1905) 22; Philip. J. Sc. 1
(1906) Suppl. 98; Sp. Blanc. (1918) 272; En Philip. 3
(1923) 101; WuHitTForD, Philip. Bur. For. Bull. 10, 2
(1911) 76, t. 81; Foxw. Philip. J. Sc. 6 (1911) Bot. 282;
ibid. 13 (1918) Bot. 196; ibid. 67 (1938) 321; REYEs,
Philip. J. Sc. 22 (1923) 320; SLoor. Bull. Jard. Bot.
Btzg III, 9 (1927) 94; in Merr. Pl. Elm. Born. (1929)
205; BROWNE, For. Trees Sarawak & Brunei (1955)
101; ANDERSON, Gard. Bull. Sing. 20 (1963) 159;
ASHTON, ibid. 20 (1963) 253; Man. Dipt. Brun. (1964)
71, f. 10; ibid. Suppl. (1968) 33; MEWER & Woop,
Sabah For. Rec. 5 (1964) 310, f. 54; ASHTON, Gard.
Bull. Sing. 31 (1978) 22.

a. ssp. mangachapoi. — Mocanera mangachapoi BLco,
Fl. Filip. ed. 1 (1837) 450. — ?V. sinensis (non GMEL.)
BLco, FI. Filip. ed. 1 (1837) 401; ibid. ed. 2 (1845) 280;
ibid. ed. 3, 2 (1878) 156, ‘chinensis’; DC. Prod. 16, 2
(1868) 623. — V. apteranthera Buco, FI. Filip. ed. 2
(1845) 281; ibid. ed. 3, 2 (1878) 156. — Dipterocarpus
mangachapoi Buco, FI. Filip. ed. 2 (1845) 313; ibid. ed.
3, 2 (1878) 216; DC. Prod. 16, 2 (1868) 614. — Shorea

1982]

mangachapoi BL. Mus. Bot. Lugd.-Bat. 2 (1852) 34;
DC. Prod. 16, 2 (1868) 632; WALP. Ann. 4 (1857) 518;
F.-VILL. Nov. App. (1880) 21. — Preranthera sinensis
BL. Mus. Bot. Lugd.-Bat. 2 (1852) 30. — Preranthera
mangachapoi BL. l.c. — Anisoptera mangachapoi DC.
Prod. 16, 2 (1868) 616.— V. scaphula (non Dyer)
F.-ViLL. Nov. App. (1880) 21. — V. bureavi Heim,
Bull. Mens. Soc. Linn. Aetna BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 133, t. 3, f. 20-21; MERR. En.
Born. (1921) 409; SLoor. Bull. ese Bot. Btzg III, 9
(1927) 76.— Sunaptea bureavi Heim, Rech. Dipt.
(1892) 114. — V. reticulata KiNG, J. R. As. Soc. Beng.
Se. 62, 2 (1893) 106, non (THw.) A. DC. 1868:
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 131; SLoor. Bull.
Jard. Bot. Btzg III, 9 (1927) 83; Foxw. Mal. For. Rec.
10 (1932) 259, p.p.; BuRK. Dict. (1935) 2224. — Coty-
lelobium philippinense HEIM ex BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 134, nomen in syn. — Synaptea reticu-
lata RIL. FI. Mal. Pen. 1 (1922) 243. — V. whitfordii
Foxw. Philip. J. Sc. 67 (1938) 322, pl. 8. — V. patula
Sym. J. Mal. Br. R. As. Soc. 19, 2 (1941) 148, pl. 5A;
Mal. For. Rec. 16 (1943) 226, f. 107.

Young twig, panicle, petiole, leaf bud, and stipule
shortly densely cream pubescent, + caducous on all
but bud and panicle. Jwig to 1.5 mm @ apically,
smooth. Leaf bud to 2 by 1.5 mm, ovoid, obtuse.
Stipule to S by 2 mm, narrowly oblong, subacute.
Leaves 6-11 by 2.7—-S cm, coriaceous, elliptic; base
cuneate; acumen to 7 mm long; nerves 7-9 pairs,
slender, hardly raised on either surface, slightly
curved, at 45°-60°; no distinct secondary nerves;
midrib rounded beneath, raised above; petiole 5S—11
mm long, short. Panicle to 14cm long, terete, singly or
doubly branched, terminal or axillary. Flower buds to
1.2 cm long; calyx shortly densely cream-buff pubes-
cent; appendage to connective sharply acute; flowers
otherwise typical. Fruit pedicel to 4mm long, slender.
Calyx glabrous; 2 longer lobes to 5.5 by 1.5 cm,
spatulate, obtuse; base to 3 mm broad, free to pedicel;
3 shorter lobes to 10 by 4mm, lanceolate, acute. Nut to
4 by 6 mm, subglobose, glabrescent; style remnant,
short, abrupt.

Distr. Peninsular Thailand, in Malesia: N. Malaya
(N. Perak and Trengganu northwards), Borneo (Sar-
awak and Brunei; E. Sabah), Philippines.

Ecol. Common, becoming gregarious, especially
on dry ridges to 800 m, in more seasonal parts of
range; confirmed to Heath forest on podsols and
shallow peats below 400 m near coast in W. and
Central Borneo.

Vern. Resak julong (Malaya), r. sale (Sabah),
narig, karig (Philippines).

b. ssp. obtusifolia (ELMER) ASHTON, Gard. Bull. Sing.
31 (1978) 23. — V. obtusifolia ELMER, Leafi. Philip. Bot.
4 (1912) 1471; Foxw. Philip. J. Sc. 13 (1918) Bot. 196:
ibid. 67 (1938) 323; MerR. En. Philip. 3 (1923) 102.

Leaf small, thickly coriaceous, obtuse to subacu-
minate. Panicle not exceeding 6 cm long.

Distr. Malesia: S.W. Philippines (Palawan), N.
Borneo (E. Sabah).

DIPTEROCARPACEAE (Ashton)

365

Ecol.
local.

Note. A species with much local variation, espe-
cially in the Philippines. Forms approaching both V.
odorata ssp. mindanensis and V. pachyphylla occur and
suggest hybridisation.

Rocky exposed ridges and plateaux, very

40. Vatica parvifolia ASHTON, Gard. Bull. Sing. 19
(1962) 316, pl. 31; Man. Dipt. Brun. (1964) 76, f. 10:
ibid. Suppl. (1968) 35.

Young twig, panicle, leaf bud, and petiole densely
ferrugineous powdery tomentose, stipule sparsely so.
Twig to 1 mm g apically, much branched, becoming
glabrous, smooth. Bud to 3 by 1.5 mm, ovoid,
subacute. Stipules to 5 by 1 mm, linear, caducous.
Leaves 2.8-6 by 1—2.3 cm, coriaceous, narrowly ovate
to lanceolate; base obtuse; acumen to | cm long,
caudate; nerves c. 8 pairs, indistinct, unraised,
strongly curved, at 60°—75°; midrib slender, promi-
nent beneath, applanate above; petiole 6-9 mm long.
Panicle to 2cm long, terminal or axillary, terete, singly
branched. Flower bud to 6 mm long, small; calyx
grey-brown pubescent; flowers otherwise typical.
Fruit pedicel to 3 mm long, slender. Fruit calyx
glabrescent but for the persistently puberulent base,
lobes free to the pedicel; longer lobes to 6 by 0.7 cm,
subequal, oblong, narrowly obtuse, revolute above
the abruptly constricted base; 3 shorter lobes to 10 by
7 cm, subequal, broadly ovate, acute, cordate at base,
prominently revolute. Nut to 5S by 3.5 mm, broadly
ovoid, obtuse, shortly fulvous pubescent.

Distr. Malesia: Borneo (Sarawak and Brunei).

Ecol. Rare, Heath forest below 600 m.

Vern. Résak kérangas padi.

41. Vatica rynchocarpa ASHTON, Gard. Bull. Sing. 22
(1967) 270. pl. 14; Man. Dipt. Brun. Suppl. (1968)
36. — V. sp. ASHTON, Man. Dipt. Brun. (1964) 80.

Young twig, raceme, leaf bud, stipule, and petiole
shortly sparsely grey puberulent, glabrescent. Twig to
1 mm @ apically, slender, terete, becoming smooth.
Bud to 1.5 by 1 mm, conical, subacute. Stipule to 2.5
mm long, narrowly hastate, acute, fugaceous. Leaves
5-8.5 by 1.3-2.5 cm, narrowly elliptic to lanceolate,
thinly coriaceous; base obtuse; acumen to | cm long,
narrow; nerves 10-14 pairs, slender, curved near the
margin, hardly raised beneath, at 75°—-85°, well
spaced, with short obscure secondary nerves; midrib
slender, raised beneath, slightly depressed, especially
towards the base, above; petiole to 4 mm long, short,
slender. Flowers unknown; panicle to 3 cm long,
terminal or axillary, singly branched, pale brown
puberulent. Fruit pedicel to 7 mm long, slender; calyx
entirely glabrous, lobes free to base; 2 longer lobes to
6.2 by 1.4 cm, chartaceous, spatulate, narrowly
obtuse, to 2.5 mm broad at base; 3 shorter lobes to 15
by 2.5 mm, hastate, acute. Nut to 18 by 8 mm, ovoid,
glabrous, drying black; style remnant to 4 mm long,
linear.

Distr. Malesia: Borneo (from Kapuas and Rejang
valleys north-eastwards to S.E. Sabah).

366

FLORA MALESIANA

[ser. I, vol. 9?

Ecol. Locally frequent on clay alluvium river
banks.

42. Vatica endertii SLooT. Bull. Bot. Gard. Btzg III,
18 (1942) 248, f. 34; ASHTON, Man. Dipt. Brun. Suppl.
(1968) 32, f. 5.

Young twig, bud, petiole and occasionally ner-
vation beneath shortly pale grey-brown caducous
puberulent. Twig c. 1 mm @ apically, terete, much
branched, at first striated, becoming smooth; stipule
scars short, obscure. Bud to 2 by 2 mm, subglobose.
Stipule fugaceous. Leaves 4.8-14 by 1.7-5.5 em,
narrowly oblong to obovate or rarely lanceolate,
chartaceous; margin revolute; base cuneate; acumen
to | cm long, slender; nerves 11—14 pairs, unraised
above, prominent beneath, at 55°-60°, without
secondary nerves; tertiary nerves scalariform, slender
but elevated beneath; midrib hardly raised above,
prominent beneath; petiole 8-20 mm long. Flowers
unknown. Panicle to 3 cm long, singly branched,
terete, striated, shortly evenly persistently grey-brown
puberulent. Fruit pedicel to 8 mm long, slender. Calyx
glabrescent; 2 longer lobes to 8 by 2 cm, oblong-spatu-
late, obtuse, c. 3 mm broad above the base; 3 shorter
lobes to 20 by 6 mm, lanceolate, acute, to c. 3 mm
broad at base, hardly recurved. Nut to 6 by 5 mm,
subglobose, buff sericeous.

Distr. Malesia: Borneo (Central Sarawak, Tawau
area, W. Kutei, Sangkulirang).

Ecol. Mixed Dipterocarp forest, especially on acid
tuffaceous rocks, in lowlands but usually 700-1000 m.

43. Vatica brevipes ASHTON, Gard. Bull. Sing. 31
(1978) 24.

Small to medium-sized tree. Buds, petioles and
panicles densely persistently pale brown scabrid
puberulent; parts of petals exposed in bud and ovary
evenly so, sepals caducously evenly so; parts otherwise
glabrous. Twigs c. 2 mm @ apically, red-brown,
prominently rugose and ribbed, becoming flaky. Buds
to 3 by 2 mm, ellipsoid, Leaves (4-)5-13 by
(1.5—)2.5—-5.5 cm, elliptic or obovate, thinly coria-
ceous; base narrowly cuneate; acumen to 6 mm long,
short but slender; nerves 7-10 pairs, ascending,
straight at first, arching and forming a + incomplete
intramarginal nerve, slender but prominent beneath,
elevated above, with a few short secondaries; tertiary
nerves distantly reticulate, clearly evident on both
surfaces though more so below; petiole 5-11 mm long,
short, rather stout. Panicles to 1.6 cm long, short, very
slender, axillary or terminal, hardly branched; buds to
3 by 2 mm, ellipsoid; sepals narrowly deltoid, lanceo-
late, subacute; anthers broadly oblong, tapering dis-
tally to the deltoid appendages; sty/e broadly colum-
nar, slightly longer than the ovary, terminating in a
prominent rim beneath the deltoid style. Fruit pedicel
to 6 mm long, slender; 2 longer calyx lobes to 5 by 1.4
cm, spatulate, subacute or obtuse; 3 shorter lobes to
12 by 3 mm, lanceolate, acute; nut to 8 mm g,
subglobose, apiculate.

Distr. Malesia: Borneo (Sarawak: Ulu Rejang).

Ecol. Local, in Mixed Dipterocarp forest, to 700 m.

44. Vatica micrantha SLoor. Bull. Bot. Gard. Btzg
III, 17 (1942) 246, f. 33; AsHton, Man. Dipt. Brun.
(1964) 73, f. 10; ibid. Suppl. (1968) 33; MEER &
Woop, Sabah For. Rec. 5 (1964) 312.

Young twig, panicle, leaf bud, stipule, and petiole
densely + persistently pale brown scabrid tomentose,
leaf nervation beneath and midrib above sparsely so.
Twig to 2 mm @ apically, terete, smooth or slightly
striated. Bud to 3.5 by 2mm, ovoid, acute. Stipule to 7
by 3 mm, hastate, acute. Leaves 4.5—16 by 1.5—6 cm,
thinly coriaceous, elliptic-oblong to lanceolate; base
obtuse; acumen to 1.5 cm long, narrow; nerves 8-11
pairs, slender, curved, at 55°—70°, raised beneath, the
lamina frequently bullate between them; secondary
nerves slender, short; midrib slender, prominent
beneath, + applanate above; petiole 5-10 mm long.
Panicle to 7.5 cm long, terete, terminal or axillary,
singly branched; flower bud to 1.3 cm long; calyx
shortly pubescent; corolla cream, suffused with violet
towards base outside at first; flowers otherwise typi-
cal. Fruit pedicel to 2.5 mm long, short; ca/yx sparsely
glabrescent, free to the impressed base; 2 longer lobes
to 5.8 by 1.5 cm, chartaceous, oblong-spatulate,
subacute, base to 3 mm broad, revolute; 3 shorter
lobes to 25 by 6 mm, ovate, caudate-acuminate,
similar at base. Nut to 14 by 8 mm, ellipsoid, obtuse,
shortly pale yellow-brown pubescent; style remnant
short or usually absent.

Distr. Malesia: Borneo.

Ecol. Widespread in Mixed Dipterocarp forest
below 600 m, especially on sedimentary rocks.

Vern. Resak hijau (Brun.), r. bulu (Sabah).

45. Vatica flavovirens SLoor. Bull. Bot. Gard. Btzg
III, 17 (1942) 252, f. 36; Reinwardtia 2 (1952) 66.

Medium-sized tree. Young twig, petiole and calyx
densely evenly + caducous pale ocherous buff pubes-
cent; panicle, ovary, fruit pedicel and part of petals
exposed in bud persistently so. Twig c. 2 mm @
apically, terete, rugulose, drying dark brown. Leaves
8-21 by 2-7 cm, narrowly elliptic to lanceolate,
coriaceous, + lustrous, margin subrevolute; base
cuneate or occasionally obtuse; acumen to | cm long
or short, tapering; nerves 12-14 pairs, arched, slender
but + distinctly and equally elevated on both surfaces
as also the midrib, short + obscure secondary nerves,
and densely reticulate tertiary nerves; petiole 13-25
mm long, slender. Panicle to 14 cm long, terminal or
axillary, + irregularly doubly branched. Flower buds
to 10 by 3 mm; sepals narrowly lanceolate, unequal;
anthers ellipsoid; appendage very small; style as long
as ovary, short, capitate; stigma deltoid; flowers
otherwise typical. Fruit pedicel to 4mm long, slender;
calyx lobes unequal, free to base; 2 longer lobes to 4.5
by 1.7 cm, spatulate, acute, c. 5 mm broad at base; 3
shorter lobes to 20 by 5 mm, lanceolate, subacu-
minate; nut to 7 mm @, subglobose.

Distr. Malesia: Celebes.

Ecol. Locally common on hill slopes below 400 m.

1982]

DIPTEROCARPACEAE (Ashton)

367

Vern. Hulodiri puteh, h. motaha, kongieh, moro
larieh (Tobela), awalasa, bolampao, dama dama, tom-
borusu (Bug.).

46. Vatica badiifolia ASHTON, Gard. Bull. Sing. 22
(1967) 266, pl. 8; Man. Dipt. Brun. Suppl. (1968) 30,
pl. 7 (bark & slash). — V. bantamensis (non B. & H. ex
Mia.) ASHTON, Man. Dipt. Brun. (1964) 266, f. 10.

Young twig, panicle, leaf bud, stipule, and petiole
+ densely evenly shortly caducous fulvous pubescent.
Twig to 3 mm @ apically, terete or ribbed, becoming
smooth. Bud to 1.5 by 1 mm, ovoid, subacute. Stipule
to 5 by 1.5 mm, hastate, acute. Leaves 7.5—15 by 3-6.5
cm, coriaceous, + elliptic; base + cuneate; acumen to
5 mm long, deltoid; nerves 9-12 pairs, curved, promi-
nent, stout, terete beneath, slightly raised above;
petiole 2-3.5 cm long, thickened in the distal half.
Panicle to 8 cm long, terminal or axillary, singly or
doubly branched. Flower buds to 1.3 cm long; calyx
densely shortly cream pubescent; flower typical. Fruit
pedicel to 5 mm long, slender. Calyx lobes caducous,
pink-brown pubescent, free almost to the pedicel,
fused into a flat plate at the base; 2 longer lobes to 8 by
2.5 cm, oblong-spatulate, chartaceous, constricted to
6 mm wide at the base; 3 shorter lobes to 30 by 8 mm,
hastate, acute, constricted to 4 mm wide at base. Nut
to 8 mm long and @, globose, shortly evenly fulvous
pubescent, crowned by the up to 2 mm long abrupt
linear style remnant.

Distr. Malesia: Borneo (Rejang valley to Brunei;
W. Kalimantan: Kapuas valley).

Ecol. Deep soils on low coastals hills, Mixed
Dipterocarp forest.

47. Vatica perakensis KING, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 103, p.p.; emend. SyM. J. Mal. Br. R. As.
Soc. 19, 2 (1941) 152; Mal. For. Rec. 16 (1943) 226;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 132, t. 3, f.
18-19, p.p.; BRUHL & KING, Ann. R. Bot. Gard. Calc.
5, 2 (1896) 149, t. 181, p.p.; SLoor. Bull. Jard. Bot.
Btzg III, 9 (1927) 86, p.p.; Foxw. Mal. For. Rec. 10
(1932) 262, p.p.; BURK. Dict. (1935) 2224; AsHTON,
Gard. Bull. Sing. 31 (1978) 24. — Synaptea perakensis
Rw. Fl. Mal. Pen. 1 (1922) 242, p.p.— V. songa
SLooT. Bull. Jard. Bot. Btzg III, 9 (1927) 93, f. 4;
HEYNE, Nutt. Pl. ed. 2 (1927) 1131.

Small to medium-sized tree. Twig endings, petiole,
panicles, and parts of perianth exposed in bud
persistently greyish pink or yellow-brown cinereous,
leaf nervation beneath and nut glabrescent. Twig c. |
mm @ apically, slender, much branched, terete, pale
brown. Leaf bud small, ovoid. Leaf 6-14 by 1.8-3.8
cm, lanceolate to oblanceolate; base cuneate, taper-
ing; acumen to 2 cm long, slender, subcaudate; nerves
11-13 pairs, slender but prominent beneath, evident
above as also the midrib, ascending; tertiary nerves
densely subreticulate, evident on both surfaces; petiole
1-2 cm long, slender. Panicle to 5 cm long, singly +
irregularly branched; flowers typical. Fruit pedicel to 6
by 1 mm, long, slender; 2 longer calyx lobes to 6 by 1.8
cm, spatulate, obtuse, c. 5 mm broad at the subrevo-

lute base; 3 shorter lobes to 15 by 6 mm, lanceolate,
acute; nut to 10 cm @, ovoid, apiculate.

Distr. Malesia: Malaya (S. Kedah, Perak), Central
Sumatra (P. Musala, Tapanuli, Indragiri); one doubt-
ful collection (SAN 41580) from Pulau Kelambu near
the N.E. Borneo coast at Tawau.

Ecol. Common in coastal hills and on low ridges to
600 m.

Vern. Resak puteh.

48. Vatica nitens KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 104; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 131;
BRUHL & KING, Ann. R. Bot. Gard. Calc. 5, 2 (1896)
150, t. 182A; Burk. J. Str. Br. R. As. Soc. 81 (1920) 61;
Dict. (1935) 2224; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 78; Foxw. Mal. For. Rec. 10 (1932) 254; Sym.
Mal. For. Rec. 16 (1943) 223, f. 107, 108; ASHTON,
Man. Dipt. Brun. (1964) 74, f. 10; ibid. Suppl. (1968)
34, f. 5. — Synaptea nitens Ripv. Fl. Mal. Pen. 1
(1922) 241; ibid. 5 (1925) 292.— V. cuspidata (non
RIDL.) BROWNE, For. Trees Sarawak & Brunei (1955)
100.

Young vegetative parts densely powdery fulvous
tomentose, fugaceous on leaf, subpersistent only on
panicle, leaf bud and stipule. Twig to 2mm g apically,
becoming smooth, glabrous. Bud to 3 by 3.5 mm,
ovoid, acute. Stipule to 20 by 3.5 mm, narrowly
oblong, subacute, caducous. Leaves 10-17 by 3.5—5.5
cm, coriaceous, narrowly oblong; base obtuse or
cuneate; acumen to | cm long; nerves 12-22 pairs,
raised beneath, curved, at 45°—70°, with short second-
ary nerves; midrib obtuse, raised beneath, applanate
or slightly raised above; petiole 1—2 cm long, slightly
geniculate. Panicle to 11 cm long, terminal, occa-
sionally axillary, densely persistently fulvous tomen-
tose, terete or angular; singly, rarely doubly,
branched; bracteoles fugaceous. Flower bud to 7 by 3
mm, fusiform; calyx densely pubescent on both
surfaces, lobes narrowly deltoid, acute; appendage to
connective short, stout; style very short, not exceeding
anthers; flowers otherwise typical. Fruit pedicel 3—5S
mm long, calyx fulvous powdery tomentose, glabres-
cent, base impressed; 2 longer lobes to 13 by 2 cm, free
to base, oblong, slightly recurved, hastate, acute; base
to 7 mm broad, + revolute. Nut to 3 cm @, globose,
shortly densely persistently pale fulvous tomentose.

Distr. Malesia: Malaya (except seasonal areas),
Borneo (Kapuas valley; Sarawak to S.W. Sabah).

Ecol. Scattered on low hills, and ridges to 600 m, in
Mixed Dipterocarp forest.

Vern. Resak daun panjang (Mal.).

49. Vatica cuspidata (RIDL.) SyM. Mal. For. 3 (1934)
200; J. Mal. Br. R. As. Soc. 19 (1941) 149; Mal. For.
Rec. 16 (1943) 219, f. 107.— Synaptea cuspidata
Ripv. J. Str. Br. R. As. Soc. 82 (1920) 172; Fl. Mal.
Pen. | (1922) 242.— Synaptea maingayi Riv. FI.
Mal. Pen. 1 (1922) 240 p.p. — V. maingayi (non DYER)
SLoot. Bull. Jard. Bot. Btzg III, 9 (1927) 85, p.p.;
Foxw. Mal. For. Rec. 10 (1932) 261, p.p.
Medium-sized to large tree. Young twigs, petioles

368 FLORA MALESIANA [ser. I, vol. 9?

uls

Fig. 47. Vatica maingayi Dyer. a. Flowering twig, b. fruiting twig, c. fruit, all x2/3 (a SF 34959, b-c ANDERSON
9159).

1982]

and panicles persistently pale ferruginous scabrid
pubescent, ovary and perianth outside densely puber-
ulent, fruit calyx glabrescent. Twig c. 2mm @ apically,
ribbed, pale brown. Bud to 3 by 2 mm, ovoid, acute;
stipule fugaceous. Leaf 7-18 by 2.8—6 cm, elliptic to
oblong-lanceolate, coriaceous; base broadly cuneate;
acumen to | cm long, slender, prominent; nerves
10-17 pairs, arched, slender but distinctly elevated
beneath, less elevated above as also midrib and the
subscalariform tertiary nerves; petiole 2—4(—S) cm
long, c. 2 mm @, slender. Panicle to 7 cm long,
terminal or axillary, irregularly branched. Flower bud
to 6 by 2 mm; appendages very short, hardly exceeding
anthers; style columnar, slightly longer than ovary,
broadening distally into the prominent long slender
conical stigma; flowers otherwise typical. Fruit pedicel
to 5 by | mm, slender, prominent; 2 longer calyx lobes
to 7 by 1.5cm, spatulate, obtuse, c. 4mm wide at base;
3 shorter lobes to 25 by 7 mm, lanceolate, acute; nut to
8 by 8 mm, ovoid, apiculate.

Distr. Malesia: Malaya.

Ecol. Common, coastal hills and inland ridges.

Vern. Resak daun runching.

50. Vatica maingayi Dyer, FI. Br. Ind. | (1874) 302;
KinGc, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 104;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 131; SLoor. Bull.
Jard. Bot. Btzg III, 9 (1927) 85, p.p. quoad desc.;
Foxw. Mal. For. Rec. 10 (1932) 261, p.p.; BURK. Dict.
(1935) 2224; Sym. J. Mal. Br. R. As. Soc. 19 (1941)
151; Mal. For. Rec. 16 (1943) 223, f. 107; MEIJER &
Woop, Sabah For. Rec. 5 (1964) 309; ASHTON, Man.
Dipt. Brun. Suppl. (1968) 33. — V. lowii (non KING
emend. SYM.) KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 103, p.p.; SLoor. Bull. Jard. Bot. Btzg III, 9
(1927) 83. — Synaptea maingayi RIDL. FI. Mal. Pen. |
(1922) 240, p.p. — Synaptea lowii RIDL. Fl. Mal. Pen.
1 (1922) 241, p.p. — V. macroptera SLOOT. ex THOR-
ENAAR, Med. Proefst. Boschw. 16 (1926) 120, t. 21;
Bull. Jard. Bot. Btzg III, 9 (1927) 83, nomen in syn. sub
V. lowii. — V. aperta SLoogT. Bull. Bot. Gard. Btzg III,
Ili, 17 (1942) 250. — Fig. 42B—B2, 47.

Twigs, petioles, buds and stipules outside shortly
fulvous flocculent tomentose, patchily caducous
except on buds. Twigs c. 2 mm @ apically, smooth.
Bud to 3 by 3 mm, ovoid, acute, prominent. Stipules to
10 by 3 mm, lorate, leaving conspicuous pale horizon-
tal scars. Leaves 5.5—12.5 by 1.2-5 cm, elliptic,
somewhat chartaceous; base cuneate; acumen to | cm
long, deltoid; nerves 9-11 pairs, with few short
secondary nerves, applanate above, slender but
prominent beneath, arched at 35°—50°; midrib promi-
nent on both surfaces; tertiary nerves slender, reticu-
late; petiole 1-2.5 cm long, prominently geniculate,
ribbed. Panicle to 4 cm long, axillary, short, ribbed,
shortly patchily caducous fulvous flocculent tomen-
tose, irregularly singly branched; bracteoles c. 2 mm
long, linear. Flower buds to 1 cm long, fusiform; calyx
rufous flocculent tomentose: corolla bright red;
flowers otherwise typical. Fruit shortly fulvous floccu-
lent tomentose, subpersistent on nut, patchily cadu-

DIPTEROCARPACEAE (Ashton)

369

cous elsewhere. Pedice/ to 3 mm long; calyx lobes free
to within 2 mm of the pedicel; 2 longer calyx lobes to 8
by 2.5 cm, lorate to narrowly ovate, obtuse, c. 7 mm
wide and somewhat revolute at base; 3 shorter lobes to
2.5 by | cm, narrowly ovate, acute, 3 mm wide at base.
Nut to 7 by 7 mm, globose, shortly mucronate.

Distr. Malesia: Malaya, Singapore, Sumatra (Palem-
bang), Borneo (Sarawak W. of Lupar, S.W. Sabah).

Ecol. Locally frequent, on low lying land and
hillsides below 500 m.

Vern. Resak lidi, kéruing babi, jenuong (Mal.),
résak daun merah (Sabah).

51. Vatica lowii KiNG, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 103, p.p.; emend. Sym. J. Mal. Br. R. As. Soc. 19
(1941) 153; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 131,
p-p.; BRUHL & KING, Ann. R. Bot. Gard. Calc. 5, 2
(1896) 150, t. 182B; Sym. Mal. For. Rec. 16 (1943) 222,
f. 107. — V. perakensis KING, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 103, p.p.; Foxw. Mal. For. Rec. 10 (1932)
262, p.p.— Synaptea lowii RipL. Fl. Mal. Pen. 1
(1922) 241 p.p.

Small to medium-sized tree. Leaf bud, petiole and
panicle persistently pale rufous scabrid somewhat
flocculent puberulent; young twigs and calyx outside
caducously so; young leaves fugaceously so; petals on
parts exposed in bud and ovary prominently evenly
pale rufous puberulent. Twig c. 2 mm @ apically,
slender, becoming dark brown, smooth, terete. Buds
to 2 by 1 mm, small, ovoid. Leaf 5-14 by 2-5 cm,
elliptic-lanceolate, coriaceous, somewhat lustrous; base
broadly cuneate to obtuse; acumen to | cm long; nerves
11-14 pairs, slender, arched, distinctly elevated beneath,
slightly so above as also the midrib; tertiary nerves
densely subreticulate, evident and slightly elevated on
both surfaces; petiole 6-16 mm long, slender. Panicle
to 3.5 cm long, terminal or axillary, shortly irregularly
branched. Flower buds to 5 by 2 mm; appendages c. +
length of anthers, small, deltoid; sty/e shorter than
ovary, columnar, expanding to the short conical style;
flowers otherwise typical. Fruit pedicel to 3 by 0.5mm,
very slender, prominent; 2 longer calyx lobes to 6 by
1.4 cm, spatulate, obtuse, c. 3 mm wide at the sub-
revolute base; 3 shorter lobes to 18 by 6 cm, lanceo-
late, acute; nut to 5 mm @, subglobose, mucronate.

Distr. Malesia: Malaya (Kelantan-Thailand
border, Perak).

Ecol. Locally abundant, coastal hills and inland
ridges below 750 m.

Vern. Resak pipit.

Not yet placed

Mature fruit of the following species are unknown

52. Vatica elliptica Foxw. Philip. J. Sc. 67 (1938) 329.

Twig endings, petioles, panicles, calyx outside, parts
of petals exposed in bud and ovary + densely
persistently ferruginous scabrous puberulent, else-
where glabrous. Twig c. 1 mm @ apically, slender, at
first rugulose, becoming smooth, pale brown. Leaves

370

FLORA MALESIANA

[ser. I, vol. 9?

Ecol. Locally frequent along river banks.

4-11 by 13-25cm, narrowly elliptic, thinly coriaceous;
tapering to a narrowly obtuse base and to a 1 cm long
acumen; margin + narrowly revolute; nerves 12-14
pairs, arched, slender but prominent beneath, ele-
vated above, with short less prominent secondary
nerves; tertiary nerves subreticulate, distinctly ele-
vated on both surfaces but more prominent beneath
as also the midrib; petiole 4-9 mm long, short. Panicle
to 5cm long, slender, terminal or axillary, irregularly
doubly branched. Flower bud to 5 by 3 mm, lanceolate,
small; sepals subequal; anthers narrowly oblong;
appendages prominent, deltoid; style columnar,
expanding distally, slightly longer than ovary; stigma
deltoid; flowers otherwise typical. Mature fruit un-
known; very young fruit with subequal reflexed calyx
lobes.

Distr. Malesia: Philippines (Zamboanga, Min-
danao), twice collected.

53. Vatica pentandra ASHTON, Gard. Bull. Sing. 31
(1978) 24.

Twigs, petioles and panicles persistently shortly
scabrid fulvous pubescent, calyx outside + cadu-
cously so, parts of petals exposed in bud and ovary
persistently evenly buff puberulent. Twigs c. 2mm @
apically, ribbed, much branched, ascending. Leaf buds
minute. Leaves 18-45 by 9-20 mm, elliptic, thinly
coriaceous, with subrevolute margins, obtuse to
broadly cuneate base and + deeply retuse apex;
nerves 4—5 pairs, ascending, at c. 45°, slender, hardly
elevated on either surface though more so below;
petiole 4-11 mm long, slender, hardly geniculate.
Panicle to 4 cm long, slender, singly branched;
branchlets bearing to 3 flowers. Flower buds to 9 by 3
mm; stamens 5, in a single whorl; flowers otherwise
typical. Fruit unknown.

Distr. Malesia: East Borneo (Central Kutei).

Ecol. Unknown, lowlands.

Note. The only Vatica with but 5 stamens; the
leaves are among the smallest and most distinctive in
the genus also.

54. Vatica cauliflora ASHTON, Gard. Bull. Sing. 31
(1978) 25.

Twigs, petioles and panicles persistently scabrid
fulvous pubescent; midrib beneath caducously so;
calyx outside, parts of petals exposed in bud and
ovary persistently evenly buff puberulent. Twigs c. 3
mm g apically, stout, ribbed; /eaf buds minute. Leaves
7-22 by 2.5-8 cm, oblanceolate, coriaceous, with
subrevolute margins, obtuse to subcordate base and
+ prominent caudate acumen; nerves 14—16 pairs,
ascending at 50°-60°, prominent beneath, evident but
depressed above, as also the many secondaries; ter-
tiaries reticulate, barely elevated though evident on
both surfaces; midrib prominent beneath, less so
above; petiole 10-18 mm long, c. 2 mm g, relatively
slender. Panicles to 6 cm long, solitary, axillary,
doubly branched, many-flowered. Flower buds to 7 by
2 mm, lanceolate, otherwise typical.

Distr. Malesia: Ulu Kapuas, W. Borneo.

Note. This species is distinguished by the nerves
which are depressed above, by the caudate acumen
and slender petiole. Very young fruit indicate a
possible affinity with V. venulosa BL.

55. Vatica glabrata ASHTON, sp. nov.

V. mangachapoi BLco simulans lamina nervis intrin-
secus 9-I1 ascendentibus petiolo longissimo partibus
glabrescentibus ovariis excepto differt.

Medium-sized tree, to 20 m tall, 1 m girth. Apical
buds and ovary densely ocherous puberulent, young
parts sparsely caducously so, otherwise glabrous.
Stipules to 12 by 8 mm, elliptic, obtuse, not at first
caducous, leaving a more or less prominent falcate
scar. Leaves 6—15(—22) by Q3254 cm, lanceolate,
coriaceous, + concave, lustrous, with obtuse base and
prominently attenuate to 15 cm long acumen; nerves
9-11 pairs, ascending, slender, elevated on both
surfaces but more so below, with + short secondaries;
tertiaries reticulate, elevated on both surfaces. Petiole
1.8-4 cm long, slender, very long. Panicles to 7 cm
long, irregular, 1—CO axillary or terminal. Flowers
white, as in other species. Fruit unknown.

Distr. Malesia: Borneo: Sarawak, S 34865, 36852
(holotype in K, in fl.), Dulit range, Ulu Tinjar, Baram;
BRUN 2526, 2533, Ulu Medamit, Limbang.

Ecol. Local, scattered in submontane forest at
1200-1500 m on acid soils overlying sandstone imme-
diately below the mossy forest zone.

Notes. I collected this species in April 1958 while
on an expedition to the Pagon range. Fallen fruit,
apparently from this species, were also collected but
were lost in a river accident on the return: they were of
the Sunaptea type, with two long and three short
sepals free to the base. The new collections from
Sarawak, and more particularly the type differ in their
concave laminae and prominent subamplexicaul sub-
persistent stipules. The unusually long slender
petioles, 9-11 pairs of ascending nerves and glabrous
young twigs and inflorescences distinguish the four
collections from those of V. mangachapoi BLCo which
it otherwise resembles and convince me that they
represent but a single species.

Doubtful species

56. Vatica obtusa BURCK, Ann. Jard. Bot. Btzg 6
(1887) 228; Merr. En. Born. (1921) 409; SLoor. Bull.
Jard. Bot. Btzg III, 9 (1927) 126, nom. illeg., non
(WALL.) STEUD.

The type, TEYSMANN HB 11352, from Karimata, is
in flower. It is indistinguishable in this state from V.
umbonata (Hook. f.) BuRCK and V. pauciflora
(KorTH.) BL.

Excluded

Vatica imbricata SLooT. Bull. Jard. Bot. Btzg III, 16
(1940) 452, f. 10, is according to KOSTERMANS,
Reinwardtia 7 (1969) 426=Mesua acuminatissima
(MERR.) KOSTERM. (Guttiferae).

1982] DIPTEROCARPACEAE (Ashton) 371

6. DRYOBALANOPS

GAERTN. f. Fruct. 3 (1805) 49; DC. Prod. 16, 2 (1868) 606; Dykr, J. Bot. 12 (1874)
98, t. 142; BuRCK, Ann. Jard. Bot. Btzg 6 (1887) 242; Heim, Rech. Dipt. (1892) 81;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 46; BRANDIS & GILG in E. & P. Pfl. Fam. ed.
1, 3, 6 (1895) 258; SLoor. Bull. Jard. Bot. Btzg III, 12 (1932) 1; Sym. Mal. For.
Rec. 16 (1943) 191, f. 93 (map); BROWNE, For. Trees Sarawak & Brunei (1955)
111; Wyatt-SmiTH, Mal. For. 18 (1955) 145; AsHTon, Man. Dipt. Brun. (1964)
48; ibid. Suppl. (1968) 20; MrveR & Woop, Sabah For. Rec. 5 (1964)
278. — Pterigium CorrEA, Ann. Mus. Hist. Nat. Paris 8 (1806) 397, p.p., quoad P.
teres CORREA. — Baillonodendron HE, Bull. Mens. Soc. Linn. Paris 2 (1890)
867. — Fig. 48-53.

Large or very large, occasionally medium-sized, trees; with tall, long, concave,
rather thick, narrowly rounded plank buttresses, spreading over the surface
apically as large sinuous roots; crown very large, hemispherical or dome-shaped,
rather diffuse; ‘cauliflower’ shaped with a few large twisted branches ascending
from the bole apex, the branchlets numerous, bunched towards the ends. Bark
surface evenly or persistently shaggily flaked; twigs ribbed owing to the decurrent
leaf trace, slender. Stipules linear, fugaceous. Leaves coriaceous, prominently
acuminate, with slender dense hardly raised nervation; nerves very many,
parallel, straight, rarely branched apically, linked at the margin by a + visible
intramarginal nerve; secondary nerves, if present, indistinct; tertiary nerves
subreticulate, generally obscure; midrib sunken above, prominent beneath;
petiole distinctly channelled above, slender, not geniculate. /nflorescences
paniculate, lax, diffuse, few flowered; bracteoles minute, fugaceous. Flower buds
narrowly ovoid, obtuse. Calyx lobes equal, imbricate, glabrous, united at base
into a + cuneate tube tapering into the pedicel. Petals broadly elliptic, subacute,
glabrous, hardly contorted in bud, united at base and falling in a rosette, white.
Stamens c. 30, glabrous, subequal, the innermost slightly longer than the
outermost, epipetalous, yellow; filaments broad and compressed, connate at
base, tapering in the distal half and filiform below the anther; anthers long, linear,
latrorse; pollen sacs subequal, the outer somewhat larger than the inner;
appendage to connective small, stout, hardly projecting above the anthers. Ovary
+ ovoid, glabrous, without distinct stylopodium; style c. 3 times length of ovary,
glabrous; stigma minute. Fruit calyx with basal cup partially enclosing, but free
from, base of nut; 5 lobes subequal, subvalvate, short and acute or long spatulate,
obtuse, + rotate. Nut large, glabrous, with short apiculate style remnant.
Pericarp splitting at germination into 3 valves; cotyledons reniform, epigeal, on
long slender hypocotyl; first 2 pairs of leaves opposite, with a very short

intermediate internode.

Distr. Malesia: 7 spp. in Malaya, Central Sumatra, Borneo and intervening islands.

Fossil wood records from S. India, Cambodia, S. Sumatra, W. Java and Amboyna in the Moluccas
(SCHWEITZER, Palaeontographica 105B, 1959, 1-66). Fig. 49.

Ecol. Semigregarious or gregarious emergent canopy trees of lowland dipterocarp, Heath and Mixed Peat
Swamp forests to 800 m; on account of their size playing an important structural role in the forests where they
occur.

Uses. The timber is an important moderately heavy and durable construction timber. D. aromatica was once
a major source of camphor, obtained as crystals in splits within the bole by felling.

372 FLORA MALESIANA [ser. I, volRg?

b

Fig. 48. Dryobalanops aromatica GAERTN. f. a. Sterile twig, b. flowering twig, c. fruit, d. nut, all x5 (a, c-d SAN
15148, b KEP 72435).

KEY TO THE SPECIES

[Peknuiticalyxalobesishortenthannutes | 2 eee 2) se eo. | 8 ele soblongiola
1. Fruit calyx lobes longer than nut, aliform.
2. Leaves usually longer than 14 cm; nerves distinctly unequal, prominent beneath, depressed above
2. D. keithii
2. Leaves shorter than 12 cm; nerves hardly raised beneath, subequal, not depressed above.
3. Fruit calyx lobes 10-12 mm broad. Calyx fused into a to 15 mm @, over 5 mm deep, cup at base.

Avlbeatélanceolatesrevolutejay bascim mee tere: econ) fs) 4 Ae eee Delancealars
4. Leaf broadly ovate, not revolute at base. . . . 4. D. aromatica
3. Fruit calyx lobes to 8 mm broad, narrow. Calyx fused into a cup at base not broader than 8 mm, not deeper
than 3 mm.
5. Leaf 5-8 by 1.7-3 cm, glabrous. . . page!) ki /hoadee Se er BRS SD Abeccani
5. Leaf 6-11 by 2-4 cm, more or less tomentose beneath.
Gyullomentum\persistentevensidarkicold uh bes Vee ae ey E10 ee OD ats

Gy lomentumi-=caducoussflocculentsrufousy s)he kee ee ee een en een ores

DIPTEROCARPACEAE (Ashton)

373

Fig. 49. Density map of Dryobalanops GAERTN. f. in
Malesia; number of endemics above the hyphen,
number of non-endemics below it.

1. Dryobalanops oblongifolia Dyer, J. Bot. 12 (1874)
100, t. 142, f. 8-12; Burck, Ann. Jard. Bot. Btzg 6
(1887) 224; BrRaNpis, J. Linn. Soc. Bot. 31 (1895) 51;
BRANDIS & GILG in E. & P. Pfi. Fam. ed. 1, 3, 6 (1895)
259: Becc. Nelle For. Born. (1902) 550, 572; MERR.
En. Born. (1921) 401; Burk. J. Str. Br. R. As. Soc. 86
(1922) 291; DEN BERGER & ENDERT, Med. Proefst.
Boschw. 11 (1925) 104; GitGin E. & P. Pfl. Fam. ed. 2,
21 (1925) 255; THORENAAR, Med. Proefst. Boschw. 16
(1926) 110; Foxw. J. Mal. Br. R. As. Soc. 5 (1927) 340;
Mal. For. Rec. 1 (1921) 76; ibid. 3 (1927) 49; ibid. 8
(1930) 17; ibid. 10 (1932) 110; HEYNgE, Nutt. Pl. ed. 2
(1927) 1105; SLoor. Bull. Jard. Bot. Btzg III, 12 (1932)
22: BurK. Dict. (1935) 867; CORNER, Ways. Trees
(1940) 212; Sym. Mal. For. Rec. 16 (1943) 196, f. 94B,
95, 98; WyaATT-SMITH, Mal. For. 18 (1955) 153, p.p.:
BROWNE, For. Trees Sarawak & Brunei (1955) 116:
AsHTON, Man. Dipt. Born. Suppl. (1968) 23, f. 4:
Gard. Bull. Sing. 31 (1978) 25.

a. ssp. oblongifolia. — Baillonodendron malayanum
Hem, Bull. Mens. Soc. Linn. Paris 2 (1891) 867; Rech.
Dipt. (1892) 38. — D. abnormis S.ooT. Bull. Jard.
Bot. Btzg III, 16 (1940) 449, f. 8; cf. ASHTON, Gard.
Bull. Sing. 22 (1967) 262, 347 (photogr. fr.).

Young twigs caducously shortly evenly densely pale
fulvous pubescent; vegetative parts otherwise gla-
brous. Twig c. 2 mm @g at apex, terete, smooth,
lustrous. Bud to 2 by 1 mm, small, conical, acute.
Stipule to 3 by 1 mm, linear, fugaceous. Leaf 6-20 by
4.5-5(-6.5) cm, narrowly oblong, frequently undu-
late, base broadly cuneate to obtuse, apex cuspidate
with 1.5 cm long slender acumen; nervation evident
but not prominent beneath, obscure above; midrib
depressed above, prominent beneath. Petiole 5-14
mm. Panicle to 14 cm long, terminal or axillary,
angular or terete, densely shortly evenly rufous pubes-

cent; singly branched, branchlets to 3 cm long, bearing
to 6 distichous flowers; bracteoles minute, fugaceous.
Flower buds to 8 by 5 mm, ovoid, glabrous. Sepals
subequal, ovate, acute, glabrous. Petals white,
oblong, glabrous. Stamens c. 40, unequal, reaching to
below the style apex; filaments connate at base,
broadly lorate, tapering below the anther insertion, c.
4 length of anther; anther linear; appendage to
connective + erect, exceeding anther apex. Ovary
ovoid, glabrous; style 2-3 times as long as ovary,
filiform, glabrous. Fruit entirely glabrous. Pedicel
short, merging with calyx. Calyx lobes to 5 by 7 mm,
deltoid, acute, incrassate, frequently reflexed, border-
ing a to 1.5 cm deep, to 1.5 cm @ massive incrassate
cup. Nut to 3.5 by 2.7 cm, ellipsoid to obovoid, obtuse
to mucronate, prominently lenticellate.

Distr. Malesia: Borneo (Kapuas hinterland, Sar-
awak W. of the Kemena, W. Kutei).

Ecol. Local on hillsides below 600 m.

Vern. Empedu (Sar.), kélansau (Iban).

b. ssp. occidentalis ASHTON, Gard. Bull. Sing. 31
(1978) 26.— D. beccariana Ripv. Fl. Mal. Pen. 1
(1922) 211.— D. ovalifolia 1. H. BuRKILL, Illustr.
Guide Bot. Gard. Sing. (1927) 22, nomen.

Differing as follows: Leaf 6-11 by 2-4.5 cm; fruit
calyx tube to 8 mm deep, funnel-shaped, with +
obscurely 5-lobed undulate margin; nut broadly ellip-
soid, striated but otherwise smooth.

Distr. Malesia: Malaya (E. coast, rare from Perak
northwards in W.); E. Sumatra (Bengkalis, Riouw,
Djambi, Palembang).

Ecol. By streams and in fresh water swamps,
locally abundant; occasionally on hillsides.

Vern. Kéladan, kapur paya, k. kuras (Mal.), k.
guras, pétanang (Sum.).

2. Dryobalanops keithii Sym. Gard. Bull. S. S. 10
(1939) 379, pl. 27; SLoor. Bull. Jard. Bot. Btzg III, 16
(1940) 449: Reinwardtia 5 (1961) 475, f. 6; WyaTT-
SMITH, Mal. For. 18 (1955) 152; MEER & Woop,
Sabah For. Rec. 5 (1964) 283, f. 3 d.

Medium-sized tree. Outside of calyx and panicle
fugaceous puberulent, parts otherwise glabrous. Twig
c. 2mm g apically, ribbed at first, verrucose lenticel-
late, becoming smooth. Leaves 14-33 by 5-10 cm,
lorate to narrowly lanceolate or oblanceolate, coria-
ceous; margin narrowly subrevolute; base obtuse or
cordate; acumen to | cm long (longer in young trees).
slender, prominent; nerves many but relatively dis-
tant, distinct and prominent, beneath, obscure but
distinctly narrowly depressed above as also the mid-
rib, arched just within the margin and then uniting
with a + straight intramarginal nerve; with shorter
less distinct parallel secondary nerves and + obscure
reticulate tertiary nerves; petiole 7-12 mm long, short,
stout. Panicles to 14 cm long, singly or doubly (if
terminal) branched, terminal or subterminal axillary,
lax; branchlets to 3.5 cm long, bearing to 4 flowers;
bracts and bracteoles to 2 by 2 mm, suborbicular-del-
toid, not at first caducous. Flower bud to 10 by 4mm,

374

ellipsoid, acute; sepals narrowly deltoid, subacute;
petals ovate, acute, glabrous; stamens subequal,
slightly shorter than the style; filaments compressed,
tapering distally, united at base, c. 5 length of the
linear tapering anthers; appendages acicular, short
but distinctly exceeding anther apex; ovary narrowly
ovoid, puberulent; style filiform, glabrous, c. 23 times
length of ovary. Fruit pedicel to 4 by 2mm; calyx lobes
to 4 by 2 cm, broadly spatulate, obtuse, c. 8 mm broad
above the to 6 by 10mm basal cup; nut to 16 by 15mm,
depressed ovoid, apiculate.

Distr. Malesia: N.E. Borneo (Sandakan bay to
Tawau and Tidung).

Ecol. Undulating land and hills to 250 m, never far
from water and often on banks of streams; locally
common.

Vern. Kapur gumpait (Kadazan, Kinabatangan), k.
daun bésar.

3. Dryobalanops lanceolata BuRCK, Ann. Jard. Bot.
Btzg 6 (1887) 244, t. 29, f. 6; BRANDIS & GILG in E. & P.
Pfl. Fam. ed. 1, 3, 6 (1895) 244; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 51, t. 1, f. 1-6; Boer. Cat. Hort. Bog. 2
(1901) 100; Merr. En. Born. (1921) 401; GitG in E. &
P. Pfl. Fam. ed. 2, 21 (1925) 238; HEYNE, Nutt. Pl. ed. 2
(1927) 1106; SLooT. Bull. Jard. Bot. Btzg HI, 12 (1932)
28, f. 3; WyaTT-SmiTH, Mal. For. 18 (1955) 115;
Browne, For. Trees Sarawak & Brunei (1955) 116;
ASHTON, Man. Dipt. Brun. (1964) 52; ibid. Suppl.
(1968) 22; Mever & Woop, Sabah For. Rec. 5
(1964) 285, pl. 21 (stem), f. 47, pl. 29 (habit). — D.
oblongifolia Dyer, J. Bot. 12 (1874) 100, p.p. — D.
kayanensis Becc. Nelle For. Born. (1902) 551; MERR.
En. Born. (1921) 401; HeyNeE, Nutt. Pl. ed. 2 (1927)
1104.

Leaf bud and stipule sometimes shortly fugaceous
tomentose, otherwise entirely glabrous. Twigs 1.5—2
mm @ apically, slender, dotted with minute pale
brown lenticels, smooth but for the prominent ribs
above the lateral bundles of the petiole. Bud c. 4 by 1
mm, narrowly lanceolate. Stipule to 12 by 1 mm,
narrowly lanceolate. Leaves 7-10 by 2—3.5 cm, lanceo-
late, thinly coriaceous, base cuneate, with revolute
margin, acumen to | cm long, slender; margin fre-
quently undulate; petiole c. 1 cm long, slender. Panicle
to 6 cm long; terminal or axillary, ribbed and angular
on drying, glabrous; simple or singly branched.
Flower bud to 12 by 5 mm, fusiform, obtuse. Calyx
glabrous; lobes subequal, lanceolate, obtuse. Corolla
white; petals lanceolate, acute, glabrescent. Stamens c.
30, subequal, shorter than the style; filaments com-
pressed, slender, tapering, united at base; anthers
linear; appendage to connective short, hardly exceed-
ing anther apex. Ovary small, ovoid, glabrous; style
filiform, c. 3 times length of ovary, glabrous. Fruit
calyx with to 2 cm g, to 5 mm deep, shallow cup at
base, impressed at the pedicel; with a shallow tubercle
at the base of each calyx lobe; lobes to 9 by 2 cm,
spatulate, equal, obtuse, tapering gradually to a 3-S
mm broad base. Nut to 2 cm long and @, ovoid to
globose, glabrous, subacute.

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 50. Dryobalanops aromatica GAERTN. f. A. Bud,

B1. outer sepal, B2. inner sepal, both from inside, C.

stamens from outside, D. pistil, all x10 (Cult. Hort.
Bog. XI-B-XVII-213).

1982]

DIPTEROCARPACEAE (Ashton)

375

Distr. Malesia: Borneo (N.E. of the Lupar to
Sabah, and S.E. to W. Kutei and Sangkulirang).

Ecol. Widespread on fertile clay-rich soils, abun-
dant on undulating land on basic volcanics and
calcareous shale to 700 m.

Uses. The most valuable kapur source in N.E.
Borneo; the timber floats.

Vern. Paji, kapur paji(M., Iban), k. daram, k. bukit
(Brun.), seswan (Murut), ngeri (Bassap), adu (Kwijau),
jalam (Dus.).

4. Dryobalanops aromatica GAERTN. /. Fruct. 3 (1805)
49, t. 186, f. 2; BL. Fl. Jav. 2 (1828) Dipt. 6 in not.;
Mus. Bot. Lugd.-Bat. 2 (1852) 38; Wap. Ann. 4
(1857) 336; Hook. f. Trans. Linn. Soc. 23 (1860) 160;
BAILLON, Hist. Pl. 4 (1872) 202, f. 211-214; Dyer, J.
Bot. 12 (1874) 100, t. 142, f. 1-5; Burck, Ann. Jard.
Bot. Btzg 6 (1887) 243; LAKE & KELSALL, J. Str. Br. R.
As. Soc. 26 (1894) 35; Agr. Bull. Str. & F. M.S. 1
(1901) 61; Fl. Mal. Pen. 1 (1922) 210; BRANDIs, J.
Linn. Soc. Bot. 31 (1895) 50; BRANDIS & GILG in E. &
P. Pfl. Fam. ed. 1, 3, 6 (1895) 259, fig; BOERL. Cat.
Hort. Bog. 2 (1901) 100; HEYNg, Nutt. Pl. ed. 1, 3
(1917) 276; Burk. J. Str. Br. R. As. Soc. 81 (1920) 56,
fig.; ibid. 86 (1922) 291; MerR. En. Born. (1921) 401;
Foxw. Mal. For. Rec. 1 (1921) 74; ibid. 2 (1921) 68;
ibid. 8 (1927) 45; ibid. 10 (1932) 105; GiLG in E. & P.
Pfl. Fam. ed. 2, 21 (1925) 255; HENDERSON, Gard. Bull.
S. S. 4 (1928) 226; Watson, Mal. For. Rec. 5 (1928)
59, 60, 184; Epwarps, Mal. For. Rec. 9 (1930) 129;
SroonmeBullavard. Bota Btze, Tk 127(1932)) 7, fe 1:
Burk. Dict. (1935) 862; CORNER, Ways. Trees (1940)
211; Sym. Mal. For. Rec. 16 (1943) 194, f. 94A, 95, 96
(map), 97; WyaTT-SMITH, Mal. For. 18 (1955) 148;
BROWNE, For. Trees Sarawak & Brunei (1955) 114;
BACKER & BAKH. f. Fl. Java 1 (1963) 330; ASHTON,
Gard. Bull. Sing. 20 (1963) 241; Man. Dipt. Brun.
(1964) 49, f. 8, pl. 15 (habit), pl. 18 (stem); ibid. Suppl.
(1968) 22: MEWER & Woop, Sabah For. Rec. 5 (1964)
278, f.45. — Arbor camphorifera RuMPuH. Herb. Amb.
cap. 82 (1755) 67. — Pterigium teres CORREA, Ann.
Mus. Hist. Nat. Paris 10 (1807) 159, t. 8, f. 1. —D.
camphora CouepR. As. Res. 12 (1816) 535, t. 8; JACK in
Hook. Comp. Bot. Mag. 1 (1835) 264; Calc. J. Nat.
Hist. 5 (1843) 117; Korn. Kruidk. (1841) 68; WALP.
Rep. 5 (1845) 125; LinDL. Veg. Kingd. (1846) 393; DE
VRIESE in Mig. Pl. Jungh. (1851) 80; Ned. Kruidk.
Arch. 3 (1851) 1-89, cum tab.; transl. in Hook. J. Bot.
Kew Misc. 4 (1852) 33, 68; Hook. f. ibid. 4 (1852) 200;
DE VRIESE, Tuinb. FI. (1856) 86; Mem. sur le Camph-
rier (1856) t. 1-2; OUDEMANS, Ann. Sc. Nat. IV, 5
(1856) 90, t. 4; HAYNE, Arzneigew. 12 (1856) 17, cum
tab.; Mia. Fl. Ind. Bat. 1, 2 (1856) 499; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2 (1868) 606;
Wacp. Ann. 7 (1868) 377; DEN BERGER & ENDERT,
Med. Proefst. Boschw. 11 (1925) 104; HEYNE, Nutt. PI.
ed. 2 (1927) 1099. — Shorea camphorifera Roxs. FI.
Ind. ed. Carey 2 (1832) 616; ibid. ed. 4 (1873) 440; DC.
Prod. 16, 2 (1868) 632. — Dipterocarpus dryobalanops
SteuD. Nomencl. Bot. ed. 2, 1 (1840) 518; DC. Prod.
16, 2 (1868) 614; WALP. Rep. 5 (1845) 124. — Diptero-

carpus teres STEUD. Nomencl. Bot. ed. 2, 1 (1840) 518;
DC. Prod. 16, 2 (1868) 614. — D. junghuhnii BEcc.
Nelle For. Born. (1902) 554. — D. vriesii Becc. Nelle
For. Born. (1902) 554. — Fig. 48, 50-52.

Young twig, raceme, stipule, petiole and leaf
beneath at first sparsely minutely lepidote, appearing
yellowish, becoming glabrous. Twig c. | mm @
apically, slender, smooth. Bud 2-S by 1—1.5 mm,
small, narrowly lanceolate, often compressed. Stipule
5-8 mm long, small, linear, fugaceous. Leaves 4-6 by
2-4 cm, + broadly ovate, coriaceous, sometimes
(excluding acumen) broader than long, base cuneate
or broadly obtuse, acumen to 1.5 cm long, prominent;
nervation indistinct; midrib depressed above, promi-
nent beneath; petiole 0.5—1 cm long, slender. Panicle
to 7 cm long, terminal or axillary, ribbed and angular
on drying, singly or doubly branched. Flower bud to 9
by 4 mm, fusiform. Calyx glabrous, lobes subequal,
lanceolate, obtuse. Corolla white, petals glabrous, +
oblong, acute. Stamens c. 30, subequal, shorter than
the style; filaments slender, tapering, united at base;
anthers linear; appendage to connective short, linear,
somewhat exceeding anther apex. Ovary small, ovoid,
glabrous; style filiform, c. 2 times length of ovary,
glabrous. Fruit calyx base 6-8 mm deep, 8-15 mm @,
cup-shaped, + constricted at the rim; lobes equal, 4-6
by 0.8—2 cm, spatulate, obtuse, to 3-5 mm broad at the
base, entirely glabrous. Nut to 3 by 1.5 cm, ovoid,
glabrous, constricted at the apex of the calyx cup,
acute, with a short style remnant.

Distr. Malesia: Malaya (down E. coast from
Trengganu southwards, rare in Selangor and Negri
Sembilan), N.W. Sumatra (Angkola, Sibolga, Kela-
san and Upper Singkil), E. Sumatra (Bengkalis, Siak),
Musala, Lingga, Singkep, Borneo (N.E. of the Rejang
to S.W. Sabah).

Ecol. Locally dominant, gregarious on dry sandy
or gravelly soils over sandstone and granite, on
subcoastal hills or (rarely) inland quartzite dikes to
400 m.

Uses. The most important source of camphor, and
kapur timber, in the genus.

Vern. Kapur (Mal.), k. barus (Sum.), k. bukit, k.
peringii, k. anggi (Brun.), kéladan (Iban), télajin
(Belait).

5. Dryobalanops beccarii Dyer, J. Bot. 12 (1874) 100,
t. 142, f. 6-7; Burck, Ann. Jard. Bot. Btzg 6 (1887)
243: BRANDIS, J. Linn. Soc. Bot. 31 (1895) 50; BEcc.
Nelle For. Born. (1902) 572; Mrrr. En. Born. (1921)
401; SLoor. Bull. Jard. Bot. Btzg III, 12 (1932) 36;
WYATT-SMITH, Mal. For. 17 (1955) 149; BROWNE,
For. Trees Sarawak & Brunei (1955) 115; AsHTON,
Gard. Bull. Sing. 20 (1963) 242; Man. Dipt. Brun.
(1964) 51, f. 8, pl. 16 (stem); ibid. Suppl. (1968) 22;
MEIER & Woop, Sabah For. Rec. 5 (1964) 280, pl. 28
(stem), f. 46. — D. beccariana RiDL. Fl. Mal. Pen. 1
(1922) 211, p.p. — D. oiocarpa SLooT. ex HEYNE, in
Den Berger & Endert, Med. Proefst. Boschw. 11
(1925) 107; Nutt. Pl. ed. 2 (1927) 1106; ENDERT, M.-O.
Born. Exp. (1927) 239; VAN DER LAAN, HARING & LIT,

376 FLORA MALESIANA [ser. I, vol. 97

Fig. 51. Crown of Dryobalanops aromatica GAERTN. f. Brunei (Photogr. ASHTON).

1982]

Bijdr. boom. Z.O. Borneo (1928) 19, nomen. — D.
oocarpa SLOOT. Bull. Jard. Bot. Btzg III, 12 (1932) 33,
f. 4; Wyatt-SmiTH, Mal. For. 17 (1955) 149.

Young parts sometimes fugaceous puberulent, all
parts otherwise entirely glabrous. Twig to 1 mm @
apically, slender, smooth. Bud 3-4 by 2 mm, narrowly
lanceolate, compressed. Stipule 5-8 mm long, linear,
fugaceous. Leaves 5—8 by 1-3 cm, ovate to lanceolate,
comparatively thin, base cuneate, acumen to 17 mm
long, narrow, margin frequently undulate; petiole
0.7—1 cm long, very slender. Panicle to 10 cm long,
terminal or axillary, terete, wrinkled on drying,
irregularly doubly branched; bracteoles and bracts
small, linear, fugaceous. Flower bud to 10 by 3.5 mm,
fusiform, acute. Sepals glabrous, equal, narrowly
deltoid, subacute. Corolla white; petals large, broadly
elliptic, obtuse, glabrous. Stamens c. 30, subequal,
almost 3 length of style; filaments united in a tube
around the base of the ovary, the tube almost half as
long as the anthers; anther narrowly oblong, the cells
tapering, acute; appendage to connective short, erect,
slightly exceeding anther. Ovary ovoid, glabrous; style
2-3 times as long as ovary, filiform, glabrous. Fruit
calyx glabrous; base with to 8 mm g, shallow or to 5
mm deep cup, unconstricted at the rim, tapering
gradually and cuneate at the pedicel; lobes equal, to
6.5 by 0.8 cm, glabrous, oblong-spatulate, narrow,
subacute, to 2 mm broad at the base, very thin and
opaque between the reticulations of the nerves. Nut to
1.4 cm long and g, ovoid to globose, glabrous, with
acute style remnant, resting on the considerably
narrower calyx cup and pushing the lobes out to a
wide angle.

Distr. Malesia: Borneo (excepting S. and S.W.).

Ecol. Locally abundant on leached sandy soils on
coastal hills and inland ridges below 700 m.

Vern. Kapur, k. mérah, k. ranggi (Mal.), kéladan
(Iban).

Note. Collections from S.E. Borneo tend to have
longer narrower leaves, often less coriaceous, than
those from elsewhere; the species generally occurs on
deeper, more fertile soils there. Nevertheless there is
no consistent way in which these plants may be
distinguished in field or herbarium.

6. Dryobalanops fusca SLooT. Bull. Jard. Bot. Btzg
III, 12 (1932) 39, f. 5; Wyatt-SMiTH, Mal. For. 17
(1955) 149: AsHton, Man. Dipt. Brun. Suppl. (1968)
22, f. 4.

Leaf undersurface shortly densely evenly persis-
tently golden tomentose; midrib beneath, petiole, leaf
bud and stipule outside (subglabrous within) more
darkly so. Twig 1-2 mm g at the apex, slender, much
branched. Bud to 4 by 2 mm, lanceolate, acute. Stipule
to 3 by 1 mm, linear, fugaceous. Leaves S—10 by 24
cm, broadly lanceolate, coriaceous, base obtuse or
broadly cuneate; acumen to 1.5 cm long, slender,
tapering, caudate; nerves obscured by tomentum;
midrib depressed above, prominent beneath; petiole
5—10 mm long, short, drying pale rufous tomentose.
Panicle to 5 cm long, terminal or axillary, terete,

DIPTEROCARPACEAE (Ashton)

cae te aoa 4
Kiger J

>.

Ss et ee

Fig. 52. Trunk-base of Dryobalanops aromatica
GAERTN. f. Brunei (Photogr. ASHTON).

densely shortly evenly pale rufous tomentose; singly
branched, branchlets to 2 cm long, bearing to 3
flowers. Flower bud to 12 by 3 mm, lanceolate,
glabrous. Sepals equal, narrowly lorate-deltoid,
obtuse, glabrous. Petals lanceolate, glabrous.
Stamens c. 30, subequal, reaching to below style apex:
filaments connate at base, tapering, c. 2/3 length of
anthers; anthers narrowly oblong, tapering; appen-
dage to connective erect, extending somewhat above
anther apex. Ovary ovoid, glabrous; style 34 times
length of ovary, filiform, glabrous. Fruit entirely
glabrous. Pedicel to 6 mm long, prominent. Calyx
lobes to 6 by 1.3 cm, spatulate, obtuse, c. 4 mm wide
above the to 3 mm deep, to 7mm g, shallow thickened
basal cup. Nut to 2 by 1.3 cm, ellipsoid-ovoid,
apiculate.

Distr. Malesia: W. Borneo (N. of the Kapuas and
W. of the Lupar).

Ecol. Locally dominant on podsols, low ridges and
especially raised beaches, in Heath forest.

Vern. Empedu, kapur émpedu (Mal.).

7. Dryobalanops rappa Becc. Nelle For. Born. (1920)
572; MerRR. En. Born. (1912) 401; SLoor. Bull. Jard.
Bot. Btzg III, 12 (1932) 41; Wyatt-SmitH, Mal. For.

378 FLORA MALESIANA [ser. I, vol. 9?

Fig. 53. Trunk and crown of Dryobalanops rappa BrEcc., kapur paya. Note shaggy bark. Sarawak (Photogr.
SMYTHIES).

18 (1955) 155; BRowneE, For. Trees Sarawak & Brunei
(1955) 116; ANDERSON, Gard. Bull. Sing. 20 (1963)
157; ASHTON, Gard. Bull. Sing. 20 (1963) 242; Man.
Dipt. Brun. (1964) 53, f. 8, pl. 19 (stem); ibid. Suppl.
(1968) 23; MEvER & Woop, Sabah For. Rec. 5 (1964)
289, f. 48: AsHton, Gard. Bull. Sing. 31 (1978)

25. — D. oblongifolia (non DYER) WYATT-SMITH, Mal.
For. 18 (1955) 155. — Fig. 53.

Young twig, raceme, stipule and bud, petiole,
midrib and leaf beneath densely powdery caducous or
persistent rufous tomentose. Twigs slender, becoming
smooth but for minute, warty lenticels. Bud 3-6 by 2-3

1982]

mm, ovoid, subacute, slightly compressed. Stipule 5-8
mm long, linear. Leaves 6-11 by 2.5—4 cm, ovate-lan-
ceolate, coriaceous, base obtuse; acumen to | cm long,
narrow; midrib prominent, grooved, beneath; petiole
6-10 mm long, stout. Panicle to 8 cm long, terminal or
axillary, ribbed apically on drying, otherwise terete;
regularly alternately doubly branched, branchlets
bearing up to 4 flowers; bracteoles linear, fugaceous.
Flower bud to 10 by 5 mm, ellipsoid, acute. Petals
cream, lanceolate, glabrous. Sepals equal, narrowly
deltoid, obtuse, glabrous. Stamens c. 30, subequal,
reaching to below style apex; filaments united in a tube
round the ovary, tapering apically to the anthers, c. 4
length of anthers; anthers narrowly oblong, tapering;
appendage to connective erect, extending somewhat
above anther apex. Ovary ovoid, glabrous; style twice
as long as ovary, filiform, tapering, glabrous. Base of
fruit calyx 5-8 mm @, 3 mm deep, shallowly cupped;
lobes to 5 by 0.6 cm, linear to spatulate, subacute,

DIPTEROCARPACEAE (Ashton)

379

2.5-3.5 mm broad at the base. Nut to 10 by 6 mm,
ovoid, glabrous, crowned by a c. 6 mm long slender
persistent style remnant.

Distr. Malesia: Borneo (Lower Kapuas, Sarawak,
Brunei, W. Sabah).

Ecol. Common, locally dominant and gregarious,
in Mixed Peat Swamp forest overlying sand terraces,
and on podsols below 200 m.

Uses. Considered of inferior quality, darker,
harder and more liable to splitting than other species.

Vern. Kapur paya (Mal.), k. rappa (Sar.), lu’an
(Dusan), /esuan (Murut), kajatan (West Borneo).

Note. SFN 32194, a flowering collection from
swamps in S.E. Johore, differs from D. oblongifolia
DYER in possessing more coriaceous leaf and rufous
flocculent pubescent innovations and inflorescence, in
this resembling this species which is otherwise un-
known outside Borneo. Fruit are required to confirm
this record.

7. PARASHOREA

KuRZ, J. R. As. Soc. Beng. Sc. 39, 2 (1870) 65; BuRcK, Ann. Jard. Bot. Btzg 6
(1887) 221; Heim, Rech. Dipt. (1892) 54; BRANpis, J. Linn. Soc. Bot. 31 (1895)
103; SLooT. Bull. Jard. Bot. Btzg III, 8 (1927) 370; Foxw. Philip. J. Sc. 67 (1938)
316; Sym. Mal. For. Rec. 16 (1943) 97, f. 58 (map), 59-62; ASHTON, Man. Dipt.
Brun. (1964) 82; ibid. Suppl. (1968) 38; MEIJER & Woop, Sabah For. Rec. 5 (1964)
190; WANG Hsre, Acta Phytotax. Sin. 15, 2 (1977) 10, 22, fig.; ASHTON, Gard. Bull.
Sing. 31 (1978) 26; SMITINAND, Thai For. Bull. (Bot.) 12 (1980) 54. — Fig. 54-59.

Large trees with large, rounded, slightly concave, hardly branched plank
buttresses. Crown dense, becoming dome-shaped or hemispherical, even, the
branchlets usually crowded towards the ends but not ‘cauliflower’ shaped; main
branches several to many, radiating from the bole apex (except in P. macro-
phylla). Bark surface distinctly mauve-grey to purplish, with narrow shallow
fissures, broad smooth or flaking flat ridges, and numerous conspicuous large
pale corky lenticels. Stipules linear to hastate, fugaceous. Leaves broadly
oblong-ovate, shortly acuminate; nerves scalariform, with subpersistent plicate
folding; no intermediate nerves; young leaves white beneath. Inflorescence
racemose (except P. macrophylla). Flower and fruit as in Shorea, but sepals in bud
+ equal, very narrowly imbricate. Petals falling separately. Stamens 15, much
longer than ovary in bud; filaments short, dilated and compressed at base,
tapering abruptly and filiform below anthers; anthers narrowly oblong, with 4
pollen sacs, the inner 2 shorter than outer 2; appendage to connective hardly
exceeding anther apex, not reflexed, stout. Ovary small, + ovoid, shortly
pubescent, with or without a slender stylopodium; style as long as or longer than
ovary, filiform, glabrous. Fruit pedicel short. Fruit calyx lobes either + spatulate,
slender, thickened and saccate at the valvate base, frequently subequal, 3
somewhat larger than the other 2, or shorter than nut, equal; pushed apart by the
ripening nut. Nut large, globose, pubescent, verrucose-lenticellate; style remnant
short, indistinct. Germination as in Shorea, but with seedling leaves at first linear,
quite unlike the peltate sapling leaves.

380

FLORA MALESIANA

(ser. I, vol. 9?

Distr. About 14 spp.: southern Burma, Thailand, Indochina and S. China (Yunnan, K wangsi); in Malesia: 10
spp., in Malaya, Sumatra, Borneo, the Philippines and intervening islands. Fig. 55.

Ecol.
occasionally hills to 1400 m.

Local, sometimes abundant, in lowland Mixed and Semi-evergreen Dipterocarp forests, and

Uses. An important pale coloured light hardwood exported from the Philippines and N.W. Borneo.

KEY TO THE SPECIES

1. Fruit sepals shorter than nut, linear-lanceolate, subequal.

2. Leaves glabrescent, silvery stellate puberulent beneath; nerves to 9 pairs (saplings excluded)

1. P. aptera

2. Leaves not as above, generally pubescent at least on nervation; nerves at least 10 pairs.
3. Leaves 7-15 by 3—3.6 cm, sparsely pubescent or glabrous beneath; tertiary nerves evident but not

prominent; petiole 9-12 mm long

. 2. P. densiflora

3. Leaves 12.5—18 by 6.5—9 cm, persistently scabrid onibesea beneath fete nerves shea prominent

beneath; petiole 14-18 mm long .

1. Fruit sepals aliform, much longer than the nut, spatulate, unequal.

3. P. globosa

4. Leaves 30-50 by 16-24 cm, very large, white beneath, with 28-36 pairs of close straight nerves

4. Leaves shorter than 20 cm, nerves less than 20 pairs.

4. P. macrophylla

5. Leaves (mature trees) glabrous, drying as dark beneath as above; plication obscure; nerves distant, arched,
elevated but slender and not prominent beneath; petiole long, slender, geniculate.

6. Leaves 6-9 by 3—4.5 cm; tertiary nerves distant, elevated beneath
6. Leaves 6-16 by 2.3-8 cm; tertiary nerves dense, obscure .

. 5. P. parvifolia
6. P. stellata

5. Leaves longer than 9 cm, silvery lepidote or puberulent, dull beneath: plication subpersistent; nerves dense,

hardly arched, prominent beneath.

7. Siipule scars amplexicaul. Leaves broadly elliptic to ovate, base unequal.
8. Leaves glabrous beneath. Connectival appendage longer than anther. Fruit sepals to 15 cm long

7. P. malaanonan

8. Leaves + persistently pubescent beneath. Connectival appendage shorter than the anther. Mature fruit

sepals to 20 cm long.

7. Stipule scars short, descending. Leaves not as above.

. 8. P. tomentella

9. Leaves 6-14 by 2.5—6 cm, glabrescent beneath as also the petiole; nerves to 12 pairs; petiole to 15 mm

long

. 9. P. lucida

9. Leaves 10-18 or aq cm; Wmenvation beneath ara satis seabrid! pubescent nerves 14~—18 pairs; petiole

15-22 mm long

1. Parashorea aptera SLooT. Bull. Jard. Bot. Btzg III,
8 (1927) 377, f. 3; Reinwardtia 5 (1961) 478.

Tall tree. Young twigs, buds, petiole, bracts out-
side, panicles, calyx parts of corolla exposed in bud
and ovary densely evenly + persistently buff pubes-
cent; nerves beneath sparsely + caducously so. Twig
c. 2 mm @ apically, much branched, + ribbed,
becoming smooth, terete, blackish; stipule scars short,
obscure. Bud to 6 by 4 mm, ovoid, acute; stipules
fugaceous, not seen. Leaves 7-17 by 2.8—-6 cm, ovate-
lanceolate, coriaceous, + distinctly persistently pli-
cate, dull and + silvery stellate puberulent beneath;
base cuneate, apex to 1.5 cm long slender acuminate;
nerves 6—9(-10) pairs, ascending, rather straight,
slender but prominent and drying blackish beneath,
applanate above; tertiary nerves very slender, hardly
elevated, densely scalariform; midrib prominent
beneath, shallowly furrowed above; petiole 9-18 mm
long, geniculate. Panicle to 9 cm long, terminal or
axillary, singly or doubly branched, branchlets with 3
prominently pedicellate flowers; bracts to 4 by 3 mm,
elliptic. Flower buds to 6 by 4mm, ovoid; calyx, petals
and ovary densely hirsute. Sepals short, broadly
ovate-deltoid. Stamens slightly exceeding style at

10. P. smythiesii

anthesis; filaments compressed, broad, short,
attenuate; anthers lorate; appendages very short,
hardly exceeding the anthers; ovary small, ovoid; style
longer than ovary, slender. Fruit pedice/ to 3 mm long,
stout; sepals to 14 by 3 mm, subequal, linear-
attenuate, appressed to nut, incrassate at base. Nut to
2.5 cm @, subglobose, apiculate, densely verrucose.

Distr. Malesia: Sumatra (Labuan Batu, E. coast:
Rawas, Palembang).

Ecol. Locally frequent below 70 m in lowland
forest on sandy soils on hills.

Vern. Meéranti horsik, ngérawan batu, mérakunyit
lawang.

Note. Some sterile Sumatran collections may
represent the Malayan P. densiflora.

2. Parashorea densiflora SLooT. & Sym. Gard. Bull. S.
S. 10 (1939) 373, pl. 24; Sym. Mal. For. Rec. 16 (1943)
100, f. S9A, 60, 61. — P. aptera (non SLooT.) Foxw.
Mal. For. Rec. 10 (1932) 243; Burk. Dict. (1935)
1664.

Large tree. Young twigs, petioles and leaves below
sparsely pubescent (West coastal Malaya) or glabrous
(E. coast), panicle and parts of perianth exposed in

1982]

DIPTEROCARPACEAE (Ashton)

38]

Bisvieccc:-

Sor.

u?
(ie,
v4

.
Ye
S*
SY:
Sy
QV
»
Ne

2 Wb NS di, LAME MIS EV SN ih

Fig. 54. Flower details in Parashorea malaanonan
(BLCO) Merk. A. Bud, B/. outer sepal, B2. inner sepal,
both from inside, C/. stamens from outside, C2.
stamens from inside, D. pistil, all x 5 (PUASA 4643).

bud densely persistently pale brown pubescent, ovary
caducously so. Twig c. 3 mm g, ribbed, pale to dark
brown; stipule scars pale, horizontal, persistent. Bud
c. 4 by 3 mm, ovoid; stipule to 12 mm long, linear,
fugaceous. Leaves 7-15 by 3-6.5 cm, elliptic to ovate,
coriaceous, + densely pale purplish stellate lepidote
beneath; base cuneate (if CO-nerved) or obtuse to
subcordate (if few-nerved); acumen to | cm long,

Fig. 55. Density map of Parashorea Kurz in Malesia:
number of endemics above the hyphen, number of
non-endemics below it.

tapering; nerves 10-20 pairs, stout and prominent
beneath, evident above, ascending, the basal pair
frequently branching laterally; tertiary nerves scalari-
form, sinuate, evident on both surfaces; midrib promi-
nent beneath, obscure and depressed above; petiole
9-12 mm long, stout. Panicle to 6 cm long, c. 3 mm g,
stout, terminal or axillary, short, congested. Flower
buds to 6 by 4mm, ovoid-lanceolate; sepals lanceolate,
the 2 inner slightly the smaller, acuminate; petals
yellow, tinged purplish at base; stamens in 3 unequal
verticils; filaments short, compressed, tapering;
anthers oblong, the outer loculi sparsely setose;
appendage hardly exceeding anthers, stout: style
filiform, + twice length of the ovoid ovary. Fruit
pedicel to 7 mm long, to 3 mm g, expanding into the
receptacle; calyx lobes to 12 by 3 mm, subequal,
narrowly lorate, acute, becoming reflexed; nut to 3 cm
@, subglobose.

Distr. Malesia: Malaya (southern half and in
Pahang and Trengganu).

Ecol. Scattered in lowland dipterocarp forest
below 500 m.

Vern. Méranti pasir, téngkawang jantong.

Note. Collections from the Malayan East coast are
glabrescent, with somewhat smaller cordate fewer-
nerved leaves; more fertile collections are needed to
establish their status, as they are indistinguishable
from P. aptera when sterile.

3. Parashorea globosa Sym. Gard. Bull. S.S. 10 (1939)
375; Mal. For. Rec. 16 (1943) 101, f. 60.

Large tree. Twig apices, leaf buds, stipules, petioles
and leaf nervation beneath + densely ocherous to
rufous scabrid puberulent. Twig c. 3 mm @, stout,
becoming rugulose; stipule scars short, somewhat
ascending, falcate. Budc. 4 by 3 mm, ellipsoid, obtuse;
stipule to 23 by 6 mm, lanceolate. Leaf 12.5-18 by
6.5-9 cm, ovate-oblong, thickly coriaceous, rufous

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 56. Stem-base and leaves of Parashorea macro-
phylla WyatT-SMITH ex ASHTON. Kuala Belalong,
Brunei (Photogr. G.H.S. Woop, SAN 17377).

beneath (mature trees); base subcordate; apex acute to
subacuminate; nerves c. 18 pairs, prominent beneath,
evident above, spreading at leaf base, ascending
towards apex; tertiary nerves slender but sharply
prominent beneath, evident above; petiole 14-18 mm
long, stout. Panicles and flowers unknown. Fruit
sepals to 6 by 3 mm, lorate, obtuse; nut to 4 cm @,
subglobose.

Distr. Malesia: Malaya (Perak), Sumatra (Labuan
Batu, E. coast).

Ecol. Rare, lowland dipterocarp forest on low-
lying land and up valleys at the margin of the plains.

Vern. Meéranti pasir daun bésar.

4. Parashorea macrophylla WyATT-SMITH ex ASH-
TON, Gard. Bull. Sing. 19 (1962) 262, pl. 5; ASHTON,
Man. Dipt. Brun. (1964) 83, f. 11; ibid. Suppl. (1968)
38, pl. 8-9 (habit, bark). — Parashorea sp. BROWNE,
For. Trees Sarawak & Brunei (1955) 128 — Fig. 56.
Young twig, leaf bud, stipule (outside, glabrous
within), petiole and inflorescence shortly densely
evenly persistently pale ocherous tomentose. Twig to
12 by 5mm @ apically, compressed, becoming terete,
smooth, amplexicaul; leaf scars large, elliptic, conspi-

cuous. Bud to 9 by 0.8 cm, linear, acute, pale
cream-brown. Stipule to 15 by 2.5 cm, linear, sub-
acute, caducous. Leaves alternate, 30-50 by 16-24 cm,
subchartaceous, oblong-elliptic, silvery to white
below, base subcordate, apex obtuse or shortly acu-
minate; nerves 28-36 pairs, dense, straight, curved
towards the margin, not silvery, with semi-persistent
interneural plicate folds; tertiary nerves slender, den-
sely scalariform, diagonal to nerves; midrib stout,
rounded beneath, slightly raised above; petiole 3-5 cm
long, terete. Panicle to 16 cm long, terminal or
axillary, doubly or trebly branched; branches alter-
nate, branchlets cymose; bracts to 4 by 2.5 cm, ovate,
acute, cupped, amplexicaul, caducous, densely shortly
tomentose outside. Petals oblong, obtuse. Stamens
subequal; filaments short, deltoid; anthers narrowly
oblong, the inner pollen sacs somewhat the shorter;
appendage prominent, tapering, c. 5 length of anther.
Ovary conical, shortly pubescent, tapering into a
prominent narrowly cylindrical stylopodium. Fruit
subsessile; calyx puberulent (more densely so at base),
glabrescent; 3 longer lobes to 22 by 1.8 cm, spatulate,
brittle, subacute, tapering to 6mm broad above the to
16 by 8 mm elliptic shallowly saccate base; fused at the
base to form an up to 8 mm deep, to | cm broad, cup at
the pedicel; 3 shorter lobes to 12 by 0.7 cm, linear,
unequal, tapering to 3 mm above the to 5 mm broad
and long more prominently saccate base. Nut to 2.5 by
1.2 cm, ellipsoid, densely buff tomentose; style rem-
nant to 6 mm long, linear.

Distr. Malesia: Borneo (Kapuas hinterland, Sar-
awak and Brunel).

Ecol. Moist clay rich soils in valleys and gulleys, on
and near river banks in inland areas, locally abundant;
to 600 m.

Vern. Péran (Mal.), bilat (Ib.).

5. Parashorea parvifolia WyATT-SMITH ex ASHTON,
Gard. Bull. Sing. 19 (1962) 264, pl. 6; ASHTON, Man.
Dipt. Brun. (1964) 85, f. 11, pl. 24 (stem); ibid. Suppl.
(1968) 38; MEIJER & Woop, Sabah For. Rec. 5 (1964)
[95seh 23%

Young twig, leaf bud, stipule, petiole and panicle
sparsely shortly pale yellow-brown tomentose, cadu-
cous on twigs and petioles. Twig 1 mm @ apically,
terete, smooth, glabrous, dotted with minute pale
round lenticels; stipule scars short, slender. Bud to 3
by 1 mm, narrowly lanceolate, acute. Stipule to 3.5
mm long, narrowly hastate, acute, fugaceous. Leaves
6-9 by 3-4.5 cm, elliptic to ovate, glabrous; base
broadly cuneate; acumen to 1.5 cm long, narrow to
caudate; nerves 8-10 pairs, slender, raised beneath,
curved, well spaced, at 50°—-60°; tertiary nerves scalari-
form to subreticulate, widely spaced, c. 90° to nerves;
midrib slender, prominent beneath, depressed above;
petiole \-1.8 cm long, geniculate, pale. Panicle to 14
cm long, terminal or axillary, singly branched,
straight, pendant; bracts and bracteoles unknown.
Flower bud to 4.5 by 3 mm, ellipsoid, obtuse. Calyx
densely shortly grey-brown tomentose outside, gla-
brous within, lobes narrowly deltoid, subequal, sub-

1982]

DIPTEROCARPACEAE (Ashton)

383

acute, not adpressed to corolla in bud. Petals elliptic,
obtuse, shortly tomentose on parts exposed in bud,
pale cream. Inner 5 stamens slightly longer than the
others, reaching 3 length of style; filaments broad at
base tapering; anther narrowly oblong; appendage to
connective short, slightly extruding above anther.
Ovary subglobose, densely tomentose; style c. 3 times
length of ovary, filiform, shortly pubescent in the
basal 4, otherwise glabrous. Fruit calyx puberulent at
base, glabrescent, becoming pushed apart by the
ripening nut; 3 longer lobes to 8.5 by 1.7 cm, broadly
spatulate, obtuse, to 3 mm broad above the to 7 mm
long unexpanded slightly thickened base; 2 shorter
lobes to 7.5 cm long, often only slightly smaller than
longer lobes, subequal. Nut to 1.5 by 1.3 cm, ellipsoid,
obtuse, verrucose with pale lenticels, buff tomentose:
style remnant to | mm long, short.

Distr. Malesia: Borneo (N.E. of Rejang to Sabah,
Bulungan and Tidung).

Ecol. Local, in Mixed Dipterocarp forests on clay
rich soil, on ridges in mountains to 1350 m, rarely on
river banks.

Vern. Urat mata bukit (Brun.), urat mata daun
kéchil (Sabah), lantan kuning (Tidung).

6. Parashorea stellata KURZ, J. R. As. Soc. Beng. Sc.
39, 2 (1870) 66; Fl. Burma 1 (1877) 117; Pierre, For.
Fl. Coch. 3 (1889) t. 224; BRANDIS & GILG in E. & P.
Pfi. Fam. ed. 1, 3, 6 (1895) 267; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 104, t. 2, f. 11-12; Indian Trees (1906)
70, fig.; GUERIN, Fl. Gen. I.-C. 1 (1910) 388, f. 38;
Ript. Agr. Bull. Str. F.M.S. 6 (1907) 170; Fl. Mal.
Pen. 1 (1922) 234; Troup, Silv. Ind. Trees 1 (1921) 134;
Foxw. Mal. For. Rec. | (1921) 70; ibid. 3 (1927) 63:
ibid. 10 (1932) 242; GAMBLE, Man. Ind. Timb. (1922)
83; BAKER f. J. Bot. 62, Suppl. (1924) 11; Crarp, FI.
Siam. Enum. | (1925) 144; Heyne, Nutt. Pl. ed. 1, 3
(1917) 310; ibid. ed. 2 (1927) 1126; LEcoMTE, Bois de
l'Indochine (1926) 113; SLoot. Bull. Jard. Bot. Btzg
III, 8 (1927) 373, f. 2, p.p.; BuRK. Dict. (1935) 1664;
ASHTON, Gard. Bull. Sing. 31 (1978) 26. — Shorea
stellata Dyer, FI. Br. Ind. 1 (1874) 304; KING, J. R. As.
Soc. Beng. 62, 2 (1893) 120.— Shorea cinerea
FISCHER, Kew Bull. (1926) 460.— P. lucida (non
(MiQ.) KuRZ) SLooTt. Kew Bull. (1927) 372. p.p.: SYM.
Mal For. Rec. 16 (1943) 102, f. 59B, 60, 62.— P.
poilanei TARDIEU, Not. Syst. 10, 3 (1942) 136.
Medium-sized or large tree. Panicles, floral ovary
and sepals outside densely greyish puberulent, ovary
and sepals glabrescent in fruit; leaf undersurface +
silvery lepidote; parts otherwise glabrous. Twig 1-2
mm @, slender, ribbed along the leaf traces, glabres-
cent; stipule scars short, pale, horizontal. Buds to 2 by
2 mm, small, ellipsoid, obtuse; stipule to 1.5 cm long,
linear, fugaceous. Leaf 6-16 by 2.3—7 cm, lanceolate,
thinly coriaceous; base cuneate; acumen to | cm long,
tapering; nerves 8-12 pairs, very slender, elevated
beneath, obscure above, steeply ascending, frequently
sinuate and obscurely branching; tertiary nerves very
slender, evident but hardly elevated, densely scalari-
form; midrib prominent beneath, shallowly depressed

and obscure above; petiole (10-)12-30 mm long,
slender, smooth. Panicle to 15 cm long. terminal or
axillary, slender, lax, many-flowered, doubly
branched; branchlets to 2 cm long, bearing to 7 secund
flowers. Flower buds to 2 by 2 mm, small, subglobose:
sepals ovate, subequal, acute: corolla white; stamens
subequal, yellow; filaments short, broad, compressed:
anthers linear-oblong; appendages short, slightly
exceeding anthers; ovary subglobose, densely pubes-
cent; style columnar, c. 15 xlength of ovary. Fruit
pedicel to 3 mm long, to 2 mm g, expanded into
receptacle: calyx lobes to 11 by 1.8 cm, subequal,
spatulate, obtuse, ascending, c. 3 mm broad above the
narrow thickened base; nut to 2 by 1.5 cm, ellipsoid.
crowned by an up to 4 mm long filiform tapering style
remnant.

Distr. Southern Burma, Thailand, Indochina, and
in Malesia: Malaya (Trengganu and Perak north-
wards).

Ecol. Frequent in lowland and hill evergreen
dipterocarp forests in seasonal areas, especially in
valleys, to 650 m.

Vern. Gérutu gérutu.

Note. Collections cited from Sumatra belong cor-
rectly to P. lucida.

7. Parashorea malaanonan (BLCO) MerR. Sp. Blanc.
(1918) 271; En. Philip. 3 (1923) 100; Reyes, Philip. J.
Se. 22 (1923) 330; SLoort. Bull. Jard. Bot. Btzg III, 8
(1927) 375; in Merr. Pl. Elm. Born. (1929) 202: Sym.
Gard. Bull. S. S. 9 (1938) 334, pl. 21; BRowng, For.
Trees Sarawak & Brunei (1955) 128: ASHTON, Man.
Dipt. Brun. (1964) 84, f. 11; ibid. Suppl. (1968) 38:
MEuUER & Woop, Sabah For. Rec. 5 (1964)
192. — Mocanera malaanonan Btico, F1. Filip. (1837)
858. — Dipterocarpus malaanonan BLico, FI. Filip. ed.
2 (1845) 312; ibid. ed. 3, 2 (1878) 214; DC. Prod. 16, 2
(1868) 614.— Shorea malaanonan BL. Mus. Bot.
Lugd.-Bat. 2 (1852) 34; WALP. Ann. 4(1857) 338; DC.
Prod. 16, 2 (1868) 631; F.-VILL. Nov. App. (1880) 21;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 103; MERR. &
Ro tre, Philip. J. Sc. 3 (1908) Bot. 115; WuHiTFoRD,
Bull. Bur. For. Philip. 10 (1911) 64; Foxw. Philip. J.
Sc. 6 (1911) Bot. 270; ibid. 13 (1918) Bot. 189. — P.
plicata BRANDIS, J. Linn. Soc. Bot. 31 (1895) 104;
Merr. & ROLFE, Philip. J. Sc. 3 (1908) Bot. 114;
Foxw. Philip. J. Sc. 6 (1911) Bot. 280; ibid. 13 (1918)
Bot. 194; ibid. 67 (1938) 317; WHITFORD, Philip. J. Sc.
6 (1911) Bot. 64. — Fig. 54, 57, 58.

Young twig, leaf bud, stipule, petiole and panicle
shortly sparsely pale brown tomentose, glabrescent on
petiole. Twig to 1.5 mm g apically, terete, becoming
glabrous, dotted with minute pale round lenticels;
stipule scars thin, pale, amplexicaul. Bud to 6 by 2mm,
lanceolate to subfalcate, acute. Leaves 9-15 by 3.5—7.5
cm, elliptic to ovate, greyish beneath, glabrous; base
obtuse to broadly cuneate, subequal; acumen to | cm
long, acute; margin undulate; nerves 9-14 pairs,
straight, with + prominent interneural plicate folds,
curved at the margin, prominent beneath; tertiary
nerves slender, scalariform, sinuate, at 90°; midrib

384 FLORA MALESIANA [ser. I, vol#92

U. W/5

Fig. 57. Parashorea malaanonan (BLCO) Merk. a. Older sterile twig, b. younger sterile twig, c. fruit, d. nut, all x 4
(a A 281, 6 A 335, c-d A 3602).

prominent beneath, slightly raised above; petiole
1.2-2 cm long, slightly geniculate. Panicle to 18 cm
long, terminal or axillary, terete; doubly branched,
branchlets to 4 cm long, bearing to 3 flowers;
bracteoles to 12 by 6 mm, oblong, obtuse, densely
pubescent outside, glabrous within. Bud to 14 by 8

mm, large. Flowers cream. Calyx densely brown-buff
pubescent outside, more sparsely so near the apex
inside; 3 outer lobes narrowly deltoid acute, 2 inner
lobes somewhat shorter, narrower. Petals hastate,
acute, densely tomentose outside, glabrous within.
Stamens subequal; filaments short, broad at base,

1982]

DIPTEROCARPACEAE (Ashton)

385

Fig. 58. Parashorea malaanonan (BLCO) Mer. a. Flowering twig, b. apex of twig with terminal bud, both xs
(NooTEBOOM 1135).

tapering; anthers linear, about twice length of fila-
ment; inner pollen sacs shorter, narrower, than outer;
appendage to connective longer than anther, reaching
to style apex, stout at base, tapering and slender in the
apical half, erect. Ovary ovoid, shortly pubescent;
stylopodium slender, tapering into style, shortly
pubescent; style filiform, glabrous, c. 4 times length of
ovary. Fruit pedicel to 4 mm long, broadening into
base of fruit. Fruit calyx puberulent to glabrescent,
more densely pubescent at base; 3 longer lobes to 16
by 1.7 cm, narrowly spatulate, subacute, to c. 3 mm
broad above the to 5 by 4 mm deltoid shallowly
saccate thickened base; shorter lobes to 10 by 0.7 cm,
linear, similar at base or somewhat narrower. Nut to
1.7 by 1.4 cm, ellipsoid, obtuse, verrucose; style
remnant to 6 mm long, linear, tomentose.

Distr. Malesia: Borneo (Brunei to Sabah, S. E. to
Berau and Nunukan), Philippines.

Ecol. Abundant in lowlands of Philippines and E.
Sabah, rare in W. Sabah, Brunei and N.E. Sarawak,
recorded to 1300 m.

Uses. The most important plywood, decking and
light construction timber exported from its region.

Vern. Urat mata, urat mata daun lichin (Borneo),
bagtican, lauan (Philippines generally), apnit (Polillo,
Tayabas, Catanduanes, Camarines, Albay, Sorso-
gon), bayucan (Nueva Ecija, Laguna), binaliuan
(Bulacan), danlig (Tayabas, Masbate, Occidental
Negros), malaanonang (Laguna), mayatas (Polillo),
tacuban (Camarines).

8. Parashorea tomentella (Sym.) MEER, Acta Bot.
Neerl. 12 (1963) 320; MEWERR & Woop, Sabah For.
Rec. 5 (1964) 199, f. 25.— P. malaanonan var.
tomentella SyM. Gard. Bull. S. S. 9 (1938) 338, pl.
22. — P. sp. THOMAS, Mal. For. 4 (1935) 131, p.p.;
Sym. ex Descu, Mal. For. Rec. 12 (1936) 33. — Fig.
59.

Large tree. Young twigs, buds, leaf undersurface,
petioles, panicles, bracts outside, calyx parts of co-
rolla exposed in bud and ovary densely evenly +
persistently pale tawny pubescent. Twig c. 3 mm @
apically, much branched, ribbed, becoming terete,
smooth, dark brown; stipule scars slender, horizontal,
amplexicaul. Bud to 10 by 4 mm, lanceolate, acute;
stipule to 16 by 6 mm, narrowly lanceolate. Leaves

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 59. Stem-base of Parashorea tomentella (SyM.) MEIER. Sabah, Tawau, Kalabakan; Gant behind buttress
(Photogr. G.H.S. Woop, Nov. 1955; SAN 17266).

10-20 by 5-10 cm, subcoriaceous, subpersistently
plicate, dull but not silvery beneath; margin fre-
quently narrowly subrevolute; base obtuse to subcor-
date, subequal (peltate in young trees and saplings);
apex subacute or to | cm long, broadly acuminate;
nerves 11-13 pairs, ascending, prominent beneath,
somewhat arched; tertiaries densely scalariform, evi-
dent and slightly elevated beneath; midrib stout and
prominent beneath, elevated above; petiole 15-25 mm
long, c. 3mm @, stout, hardly geniculate. Panicle to 13
cm long, singly or doubly branched, the branchlets
bearing to 3 flowers; bracts to 10 by 4 mm, lanceolate.
Flower buds to 15 by 10 mm, broadly lanceolate,
flowers very large; sepals narrowly deltoid; stamens

somewhat shorter than style; filaments compressed,
tapering; anthers oblong-linear; appendages some-
what longer than anthers, stoutly acicular; ovary
small, ovoid; style columnar, c. 5 times length of
ovary, stout, puberulent but for the apical 4. Fruit
pedicel c. 3 mm long; sepals aliform, 3 longer lobes to
20 by 2.3 cm, spatulate, obtuse; 2 shorter lobes to 10
by 0.8 cm, lorate, acute; nut to 2 cm @, subglobose,
verrucose, apiculate.

Distr. Malesia: N. E. Borneo (Sangkulirang to
Sandakan).

Ecol. Mixed Dipterocarp forest on flat and undu-
lating land below 200 m. Common.

Vern. Urat mata beludu.

1982]

Note. Sterile and fruiting collections from the
Philippines resembling this species were cited by
Foxwortny under P. warburgii BRANDIS, regarded
here as species dubium. Without flowering collection
their identity remains obscure.

9. Parashorea lucida (MiQ.) Kurz, J. R. As. Soc.
Beng. Sc. 39, 2 (1870) 66; BurcK, Ann. Jard. Bot. Btzg
6 (1887) 221; BRANbDis, J. Linn. Soc. Bot. 31 (1895)
104; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 310; ed. 2 (1927)
1126; SLoor. Bull. Jard. Bot. Btzg III, 8 (1927) 372,
p.p., ASHTON, Man. Dipt. Brun. Suppl. (1968) 38, in
nota sub P. smythiesii; Gard. Bull. Sing. 31 (1978)
27. — Shorea lucida Mig. Sum. (1862) 487, 191; DC.
Prod. 16, 2 (1868) 631; Warp. Ann. 7 (1868)
379. — Shorea subpeltata MiqQ. Sum. (1862) 488, 191;
DC. Prod. 16, 2 (1868) 632; WALP. Ann. 7 (1868) 379;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 219; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 103.— P. stellata (non
KurZ) SLoot. Bull. Jard. Bot. Btzg III, 8 (1927) 373,
P-p-

Large tree. Young twigs, buds, petioles, panicles,
bracts outside, calyx, parts of corolla exposed in bud
and ovary densely evenly + persistently buff pubes-
cent; nerves beneath sparsely + caducously so. Twig
c. 2 mm @g apically, + ribbed, becoming smooth,
blackish, terete: stipule scars short, pale, horizontal.
Bud to 4 by 2 mm, ovoid-apiculate; stipules to 5 by 3
mm, narrowly ovate, acute. Leaves 6-14 by 2.5-6.5
cm, ovate-lanceolate to elliptic, thinly coriaceous, +
distinctly persistently plicate, dull and + silvery
stellate beneath; base broadly cuneate to occasionally
cordate, subequal; acumen to | cm long; nerves 9-12
pairs, slender but prominent beneath, rather straight,
dense; tertiary nerves densely scalariform, very
slender but distinct beneath; midrib prominent
beneath, shallowly furrowed to elevated above; petiole
10-20 mm long, hardly geniculate. Panicle to 12 cm
long, slender, terminal or axillary, singly branched,
the branchlets bearing to 6 + secund flowers; bracts to
4 by 3 mm, elliptic. Flower bud to 7 by 4 mm,
lanceolate; sepals narrowly deltoid; stamens shorter
than style; filaments broad, compressed, tapering,
short; anthers lorate-oblong; appendages acicular,
prominent and longer than anthers; ovary small,
ovoid, surmounted by a distinct tapering somewhat
longer stylopodium; style twice as long as ovary and
stylopodium, filiform, somewhat expanding distally.
Fruit pedicel to 3 mm long, broadening into recepta-
cle; sepals aliform, unequal; 3 longer lobes to 8 by 1.7
cm, spatulate, obtuse; 2 shorter lobes to 7.5 by 0.8 cm,
narrow; nut to 2.5 cm g@, subglobose or ovoid,
apiculate, densely verruculose.

Distr. Malesia: Sumatra (central E. & W. of
Barisan range), Borneo (Central Kalimantan, Central
and N. E. Sarawak).

Ecol. Mixed Dipterocarp forest on hills, to 700 m.

Vern. Itjap, katoeka, damar laut, d. surantih, d. tyirik
ayam, méranti hitam (Sumatra), méruyun (Iban).

10. Parashorea smythiesii WYATT-SMITH ex ASHTON,

DIPTEROCARPACEAE (Ashton)

387

Gard. Bull. Sing. 19 (1926) 266, pl. 7; ASHTON, Man.
Dipt. Brun. (1964) 86, f. 11, pl. 23; ibid. Suppl. (1968)
38, p.p.; MEUER & Woop, Sabah For. Rec. 5 (1964)
WO Tet 24

Young twig, leaf bud, stipule, petiole and panicle
sparsely shortly pale yellow-brown tomentose, cadu-
cous on twigs and petioles. Twig 1 mm g apically,
terete, smooth, glabrous, dotted with minute pale
round lenticels; stipule scars short, slender. Bud to 3
by | mm, narrowly lanceolate, acute. Stipule to 3.5
mm long, narrowly hastate, acute, fugaceous. Leaves
6-9 by 3-4.5 cm, elliptic to ovate, glabrous; base
broadly cuneate; acumen to 1.5 cm long, narrow to
caudate; nerves 8-10 pairs, slender, raised beneath,
curved, well spaced, at 50°—60°; tertiary nerves scalari-
form to subreticulate, widely spaced at c. 90° to
nerves; midrib slender, prominent beneath, depressed
above; petiole 1—-1.8 cm long, geniculate, pale. Panicle
to 14 cm long, terminal or axillary, singly branched,
straight, pendent; bracts and bracteoles unknown.
Flower bud to 4.5 by 3 mm, ellipsoid, obtuse. Calyx
densely shortly grey-brown tomentose outside, gla-
brous within; lobes narrowly deltoid, subequal, suba-
cute, not adpressed to corolla in bud. Petals elliptic,
obtuse, shortly tomentose on parts exposed in bud,
cream. /nner 5 stamens slightly longer than the others,
reaching ? length of style; filaments broad at base,
tapering; anther narrowly oblong; appendage to
connective short, slightly protruding above anther;
ovary subglobose, densely tomentose; style c. 3 times
length of ovary, filiform, shortly pubescent in the
basal 3, otherwise glabrous. Fruit calyx glabrescent,
puberulent at base, becoming pushed apart by the
ripening nut; 3 longer lobes to 8.5 by 1.7 cm, broadly
spatulate, obtuse, to 3 mm broad above the to 7 mm
long unexpanded slightly thickened base; 2 shorter
lobes to 7.5 cm long, often only slightly smaller than
longer lobes, subequal. Nut to 1.5 by 1.3 cm, ellipsoid,
obtuse, verrucose with pale lenticels, buff tomentose;
style remnant to | mm long, short.

Distr. Malesia: Borneo (Rejang hinterland east-
wards and northwards to Sabah and Tidung).

Ecol. Scattered in Mixed Dipterocarp forest on
clay rich soils, on moist lower slopes, and hillsides to
1000 m.

Vern. Méruyun (Iban), urat mata batu, urat mata
daun puteh.

Doubtful

Parashorea warburgii BRANDIS, J. Linn. Soc. Bot. 31
(1895) 105; Foxw. in Elmer, Leafl. Philip. Bot. 6
(1913) 1954; Philip. J. Sc. 13 (1918) Bot. 194; ibid. 67
(1938) 318; Merr. En. Philip. (1923) 100.

Described from a collection of WARBURG from
Mindanao, Philippines, consisting of a single fruit
undistinguishable from those of P. malaanonan. Fox-
WORTHY associated the name with pubescent-leaved
forms from the Philippines otherwise resembling P.
malaanonan though these may represent P. tomentella
of Borneo (g.v.).

388 FLORA MALESIANA [ser. I, vol. 9?

Fig. 60. Neobalanocarpus heimii (KING) ASHTON. a. Habit, b. immature fruits, both x 5 (a KEP 69424, b KEP
69415).

8. NEOBALANOCARPUS, gen. nov.

ASHTON, Gard. Bull. Sing. 31 (1978) 27. — Balanocarpus (non BEDD.) KING, p.p.,
BRANDIS, p.p., RIDL., p.p., HEYNE, p.p., FOXW., p.p., SYM. (1934) 27, p.p., (1943)
147. — Fig. 60, 61.

Calyx in fructu ut in Balanocarpus BEDD. (Hopea Roxs. pro parte), floribus
maximus antheris lineare-oblongis appendiculatis facile differt.

Large tree with tall stout buttresses. Leaves penninerved, unequal-based, with
scalariform tertiary nerves. Inflorescence paniculate. Flowers medium-sized,
secund; stamens 15, glabrous, with slender tapering filaments and linear-oblong
anthers bearing rudimentary appendages; ovary ovoid, with long slender style.
Fruit sepals short, subequal; pericarp splitting into 3 equal valves at germination;
cotyledons very unequal; first 4-5 seedling leaves in a whorl.

Distr. Monotypic. Peninsular Thailand (Pattani) and Malesia: Malaya.

Note. The possession of short equal fruit sepals, in the presence of a unique androecium structure deprives
this single species from the sole character by which it could be allotted to the genera Shorea or Hopea,
underlining the close affinity between these genera. The general appearance of tree and foliage and especially the
inflorescence, fruit embryo and mode of germination, suggests that this unsatisfactory genus bears very close
affinity with Hopea sect. subsect. Hopea; the linear anthers are approached by those of H. plagata (BLCO) VIDAL,
though there the appendage is acicular and prominent.

1. Neobalanocarpus heimii (KING) ASHTON, Gard. Bull. Str. & F.M.S. 7 (1908) 386; Trees and Timbers
Bull. Sing. 31 (1978) 27. — Balanocarpus heimiiK1NG, (1911) 3, fig.; BURK. J. Str. Br. R. As. Soc. 81 (1920) 3;
J. R. As. Soc. Beng. Sc. 62, 2 (1893) 133; BRANDIS, J. J. Mal. Br. R. As. Soc. 1 (1923) 218; Dict. (1935) 204;
Linn. Soc. Bot. 31 (1895) 110; BURN-Murpocu, Agr. — RIDL. Fl. Mal. Pen. 1 (1922) 247; HEYNE, Nutt. PI. ed.

1982]

DIPTEROCARPACEAE (Ashton)

389

Fig. 61. Neobalanocarpus heimii (KING) ASHTON. A. Bud, B/. outer sepal, B2. inner sepal, C. petal, D. stamens
from outside, E. pistil, all x 10 (KEP 94605).

2 (1927) 1128; Foxw. Mal. For. Rec. 1 (1921) 64; ibid.
3 (1927) 53; ibid. 8 (1930) 10; ibid. 10 (1932) 149; J.
Mal. Br. R. As. Soc. 5 (1927) 399: STRUGNELL, ibid. 9
(1931) 24; Sym. Gard. Bull. S. S. 8 (1934) 27; Mal. For.
Rec. 16 (1943) 147, f. 80-82; CoRNER, Ways. Trees
(1940) 210; SMITINAND, Thai For. Bull. (Bot.) 12
(1980) 23. — Balanocarpus wrayi KiNG, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 134.— Balanocarpus acu-
minatus (non BRANDIS) HEIM, Ass. Fr. Adv. Sc.
Besangon 1893 (1894) 560, t. 4. — Pierrea penangiana
HEIM ex BRANDIS, J. Linn. Soc. Bot. 31 (1895) 110,
nomen in syn. — Fig. 60, 61.

Very large flaky barked buttressed tree. Twigs,
midrib above, petioles and sepals outside caducous
puberulent, leaf buds and panicles persistently so,
parts of petals exposed in bud densely buff pubescent.
Twig c. 2 mm @ apically, ribbed, becoming smooth,
dotted with minute pale lenticels; stipule scars linear,
horizontal. Buds small, ovoid; stipules to 12 mm long,
narrowly lorate, spreading, fugaceous. Leaves 7-17 by
2.3-5 cm, lanceolate-falcate, coriaceous; base un-
equal, cuneate to obtuse; acumen to 1.5 cm long,
tapering; nerves 9-12 pairs, ascending, arched, promi-
nent beneath, narrowly depressed above, the basal
nerve on the broader (adaxial) side frequently with
prominent lateral branchlets; tertiary nerves subreti-
culate, slender but prominently elevated beneath,
obscure above; petiole S~10 mm long, short. Panicle to

9 cm long, terminal or axillary, singly branched;
branchlets to 2.5 cm long, bearing to 7 flowers. Flower
buds to 4 by 3 mm, ovoid; sepals broadly ovate, acute,
subequal, corolla pale greenish yellow; stamens 15,
subequal; filaments tapering, long; anthers linear-
oblong, appendages rudimentary; ovary ovoid, gla-
brous, surmounted by a filiform style twice its length.
Fruit pedicel to 2 mm long, to 3 mm @, stout, inserted
ona + impressed receptacle base; calyx lobes to 20 by
18 mm, subequal, ovate, thickened, saccate, adpressed
to the base of the nut; nut to 5.5 by 2.5 cm,
oblanceolate, cylindrical, shortly apiculate, lustrous.

Distr. Peninsular Thailand (Pattani) and in Male-
sia: Malaya.

Ecol. Widespread in Mixed Dipterocarp forest
below 1000 m, especially on undulating land on well
drained friable soils.

Uses. The heavy durable timber used to be the best
known in Malaya and the ‘standard by which other
timbers are judged’ (SYMINGTON). Now largely re-
placed owing to the introduction of modern preserva-
tive techniques.

Vern. Chéngal, chéngai, c. témbaga, c. batu, c.
bunga, c. dédap, c. mas, c. kémunting, c. labu, c. sabut,
c. siput, c. tembaga, c. témpurang.

Note. The young leaves are bronze. Abnormal
meiosis was observed by JONG & LETHBRIDGE (Notes
R. Bot. Gard. Edinb. 27, 1967, 175).

390 FLORA MALESIANA [ser. I, vol. 9?

Fig. 62. Hopea beccariana BURCK. a. Flowering branch, x 2/3. — H. dryobalanoides Mia. b. Fruiting branch, c.
fruit, d. nut, all x 2/3 (a SAN 30641, b-d BRUN 3179).

1982] DIPTEROCARPACEAE (Ashton) 39]

9. HOPEA

Roxs. Pl. Corom. 3 (1811) 7, mom. gen. cons., non L. 1767; ENDL. Gen. Pl. (1840)
1014, ‘Hoppea’; DC. Prod. 16, 2 (1868) 632; Dykr, Fl. Br. Ind. | (1874) 308;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 235; HEI, Rech. Dipt. (1892) 59, incl. sect.
Hancea (PIERRE) HEM, /.c. 62; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 53; Foxw.
Philip. J. Sc. 67 (1938) 273; Sym. Mal. For. Rec. 16 (1943) 108, f. 67 (maps);
ASHTON, Gard. Bull. Sing. 20 (1963) 254; Man. Dipt. Brun. (1964) 89; ibid. Suppl.
(1968) 37; MEWER & Woop, Sabah For. Rec. 5 (1964) 203; GUTIERREZ, Act.
Manill. 4A, 2 (1968) 3; ASHTON, Blumea 20 (1972) 359; Gard. Bull. Sing. 31 (1978)
28; SMITINAND, Thai For. Bull. (Bot.) 12 (1980) 42. — Neisandra RaFin. Sylv.
Tellur. (1838) 163. — Petalandra Hassk. Cat. Hort. Bog. (1858) 104. — Balano-
carpus BEDD. For. Man. Bot. (1873) 236 bis; HEIM, Rech. Dipt. (1892) 77, pro sect.
Sphaerocarpae HEIM, /.c.; SYM. Mal. For. Rec. 16 (1943) 147, p.p.— Hancea
PIERRE, For. Fl. Coch. 4(1891) sub t. 244. — Pierrea HEIM, Bull. Mens. Soc. Linn.
Paris 2 (1891) 958, nom. gen. cons., non HANCE, 1877. — Dioticarpus DUNN, Kew
Bull. (1920) 337. — Pierreocarpus RIDL. ex SYM. Gard. Bull. S. S. 8 (1934) 30,
nomen in syn. — Fig. 12-13, 62-76.

Small or medium-sized, occasionally large, trees; bole usually tapering,
frequently branching low; buttresses usually thin, sometimes thick; stilt roots and
flying buttresses sometimes present; crown, in small species, persistently lanceo-
late, monopodial, the branches + horizontal and pendent; becoming densely
evenly hemispherical in large trees, with many small straight branches radiating
from the bole apex. Bark surface at first smooth, chocolate and grey mottled,
hoop-marked; remaining so or becoming cracked and flaked, or fissured. Parts
with or without indumentum of broad or narrow lobed hairs. Twigs slender,
usually branching horizontally; stipule scars small, inconspicuous. Stipules
linear, fugaceous (subpersistent in saplings). Leaves small or medium-sized, or
narrowly oblong, large; nerves (see Fig. 64a) either scalariform, with scalariform
tertiaries, or ‘dryobalanoid’ with + indistinct nerves, and with many equally
prominent, but more or less shorter, secondaries, and indistinct reticulate
tertiaries, superficially resembling those of Dryobalanops; or intermediate
between these two types: ‘subdryobalanoid’, with more prominent reticulate or
partially scalariform tertiaries, more prominent nerves, and fewer, shorter,
intermediates. Petiole never geniculate. Inflorescence paniculate, slender, ter-
minal or axillary. Flower buds small, ovoid or rarely globose. Sepals imbricate; 2
outer ovate, + obtuse, thickened; 3 inner suborbicular, frequently mucronate,
thin at the margins. Petals oblong, connate at base and falling in a rosette.
Stamens 10, 15 or up to 38 (H. plagata), in 1-3 verticils or irregular, falling with
the petals; filaments broad and compressed at base, tapering medially and
filiform below the anthers, anthers subglobose, tapering apically, latrorse; with 4
pollen sacs, the outer pair somewhat the larger; appendage to connective usually
at least twice as long as anther, slender, glabrous or minutely glandular
tuberculate. Ovary glabrous or tomentose, ovoid and with indistinct stylopodium
marked by a ring of hairs at the apex of the ovary, or with a distinct stylopodium
and hence pyriform, hour-glass-shaped, or cylindrical; style long or short,

392 FLORA MALESIANA [ser. I, vol. 9?

e
Fig. 63. Flower details in Hopea sect. Dryobalanoides Mia. All x 10. Sepals drawn from inside. — H. myrtifolia
Mia. A. Bud, A/. outer sepal, A2. inner sepal, 43. stamens from outside, 44. pistil. — H. ferruginea PaRuUs. B/.
Outer sepal, B2. inner sepal, B3. stamens from inside, B4. pistil. — H. beccariana BuRCK. C1. Outer sepal, C2.

inner sepal, C3. stamens from outside, C4. pistil. — H. dyeri HEM. D3. Stamens, from outside, D4. pistil(A KEP
99627, B A 4342, C KEP 76753, D BECCARI 2504).

[Np Se A2

glabrous; stigma minute (except H. ferrea). Fruit relatively small: 2 outer fruit
calyx lobes prolonged, spatulate; 3 inner lobes short, or 5 short, subequal; lobes
thickened and saccate at base. Nut ovoid, usually glabrous, with a distinct apical
stylopodium if present in the flower. Pericarp splitting at germination into 3 valves
(H. pachycarpa) or irregularly; cotyledons photosynthetic, subequal; first pair of
leaves opposite, followed by spiral leaves or an initial whorl of 3; branching of
sapling mainly at initiation of each period of growth by leader, hence appearing
pagoda-like.

Distr. About 102 spp. in Ceylon, Andamans, South and East India, Burma, Thailand, Indochina, continental
S. China (Yunnan, K wangsi, S. Kwantung), Hainan, and 84 spp. throughout Malesia except the Lesser Sunda
Islands. Fig. 65.

Ecol. Main canopy or understorey, rarely emergent, trees of lowland evergreen forests; and also
semi-evergreen forests where there are more species, many of them local endemics, than any other dipterocarp
genus. Several are semi-gregarious, several riparian.

Uses. Though some of the larger species provide a heavy durable construction timber few are common
enough to be important economically. Several species in sect. subsect. Dryobalanoides produce a clear crystalline
resin, damar mata kuching, that was formerly a valuable article of commerce.

Notes. Though apparently natural groupings whose typical members are at once recognisable, the

1982] DIPTEROCARPACEAE (Ashton) 393

subsections and even sections of this genus are ill-defined in that several species share certain characters from
more than one section, in marked contrast with the infrageneric groupings of the closely allied genus Shorea. See
for a discussion about the subdivision of the genus accepted here ASHTON, Gard. Bull. Sing. 20 (1963) 254.

Pollination in those examined appear to be effected by thrips. Triploidy is known in both emergent (H.
odorata), main canopy (H. beccariana) and understorey species (H. subalata). Either or both these factors may
explain the high degree of local endemism in the understorey subsections Sphaerocarpae and Pierrea, and the
curious local diversification in New Guinea.

SUBDIVISION OF HOPEA IN MALESIA

1. Leaf nervation truly dryobalanoid. Spp. 1-26. . . . Sect. Dryobalanoides subsect. Dryobalanoides
1. Not so.
2. Ovary and stylopodium not constricted between.
3. Flowers remote on raceme; bracts subpersistent. Spp. 27-39
Sect. Dryobalanoides subsect. Sphaerocarpae
a) Flowers dense on raceme; bracts fugaceous. Spp. 40-70 . . . . Sect. Hopea subsect. Hopea
2. Ovary and stylopodium hour-glass-shaped, distinctly constricted. Spp. 71-84
Sect. Hopea subsect. Pierrea

KEY TO THE SPECIES

1. Leaf nervation truly dryobalanoid. Fig. 67. Spp. 1-26.
1. Sect. Dryobalanoides 1a. subsect. Dryobalanoides
2. Ovary without distinct stylopodium.
3. Midrib obscurely depressed above.
4. Ovary glabrous.

eM VIPWPUDESCED tite Wa esii ier sw KL pe. oksweulce « owe) al eegve qerlese SIRE pubescens
Se WiostolADTOUS eyes -ens ica cewek br st ceeictitad pu oe Selec 0c. eZ eRfoxworthyi
4. Ovary pubescent. . Soe 2 8 68 « « © » « «» « «» 3: H quisumbingiana

3. Midrib evident, + elevated above.
6. Shorter fruit sepals exceeding nut and + enclosing it.
7. Leaf margin revolute.

8. Leaf S—9 by 2-7 cm, thinly coriaceous; stamens 15-18. . . . . . . +. +. #.4.H.cernua

8. Leaf 11-16 by 6-10 cm, thickly coriaceous; stamens 15. . . . . . +. +. 5. H. coriacea
7. Leaf margin applanate.

9. Fruit pedicel to 7 mm long, calyx lobestuberculate . . . . . . +. . 6. H. longirostrata

9. Fruit pedicel to 2 mm long; calyx lobes not as above.
10. Main nerves with axillary domatia, the basal pair not longer than the rest; fruit pendent
7. H. sulcata
10. Main nerves without axillary domatia, the basal pair very long and continuing along 2/3 of the
margin; fruit erect on panicle .. Sik Wool eno ere Oo) aes EL sfluvialis
6. Shorter fruit sepals shorter than nut and not concealing it
11. Nerves very many, indistinct.

I2eeBeatibaseicuneates 5. 2] 3 = = 5 «© 5 = paeeeeee eedp ies eo eH menserawan
IDS reat basevobtuse..o-s.). 28 a oe ee ee See 5 cee ed Oe Elemicrantha
11. Nerves less than 13 pairs.

13. Nerves c. 6 pairs, fruit calyx lobes short, subequal . . . . . . 11. H. Kerangasensis
13. Nerves more than 8 pairs, fruit calyx lobes unequal, 2 long and aliform.

L455Nut.tolSmmilongy cylindrical S| a) sete en 2 ee escue)

14. Nut shorter than 11 mm, ovoid.
15. Fruit sepals to 6.5 by 1.5 cm; midrib acutely elevated on both surfaces, drying black
13. H. dryobalanoides
15. Fruit sepals to 4 by 0.8 cm; midrib not acute below, not drying black.
16. Leaf base unequal; twig apices glabrescent.

17. Flower bud to 6 by 3 mm, relatively large; panicle glabrous . . . . .414. H. malibato
17. Flower bud less than 3 mm; panicle + puberulent.
18. Leaves small, petiole less than 13 mm long; panicle less than 2cmlong . 15. H. johorensis
18. Leaves broadly ovate, petiole 12-17 mm long; panicleto6cmlong . . . 16. H. latifolia

16. Leaf base equal; twig apices persistently pubescent . . . . . +. +. 17. Hz. ferruginea

394 FLORA MALESIANA (ser. 1, wore?

Fig. 64. Three venation types in Hopea. — a. Scalariform in H. acuminata MeERR., b.dryobalanoid in H.
beccariana BURCK, c. subdryobalanoid in H. subalata SYM.

2. Ovary with distinct stylopodium.
19. Midrib above obscure, depressed.

PieNainmnervesivery.many.¢s [8ipairs) 2) 5) Ree ee tee ee ae 18. H. pierrei
20. Main nerves at most 10 pairs.
Die VOunespants|sckiccOUS, \-p 62... 2 ey ee he ta 19. H. inexpectata
Dibaviounpepants glabrous). 500. 9e ie, ne ee eel rs 20. H. griffithii
19. Midrib evident above.
DOL AS GATTI OMEID bite. BEL TRG! WORK HR, Uh eT ELITE Aithig 95) 0) OT A treubii

22. Stamens 15.
23. Ovary and stylopodium pyriform; nerves at most 12 pairs.

Aw Petioleyc:Simmeong, leaf broadlysovatei =.) = 9.92 a8) 1G a 22. H. beccariana

24. Petiole shorter than 10 mm, leaf narrowly ovate-lanceolate. . . . . . . . 23. H. dyeri
23. Ovary and stylopodium cylindrical; nerves at least 13 pairs, 13-16 pairs.

DS wies andipetiolespubescemt hs) at aD eR Peis 1. es 24. H. myrtifolia

25. Twigs and petioles glabrous.
2 Guleeaves applamaten ee) Py Bee ee eg 25. H. pedicellata
26. Leaves prominently revolute atleast at base. . . . - . - + = - 26. H. altocollina
1. Leaf nervation not truly dryobalanoid. Compare fig. 64a & c.
27. Ovary and stylopodium not constricted between.
28. Flowers remote on raceme; bracts subpersistent, corolla dark coloured. Spp. 27-39.
1. Sect. Dryobalanoides 1b. subsect. Sphaerocarpae
29. Leaf nervation scalariform. Fig. 64a.

BOs bruiticalyxdlobesishorts subequal rey) res! "nF ets nrc iee.” Oc Bie 27. H. aequalis
30. Fruit calyx lobes unequal: 2 long, aliform.
BilkeiNerves: W113) pairs: leaves‘paleibencaths "= = a 28. H. rudiformis
31. Nerves (13—)16-19 pairs; leaves not pale beneath.
a2 alwolongemirait sepals toli2icm! longs 9) Ga a ae) Eee 29. H. nervosa
32. Two longer fruit sepals not exceeding 8cm . . . . .- .- + = - 30. H. sublanceolata

29. Leaf nervation subdryobalanoid. Fig. 64a.

1982] DIPTEROCARPACEAE (Ashton)

33. Lamina base obtuse.
34. Fruit sepals unequal, 2 aliform .
34. Fruit sepals short, subequal.
35. Petiole 3-6 mm long
35. Petiole 7-10 mm long
33. Lamina base cuneate.
36. Fruit sepals unequal.
37. A single fruit sepal exceeding the nut, shortly lorate.
37. Two fruit sepals aliform, spatulate.
38. Base of fruit sepals auriculate .
38. Base of fruit sepals not auriculate
36. Fruit sepals all shorter than the nut, subequal.
39. Stamens 10 .
39. Stamens 15.
40. Ovary and stylopodium puberulent .
40. Ovary and stylopodium glabrous :
28. Flowers dense on raceme; bracts fugaceous; corolla pale. ‘Sop. 40-69.
2. Sect. Hopea 2a. subsect. Hopea

395

31. H. nigra

32. H. sphaerocarpa

33. H. mesuoides

. 34. H. subalata

35. H. auriculata
. 36. H. montana

37. H. vaccinifolia

38. H. bracteata

. 39. H. brachyptera

41. Nerves united into a prominent continuous intramarginal nerve midway between midrib and margin

. Nerves without intramarginal nerve.
a5) Fruit sepals suborbicular; stamens 10 3
42. Fruit sepals spatulate or, if suborbicular, then stamens ils
43. Nerves on average at least 13 pairs.
44. Leaf base obtuse, + equal.

40. H. celtidifolia

41. H. dasyrrhachis

45. Leaf at least 10 by 3.5 cm; petiole at least 11 mm long; shorter fruit sepals shorter than nut

42. H. similis

45. Leaf at most 11 by 4 cm; petiole at most 8 mm eee 1 or the 3 shorter fruit sepals frequently lorate

and exceeding nut .

43. H. forbesii

44. Leaf base prominently unequal, generally cordate on one side: or - subequal, subcordate on both

sides.
46. Leaf glaucous beneath; stylopodium truncated, pubescent.
46. Leaf not glaucous; stylopodium tapering, glabrous.
47. Fruit sepals short, subequal . Stale
47. Fruit sepals unequal, 2 aliform.
48. Base of leaf subequal, subcordate .
48. Base of leaf prominently unequal.

44. H. helferi
45. H. aptera

46. H. ultima

49. Young parts evenly pubescent, leaf beneath glabrous, drying dull greyish

47. H. novoguineensis

49. Young parts scabrid pubescent; leaf nervation beneath + pubescent; leaf drying coppery brown

to lustrous beneath.
50. Ripe nut to 8 by 6 mm; tertiary nerves beneath scabrous.

50. Ripe nut to 16 by 9 mm, tertiary nerves beneath glabrescent

43. Nerves 12 pairs or less.
51. Stamens 10.
52. Leaf elliptic-lanceolate, nerves ascending.

. 48. H. scabra
. 49. H. papuana

53. Leaf drying coppery-brown; nerves 9-12 pairs, domatia small but distinct, puberulent

53. Leaf drying pale grey-brown; nerves 6-8 pairs; domatia obscure or absent
51. H. depressinerva
52. Leaf ovate, drying dark grey-brown; nerves patent; domatia pubescent .

51. Stamens 15 or more.

54. Connectival appendages shorter than anthers; sapling leaves peltate .

50. H. acuminata

52. H. sangal

53. H. ferrea

54. Connectival appendages at least as long as anthers; sapling leaves not peltate.
55. Ovary and stylopodium cylindrical or pyriform, the stylopodium stout, no longer than the ovary.

56. Style prominent, as long as ovary.

57. Nerves c. 11 pairs, very slender, distinct; midrib evident, applanate, above . 54. H. odorata
57. Nerves 7-9 pairs, very slender, distinct; midrib evident, applanate, above 55. H. centipeda

56. Style very short.
58. Stamens 32-38 .
58. Stamens 15

56. H. plagata

396 FLORA MALESIANA

59. Leaf base obtuse; domatia pore-like, prominently swollen.
60. Leaf thickly coriaceous, undersurface greyish lepidote
60. Leaf chartaceous, undersurface shagrened

59. Leaf base cuneate; domatia not as above.

61. Nerves c. 5 pairs.
61. Nerves at least 6 pairs.
62. Leaf base distinctly unequal.
63. Midrib above and young twigs pubescent .
63. Midrib above and young twigs glabrous
62. Leaf base + equal.

64. Panicle densely persistently buff pubescent.
65. Ovary and stylopodium ovoid . Aye
65. Ovary and stylopodium cylindrical, truncate .

64. Panicle glabrous.

(ser. L, voRe9?

. 57. H. nutans
. 58. H. bancana

. 59. H. pentanervia

60. H. basilanica
61. H. andersonii

62. H. ovoidea
. 63. H. semicuneata

66. Panicles axillary. Ovary and stylopodium cylindrical-truncate. Leaf nerves to 7 pairs

64. H. megacarpa

66. Panicles ramiflorous. an and stylopodium narrowly pyriform. Leaf nerves at least 9

pairs

65. H. samarensis

55. Gynoecium glabrous, narrow; stylopodium longer than ovary, slender, merging with the very

short style.
67. Leaves equal at base, petiole (7—)10—-16 mm long.
68. Nerves + depressed above, without domatia

66. H. nodosa

68. Nerves applanate to somewhat elevated above, with dematia towards leaf base

67. Leaves distinctly unequal at base, petiole S—9(—10) mm long.

69. Leaves mostly with prominent pore-like glabrous axillary domatia

69. Leaves without distinct domatia.
70. Leaf undersurface glabrous; fruit sepals aliform

67. H. celebica
68. H. iriana

69. H. glabrifolia

70. Leaf undersurface stellate lepidote; fruit sepals short, becoming reflexed . 70. H. gregaria
27. Ovary and stylopodium hour-glass-shaped, distinctly constricted medially; stylopodium tapering into the

short style. Spp. 70-83 2. Sect. Hopea 2b. subsect. Pierrea
71. Leaves broad, base equal, cuneate on both sides; panicles solitary.
72. Fruit calyx lobes unequal, 2 aliform, spatulate
age Fruit calyx lobes shorter than the nut, subequal .

. 71. H. glaucescens
72. H. wyatt-smithii

. Leaves narrow, base + unequal, obtuse to cordate at least on one side; panicles usually more than

l-axillary.

73. Tertiary nerves remotely subreticulate; petiole short, stout (flowers and fruit unknown)

73. Tertiary nerves not as above or, if so, then petiole slender.
74. Fruit sepals short, subequal.
75. Nerves 9-12 pairs; panicles shorter than 5 cm.
76. Tertiary nerves densely scalariform
76. Tertiary nerves remotely scalariform ..
75. Nerves at least 12 pairs; panicles exceeding 6 cm long.
77. Tertiary nerves subreticulate :
77. Tertiary nerves scalariform. Fig. 64a .
74. Fruit sepals unequal, aliform, spatulate.
78. Nerves 6-8 pairs, tertiaries remote .
78. Nerves at least 12 pairs
79. Leaves prominently bullate between the tertiary nerves .
79. Leaves not as above.
80. Base of fruit calyx lobes auriculate
80. Base of fruit calyx lobes not auriculate.
81. Leaves with prominent pubescent axillary domatia.
82. Nerves at least 16 pairs.
83. Leaves at most 15 cm long; fruit sepals to 7.5 cm long.
83. Leaves at least 15 cm long; fruit sepals exceeding 8 cm long
82. Nerves at most 12 pairs. ,
81. Leaves without prominent axillary domatia.
84. Leaves typically 13—22 by 4-7 cm; nerves 12-15 pairs
84. Leaves typically 27-46 by 8-15 cm; nerves 16-30 pairs .

73. H. polyalthioides

74. H. cagayanensis
75. H. paucinervis

.76. H. apiculata
. 77. H. pachycarpa

78. H. bilitonensis
79. H. bullatifolia

80. H. pterygota

81. H. philippinensis

. 82. H. mindanensis

65. H. samarensis

. . 83. H. tenuineryula
. 84. H. enicosanthoides

1982] DIPTEROCARPACEAE (Ashton) 397

1. Section Dryobalanoides

Mig. Sum. (1861) 491, 192 as subgenus; BuRrcK, Ann. Jard. Bot. Btzg 6 (1887)
239; HEM, Rech. Dipt. (1892) 62; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 66; Foxw.
Philip. J. Sc. 6 (1911) Bot. 265; Mal. For. Rec. 10 (1932) 132; Sym. Mal. For. Rec.
16 (1943) 108; AsHToN, Gard. Bull. Sing. 20 (1963) 258; Man. Dipt. Brun. (1964)
90; GUTIERREZ, Act. Manil. 4A, 2 (1968) 25. — Fig. 63, 70.

Nervation dryobalanoid or subdryobalanoid (H. nervosa, H. sublanceolata
excepted). Bark surface smooth, fissured or cracked, not evenly flaky. Wood with
numerous chambered parenchyma strands; rays not markedly heterogeneous.

la. Subsection Dryobalanoides

Hancea PIERRE. — Hopea sect. Hancea (PIERRE) HEIM.

Nervation dryobalanoid. Bracts fugaceous. Corolla pale (H. griffithii excl.);
panicles regularly branched, branchlets short; flowers many. Ovary and stylopo-
dium ovoid or pyriform, rarely truncate.

Distr. Cochinchina, S.E. and Peninsular Thailand and Burma; in Malesia: Malaya, Sumatra, Borneo,

Philippines, West New Guinea.

1. Hopea pubescens RIDL. FI. Mal. Pen. | (1922) 239:
Foxw. Mal. For. Rec. 10 (1932) 139; Burk. Dict.
(1935) 1194; Sym. Mal. For. Rec. 16 (1943) 140, f. 69.

Medium-sized fissure-barked tree with small +
stilted buttresses. Twigs, petioles, midrib above and
panicle densely persistently tawny puberulent, petals
outside cream pubescent, otherwise glabrous. Twig c.
1 mm @ apically, much branched, terete, becoming
dark brown; internodes 5-12 mm long, short; stipule
scars obscure. Leaf bud minute; stipules fugaceous.
Leaves 2.5—6 by 1.3—2.8 cm, small, lanceolate, coria-
ceous; base broadly abruptly cuneate; acumen to | cm
long, caudate; nervation dryobalanoid, main nerves c.
12 pairs with + shorter secondaries, hardly evident
beneath, obscure above; midrib sharply prominent
beneath, obscure and depressed above; petiole 4-6
mm long, c. | mm g, slender. Panicles to 3 cm long,
axillary or sometimes terminal, short, singly
branched; branches to 8 mm long, bearing to 4 secund
flowers. Flower buds to 3 by 1.5 mm, ellipsoid, small.
Sepals ovate, acute, the outer 2 somewhat the longer
and narrower. Petals cream. Stamens 15; filaments
slender, tapering, compressed; appendages slender,
somewhat papillose, c. 2 times length of the broadly
oblong anthers. Ovary ovoid, glabrous, crowned by a
somewhat longer columnar style. Fruit pedicel to 1.5
mm long, very slender. 2 longer calyx lobes to 30 by 6
mm, spatulate, obtuse, c. | mm broad above the to 3
by 1.5 mm narrowly elliptic saccate base; 3 shorter
lobes to 3 by 2 mm, ovate, acuminate. Nut to 5 by 4
mm, ovoid, very shortly apiculate.

Distr. Malesia: Malaya (Kelantan, Pahang).

Ecol. Frequent, sometimes abundant, on well-
drained flat land and low hills.

Vern. Mérawan bunga, m. pipit, pengarawan.

2. Hopea foxworthyi ELmer, Leafi. Philip. Bot. 4
(1912) 1469; Foxw. Philip J. Sc. 67 (1938) 282, p.p.;
GUTIERREZ, Act. Manil. 4A, 2 (1968) 32, f. 6, pl.
2. — H. pierrei (non HANCE) Foxw. Philip. J. Sc. 6
(1911) Bot. 265, p.p.; ibid. 13 (1918) Bot. 184;
WHITFORD, Bull. Bur. For. Philip. 10 (1911) 76;
BROWN & MATTHEWS, Philip. J. Sc. 9 (1914) Bot. 439,
481; MerR. En. Philip. 3 (1923) 94; Reyes, Philip. J.
Sc. 22 (1923) 339.—H. glutinosa ELMER, Leafl.
Philip. Bot. 4 (1912) 1470; Foxw. Philip. J. Se. 13
(1918) Bot. 184.

Medium-sized smooth barked narrowly buttressed
tree. Twig apices, leaf buds, petioles, base of peduncle,
ovary and stylopodium fugaceous puberulent, other-
wise glabrous. Twigs c. 1 mm g apically, slender,
much branched, terete, rugulose, dark brown. Leaves
2.7-6.5 by 1-2.5 cm, lanceolate, thinly coriaceous,
lustrous; margin subrevolute; base + equal, cuneate,
apex with to 1.5 cm long slender caudate acumen:
nerves c. 10 pairs, very slender and + obscure on both
surfaces or slightly elevated beneath, slightly
depressed above, ascending, arched, with a few
shorter obscure secondary nerves; tertiary nerves
reticulate, obscure; midrib sharply prominent
beneath, obscure and depressed above; petiole 5-8
mm long, very slender. Panicles to 1.5 cm long, short,
slender, axillary or terminal, singly branched; branch-
lets few, bearing to 5 flowers, bracts and bracteoles
fugaceous. Flower bud to 5 by 3 mm, ellipsoid, rather
large. 2 outer sepals ovate, acute; 3 inner suborbicular,
subacute. Stamens 15; filaments compressed at base,
tapering and filiform below the subglobose anthers;

398

FLORA MALESIANA

[ser. I, vol. 92

Fig. 65. Density map of Hopea Roxs. in Malesia:
number of endemics above the hyphen, number of
non-endemics below it.

appendage about twice as long as anther, slender.
Ovary ovoid, glabrous, without stylopodium: style
about twice as long as ovary, filiform, tapering. Fruit
pedicel c. | mm long, short; 2 longer calyx lobes to 3.5
by | cm, broadly spatulate, obtuse, tapering to 2mm
broad above the to 5 by 4mm ovate saccate thickened
base; 3 shorter lobes to 5 by 5 mm, broadly ovate,
acute; nut to 15 by 5S mm, narrowly ovoid, resinous,
tapering to a prominent attenuate apiculus.

Distr. Malesia: Philippines (Sibuyan).

Ecol. Locally common on red sticky volcanic soil
along ridges at 600—700 m, in seasonal semi-evergreen
forest.

Vern. Mangachapuy.

Note. Collections cited by FoxwortHy from
islands other than Sibuyan belong to H. malibato.

3. Hopea quisumbingiana GUTIERREZ, Act. Manil. 4A,
21968) 81. fe 5; plo:

Twig apices caducous grey-brown puberulent,
ovary and parts of petals exposed in bud persistently
so. Twig c. 1mm g apically, slender, much branched,
becoming dark chocolate-brown; stipules minute.
Leaves 5—7 by 2.5—3.2 cm, ovate-lanceolate, subcoria-
ceous; base broadly cuneate; acumen to 8 mm long,
caudate; nervation dryobalanoid; nerves 9-13 pairs,
very slender but evident beneath, sometimes with a
few tomentose domatia, with shorter secondaries,
tertiary nerves obscure; midrib slender but prominent
beneath, obscure and depressed above; petiole 6-8
mm long, slender. Panicles to 3.5 cm long, terminal or
to 2-axillary, singly branched, branchlets to | cm long,
bearing to 5 secund flowers. Flower buds to 3 by 2 cm,
ellipsoid. Sepals broadly ovate, subacute, subequal.
Stamens 15, subequal; filaments broadly compressed
at base, tapering and filiform distally; appendage
aristate, c. twice length of subglobose anthers. Ovary
ovoid, tapering into the equally long columnar taper-

ing style; ovary and basal 5 of style pubescent. Fruit
unknown.

Distr. Malesia: Philippines (Samar, once col-
lected).

4. Hopea cernua T. & B. Nat. Tijd. N. I. 29 (1867) 252:
Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 4, fig.;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 241; BRANDIS &
GitGc in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 244;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 71, t. 2, f. 8-9:
Merr. En. Born. (1921) 402; Gitc in E. & P. Pfl. Fam.
ed. 2, 21 (1925) 238; ASHTON, Man. Dipt. Brun. Suppl.
(1968) 49, f. 6; Gard. Bull. Sing. 31 (1978) 28. — H.
microcarpa HeIM, Bull. Mens. Soc. Linn. Paris 2
(1891) 954. — Hancea cernua PIERRE, For. Fl. Coch. 4
(1891) sub t. 244. — H. argentea MEIER, Acta Bot.
Neerl. 12 (1963) 348, pl. 13; MEIER & Woop, Sabah
For. Rec. 5 (1964) 207; ASHTON, Man. Dipt. Brun.
Suppl. (1968) 47, f. 6.

Medium-sized buttressed tree with flying but-
tresses, bark becoming thickly flaky. Leaf bud, young
twig and petiole caducous grey-brown pubescent.
Twig c. 1 mm @, terete, smooth; stipule scars small,
obscure. Bud to 3 by 2mm, ellipsoid, obtuse. Stipule to
4 by 2 mm, lorate, elliptic, caducous. Leaves 5-15 by
2-5 cm, elliptic to ovate, subcoriaceous, + distinctly
silvery lepidote beneath; base cuneate, equal, acumen
to 6 mm long; nerves 10-12 pairs, slender but
distinctly elevated and even prominent beneath, often
with a few axillary pore-like pubescent domatia; with
stout secondaries; midrib slender, slightly raised on
both surfaces; petiole 7-9 mm long, slender. Panicle to
3 cm long, glabrous, terminal or axillary, terete; singly
branched, branchlets bearing to 4 flowers: bracteoles
c. 1 mm long, small, linear, glabrous, fugaceous.
Flower bud to 5 by 4 mm, fusiform; sepals narrowly
ovate, acute to subacuminate, subequal, glabrous,
usually patent; petals lanceolate, densely pubescent on
parts exposed in bud. Stamens 15-18, in 3 unequal
verticils; filaments compressed at base, tapering and
filiform below the subglobose anthers; appendage to
connective slender, 2-3 times length of anther, minu-
tely papillose towards base. Ovary ovoid, glabrous.
Style c. 14 times length of ovary, sometimes slightly
swollen in the villous basal 4. Fruit glabrous; pedicel to
2 mm long; 2 longer calyx lobes to 6.5 by 12 cm,
spatulate, obtuse, c. 4 mm broad above the to 5 by 5
mm ovate thickened saccate base; 3 shorter lobes to
1.5cm long, lanceolate, acute, similar at base. Nut to 7
by 5 mm, ovoid, glabrous, apiculate.

Distr. Malesia: Banka, ? Sumatra, N. and E.
Borneo.

Ecol. Local, in Mixed Dipterocarp forest on fertile
soils especially on intermediate and basic igneous
rocks including limestone, to 1650 m.

Vern. Sélangan urat (Sabah), luis timbul (Sar.),
témang djankar, damar putih, d. puteh (Indon. Bor-
neo).

5. Hopea coriacea BuRCK, Ann. Jard. Bot. Btzg 6
(1887) 237; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 64;

1982]

DIPTEROCARPACEAE (Ashton)

399

Merr. En. Born. (1921) 402; AsHToNn, Gard. Bull.
Sing. 31 (1978) 28. — H. kelantanensis SyM. J. Mal.
Br. R. As. Soc. 19 (1941) 144, pl. 3; Mal. For. Rec. 16
(1943) 130, f. 69. — H. garangbuaya ASHTON, Gard.
Bull. Sing. 19 (1962) 256, pl. 2; Man. Dipt. Brun.
(1964) 101, f. 12; ibid. Suppl. (1968) 51.

Tall flaky or fissure-barked tree with + prominent
buttresses. Glabrous but for petals. Twig to 2mm @
apically, stout, terete, smooth; bud ovoid, to 2 by |
mm. Stipule to 4 mm long, linear, fugaceous. Leaves
11-16 by 6-10 cm, broadly ovate, thickly coriaceous,
base obtuse; acumen to 1.2 cm long, narrow, margin
slightly revolute; nerves 8-11 pairs, dryobalanoid but
relatively prominent, strongly arched, at 60°—70° with
short secondaries; tertiary nerves rather distinct,
densely scalariform, at 90°; midrib broad, promi-
nently rounded beneath, slightly raised above; petiole
2-2.5 cm long, stout. Panicle to 9 cm long, terminal or
axillary, to 2-axillary, terete, glabrous; regularly
singly branched, branchlets to 3 cm long, bearing to 6
secund flowers; bracts and bracteoles unknown.
Flower bud to 3 by 2.5 mm, ellipsoid, relatively large,
distinctly pedicellate. Calyx glabrescent outside, fim-
briate; 2 outer lobes ovate, acuminate, 3 inner lobes
thin, suborbicular, terete. Pefa/s to 1.3 cm long,
narrowly lanceolate, acute, pubescent in parts
exposed in bud, otherwise glabrous. Stamens 15, in 3
whorls, pairs alternating with single stamens; fila-
ments slender, tapering; anthers oblong, the anterior
cells slightly the larger; appendage to connective 2—3
times length of anther, prominently glandular tuber-
culate towards base. Ovary ovoid, glabrous at base,
tapering gradually into style; apex of ovary and basal
half of style densely setose; style filiform, tapering, as
long as ovary; no distinct stylopodium. Fruit calyx
glabrous, tapering into pedicel, lobes closely imbricate
at base, hiding the nut, 2 longer to 7 by 1.5 cm,
spatulate, somewhat tapering above the auriculate
obtuse shallowly saccate unthickened base; 2 shorter
to 2 by 1.2 cm, ovate, acute, similar at base. Nut to 18
by 9 mm, narrowly ovoid, with to 2mm long apiculus;
frequently exuding resin.

Distr. Malesia: Malaya (E. coast: Kelantan to
Pahang); Borneo (Kapuas valley, Sarawak and
Brunel).

Ecol. Local, on or near sandy river banks, rarely (in
Brunei) on hills to 200 m.

Vern. Giam hantu (Mal.), damar mélapi (Kapuas),
garang buaya, arang bayar (Brun.).

6. Hopea longirostrata ASHTON, Gard. Bull. Sing. 22
(1967) 277, pl. 23; Man. Dipt. Brun. Suppl. (1968) 52,
lish

Smooth or patchily flaky-barked tree of medium
size. All parts apparently glabrous. Twig c. 2 mm @
apically, much branched, terete, smooth; stipule scar
short, obscure. Bud to 2 by 2mm, subglobose, obtuse.
Stipule caducous, unknown. Leaves 7-9 by 3-5 cm,
ovate-elliptic, coriaceous; base obtuse; acumen to 1.5
cm long, subcaudate; nervation dryobalanoid; main
nerves c. 12 pairs, obscure, unraised, with subequal

long intermediates; midrib slender but prominent
above, acute, elevated beneath. Petiole 7-10 mm long,
geniculate, blackish. Flower unknown. Panicle and
fruit glabrous. Panicle to 4 cm long, terminal or
3-axillary; singly branched, branchlets bearing to 5
flowers. Pedicel c. 5mm long, uniquely long. 2 longer
calyx lobes to 24 by 6mm, spatulate, tapering to 3 mm
and terminating abruptly in a small incrassate central
tubercle; 3 shorter lobes to 15 mm long, linear to
spatulate, acute or obtuse, similar at the base. Nut to 6
by 4 mm, ovoid; style remnant to 2 mm long.

Distr. Malesia: Borneo (Central Sarawak).

Ecol. Rare, lowland Mixed Dipterocarp forest.

7. Hopea sulcata Sym. Gard. Bull. S. S. 10 (1939) 358,
pl. 20; Mal. For. Rec. 16 (1943) 145, f. 68G, 69,
79. — H. micrantha (non Hook. f.) Foxw. Mal. For.
Rec. 10 (1932) 135, p.p.

Medium-sized fissure-barked small-buttressed tree,
at first with stilt roots. Twigs, petioles and panicles
silvery lepidote, domatia and petioles outside cream
pubescent, parts otherwise glabrous. Twig c. 1 mm @
apically, slender, dark brown, + prominently ribbed;
stipule scars obscure. Leaf buds minute, ovate, stipules
minute, linear, fugaceous. Leaves 4-10 by 1.7—-4 cm,
ovate; lanceolate, falcate, relatively small; base
cuneate, shortly decurrent, subequal; acumen to 2 cm
long, slender, caudate; nervation dryobalanoid, main
nerves c. 10 pairs with many shorter unequal secon-
daries, arched, + distinctly elevated beneath, +
depressed or obscure above, the main nerves with +
prominent axillary tomentose domatia; midrib ele-
vated on both surfaces; petiole 5-11 mm long, slender.
Panicle to 6 cm long, terminal or axillary, singly
branched; branchlets to 16 mm long, bearing to 6
secund pale yellow flowers. Flower bud to 2 by 1 mm,
ovoid. Sepals ovate, acute, the outer 2 narrower,
longer than inner 3. Stamens 15, filaments broadly
compressed at base, tapering, appendage aristate, c.
3x length of subglobose anthers. Ovary ovoid,
tapering into the equally long filiform style. Fruit
pedicel c. 2 mm long, slender; base of fruit frequently
impressed. 2 longer calyx lobes to 5.5 by 1.2 cm,
spatulate, narrowly obtuse, c. 5mm broad above the
c. 7mm broad subauriculate centrally saccate thick-
ened base; 3 shorter lobes to 20 by 7 mm, similarly
subauriculate, completely enclosing the nut. Nut to 10
by 7 mm, ovoid, acute.

Distr. Malesia: Malaya (Perak, Selangor, Treng-
ganu, Johore).

Ecol. Locally abundant on ridges, at 100-400 m.

Vern. Mérawan meéranti, péngérawan bukit.

8. Hopea fluvialis ASHTON, Gard. Bull. Sing. 19 (1962)
254, pl. 1; Man. Dipt. Brun. (1964) 100, f. 12; ibid.
Suppl. (1968) 51. — Fig. 64c.

Medium sized, usually leaning, tree with smooth
bark. Young parts shortly densely pale grey-brown
tomentose, persistent on twig, leaf bud, stipule,
panicle and petiole, fugaceous elsewhere. Twig to 1.5
mm @ apically, terete, becoming smooth, glabrous.

400

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 66. Habit of a 60 m high Hopea mengerawan
MiQ., ngarawan. Palembang (Photogr. THORENAAR,
1923).

Bud to 1 mm long, ovoid, obtuse. Stipule to 2 mm
long, linear, fugaceous. Leaves 7-12 by 2.8-4.8 cm,
chartaceous to thinly coriaceous, lanceolate to ovate;
base narrowly or broadly cuneate; acumen subequal,
to 1.5 cm long, caudate; nerves many, c. 10 pairs with

long secondaries, slender, at 60°—80°, arched, the
basal pair continuing as intramarginal nerves to 3 up
the margin; tertiaries + reticulate, indistinct; midrib
flat or slightly raised beneath, prominently raised
above; petiole 7-10 mm long. Panicle to 6 cm long,
axillary, rarely terminal, borne singly or to 3-axillary,
terete; regularly singly or doubly branched; branchlets
to 1.5cm long, bearing to 7 flowers; bracteoles to 2mm
long, ovate, acuminate, 3 inner lobes thinner, elliptic,
mucronate. Petals narrowly lanceolate, densely
pubescent outside, glabrous within. Stamens 15; fila-
ments broad at base, tapering and filiform distally;
anthers subglobose, the posterior cells somewhat
smaller than the anterior cells; appendage to connec-
tive c. 2 times length of anther. Ovary ovoid, glabrous;
style filiform, tapering, as long as ovary. Fruit calyx
glabrous, tapering into 7 by 5.5 mm expanded, but
unthickened and hardly saccate, base; 3 shorter lobes
unequal, 1—2.5 cm long, acute, tapering, cupped and
enveloping the nut. Nut to 11 by 6 mm, narrowly
ovoid, tapering to a short acute style remnant.

Distr. Malesia: Borneo (S. E. Borneo, S. E. Sabah,
North-East Sarawak and Brunei).

Ecol. Locally abundant on clay rich river banks.

Vern. Mérawan ayer.

9. Hopea mengerawan Mia. Sum. (1860) 492, 192; DC.
Prod. 16, 2 (1868) 635; WaALpP. Ann. 7. (1868) 379:
ScHEFF. Nat. Tijd. N. I. 1 (1870) 351; HANCE, J. Bot.
14 (1876) 308; Burck, Ann. Jard. Bot. Btzg 6 (1887)
240, excl. syn. H. cernua T. & B.; KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 125; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 70, p.p.; BuRK. J. Str. Br. R. As. Soc. 81
(1920) 59, fig.; Rip_. Fl. Mal. Pen. 1 (1922) 238;
THORENAAR, Med. Proefst. Boschw. 16 (1926) 112;
HeEyne, Nutt. Pl. ed. 1, 3 (1917) 1190; ed. 2 (1927)
1107, 1110; Foxw. Mal. For. Rec. 10 (1932) 137;
Burk. Dict. (1935) 1190; Sym. Gard. Bull. S. S. 10
(1939) 361; Mal. For. Rec. 16 (1943) 132, f. 69, 73;
ASHTON, Man. Dipt. Brun. Suppl. (1968) 53, f.
7. — Hancea mengerawan PIERRE, For. Fl. Coch. 3
(1891) sub t. 244. — Fig. 66.

Tall buttressed tree with dark fissured bark. Twigs,
petiole and leaf beneath caducous lepidote, parts
otherwise glabrous. Twig c. 2mm @ apically, slender;
stipule scars short, pale. Leaf bud to 2 by 1 mm, ovoid.
Stipule fugaceous. Leaf 6-12 by 2.5—5 cm, lanceolate,
thickly coriaceous; base cuneate; acumen to 1.5 cm
long, slender but evident beneath, with many short to
subequal secondaries; midrib stout, prominent, on
both surfaces; petiole 9-11 mm long, relatively short.
Panicle to 3 cm long, terminal or axillary, terete, singly
branched; branchlets bearing to 6 secund flowers;
bracteoles c. 2 mm long, acicular, fugaceous. Flower
pale yellow; bud to 3 by 2 mm, ovoid. Calyx lobes
ovate, the 2 outer narrower, more coriaceous than the
frequently suborbicular 3 inner. Petals sericeous on
parts exposed in bud; stamens 15, in 3 unequal
verticils; filaments compressed at base, tapering and
filiform below the subglobose anthers; appendage to
connective slender, 2—3 times length of anther. Ovary

1982]

ovoid, glabrous; style c. 2 times length of ovary,
villous in the basal 4}. Fruit pedicel to 2 mm long,
slender. 2 longer calyx lobes to 7 by 1.2 cm, narrowly
spatulate, narrowly obtuse, c. 3 mm broad above the
to 7 by 4mm narrowly ovate saccate thickened base; 3
shorter lobes to 6 by 5 mm, ovate, acute, saccate. Nut
to 10 by 5mm, narrowly ovoid; style remnant slender.

Distr. Malesia: Malaya (Negri Sembilan, Pahang
southwards in east), Singapore, Billiton, Banka,
Sumatra (Palembang, Lampong, Riouw, West Coast
Res. at Tapanuli), Borneo (Central Sarawak, Melawi,
Sampit N.E. to Nunukan).

Ecol. Local, on soils with impeded drainage on flat
land and the base of hills.

Use. The resin was considered one of the best
varieties in Sumatra and Malaya.

Vern. Mérawan pénak, m. hitam, péngérawan, p.
pénak (Mal.), mérawan banglai, chéngal, c. bulu
(Sumatra), njérakat, émang, bangkirai témbaga, b.
telor (S. E. Borneo).

10. Hopea micrantha Hook. f/. Trans. Linn. Soc. 23
(1860) 161; DC. Prod. 16, 2 (1868) 634; WaLp. Ann. 7
(1868) 379; Dyer, Fl. Br. Ind. 1 (1874) 310, p.p., quoad
spec. Born.; BurRcK, Ann. Jard. Bot. Btzg 6 (1887) 239;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 70, p.p.; MERR.
En. Born. (1921) 402, p.p.; Sym. Gard. Bull. S. S. 9
(1938) 323-329; ibid. 10 (1939) 355, pl. 19; BROWNE,
For. Trees Sarawak & Brunei (1955) 121, p.p.;
ASHTON, Man. Dipt. Brun. (1964) 103, f. 12; ibid.
Suppl. (1968) 54; MEER & Woop, Sabah For. Rec. 5
(1964) 213.— Hancea micrantha Pierre, For. FI.
Coch. 4 (1891) t. 243.

Small to medium-sized smooth barked tree with
small narrow buttresses and abundant stilt roots.
Young twig, leaf bud, panicle, petiole, and stipule pale
brown fugaceous pubescent. Twig to 1.5 mm @
apically, slender, glabrous apart from the apices,
terete, smooth. Bud to 2 by 1.5 mm, ovoid, obtuse.
Stipule to 2 mm long, narrowly deltoid, acute, fuga-
ceous. Leaves 6-8 by 2.5—3 cm, oblong-lanceolate,
coriaceous; base obtuse or broadly cuneate, equal:
acumen to 1.5 cm long, caudate; nerves 10-12 pairs,
indistinct, curved, with short secondaries; midrib
straight, slightly raised above, prominently so
beneath; petiole 7-10 mm long, short. Panicle to 1 cm
long, terminal or axillary, terete; singly branched,
branchlets short, bearing up to 5 + secund flowers:
bracteoles small, narrowly deltoid, glabrescent, fuga-
ceous. Bud small, ovoid. Calyx puberulent outside,
glabrous within; 2 outer lobes narrowly ovate, acu-
minate, 3 inner lobes thin, suborbicular, mucronate;
petals linear, densely pubescent on parts exposed in
bud. Stamens 15, of 3 lengths; filaments slender,
tapering; anthers broadly oblong; appendage to con-
nective c. 15 times length of anther. Ovary ovoid,
glabrous, without distinct stylopodium; style c. 15
times length of ovary, filiform, glabrous. Fruit calyx
glabrous; 2 longer lobes to 5 by 1.2 cm, spatulate,
obtuse, to 2 mm wide above the to 4 by 3 mm saccate
deltoid thickened base; 3 shorter lobes to 5 by 5 mm,

DIPTEROCARPACEAE (Ashton)

40]

broadly ovate to suborbicular, subacute to obtuse,
thin, saccate, shorter than the nut. Nut to 10 by 6mm,
ovoid; style remnant to 1.5 mm long, filiform.

Distr. Malesia: Borneo (northern coast between
Limbang and Bintulu, including Labuan).

Ecol. Mostly on kerangas and white-sand terraces,
or in heath forest, sometimes associated with A gathis,
rarely on sandy clay soil, or on hillsides, at low
altitude.

Vern. Mérawan kérangas.

11. Hopea kerangasensis ASHTON, Gard. Bull. Sing.
22 (1967) 277, pl. 22; Man. Dipt. Brun. Suppl. (1968)
Sy) Sis Th

Medium-sized smooth barked tree with thin flying
buttresses. Twigs, petiole, buds, midrib above and
domatia beneath shortly evenly persistently pale
tawny pubescent, sometimes glabrescent. Twig c. |
mm g@ apically, much branched, terete, becoming
smooth. Bud to 1 mm, minute, globose. Stipule
fugaceous. Leaves 1.5—4.5 by 1-3 cm, small, ovate,
chartaceous, with broadly cuneate base; acumen to 1
cm long, caudate; nervation dryobalanoid, obscure,
main nerves c. 6 pairs: midrib slightly depressed
above, slender but prominent beneath, with to 6 pairs
of prominent large pale fulvous pubescent domatia;
petiole 3-S mm long, slender. Panicle to 12 mm long,
axillary, small, terete, sparsely buff puberulous; singly
branched, branchlets to 4 mm long, bearing to 3
distichous flowers; bracteoles minute, linear, fuga-
ceous. Flower bud c. 1.5 by 1 mm, ovoid. Sepals ovate,
acuminate, glabrous; the inner 3 shorter, relatively
narrower at apex, relatively broader medially, than
the outer. Petals lanceolate, puberulent on parts
exposed in bud. Stamens 15, in 3 unequal verticils;
filaments compressed at base, tapering and filiform
below the subglobose anthers; appendage to connec-
tive slender, 2-3 times length of anther, reaching
almost to style apex at anthesis. Ovary ovoid, gla-
brous; style as long as ovary, columnar, tapering.
Fruit glabrous. Pedicel to | mm long, short. Calyx
lobes to 6 by 5 mm, subequal, ovate, acute, saccate,
thickened, the 2 outer shorter, narrower and more
incrassate than the 3 inner. Nut to 8 by 5 mm, ovoid,
subacute.

Distr. Malesia: Malaya (Pahang, Trengganu);
Sumatra (Indragiri); Borneo (Sarawak west of the
Lupar; Central Kalimantan: Upper Barito).

Ecol. Very local; there abundant, on leached soil in
Mixed Dipterocarp and Heath forests on low hills.

Vern. Sélangan kérangas (Sar.).

12. Hopea vesquei HEIM, Bul!. Mens. Soc. Linn. Paris
2 (1891) 971; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 68;
Merr. En. Born. (1921) 403; BROwNE, For. Trees
Sarawak & Brunei (1955) 122; MEER & Woop,
Sabah For. Rec. 5 (1964) 226; ASHTON, Man. Dipt.
Brun. Suppl. (1968) 57, f. 7.

Medium-sized tree with patchily cracked bark and
thin buttresses. Young twigs, leaf bud, stipule and
petiole + caducous grey-brown puberulent. Twig c. |

FLORA MALESIANA

[ser. I, vol. 9?

5 ¥
a. 9
tee)

AYA
re
*<

,

rs

Fig. 67. Stilt-rooted stem-base of Hopea malibato Foxw. Basilan, Philippines (Photogr. GUTIERREZ).

mm @ apically, terete, much branched, straight,
smooth; stipule scars short, obscure. Bud to 2 by 1
mm, ellipsoid, obtuse. Stipule to 3 mm long, linear,
caducous. Leaves 3.5—6 by 1.5—3.5 cm, broadly ovate,
coriaceous; base cuneate to obtuse; acumen to | cm
long, slender; nervation dryobalanoid, main nerves c.
10-13 pairs, slender, hardly raised, with shorter
secondaries; tertiary nerves obscure, reticulate; midrib
somewhat raised on both surfaces; petiole 6-7 mm
long, slender. Panicle to 3 cm long, + terete, ribbed,
greyish tawny puberulent; singly branched, branchlets
to 1 cm long, bearing to 5 flowers; bracteoles c. 1 mm
long, short, linear, fugaceous. Flower bud to 3 by 2
mm, ellipsoid. Sepals ovate, pubescent on parts
exposed in bud, 3 outer acute, 2 inner relatively
broader, narrower at base, mucronate. Petals lanceo-
late, pubescent on parts exposed in bud; stamens 15,
the inner 5 taller than the rest; filaments broad and
compressed at base, tapering and filiform below the
subglobose anthers; appendage to connective slender,
slightly longer than anther. Ovary ovoid, glabrous;
style c. 14 times length of ovary, columnar. Fruit
glabrous. Pedicel 1-2 mm long, short, slender. Calyx

lobes sparsely setose in the basal half; 2 longer to 3.4
by 0.8 cm, spatulate, obtuse, c. 2mm broad above the
to 4 by 3 mm narrowly ovate somewhat thickened and
saccate base; 3 shorter lobes to 4 by 4 mm, ovate,
acute, saccate. Nut to 15 by 3 mm; cylindrical, style
remnant c. | mm long, short.

Distr. Malesia: Borneo (W. and N. E. Sarawak).

Ecol. Locally abundant on leached yellow sandy
soils in Mixed Dipterocarp forest on coastal hills.

Vern. Luis tébal.

13. Hopea dryobalanoides Mia. Sum. (1860) 492; DC.
Prod. 16, 2 (1868) 634; WALP. Ann. 7 (1868) 379;
SCHEFF. Nat. Tijd. N. I. 31 (1870) 351; BuRCK, Ann.
Jard. Bot. Btzg 6 (1887) 240; KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 125, 126; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 69; BoERL. Cat. Hort. Bog. (1901) 104:
HeEyYNE, Nutt. Pl. ed. 2 (1927) 1107; ENDERT, Tectona
28 (1935) 248: RAPPARD, Tectona 30 (1937) 897; Sym.
Gard. Bull. S.S. 10 (1939) 345, pl. 15; Mal. For. Rec.
16 (1943) 123, f. 69; BROWNE, For. Trees Sarawak &
Brunei (1955) 120; AsHTON, Gard. Bull. Sing. 20
(1963) 259: Man. Dipt. Brun. (1964) 50, f. 12, pl. 29

1982]

DIPTEROCARPACEAE (Ashton)

403

(seedlings); ibid. Suppl. (1968) 50; MEVER & Woop,
Sabah For. Rec. 5 (1964) 209.— Hancea dryoba-
lanoides PieERRE, For. Fl. Coch. 4 (1891) sub t.
244. — H. sarawakensis HEM, Bull. Mens. Soc. Linn.
Paris 2 (1891) 971; BraNois, J. Linn. Soc. Bot. 31
(1895) 69; Merr. En Born. (1921) 403. — H. bor-
neensis HEIM, Bull. Mens. Soc. Linn. Paris 2 (1891)
972; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 66, 69;
Merr. En. Born. (1921) 402. — H. micrantha (non
Hook. f.) KiNG, J. R. As. Soc. Beng. Sc. 62, 2 (1893)
126; Foxw. Mal. For. Rec. 10 (1932) 135; MERR. En.
Born. (1921) 402, p.p. — Fig. 12, 13, 62b-d.

Tall tree with flaky bark and prominent thin
buttresses and a few stilt roots. Young twig, leaf bud,
petiole and leaf beneath shortly densely greyish brown
fugaceous pubescent; domatia persistently so. Twig to
1 mm @ apically, terete, slender, glabrous, smooth.
Leaf bud to 1 mm long, minute. Stipule to 2 mm long,
fugaceous. Leaves S—12 by 1.54.5 cm, ovate-lanceo-
late, thinly coriaceous; base cuneate, equal or sub-
equal; acumen to 2 cm long, narrow, caudate; nerves
8-12 pairs, slender but distinct beneath, curved, with
or without scattered axillary domatia; with few short
secondaries; midrib slender, raised and frequently
sharp on both surfaces, more prominently so beneath;
petiole 5-10 mm long, short, slender. Panicle to 5 cm
long, terminal or axillary, 1—2-axillary, lax, terete,
puberulent to glabrous; singly branched, branchlets
bearing to 6 flowers; bracteoles small, linear. Bud
small, broadly ovoid. Calyx puberulent outside, fim-
briate; 2 outer lobes ovate, acuminate; 3 inner lobes
suborbicular, shortly mucronate. Petals narrowly
lanceolate, shortly tomentose on parts exposed in
bud, pale yellow. Stamens 15, in 3 whorls; filaments
slender, tapering; anthers subglobose; appendage to
connective c. 2 times length of anther, sometimes
sparsely glandular papillose. Ovary ovoid, glabrous,
without distinct stylopodium; style as long as ovary,
setose in the basal half, glabrous distally. Fruit calyx
glabrous; 2 longer lobes to 6.5 by 1.5 cm, long,
spatulate, subacute, pronouncedly twisted, tapering
to 5 mm broad above the to 6 by 6 mm deltoid
thickened saccate base; 3 shorter lobes to 8 by 6 mm,
shorter than the nut, broadly ovate, obtuse or sub-
acute, saccate but thinner than base of outer lobes.
Nut to 10 by 8 mm, broadly ovoid, glabrous; style
remnant to 1.5 mm long, filiform.

Distr. Malesia: Malaya, Sumatra, Borneo.

Ecol. Widespread, locally frequent, clay-rich fertile
soils on undulating or well drained flat land, or ridges
below 600 m; common on basalt and intermediate
igneous and volcanic rocks on slopes and ridges.

Uses. Formerly an important producer of damar
mata kuching.

Vern. Mata kuching hitam, mérawan mata kuching
(Mal.), damar m.k., bayang gunong, séluai hitam
(Sum.), mang bési, (emang) bérjangkar, ménséga
(Indon. Borneo).

14. Hopea malibato Foxw. in Elmer, Leafi. Philip.
Bot. 6 (1913) 1953; Philip. J. Sc. 13 (1918) Bot. 184;

ibid. 67 (1938) 281; MerRR. En. Philip. 3 (1923) 93;
GUTIERREZ, Act. Manil. 4A, 2 (1968) 39, f. 7, pl. 3;
ASHTON, Gard. Bull. Sing. 31 (1978) 28. — H. pierrei
(non HANCE) Foxw. Philip. J. Sc. 6 (1911) Bot. 265,
p.p.; ibid. 13 (1918) Bot. 184, p.p. — H. foxworthyi
(non ELMER) Foxw. Philip. J. Sc. 67 (1938) 282,
p.p. — H. woodiana GUTIERREZ, Act. Manil. 4A, 2
(1968) 42, f. 8, pl. 4— H. dalingdingan GUTIERREZ,
Kalikasan 5 (1976) 92, f. 1. — Fig. 67, 68.

Medium-sized smooth barked tree. Domatia persis-
tently greyish puberulent; young twigs, petioles and
sepals fugaceously so, parts otherwise glabrous. Twigs
c. 1 mm g apically, slender, terete, smooth, dark
brown; stipules fugaceous. Buds minute. Leaves 5-9
by 1.5-4 cm, lanceolate-falcate; base cuneate, +
shortly decurrent; acumen to 2 cm long, caudate, very
slender; nervation dryobalanoid, main nerves c. 11
pairs, ascending, arched, with + shorter secondaries,
very slender but evident and elevated beneath, +
obscure above, with or without prominent pubescent
axillary domatia; tertiary nerves densely reticulate, +
obscure; midrib slender but distinctly and + equally
elevated on both surfaces; petiole 8-16 mm long,
slender. Panicles to 2.5 cm long, axillary, slender,
singly branched; branchlets to 12 mm long, bearing to
5 flowers. Flower buds to 6 by 3 mm, ellipsoid. Sepals
ovate, subacute, fimbriate apically, the inner 3 some-
what broader at base. Petals glabrous. Stamens 15;
filaments lorate, tapering; appendages c. 15 times as
long as the subglobose anthers, shorter than style.
Ovary ovoid, glabrous, without stylopodium, with
columnar style 14-2 times its length. Fruit pedicel to 2
mm long, short, slender; 2 longer calyx lobes to 35 by 9
mm, spatulate, obtuse, to 2mm wide above the 4 by 3
mm ovate saccate thickened base; 3 shorter lobes to 4
by 4 mm, ovate, subacute, shorter than the to 7 by 4
mm apiculate nut.

Distr. Malesia: Philippines.

Ecol. Widespread and locally common in ever-
green non-seasonal dipterocarp forests.

Vern. Malibato, barakbakau, danginginan (Mbo.),
dadingdingan (Tag.), dala (Neg.), dalingdingan (Bik..
Tag.. Dum.), dalingdinganisak (Tag., Bik.), isak
(Tag.), kaliot (Pang.), lito (Ibn.), malatagum (Bik.),
manggachapuy (Bik., Tag., Mbo., Pang.), mululagum,
pisak (Bik.), pisak (Ibn.), sarabsaban (Mang.), siyan,
sugkad (S.L. Bis.), yakal-keliot (official name).

Notes. This species is variable in tree size, leaf size
and shape, and flower size and colour. On the basis of
these differences GUTIERREZ has recognized two addi-
tional species for small-leaved collections formerly
confused with H. pierrei HANCE and H. foxworthyi
ELMER. These latter species conspicuously differ in
their obscure depressed midribs on the leaf above as
well as other characters. Though further study may
prove him correct, I do not uphold these as the same
level of variation occurs within some other diptero-
carp species as here recognized, e.g. H. dryobalanoides
and also in Shorea ovata and S. curtisii.

I define H. malibato by its dryobalanoid, unequal-
based, ovate-lanceolate leaf with c. 11 pairs of slender

404

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 68. Flowering twig of Hopea malibato Foxw.
Basilan (Photogr. GUTIERREZ).

but elevated nerves with shorter secondaries, its
evident and elevated midrib above, and by the absence
of a floral stylopodium, small fruit and glabrous parts
except for the petals and more or less caducous
puberulent innovations. It differs therefore from H.
vesquei HEIM in that the ripe nut does not exceed 11
mm length, from H. dryobalanoides Mia. in that the
fruit sepals do not exceed 4 cm length (as well as in
characters of the leaf midrib and petiole), and from H.
johorensis SYM., H. latifolia SyM. and H. ferruginea
Parus notably in its glabrous panicle.

15. Hopea johorensis Sym. J. Mal. Br. R. As. Soc. 19, 2
(1941) 139, pl. 1B; Mal. For. Rec. 16 (1943) 130, f. 69.

Medium-sized trees with stilt roots and reddish
powdery bark. Panicles and petals outside persistently
greyish buff pubescent, twig apices, petioles and calyx
outside caducously so. Twig c. 1 mm g, ribbed along
the leaf traces, becoming blackish; stipule scars small,
pale. Leaf bud minute; stipule fugaceous. Leaves 3—7
by 1.5-3.5 cm, ovate, coriaceous; base + abruptly
cuneate, subequal; acumen to 12 mm long, prominent,
slender, caudate; nerves many, main nerves c. 16 pairs
with many subequal secondary nerves, very slender
and hardly elevated beneath, obscure above; midrib
slender but distinctly and equally elevated on both
surfaces; petiole 7-13 mm long, slender. Panicle to 15
mm long, short, axillary, with short branchlets bear-
ing to 3 flowers. Mature flowers unknown. Stamens

15. Fruit pedicel c. 2 mm long, very slender; 2 longer
calyx lobes to 4 by 0.8 cm, spatulate, subacute,
tapering to c. 1 mm wide above the to 3 by 3 mm ovate
saccate thickened base; 3 shorter lobes to 3 by 3 mm,
ovate, subacuminate. Nut to 8 by 4 mm, lanceolate,
minutely apiculate.

Distr. Malesia: Malaya (E. Johore).

Ecol. Local, on hill ridges.

Vern. Mata kuching pipit, mérawan.

16. Hopea latifolia Sym. Gard. Bull. S. S. 10 (1939)
360; Mal. For. Rec. 16 (1943) 131, f. 69; ASHTON,
Man. Dipt. Brun. (1964) 102, f. 12; ibid. Suppl. (1968)
52. — H. intermedia (non KiNG) Foxw. Mal. For.
Rec. 10 (1932) 134, p.p. — H. beccariana (non BURCK)
Sym. Gard. Bull. S. S. 9 (1938) 325, p.p.
Medium-sized smooth barked tree with small thin
buttresses and stilt roots. Twig, leaf bud, stipule and
leaf as H. beccariana. Panicle to 4 cm long axillary,
rarely terminal, shortly grey-brown pubescent or
glabrescent, terete, borne singly; singly or doubly
branched, the branchlets bearing up to 5 distichous
flowers. Leaf bud small, ovoid. Calyx shortly pubes-
cent outside, glabrous within; 2 outer lobes ovate,
acuminate, 3 inner lobes suborbicular, mucronate,
thinner. Petals small, ovate, obtuse, pubescent on
parts exposed in bud. Stamens 15, in 3 whorls;
filaments broad at base, tapering somewhat abruptly
distally; anthers subglobose; appendage to connective
as long as anther, slender. Ovary ovoid, glabrous; style
as long as ovary, without distinct stylopodium, fili-
form, + villous towards base. Fruit calyx glabrous; 2
longer lobes to 6 by 1.4 cm, spatulate, narrowly
obtuse, tapering to 3 mm broad above the to 5 by 4
mm deeply saccate thickened base; 3 shorter lobes to 9
by 7 mm, ovate, acute, saccate, thickened, frequently
hiding the nut. Nut to 8 by 7 mm, broadly ovoid, style
remnant to 2 mm long, filiform.
Distr. Malesia: Malaya,
Brunei).
Ecol. Rare, low lying land in Mixed Dipterocarp
forest; once from deep soil over limestone in Perlis.
Vern. Mérawan daun bulat, m. jongkang, m. batu,
chéngal mata kuching (Mal.).

Borneo (Sarawak,

17. Hopea ferruginea PARus in Fedde, Rep. 33 (1933)
243; Bijdr. O-I. Damarhars (1933) 89; Sym. Gard.
Bull. S. S. 10 (1939) 349, pl. 17; Mal. For. Rec. 16
(1943) 125; MEIER & Woop, Sabah For. Rec. 5 (1964)
211, f. 27, pl. 23a (stem). — H. micrantha (non HOOK.

jf.) Dyer, FI. Br. Ind. 1 (1874) 310, p.p.; KING, J. R. As.

Soc. Beng. Sc. 62, 2 (1893) 124, p.p.; BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 70, p.p.; RipL. Fl. Mal. Pen. 1
(1922) 237, p.p.; Foxw. Mal. For. Rec. 10 (1932) 136,
f. 68D, 69; BucKLEy, Mal. For. Rec. 11 (1932)
21. — H. myrtifolia (non MiQ.) HEYNE, Nutt. Pl. ed. 2
(1927) 1107; ENDERT, Tectona 28 (1935) 248. — H.
intermedia (non KING) Foxw. Mal. For. Rec. 3 (1927)
74, p.p.; ibid. 10 (1932) 134, p.p.— H. pierrei (non
Hance) Foxw. Mal. For. Rec. 10 (1932) 133,
p.p. — Fig. 63 B1-B4.

1982]

Medium-sized flaky barked tree with stilt roots and
flying buttresses. Twig endings, petioles, midrib above
and panicles densely + persistently greyish puberu-
lent, petals outside densely cream pubescent. Twig c. |
mm @ apically, terete, becoming dark brown, smooth;
internodes short; stipule scars obscure. Leaf bud
minute; stipules minute, fugaceous. Leaves 4.5—7.5 by
1.54 cm, ovate to lanceolate, subcoriaceous; base
cuneate, shortly decurrent; acumen to 1.5 cm long,
slender, caudate; nervation dryobalanoid, main
nerves c. 14 pairs with many subequal secondaries,
distinct and frequently with + prominent axillary
pubescent domatia especially in young trees; midrib
slender but distinctly elevated on both surfaces;
petiole 6-10 mm long, slender. Panicles to 2 cm long,
axillary, slender, short, with to 8 mm long short
branchlets bearing to 4 secund pale yellow flowers.
Flower bud to 3 by 2mm, ellipsoid. Sepals ovate, the 3
outer somewhat longer, acute or subacuminate, the
inner acute. Stamens 15, unequal; filaments broadly
compressed at base, tapering and filiform distally;
appendage very slender, c. twice length of the subglo-
bose anthers. Ovary ovoid, style somewhat longer,
columnar, tapering, villous towards base. Fruit pedicel
to 2 cm long, very slender; 2 longer fruit calyx lobes to
3 by 0.5 cm, spatulate, narrowly obtuse, c. 1.5 cm
broad above the to 4 by 2.5 mm ovate saccate
thickened base; 3 shorter lobes to 8 by 4 mm,
ovate-acuminate, closely enveloping nut. Nut to 7 by 4
mm, ovoid, apiculate.

Distr. Malesia: Malaya (Perak and Pahang south-
wards), Riouw Arch., E. and Central Sumatra
(Tapanuli, Djambi), E. Borneo (Kudat to Pleihari and
Martapura); wrongly recorded from Sarawak by
BROWNE (Forest Trees of Sarawak and Brunei, 1955,
21).

Ecol. Deep fertile soils in Mixed Dipterocarp forest
below 750 m; locally frequent.

Uses. A minor source of damar mata kuching.

Vern. Mata kuching merah, mérawan m.k., mér-
awan jangkang, m. meérah, m. pasir (Malaya), gangsal
(Djambi), méranti emeh (Tapanuli), mérawan puteh
(Kalimantan), sélangan mata kuching (Sabah).

18. Hopea pierrei HANcE, J. Bot. 15 (1876) 308; ibid.
16 (1877) 329: BRANDIS & GILG in E. & P. Pfl. Fam. ed.
1, 3, 6 (1894) 263; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 67, p.p.; GUERIN, Fl. Gen. I.-C. 1 (1910) 372;
Gwen Be éeP, Pil, Fam. ed) 2) 2171(1925)) 238:
Dakkus, Bull. Jard. Bot. Btzg III, Suppl. 1 (1930) 162;
Sym. Gard. Bull. S. S. 9 (1938) 323, pl. 17; Mal. For.
Rec. 16 (1943) 139, f. 69: SMITINAND, Thai For. Bull. 1
(1954) 19; AsHToN, Gard. Bull. Sing. 31 (1978)
29. — H. micrantha (non Hook. f.) HANCE, J. Bot. 15
(1876) 242. — Hancea pierrei PIERRE, For. Fl. Coch. 4
(1891) t. 248. — H. avellanea Hem, Bot. Tidsskr. 25
(1902) 46; GuERIN, FI. Gen. I.-C. 1 (1910) 375; CRAB,
Fl. Siam. Enum. 1 (1925) 147; SMITINAND, Thai For.
Bull. 1 (1954) 10; Nat. Hist. Bull. Siam Soc. 19 (1958)
63.

Medium-sized smooth barked tree, with thin but-

DIPTEROCARPACEAE (Ashton)

405

tresses and sometimes stilt roots. Twig apex fugaceous
puberulent, ovary caducously so, petals outside den-
sely cream pubescent, parts otherwise glabrous. Twig
c. | mm @ apically, very slender, straight, terete,
horizontal, dark brown; stipule scars obscure. Leaf
bud minute; stipules minute, linear, fugaceous. Leaves
4-8 by 1.64 cm, lanceolate, subcoriaceous; base
abruptly broadly cuneate; acumen to 12 mm long,
slender, caudate; nervation dryobalanoid, main
nerves c. 18 pairs with many + equal secondaries,
very slender and hardly evident beneath, obscure
above; midrib slender but prominent, terete beneath,
obscurely depressed above; petiole 6-11 mm long,
slender. Panicle to 2 cm long, slender, axillary or
sometimes terminal, singly branched; branchlets to 8
mm long, bearing to 4 pale yellow flowers. Buds to 3
by 2 mm. Sepals broadly ovate, acute to subacu-
minate. Stamens 15, compressed at base, tapering;
appendages very slender, c. 3 x length of the subglo-
bose anthers. Ovary and stylopodium hour-glass-
shaped, equal, punctate beneath the short tapering
style, with prominent median constriction. Fruit pedi-
cel to 2 mm long, very slender. 2 longer calyx lobes to
27 by 7 mm, small, spatulate, obtuse, c. 1 mm broad
above the to 3 by 2mm ovate saccate thickened base; 3
shorter lobes to 3 by 2 mm, ovate. Nut to 6 by 5mm,
ovoid, shortly apiculate.

Distr. Vietnam, Cambodia, S.E. Thailand, and in
Malesia: Malaya (Pahang, Selangor, Negri Sembi-
lan).

Ecol. Ridges at 300-700 m in Malaya, sometimes
locally gregarious (abundant in Heath forest in
Cambodia, and widespread on sandy soils in ever-
green forest in Indochina).

Vern. Mérawan palong.

19. Hopea inexpectata ASHTON, Gard. Bull. Sing. 31
(1978) 29.

Medium-sized smooth barked tree. Young twigs,
petioles and base of peduncle caducous tawny puberu-
lent. Twigs c. 2 mm @ apically, slender, much
branched, somewhat ribbed, becoming blackish,
smooth. Leaves 5—9 by 2.4-4.5 cm, ovate, coriaceous;
margin subrevolute; base + equal, broadly cuneate;
acumen to 15 mm long, slender; nervation dryoba-
lanoid, main nerves 8-10 pairs, slender but distinct
elevated beneath, arched, with | or a few shorter less
distinct secondary nerves; tertiary nerves reticulate,
evident beneath; midrib sharply prominent beneath,
depressed and obscure above; petiole 6-7 mm long,
short, slender. Flowers unknown. Panicle to 2.5 cm
long, terminal or axillary to ramiflorous, slender,
singly branched; bracts unknown, caducous. Fruit
pedicel to 2 mm long, slender. 2 longer calyx lobes to 7
by 1.5 cm, spatulate, subacute, tapering to 2 mm
broad above the 4 by 2 mm ovate saccate thickened
base; 3 shorter lobes to 6 by 3 mm, ovate, acuminate,
shorter than nut; nut to 8 by 4 mm, ovoid-acuminate,
tapering to a short apiculus bearing the vestiges of a
distinct stylopodium.

Distr. Malesia: West New Guinea (Kebar valley).

406

FLORA MALESIANA

[ser. I, vol. 9?

' ho
a

Fig. 69. Trunk-base of Hopea beccariana BuRCK.
Brunei, Andalau For. Res. (Photogr. G.H.S. Woop,
SAN 17534).

Ecol. Locally frequent in lowland forest.

Vern. Bu-aan, arais.

Notes. This unexpected record of the first member
of sect. Dryobalanoides east of Wallace’s Line suggests
the possibility of polyphyletic origin, presumably
from sect. Hopea, which is otherwise the only section
occurring in New Guinea and already shows remark-
able plasticity there, as in H. celtidifolia.

H. inexpectata shows a strong superficial resem-
blance to H. griffithii KURZ, but differs in its lustrous
leaf undersurface, sericeous young parts and short
petiole; flowers are required for critical comparison of
androecium and gynoecium.

20. Hopea griffithii KURZ, J. R. As. Soc. Beng. Sc. 42,
2 (1873) 60; Fl. Burma | (1877) 122; Dyer, FI. Br. Ind.
1 (1874) 310; BRANpis, J. Linn. Soc. Bot. 31 (1895) 69
(var. pedicellata excl.); Indian Trees (1906) 68; HEYNE,
Nutt. Pl. ed. 2 (1927) 1107; Sym. Gard. Bull. S. S. 9
(1938) 324, 329; ibid. 10 (1939) 343, pl. 14; Mal. For.
Rec. 16 (1943) 127, f. 69; AsHTon, Man. Dipt. Brun.
Suppl. (1968) 51, f. 7. — Hancea griffithii PrERRE, For.
Fl. Coch. 4 (1891) sub t. 248.— H. pierrei (non
HANCE) RIDL. Fl. Mal. Pen. 1 (1922) 238, p.p.
Medium sized smooth-barked tree with thin but-
tresses and a few stilt roots. Parts glabrous but for the

petals. Twig 1 mm @ apically, terete, slender, much
branched. Bud to | by | mm, small, conical. Stipule to
2 mm long, linear, fugaceous. Leaves 4-9 by 1.7-4.5
cm, ovate to lanceolate, coriaceous; base narrowly or
broadly cuneate; acumen to 1.5 cm long, caudate;
margin frequently subrevolute; nerves c. 9 pairs,
dryobalanoid, hardly raised beneath; tertiary nerves
reticulate; midrib depressed above, prominent
beneath; petiole 8-15 mm long, slender. Panicle to 2.5
cm long, terminal or axillary, ribbed, glabrous, singly
branched; branchlets short, bearing to 5 secund
flowers; bracteoles to 2 mm long, linear, fugaceous.
Flower bud to 2 mm long, ellipsoid. Sepals broadly
ovate, subacuminate, fimbriate, otherwise glabrous,
subequal; the inner 3 narrower at base, thinner at
margin, than the outer 2. Petals lanceolate, erose,
densely pubescent on parts exposed in bud, dark red.
Stamens 15, in 3 unequal verticils; filaments com-
pressed at base, tapering and filiform below the
broadly ellipsoid anthers; appendage to connective c.
2 times length of anther, slender. Ovary and stylopo-
dium stoutly pyriform, papillose towards apex, other-
wise glabrous, tapering somewhat abruptly below the
shortly columnar glabrous style. Fruit glabrous. Pedi-
cel c. 1 mm long, slender. 2 longer calyx lobes to 3 by
0.5 cm, spatulate, obtuse, c. 2mm wide above the to 4
by 3 mm deltoid thickened saccate base; 3 shorter
lobes to 8 by | mm, linear, similarly expanded at base.
Nut to 7 by 5 mm, ovoid, with to 1 mm long, slender
apiculus.

Distr. Lower Burma, and in Malesia: Malaya (E.
coast, W. Johore), Borneo (Sarawak W. of the Lupar,
? Rejang hinterland; W. & S. Kalimantan).

Ecol. Leached soils in Mixed Dipterocarp forest on
low hills, locally common.

Vern. Mérawan jantan, pérawan, péngérawan bunga
(Mal.).

21. Hopea treubii HEI, Bull. Mens. Soc. Linn. Paris 2
(1891) 955; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 68:
Merr. En. Born. (1921) 403; ASHTON, Man. Dipt.
Brun. (1964) 111, f. 12, pl. 30 (bark); ibid. Suppl.
(1968) 57. — Fig. 70 B-B3.

Medium-sized, fissure-barked tree with low but-
tresses and flying buttresses. Glabrous but for petals
and stylopodium. 7wig to 1.5 mm @ apically, slender,
terete, smooth. Bud to 2 mm long. Stipule to 8 mm
long, linear, fugaceous. Leaves 5-8 by 3-—S5.5 cm,
broadly elliptic to obovate, coriaceous, base cuneate;
acumen to 5 mm long, short, broad; margin subrevo-
lute; nerves c. 7 pairs, strongly curved, dryobalanoid,
at 60°—70° with secondaries running almost to margin;
tertiary nerves reticulate, indistinct; midrib broad,
rounded, slightly raised on both surfaces; petiole c. |
cm long, short. Panicle to 8 cm long, terminal or
axillary to ramiflorous, to 2-axillary, slender, rigid,
strongly ascending, terete or + compressed, glabrous;
regularly singly branched, branchlets to 2.5 cm long,
straight, bearing to 7 secund flowers; bracteoles to |
mm long, narrowly deltoid, glabrous, fugaceous.
Flower bud to 2.5 mm long, ellipsoid, on to 2.5 mm

1982]

long pedicel. Ca/yx glabrous but for the fimbriate
margin; outer lobes ovate-acuminate, inner lobes
shorter, thinner, broadly ovate-mucronate. Petals
narrowly lanceolate, acute, fimbriate, shortly pubes-
cent on parts exposed in bud, lemon yellow. Stamens
10, subequal, in a single row; filaments slender,
tapering from the base; anthers broadly oblong,
subequal: appendage to connective about twice length
of anther. Ovary and stylopodium cylindrical, gla-
brous but for the glandular papillose apex, subtrun-
cate; style short, abrupt, glabrous. Fruit pedicel to 3
mm long, slender. Ca/yx glabrous; 2 longer lobes to
3.5 by 0.7 cm, spatulate, obtuse, to 2 mm broad at the
narrow saccate unexpanded base; 3 shorter lobes to 8
by 2mm, similar at base. Nut to 11 by 5mm, narrowly
ovoid, glabrous; style remnant short, acute.

Distr. Malesia: Borneo (Sarawak and Brunei).

Ecol. Local, on deep yellow sandy soils in Mixed
Dipterocarp forest on coastal hills.

Vern. Mérawan daun tébal.

22. Hopea beccariana BurckK, Ann. Jard. Bot. Btzg 6
(1887) 240: BrRANDis, J. Linn. Soc. Bot. 31 (1895) 68;
Merr. En. Born. (1921) 401; Sym. Gard. Bull. S.S.8
(1934) 28, pl. 18; ibid. 9 (1938) 325, p.p.; Mal. For.
Rec. 16 (1943) 122, f. 69, 71; SMITINAND, Thai For.
Bull. 1 (1954) 19; BRowng, For. Trees Sarawak &
Brunei (1955) 120; ASHTON, Man. Dipt. Brun. (1964)
95, f. 12, pl. 26 (stem): ibid. Suppl. (1968) 48; MEVER &
Woop, Sabah For. Rec. 5 (1964) 207, f. 26, pl.
22. — Hancea beccariana PIERRE, For. Fl. Coch. 4
(1891) sub t. 244, in tab. also partly (by error)
‘beccarii.— H. nicholsoni Heim, Bull. Mens. Soc.
Linn. Paris 2 (1891) 973. — H. intermedia KING, J. R.
As. Soc. Beng. Sc. 62, 2 (1893) 126, p.p.; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 67, p.p.; BRUHL & KING,
Ann. R. Bot. Gard. Calc. 5, 2 (1896) 156, t. 189, p.p.;
HEyYneE, Nutt. Pl. ed. 2 (1927) 1107; Foxw. Mal. For.
Rec. 10 (1932) 134; DURANT, Rep. For. Brunei (1932)
40; Burk. Dict. (1935) 1190. — Balanocarpus ovalifo-
lius Rip. J. Fed. Mal. Str. Mus. 10 (1920) 130, p.p.;
Fl. Mal. Pen. | (1922) 247, p.p.; Foxw. Mal. For. Rec.
10 (1932) 143, p.p. — H. pierrei (non HANCE) RIDL. FI.
Mal. Pen. 1 (1922) 238, p.p. — Fig. 62a, 63 C1—C4,
64b, 69.

Large, smooth or fissure-barked buttressed tree.
Young twig, bud and petiole waxy glaucescent. Twig
to 1 mm @g apically, slender, terete, smooth. Bud to 2
by 1 mm, ovoid. Stipule to 2.5 mm long, linear,
fugaceous. Leaves 5-8 by 2.2-4.5 cm, ovate, thinly
coriaceous; base cuneate, frequently subequal;
acumen to 1.5 cm long, caudate; margin not revolute;
nerves c. 8 pairs, indistinct, dryobalanoid; main
secondaries almost reaching margin; tertiary nerves
reticulate, indistinct; midrib slender, slightly raised on
both surfaces; petiole 1.2-1.7 cm long, slender. Panicle
to 7 cm long, axillary (rarely terminal), terete, puberu-
lent or glabrous; singly branched, branchlets bearing
to 5 secund cream flowers; bracteoles small, linear,
fugaceous. Flower bud small, ellipsoid. Calyx shortly
pubescent outside, glabrous within; 2 outer lobes

DIPTEROCARPACEAE (Ashton)

407

ovate-acuminate, 3 inner lobes ovate to suborbicular,
obtuse or subacute. Petals linear, densely pubescent
on parts exposed in bud. Stamens 15; filaments
slender, tapering; anthers broadly oblong; appendage
to connective about twice as long as anther, slender.
Ovary and stylopodium hour-glass-shaped, puberu-
lent towards the apex, tapering into the short glabrous
style. Fruit calyx glabrous; 2 longer lobes to 3.5 by |
cm, obtuse, to 2mm broad above to 4 by 3.5 mm ovate
saccate slightly thickened base, twisted; 3 shorter
lobes to 7mm long, acute, similar at base. Nut to 9 by 5
mm, broadly ovoid, glabrous; style remnant to 1 mm
long.

Distr. Peninsular Thailand (Pattani) and in Male-
sia: Malaya, E. Sumatra (Karimun), Borneo.

Ecol. Widespread, on coastal hills and on deep soils
on inland ridges (especially in Malaya), occasionally
to 1200 m.

Vern. Mérawan batu, m. jangkang, m. pénak (Mal.),
sélangan pénak (Sab.), s. hijau, garang buaya daun
kéchil (Brun.), chéngai pasir (Iban), témang bési,
bangkirai, nuas njérakat hitam, bélang kémai (S.E.
Borneo).

23. Hopea dyeri HEI, Bull. Mens. Soc. Linn. Paris 2
(1891) 972; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 68;
Me_rr. En. Born. (1921) 402, p.p.; Sym. Gard. Bull. S.
S. 8 (1932) 53; ibid. 9 (1938) 323, 324; ibid. 10 (1939)
353: Mal. For. Rec. 16(1943) 124, f. 68F, 69; BROWNE,
For. Trees Sarawak & Brunei (1955) 120; ASHTON,
Man. Dipt. Brun. (1964) 99, f. 12, pl. 28 (stem-base);
ibid. Suppl. (1968) 50; MEER & Woop, Sabah For.
Rec. 5 (1964) 210.— H. micrantha (non Hook. f.)
Burck, Ann. Jard. Bot. Btzg 6 (1887) 239, p.p.; KING,
J.R. As. Soc. Beng. Sc. 62, 2 (1893) 124, p.p.; BRANDIS,
J. Linn. Soc. Bot. 31 (1895) 70; BurK. J. Str. Br. R. As.
Soc. 81 (1920) 58; HEYNE, Nutt. Pl. ed. 2 (1927) 1107;
Burk. Dict. (1935) 1190.— Hancea microptera
PrerRE, For. Fl. Coch. 4 (1891) sub t. 244,
nomen. — H. microptera DYER ex BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 68, nomen in syn. — H. intermedia
Kina, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 126, p.p.;
Ann. R. Bot. Gard. Calc. 5, 2 (1896) 156, p.p.;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 67, p.p.; Foxw.
Mal. For. Rec. 10 (1932) 134, p.p. — H. pierrei (non
HANCE) BRANDIS, J. Linn. Soc. Bot. 31 (1895) 67, t. 2,
f. 10, p.p.; MERR. En. Born. (1921) 403, p.p.; RIDL. FI.
Mal. Pen. 1 (1922) 238, p.p.; Foxw. Mal. For. Rec. 10
(1932) 133, p.p. — Fig. 63 D3-D4.

Medium-sized tree with flaking bark and thin low
buttresses and stilt roots. Bud, stipule, panicle and
domatia persistently grey-brown pubescent, twig and
petiole caducously so. Twig to 1 mm @ apically,
slender, terete, much branched, becoming smooth.
Bud minute, ovoid, obtuse. Stipule to 1.5 mm long,
linear, caducous. Leaves 2.7—7 by 1.2—2.5 cm, ovate-
lanceolate, somewhat coriaceous, frequently lepidote
beneath; base cuneate, subequal; acumen to 1.5 cm
long, narrow, caudate; margin frequently slightly
revolute; nerves 8-12 pairs, slender, slightly raised,
distinct beneath with many distinct secondaries fre-

408

quently subequal to them; midrib stout, raised on
both surfaces, more prominently raised above than
beneath; petiole 5-8 mm long, short, slender. Panicle
to 3 cm long, terminal or axillary, borne singly, terete;
singly or doubly irregularly branched, the branchlets
short, bearing to 4 flowers; bracteoles minute, oblong,
obtuse, pubescent, fugaceous. Flower cream; bud
small, ovoid, Calyx puberulent to glabrous outside,
glabrous within; 2 outer lobes narrowly ovate, sub-
acuminate; 3 inner lobes thin, suborbicular, mu-
cronate. Petals narrowly lanceolate, densely pubes-
cent on parts exposed in bud. Stamens 15, in 3 whorls;
filaments broad at base, tapering; anthers subglobose;
appendage to connective about twice length of anther.
Ovary and stylopodium pyriform, glabrous but for the
puberulent apex, crowned by a short glabrous style.
Fruit calyx glabrous; 2 longer lobes to 2.5 by | cm,
small, spatulate, narrowly obtuse, to 2.5 mm broad
above the to 5 by 3 cm narrowly deltoid slightly
thickened saccate base; 2 shorter lobes to 7 by 4mm,
ovate, glabrous, acuminate, acute, thin. Nut to 9 by 4
mm, narrow, ovoid; style remnant short, acute.

Distr. Malesia: Malaya, Borneo (Sarawak,
Sabah).

Ecol. Well drained soils on low hills and ridges to
1000 m; locally abundant.

Vern. Mérawan palit, m. hitam, péngawan pasir
(Mal.), sélangan palit (Sab.), emang besi (Kapuas).

24. Hopea myrtifolia Mig. Sum. (1860) 49, 192; DC.
Prod. 16, 2 (1868) 635; WaLp. Ann. 7 (1868) 379;
SCHEFF. Nat. Tijd. N. I. 31 (1870) 551; Burck, Ann.
Jard. Bot. Btzg 6 (1887) 242; BRANpis, J. Linn. Soc.
Bot. 31 (1895) 71, p.p.; MERR. En. Born. (1921) 403,
p.p-; HEYNE, Nutt. Pl. ed. 2 (1927) 1107; Sym. Gard.
Bull. S. S. 10 (1939) 347, pl. 16; Mal. For. Rec. 16
(1943) 134, f. 69, 74. — Fig. 63 A—A4.
Medium-sized, sometimes large, smooth barked
tree with stilt roots and thin buttresses. Twigs, leaf
buds, petioles, midrib above, and domatia densely +
persistently tawny puberulent, panicles sparsely so,
petals outside densely golden pubescent. Twig c. 1 mm
@ apically, slender, straight, + horizontal, terete,
becoming dark brown; stipule scars minute, pale.
Buds minute; stipules small, linear, fugaceous. Leaves
(4.5—)6-12 by (1.8—)2.2—5 cm, lanceolate, coriaceous;
base broadly cuneate, subequal; acumen to 1.8 cm
long, prominent; nervation dryobalanoid; main
nerves c. 13 pairs, with many + shorter secondaries,
slender but evident beneath, distinctly narrowly
depressed above, frequently with pubescent axillary
domatia; tertiary nerves usually evident on both
surfaces, minutely reticulate; midrib slender but dis-
tinctly elevated on both surfaces; petiole 6-12 mm
long, slender. Panicle to 4 cm long, slender, axillary to
ramiflorous, singly branched; branchlets to 1.5 cm
long, bearing to 7 secund flowers. Flower buds c. 2.5 by
2 mm, ovoid. Sepals broadly ovate to suborbicular,
mucronate or subacuminate. Stamens 15, filaments
broadly compressed at base, tapering and filiform
distally; appendages slender, c. twice as long as

FLORA MALESIANA

[ser. I, vol. 9?

subglobose anthers; ovary and stylopodium narrowly
subcylindrical, equal in height but the ovary broader,
without median constriction; style shorter than either,
columnar, tapering. Fruit pedicel to 2 mm long,
slender. 2 longer fruit calyx lobes to 5.5 by 1.2 cm,
spatulate, narrowly obtuse, tapering to c. 2mm broad
above the to 7 by 3 mm ovate saccate thickened base; 3
shorter lobes to 4 by 3 mm, ovate, saccate, thickened.
Nut to 9 by 7 mm, ovoid, bluntly apiculate.

Distr. Malesia: Malaya (S. Perak and Pahang
southwards), Karimun, S. Sumatra (Lampong), S.E.
Borneo (Pulau Laut, Muara Teweh to Balikpapan
and Tarakan), W. Borneo (Melawi).

Ecol. Well-drained undulating land with deep
fertile soil.

Vern. Mata kuching béludu, mérawan jangkang
(Malaya), mata kuching (Sumatra), njérakat jangkar
(Kutei), bangkirai batu, lampéngwea (Muaratewe).

25. Hopea pedicellata (BRANDIS) SyM. Gard. Bull. S.
S. 9 (1938) 327, pl. 19; Mal. For. Rec. 16 (1943) 138, f.
68E, 69; SMITINAND, Thai For. Bull. 1 (1954) 10, 19;
ASHTON, Man. Dipt. Brun. Suppl. (1968) 54, f. 7;
Gard. Bull. Sing. 31 (1978) 30. — H. micrantha (non
Hook. f.) KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893)
124, p.p.; RIDL. Fl. Mal. Pen. 1 (1922) 237, p.p. — H.
griffithii var. pedicellata BRANDIS, J. Linn. Soc. Bot. 31
(1895) 69, p.p. quoad specim. Malay.; Riv. Fl. Mal.
Pen. 1 (1922) 238.—H. intermedia (non KING)
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 67, p.p. — H.
mengerawan (non MiqQ.) BRANDIS, J. Linn. Soc. Bot. 31
(1895) 70, p.p. —H. pierrei (non HANCE) RIDL. FI.
Mal. Pen. | (1922) 238, p.p.; Foxw. Mal. For. Rec. 10
(1932) 133, p.p.; BURK. Dict. (1935) 1193, p.p. — H.
siamensis HEIM, Bot. Tidsskr. 25 (1902) 46; WILLIAMS,
Bull. Herb. Boiss. 2, 5 (1905) 147; ScHMipT, Bot.
Tidsskr. 7 (1907) 46; GUERIN, Fl. Gen. I.-C. 1 (1910)
376; CRAIB, Fl. Siam. Enum. | (1925) 147; SMITINAND,
Nat. Hist. Bull. Siam Soc. 19 (1958) 63.
Medium-sized tree with flaky bark, thin buttresses
and sometimes a few stilt roots. Young twig and
domatia grey-brown puberulent, glabrescent; petiole,
leaf bud and stipule persistently so. Twig c. 1 mm @
apically, slender, much branched, terete, smooth;
stipule scars short, obscure. Bud to | by | mm,
ellipsoid, obtuse, minute. Stipule small, linear, fuga-
ceous. Leaves 4-9 by 1-3.5 cm, ovate-lanceolate, base
cuneate; acumen to 1.5 cm long, subcaudate, slender;
nervation dryobalanoid, main nerves c. 8-12 pairs,
with subequal secondaries, indistinct, slender, hardly
raised; midrib raised on both surfaces; petiole 6-8 mm
long. Panicle to 2 cm long, terminal or axillary, terete,
caducous puberulent; singly branched. Calyx gla-
brous, 2 outer lobes ovate, acute, 3 inner suborbicular,
mucronate. Petals sericeous on parts exposed in bud,
pale yellow. Stamens 15, unequal; filaments com-
pressed at base, tapering abruptly medially and
filiform below the oblong anthers; appendage to
connective filiform, c. twice length of anthers. Ovary
with stylopodium, cylindric-conical, attenuate, trun-
cate, punctate in the distal 5, surmounted by a short

1982]

style. Fruit glabrous. Pedice/ c. 2 mm long. 2 longer
calyx lobes to 3 by 0.5 cm, spatulate, c. 2 mm broad
above the c. 5 by 3 mm ovate saccate thickened base; 3
shorter lobes to 3 by 3 mm, ovate, saccate, adpressed
to the nut. Nut to 6 by 4 mm, ovoid, abruptly acute.
Distr. Southern Indo-China, Peninsular Thailand
and in Malesia: Malaya (Trengganu and Perak and
northwards), Borneo (Sarawak, Sabah, Nunukan).
Ecol. Rare, in hill forests to 750 m.
Uses. A minor source of damar mata kuching.
Vern. Mata kuching bukit (Mal.).

26. Hopea altocollina ASHTON, Gard. Bull. Sing. 22
(1967) 271, pl. 16, 348 (phot. habit); Man. Dipt. Brun.
Suppl. (1968) 46, f. 6, pl. 10 (stem-base).

Tall tree with prominent thin buttresses and pale
thinly flaking bark. Sepals and petals sericeous out-
side; parts otherwise glabrous. Twigs c. 2 mm @
apically, terete; stipule scars short, obscure. Bud to |
by | mm, subglobose, small. Stipule unknown. Leaves
7-10 by 34.5 cm, lanceolate; base obtuse, appearing
cuneate owing to the prominently revolute margin:

DIPTEROCARPACEAE (Ashton)

409

acumen to | cm long, caudate; nervation dryoba-
lanoid, nerves c. 16 pairs, + obscure, with short
secondaries; midrib slender, slightly elevated on both
surfaces; petiole 10-13 mm long, somewhat thickened
distally and geniculate, slender. Panicle to 8 cm long,
terminal or axillary, frequently 2-axillary, terete,
glabrous, singly branched. Flower bud to 4 by 3 mm.
Sepals broadly ovate, acute, the inner 3 broader and
thinner. Stamens 15; filaments rather slender, lorate at
base, tapering medially; anthers ellipsoid; appendages
c. 2xlength of anthers, very slender. Ovary and
stylopodium cylindrical, punctate at the rounded apex;
style short. Fruit entirely glabrous. Pedicel to 2 mm
long. 2 longer calyx lobes to 4.5 by 0.8 cm, spatulate,
obtuse, 2-3 mm broad above the 6 by 3 mm narrowly
ovate thickened saccate base; 3 shorter lobes to 4 by 4
mm, suborbicular, obtuse, shorter than nut. Nut to 7
by 5 mm, ovoid; style remnant to 2 mm long, slender.

Distr. Malesia: Borneo (Rejang hinterland, Sar-
awak to Crocker range, S.W. Sabah).

Ecol. Locally frequent, clay rich soils, hillsides and
spurs, 800—1000 m.

1b. Subsection Sphaerocarpae

(Hem) AsHTon, Gard. Bull. Sing. 20 (1963) 258; Man. Dipt. Brun. (1964) 90;
GUTIERREZ, Act. Manil. 4, A, 2 (1968) 26. — Balanocarpus sect. Sphaerocarpae
Hem, Rech. Dipt. (1892) 77. — Hopea, Bracteata group SyM. Gard. Bull. Sing.
10 (1939) 338; Mal. For. Rec. 10 (1943) 108.

Leaf nervation subdryobalanoid (except spp. 27-30 with scalariform venation);
bracts subpersistent; panicles irregularly branched, with long branchlets and few
flowers (H. nervosa, H. sublanceolata excl.); corolla dark coloured; ovary and
stylopodium truncate. Small or occasionally medium-sized trees; bark smooth or
rarely irregularly cracked; buttresses thin small; usually with stilt roots.

Distr. Malesia: Malaya, Sumatra, Borneo, Mindanao.

27. Hopea aequalis ASHTON, Gard. Bull. Sing. 22
(1967) 271, pl. 15; Man. Dipt. Brun. Suppl. (1968) 46,
f. 6. — H. nov. spec. aff. H. pachycarpa (non SYM.)
MEUWER & Woop, Sabah For. Rec. 5 (1964) 229.

Small to medium-sized tree. Apparently glabrous
apart from the puberulent midrib above. Twig c.2mm
@ apically, terete, smooth; stipule scars short, hori-
zontal, obscure. Bud c. 2 by | mm ellipsoid, obtuse.
Stipule unknown. Leaves 13-25 by 5.5—-8 cm, lanceo-
late, chartaceous; base obtuse; acumen to | cm long;
nerves 16-20 pairs, slender, elevated beneath, at
65°-70°; tertiary nerves densely scalariform, very
slender, diagonal to the nerves; midrib shallowly
depressed above, prominent beneath, slender; petiole
15-18 mm long. Flower unknown. Panicle to 9 cm
long, axillary, terete, glabrous, unbranched or singly
branched. Fruit glabrous. Calyx lobes to 18 by 16 mm,
subequal, ovate, subacute, saccate, incrassate. Nut to
23 by 15 mm, ovoid, acute, frequently coated with
resin.

Distr. Malesia: Borneo (Central Sarawak, San-
dakan Distr.).
Ecol. Rare, low hills in Mixed Dipterocarp forest.

28. Hopea rudiformis ASHTON, Gard. Bull. Sing. 31
(1978) 30.

Medium-sized tree. Twigs, leaf buds and parts of
petals exposed in bud densely + persistently pale
tawny puberulent; panicles sparsely so; calyx outside
and nerves and midrib beneath sparsely caducously
so. Twigs c. 2 mm @ apically, rather straight, ribbed,
becoming smooth, dark brown. Bud small, ovoid,
acute; stipules fugaceous, unknown. Leaves 6.5—14 by
3.5—7.5 cm, ovate to broadly lanceolate, thinly coria-
ceous, minutely stellate lepidote and appearing pale
and dull beneath; margin subrevolute; base broadly
cuneate; acumen to 1.5 cm long, broad, tapering,
down curved and twisted over on pressing; nerves
11-13 pairs, slender but prominent beneath, + ob-
scurely depressed above, arched, at 55°—65°; second-

410

FLORA MALESIANA

[ser. I, vol. 9?

A Ge

—

D3

A3

Fig. 70. Flower details in Hopea sect. Dryobalanoides Miq. All x 10. Sepals drawn from inside. — H. bracteata
Burck. A. Bud, A/. outer sepal, A2. inner sepal, 43. stamens from inside, 44. stamens and pistil. — H. treubii
Heim. B. Bud, B/. outer sepal, B2. inner sepal, B3. stamens and pistil. — H. vaccinifolia RIDL. ex ASHTON. C.
Bud, C/. outer sepal, C2. inner sepal, C3. stamens from inside, C4. pistil. — H. nervosa KiNG. D3. Stamens from
outside, D4. pistil (A HAVILAND 2225, B ANDERSON S 9482, C Hose 583, D KOSTERMANS 13981).

ary nerves few, short, hardly elevated beneath; tertiary
nerves densely scalariform, obscure; petiole 8-13 mm
long. Panicle to 3.5 cm long, to 2-axillary; singly
branched, branchlets bearing to 3 flowers; bracts
unknown, fugaceous. Flower bud to 3 by 2mm, ovoid,
small. Sepals subequal, broadly ovate, subacuminate.
Stamens 15, in 3 subequal verticels; filaments com-
pressed at base, tapering medially and filiform below
the shortly oblong anthers; appendages slightly longer
than anthers, short, slender. Ovary ovoid, sur-
mounted by an indistinct tapering ciliate stylopo-
dium, and short columnar style c. + length of ovary
and stylopodium. Fruit pedicel to 2 mm long, short,
base of ripe fruit + impressed. 2 longer calyx lobes to
9 by 2 cm, broadly spatulate, obtuse, c. 3 mm broad
above the to 6 by Smm ovate deeply saccate thickened
base; 3 shorter lobes to 8 by 8 mm, ovate, chartaceous
at margin, subacute, reaching apex of nut and
adpressed to it; nut to 8 by 8 mm, ovoid, terminating in
the persistently truncate stylopodium.

Distr. Malesia: S.E. Borneo (Tawau to Pulau
Laut); Ulu Kapuas, W. Borneo.

Ecol. Undulating land on deep well drained soil in
lowlands; sometimes in freshwater swamps.

Vern. Sélangan jangkang, emang bahau, putang
léman, damar jangkar.

29. Hopea nervosa KING, J. R. As. Soc. Beng. Sc. 62,2
(1893) 124; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 62;
Rip. Fl. Mal. Pen. 1 (1922) 236; Foxw. Mal. For.
Rec. 10 (1932) 129, pl. 10 (hab.), p.p.; BURK. Dict.
(1935) 1191; Sym. Mal. For. Rec. 16 (1943) 135, f. 69,
75; ASHTON, Man. Dipt. Brun. (1964) 104, f. 12; ibid.
Suppl. (1968) 54; MEVER & Woop, Sabah For. Rec. 5
(1964) 215, f. 28, pl. 23b (stem). — Fig. 70 D3—D4,
TA

Medium-sized tree. Twig, leaf bud, stipule, panicle,
petiole and midrib on both surfaces puberulent,
glabrescent. Twig to | mm g@ apically, slender,
becoming smooth, glabrous. Bud to 2 by | mm, ovoid,
acute. Stipule to 2 mm long, narrowly oblong, acute.
Leaves 9-18 by 4-7 cm, thinly coriaceous, narrowly
ovate to lanceolate; base broadly cuneate; acumen to
1.4 cm long, narrow; nervation scalariform, nerves

1982]

13-15 pairs, dense, slender, prominent beneath,
slightly curved, at 40°-60°, with or without short
slender secondaries; tertiary nerves dense, slender,
scalariform, slightly sinuate, diagonal to nerves; mid-
rib slender, prominent beneath, slightly depressed
above; petiole 1-1.2 cm long. Panicle to 9 cm long,
terminal or axillary, to 2-axillary, terete; singly
branched, branchlets to 1.5 cm long, bearing to 5
flowers; bracteoles small, ovate, acute, puberulent
outside, glabrous within. Flower bud to 3 mm long,
subglobose. Petals narrowly oblong, densely tomen-
tose on parts exposed in bud, purple with pale tips.
Stamens 15; filaments of inner 2 rows broad at base,
abruptly tapering, 5 outer filaments slender; anthers
subglobose; appendage to connective slender, c. 2
times length of anther. Ovary and stylopodium
broadly cylindrical, truncate, glabrous; style short.
Fruit subsessile. Calyx glabrous, impressed at base; 2
longer lobes to 12 by 1.8 cm, thinly coriaceous,
spatulate, narrowly obtuse, to 2.5 mm broad above
the to 13 by 8 mm broadly ovate saccate thickened
base; 3 shorter lobes to 1.5 cm long, acute, similar at
base, closely adpressed to nut. Nut to | cmlongand @,
ovoid, style remnant short.

Distr. Malesia: Malaya, Borneo (Sarawak to
Sabah, S. E. Borneo to Balikpapan).

Ecol. Low lying flat land and hill slopes below 400
m on clay rich fertile soils; locally frequent.

Vern. Mérawan jangkang, m. pénak, chéngal rawan,
c. pasir (Mal.), sélangan puteh, s. bértunjang, damar
jangkar (S.E. Borneo).

30. Hopea sublanceolata Sym. Gard. Bull S. S. 10
(1939) 341, pl. 13; Mal. For. Rec. 10 (1943) 144, f.
69. — H. nervosa (non KING) Foxw. Mal. For. Rec. 10
(1932) 129, p.p.

Medium-sized, sometimes large, tree with reddish
flaky bark. Young twigs and petioles fugaceous
puberulent, calyx outside caducously so, panicles
sparsely persistently so; petals densely cream pubes-
cent outside. Twig c. 2 mm g, dark brown, with a
short rib following the leaf trace otherwise smooth,
terete; internodes 1—2 cm long; stipule scars obscure.
Leaf bud minute, ovoid; stipule fugaceous. Leaves
6.5-15 by 3.5-6.5 cm, elliptic-lanceolate, subcoria-
ceous; base cuneate, typically shortly decurrent:
acumen to 1.5 cm long; nerves 13-18 pairs, arched,
slender but prominent beneath, depressed above as
also the midrib; occasionally with a few short second-
ary nerves; tertiary nerves densely scalariform, evi-
dent beneath, obscure above; petiole 7-20 mm long.
Panicle to 5 cm long, terminal or axillary, ramifiorous,
relatively short, singly branched; branchlets to 2.5 cm
long, bearing to 8 flowers. Flower buds to 3 by 2 mm,
ovoid. 2 outer sepals ovate, deltoid, acute; 3 inner
broadly ovate, subacuminate. Corolla dark red.
Stamens 15. Ovary and stylopodium cylindrical, trun-
cate, with slight median constriction, the stylopodium
the shorter and narrower; style short, stout. Fruit
pedicel c. 1 mm long, short, stout, base of fruit
shallowly depressed. 2 longer fruit calyx lobes to 11.5

DIPTEROCARPACEAE (Ashton)

411

by 2 cm, spatulate, obtuse, c. 4mm broad above the to
8 by 8 mm ovate saccate thickened base; 3 shorter
lobes to 17 by 12 mm, ovate-acuminate, appressed to
nut and enclosing it except at apex. Nut to 2 by 1.3. cm,
shortly apiculate.

Distr. Malesia: Malaya (S. Kedah, N. Perak,
Kelantan, Pahang).

Ecol. Locally common on undulating land and low
spurs.

Vern. Mérawan jéruai, chéngal karang, mata puteh,
panah, pau yang, pahi yang.

31. Hopea nigra Burck, Ann. Jard. Bot. Btzg 6 (1887)
238; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 64; MERR.
En. Born. (1921) 403.

Small tree. Twigs, leaf buds, stipules and parts of
petals exposed in bud densely persistently tawny
pubescent; panicles sparsely + caducously so. Twigs
c. 2mm g apically, rather straight, hardly branched,
becoming smooth, terete. Buds minute, subglobose:
stipules c. 2.5 mm long, linear, not at first caducous.
Leaves 3-10 by 14 cm, broadly lanceolate, subcoria-
ceous; base obtuse; acumen to 1.5 cm long, slender;
nerves 8—10 pairs, very slender, + elevated or obscure
beneath, applanate or narrowly depressed above.
arched, sometimes with small pubescent domatia:
secondary nerves 00, short, obscure; tertiary nerves
subscalariform, obscure; midrib sharply prominent
beneath, + obscure and depressed above: petiole 3-6
mm long, short, slender. Panicle to 3 cm long, short,
slender. axillary, usually solitary; singly branched,
branchlets to 1 cm long, bearing c. 4 secund flowers:
bracts and bracteoles to 2 mm long, linear, subpersis-
tent. Flower bud to 2 by 1 mm, small, ovoid. 2 outer
sepals narrowly ovate-acuminate, 3 inner suborbicu-
lar, mucronate, sparsely fimbriate; stamens 15, in 3
subequal verticils, exceeding style; filaments broadly
compressed at base, tapering and filiform below the
small subglobose anthers; appendages 2-3 times
length of anthers, very slender. Ovary and stylopodium
cylindrical, truncate, punctate distally; style c. 5 length
of ovary and stylopodium, filiform. Fruit pedicel
short. 2 longer calyx lobes to 5.5 by 1 cm, spatulate, +
subacute, tapering to 2 mm broad above the to 6 by 4
mm ovate saccate somewhat thickened base; 3 shorter
lobes to 4 by 5 mm, suborbicular, obtuse or mu-
cronate. Nut to 8 by 6 mm, ovoid, retaining the
prominently truncate stylopodium.

Distr. Malesia: E. Sumatra, Banka, Billiton.

Ecol. Lowland forests.

Vern. Médémut, sasak lingga (Banka), mang (Billi-
ton).

32. Hopea sphaerocarpa (HEIM) ASHTON, Gard. Bull.
Sing. 20 (1963) 258; Man. Dipt. Brun. Suppl. (1968)
56, f. 8. — Balanocarpus sphaerocarpus HEIM, Rech.
Dipt. (1892) 77; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
113; Merr. En. Born. (1921) 407.

Small tree. Twigs, petioles, buds and midrib on
both surfaces + densely persistently evenly tawny
pubescent, leaf nervation sometimes sparsely so. Twig

FLORA

MALESIANA

[ser. I, vol. 9?

Fig. 71. Stilt-rooted Hopea nervosa KING. Sabah (Photogr. G.H.S. Woop).

c. 1mm @ apically, terete, straight, lax, smooth. Bud c.
1 by 1 mm, small. Stipule c. 4 by 2mm, oblong, obtuse,
caducous. Leaves 3.5-10 by 1.4-4.5 cm, ovate to
broadly lanceolate, chartaceous, undulate; base
obtuse to subcordate; acumen to | cm long; nervation
subdryobalanoid, main nerves 9-11 pairs, slender,
elevated beneath, with obscure short secondaries;
tertiary nerves densely scalariform, slender; midrib
slender, elevated beneath, applanate above; petiole
3-6 mm long, short. Panicle to 10 cm long, axillary,
sometimes terminal, lax, slender, glabrous; singly
branched, branchlets zig-zag, bearing to 6 + disti-
chous flowers; bracteoles c. 2 mm long, linear, gla-
brous, subpersistent. Flower bud to 2 by 2 mm,
subglobose. Sepals broadly ovate, subacute, sub-

equal, densely shortly pubescent on parts exposed in
bud. Petals elliptic-oblong, subacute, puberulent on
parts exposed in bud, dark crimson. Stamens 15, in 3
unequal rows; filaments broad and compressed at
base, tapering abruptly and filiform beneath the
subglobose anthers; appendage to connective 3-5
times as long as anther, slender. Ovary and stylopo-
dium cylindrical, truncate, somewhat constricted
medially, glabrous; style short, conical. Fruit gla-
brous. Pedice/ to 4mm long, very long. Calyx lobes to
8 by 7 mm, subequal, ovate, acute, saccate, thickened.
Nut to 10 by 8 mm, ovoid; stylopodium to 2 mm tall,
prominent, truncate.

Distr. Malesia: Borneo (W. and N. E. Sarawak;
N.W. Kalimantan).

1982]

Ecol. Local; Mixed Dipterocarp forest on leached
clay-rich soils on low hills.

33. Hopea mesuoides ASHTON, Gard. Bull. Sing. 22
(1967) 279, pl. 25; Man. Dipt. Brun. Suppl. (1968)
54. — H. subalata (non SYM.) ASHTON, Man. Dipt.
Brun. (1964) 110, f. 12, pl. 27 (stem-base).

Medium-sized tree. Leaf bud and stipule shortly
persistently pale yellow-brown pubescent, other parts
glabrous. Twig to 1.5mm @ apically, slender, smooth,
terete. Bud to | mm long, subglobose. Stipule to 2.5 by
1 mm, hastate, acute, fugaceous. Leaves 8-14 by 2.5—S
cm, thinly coriaceous, ovate; base obtuse; acumen
5-15 mm long, caudate; margin subrevolute; nerves c.
11 pairs, subdryobalanoid, distinct, slightly raised
beneath, with shorter secondaries, at 55°—75°,
strongly curved; tertiary nerves reticulate; midrib
prominently raised beneath, narrow, depressed
above, petiole 7-10 mm long, slender. Panicle to 4 cm
long, axillary, short, terete, glabrous, singly branched;
bracteoles to 1 mm long, deltoid, glabrous, subpersis-
tent. Flower bud small, ovoid to subglobose. Calyx
sparsely pubescent outside, glabrous within; 2 outer
lobes suborbicular, subacute or obtuse; 3 inner lobes
suborbicular, + shortly mucronate. Petals oblong,
obtuse, pubescent on parts exposed in bud. Stamens
15; filaments broad at base, tapering; anthers subglo-
bose; appendage to connective 2-3 times length of
anther, very slender. Ovary and stylopodium glabrous
but for the puberulent apex, cylindrical, truncate,
ovary slightly the broader; style short, glabrous. Fruit
pedicel c. 1 mm long, short, impressed in the base of
the calyx. Calyx lobes to 14 by 12 mm, subequal,
ovate, chartaceous, + erose at the subacute apex,
broadly imbricate, saccate, completely enclosing the
nut and closely adpressed to it. Nut to 14 by 14mm,
subglobose, crowned by the persistent truncate stylo-
podium.

Distr. Malesia: Borneo (Sarawak N. E. of Bal-
ingian, Brunei).

Ecol. Mixed Dipterocarp and Heath forests, deep
leached soil, lowlands.

34. Hopea subalata Sym. Gard. Bull S. S. 10 (1939)
339, pl. 12; Mal. For. Rec. 16 (1943) 143, f. 68A, 69,
78. — Balanocarpus ovalifolius (non RIDL.) Foxw.
Mal. For. Rec. 10 (1932) 143, p.p.

Small tree. Twigs and petioles sparsely fugaceous
puberulent, petals outside cream pubescent, parts
otherwise glabrous. Twig c. | mm @ apically, slender,
dark brown, at first ribbed along the leaf traces;
internodes 1.5—2.5 cm long; stipule scars obscure. Leaf
bud minute, ovoid; stipule to 5 mm long, linear,
caducous. Leaves 4.5—10 by 2.3—5 cm, ovate-lanceo-
late, thinly coriaceous; base broadly cuneate; acumen
to 1.5 cm long, subcuspidate; nervation subdryoba-
lanoid; main nerves c. 12 pairs, arched, frequently
with small pubescent axillary domatia, slender but
distinctly elevated beneath, obscurely depressed
above as also the many shorter secondaries; tertiary
nerve subscalariform, obscure; midrib sharply promi-

DIPTEROCARPACEAE (Ashton)

413

nent beneath, narrowly depressed above; petiole 5—7
mm long, short. Panicle to 5 cm long, axillary ot
terminal, slender, lax, with to 2.5 cm long branchlets
bearing to 5S flowers; bracteoles to 1 mm long, deltoid,
apiculate, subpersistent. Flower buds to 3 by 2 mm,
ovoid; sepals suborbicular, obtuse or subacute, sub-
equal; petals purple; stamens 15, filaments short,
compressed, tapering immediately beneath the sub-
globose anthers; appendages very slender, 2—4 times
length of anthers; ovary and stylopodium cylindrical,
equal, subtruncate, punctate distally, with short stout
tapering style. Fruit pedicel to 2mm long and @. Fruit
calyx lobes to 10 by 8 mm, ovate, subequal, saccate,
thickened, enclosing the nut (the apex excepted), 1(—2)
lobes generally with a to 10 by 3 mm aliform apical
extension. Nut to 10 by 8 mm, ovoid, crowned by a
truncate conical stylopodium with central terminal
apiculus.

Distr. Malesia: Malaya (Selangor).

Ecol. Locally common in one locality, on low hills.

Vern. Mérawan kanching, m. jangkang.

35. Hopea auriculata Foxw. Mal. For. Rec. 10 (1932)
125, pl. 9; BurRK. Dict. (1935) 1189; Sym. Mal. For.
Rec. 16 (1943) 121, f. 69.

Small tree. Young twigs and petioles caducous pale
grey-tawny puberulent. Twig c. 2 mm @ apically,
slender, horizontal; stipule scars minute. Leaf bud
minute; stipule acicular, fugaceous. Leaf 5—10 by 2-5
cm, ovate-lanceolate, thinly coriaceous; base cuneate;
acumen to 1.5 cm long, caudate; nervation subdryo-
balanoid; nerves c. 9 pairs, arched, slender but
distinctly elevated beneath, obscure above, with +
prominently buff pubescent axillary domatia and
short slender secondary nerves; tertiary nerves densely
subscalariform, + obscure; midrib slender but promi-
nent beneath, obscurely depressed above; petiole 5—11
mm long, slender. Panicle to 4 cm long, slender,
hardly branched; flowers unknown. Fruit pedicel to 2
mm long; 2 longer calyx lobes to 7 by 1.5 cm,
spatulate, obtuse, to 5mm broad above the to 15 by 10
mm auriculate saccate thickened base; 3 shorter lobes
to 2 cm long, ovate-auriculate, similarly thickened
centrally; nut to 10 by 7 mm, ovoid, apiculate,
completely enclosed in calyx.

Distr. Malesia: Malaya (N. E. Johore, E. Pahang).

Ecol. Rare, between 250-700 m on ridges.

36. Hopea montana Sym. J. Mal. Br. R. As. Soc. 19, 2
(1941) 141, pl. 1A; Mal. For. Rec. 16 (1943) 133, f. 69;
MEIJER & Woop, Sabah For. Rec. 5 (1964) 214.
Medium-sized tree. All known parts of mature tree
glabrous. Twig c. | mm @ apically, very slender,
straight, blackish; stipule scars obscure. Leaf bud
minute; stipules fugaceous. Leaves 6.5—9 by 2.2—-4 cm,
ovate-lanceolate, thinly coriaceous; base abruptly
cuneate; acumen to | cm long, cuspidate; nervation
subdryobalanoid; main nerves c. 14 pairs, arched,
ascending, with many + shorter secondaries, elevated
beneath, obscurely depressed above as also the mid-
rib; tertiary nerves obscure; petiole 9-11 mm long,
slender. Flowers unknown. Panicle to 2 cm long,

414

slender, hardly branched or unbranched. Fruit pedicel
to 2 mm long, broadening into receptacle. 2 longer
calyx lobes to 5 by 1.2 cm, spatulate, subacute,
tapering to c. 2 mm broad above the to 5 by 3 mm
elliptic saccate thickened base; 3 shorter lobes to 10 by
3 mm, ovate, frequently shortly winged apically. Nut
to 7 by 5mm, ovoid, the exposed apex surmounted by
a truncate stylopodium and central apiculus.

Distr. Malesia: Malaya (Perak, Kelantan); Borneo
(G. Kinabalu).

Ecol. Rare, hill slopes in upper dipterocarp forest,
to 1200 m.

37. Hopea vaccinifolia RIDL. ex [BROWNE, For. Trees
Sarawak & Brunei (1955) 122, nomen] ASHTON, Gard.
Bull. Sing. 19 (1962) 258, pl. 3; Man. Dipt. Brun.
(1964) 112, f. 12; ibid. Suppl. (1968) 57. — Fig. 70
C-C4.

Small tree. Young twig, leaf bud, stipule and petiole
densely shortly pale grey-brown pubescent. Twig to
0.5 mm @ apically, slender, much branched, horizon-
tally, becoming smooth, glabrous. Bud minute. Sti-
pule to 1.2 mm long, narrowly hastate, acute, fuga-
ceous. Leaves 1-2.5 by 0.4-1.2 cm, chartaceous,
elliptic to broadly ovate; base cuneate; apex obtuse or
with to 1.5 mm long obtuse acumen in mature tree;
nervation subdryobalanoid; main nerves c. 6 pairs,
very indistinct with slender secondaries; tertiary
nerves reticulate; midrib slender, applanate or slightly
raised beneath, narrow and depressed above; petiole
to 2 mm long, slender. Panicle to 1.3 cm long, to
2-axillary, short, terete, glabrous; irregularly singly
branched, branchlets short, bearing to 3 flowers;
bracteoles minute, deltoid, subpersistent, glabrous.
Flower bud small, ovoid, subsessile. Calyx glabrous
but for a fimbriate distal margin; 2 outer lobes ovate,
acute; 2 inner lobes broadly suborbicular, obtuse,
thinner. Petals oblong, obtuse, glabrous, strongly
contorted, imbricate and forming a tube at base on
opening, the distal half becoming abruptly rotate or
recurved, dark wine-red. Stamens 10, subequal; fila-
ments broad at base, abruptly narrowing and filiform
distally; anthers subglobose, the interior cells some-
what the longer; appendage to connective slender, c. 2
times length of anther, reaching apex of stylopodium.
Ovary and stylopodium cylindrical, truncate, glabrous
but for the puberulent apical platform; style short.
abrupt, glabrous. Fruit calyx glabrous; lobes sube-
qual, to 4 by 3.5 mm, ovate, acute or obtuse,
thickened, shallowly saccate. Nut to 8 by 6mm, ovoid,
glabrous; apex minutely truncate.

Distr. Malesia: Borneo (Brunei, N. E. Sarawak at
Marudi).

Ecol. Locally abundant; Heath forest on giant
podsols on raised beaches, and on sandstone cuestas.

Vern. Mérawan ribu.

38. Hopea bracteata BurcK, Ann. Jard. Bot. Btzg 6
(1887) 239; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 111;
Sym. Gard. Bull. S. S. 10 (1939) 337; SLoor. Rein-
wardtia 3 (1956) 317; AsHToN, Man. Dipt. Brun.

FLORA MALESIANA

[ser. I, vol. 97

(1964) 97, f. 12; ibid. Suppl. (1968) 48; MEUER &
Woop, Sabah For. Rec. 5 (1964) 229: ASHTON, Gard.
Bull. Sing. 31 (1978) 31.— Balanocarpus curtisii
KiNG, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 131; BRUHL
& Kina, Ann. R. Bot. Gard. Calc. 5, 2 (1896) 158, t.
191; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 111; BurK.
J. Str. Br. R. As. Soc. 81 (1920) 59, fig.; RIDL. F1. Mal.
Pen. 1 (1922) 246; Sym. Gard. Bull. S. S. 8 (1934)
27. — Balanocarpus bracteatus MeRR. En. Born.
(1921) 407; Foxw. Mal. For. Rec. 10 (1932) 142;
Burk. Dict. (1935) 284. — H. minima Sym. Gard.
Bull. S. S. 10 (1939) 337, pl. 11, incl. var. penangiana
Sym. et var. perakensis SYM. I.c. 338; Mal. For. Rec. 16
(1943) 133, f. 69. — Fig. 70 A-A4.

Small, occasionally medium sized, tree. Twig, leaf
bud, stipule and petiole shortly persistently grey-
brown puberulent. Twig to 0.7 mm@g apically, slender,
terete, much branched, becoming smooth, glabrous.
Bud to 2.5 mm long, lanceolate. Stipule to 3 mm long,
linear, fugaceous. Leaves 2.5-6 by 0.7-2 cm, thin,
narrowly ovate, base cuneate; acumen to 1.5 cm long;
nerves c. 11 pairs, slender, hardly more distinct than
secondaries, at 55°-65°, strongly curved; tertiary
nerves slender, reticulate; midrib slender but raised
beneath, narrow, depressed above; petiole 2-4 mm
long, slender. Panicle to 9 cm long, terminal or
axillary, very slender, lax, terete, glabrous; remotely
irregularly singly or doubly branched, branchlets to 3
cm long, zigzag, bearing to 5 flowers; bracteoles to |
mm long, deltoid, glabrous. Bud to 3 mm long,
subglobose, distinctly pedicellate. Calyx lobes fim-
briate, otherwise glabrous; outer lobes suborbicular,
thin; inner lobes suborbicular, mucronate, thin. Petals
oblong, acute, densely shortly tomentose on parts
exposed in bud, dark red with pale tips. Stamens 15;
filaments broad at base, tapering; anthers subglobose;
appendage to connective c. 2 times length of anther,
slender, joining abruptly with the distinctly broader
connective and filament. Ovary and stylopodium gla-
brous, cylindrical, truncate, ovary only slightly the
broader; style short, abrupt, glabrous. Fruit entirely gla-
brous. Calyx lobes to 5 by 5mm, subequal, broadly ovate,
obtuse, incrassate at base but for a chartaceous margin.
Nut to 9 by 6 mm, ovoid; apex minutely truncate.

Distr. Malesia: Malaya (Penang, Perak), Borneo
(Sarawak, Brunei, Sandakan, S.E. Borneo to Purukt-
jau).

Ecol. Locally abundant on spurs and ridges below
650 m.

Vern. Mérawan ungu, damar mata kuching (Mal.),
m. padi (Brun.), bangkirai (S.E. Borneo).

Note. The rare H. brachyptera (Foxw.) SLOOT. of
Zamboanga (Mindanao, Philippines) closely resem-
bles H. bracteata though the leaf is somewhat larger.
According to Foxworthy (Philip. J. Sc. 13, 1918, Bot.
195; ibid. 67, 1938, 285) the ovary differs in being
densely pubescent. I have seen neither flowers nor
fruit, which may all have been lost in the destruction
of the Manila Herbarium in 1945.

39. Hopea brachyptera (Foxw.) SLoot. Reinwardtia 3

1982]

DIPTEROCARPACEAE (Ashton)

415

(1956) 317. — Balanocarpus brachypterus Foxw. Phi-
lip. J. Sc. 13 (1918) Bot. 195; ibid. 67 (1938) 285; MERR.
En. Philip. 3 (1923) 101; Sym. Gard. Bull. S.S. 8 (1934)
D238:

Medium-sized tree. Leaf buds, twigs and petiole,
midrib above and outside of calyx sparsely + cadu-
cous puberulent; petals outside, ovary and domatia
densely persistently pubescent. Twigs c. 1 mm @
apically, terete, dark brown, + rugose. Leaves 3.5—7
by 2.3-3.5 cm, ovate-lanceolate, chartaceous; base
unequal, cuneate; acumen to 12 mm long, slender,
subcaudate; nerves 9-11 pairs, slender but prominent
beneath, obscure and adpressed above, arched;
secondary nerves many but very short; both +
continuously furnished with pubescent domatia; ter-
tiary nerves subscalariform, distinct but hardly ele-
vated; petioles 3-5 mm long, very slender. Flowers to 3

mm long, dull, violet. Sepals ovate, acute. Stamens 15;
filaments filiform, slightly expanded at base; appen-
dages aristate, considerably longer than oblong
anthers. Ovary and stylopodium cylindrical, truncate,
with short style. Fruit calyx lobes short, subequal,
ovate, adpressed to and hiding the 10 by 9 mm nut
except at apex.

Distr. Malesia: Philippines (Zamboanga, Min-
danao).

Ecol. Rare, lowland forests.

Vern. Babase (Sub.).

Note. The flower and fruit description are taken
from Foxwortny; both material seems to have been
completely destroyed during the burning of the
Philippine herbarium. See the note under the preced-
ing species H. bracteata.

2. Section Hopea

ASHTON, Gard. Bull. Sing. 20 (1963) 258; Man. Dipt. Brun. (1964) 91; GUTIERREZ,
Act. Manil. 4, A, 2 (1968) 25, 26. — Hopea sect. Euhopea Mia. Sum. (1862) 192,
as subgenus, 489, as section; BURCK, Ann. Jard. Bot. Btzg 6 (1887) 237, p.p.;
BRANDIS, J. Linn. Soc. Bot. 3 (1895) 57; Foxw. Philip. J. Sc. 6 (1911) Bot. 260;
Mal. For. Rec. 10 (1932) 118. — Hopea, Euhopea group Sym. Gard. Bull. S. S. 8
(1934) 72; Mal. For. Rec. 16 (1943) 108. — Fig. 72, 74.

Nervation scalariform; panicles irregularly branched, many flowered, flowers
pale; bark surface smooth or evenly flaky. Wood with markedly heterogeneous
rays; without chambered parenchyma strands (excepting H. pachycarpa).

2a. Subsection Hopea

Neisandra RaA¥FIN. — Petalandra Hassk. — Balanocarpus BEDD. — Dioticarpus
DuNN. — Hopea sect. Petalandra HEm, Rech. Dipt. (1892) 63; BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 65; Foxw. Philip. J. Sc. 6 (1911) Bot. 264; Mal. For. Rec. 10
(1932) 130. — Balanocarpus sect. Pachynocarpoides HEM, Rech. Dipt. (1892) 75
et sect. Microcarpae HEIM, /.c. 76. — Fig. 72.

Panicles generally tomentose, rarely fascicled; ovary and stylopodium +
ovoid, or, if elongate, without median constriction; style evident. Large, less often
small, trees, usually with flaky bark and buttresses, but rarely stilt-rooted.

Distr. Throughout the range of the genus.

40. Hopea celtidifolia KosTERM. Gard. Bull. Sing. 22
(1968) 443; AsHToN, Gard. Bull. Sing. 31 (1978) 32.

Medium-sized scaly barked buttressed tree. Young
twigs, leaf buds and stipules densely + persistently
tawny puberulent, petioles caducously so. Twig c. |
mm @ apically, much branched; internodes short.
Buds small, ovoid, subacute; stipules small, fugaceous.
Leaves 5-10 by 3-4.5 cm, elliptic to lanceolate,
coriaceous, margin subrevolute; base obtuse to

broadly cuneate; acumen to 8 mm long, slender;
nerves 4-5 pairs, frequently all arising from the
proximal 4 of the leaf, arched and coalescing midway
to the margin forming an almost unlooped continuous
prominent intramarginal nerve continuing to the
acumen, the whole slender but prominent beneath,
distinctly elevated above; intramarginal nerve with
indistinct lateral branches to the margin; tertiary
nerves scalariform, evident beneath; midrib stout,

416 FLORA MALESIANA [ser. I, vol. 92

A A2 i

Fig. 72. Flower details in Hopea sect. Hopea subsect. Hopea. All x 10. Sepals drawn from inside. — H. plagata

(BLco) VipaL. A. Bud, A/. outer sepal, A2. inner sepal, 43. stamens from inside, A4. pistil. — H. forbesii

(BRANDIS) SLooT. B. Bud, B/. outer sepal, B2. inner sepal, B3. stamens from inside, B4. pistil. — H. ferrea

Laness. C. bud, C/. outer sepal, C2. inner sepal, C3. stamens from outside, C4. pistil. — H. centipeda ASHTON.

D1. Outer sepal, D2. inner sepal, D3. stamens from inside, D4. pistil. — H.sangal KorTu. E/. Outer sepal, E2.

inner sepal, E3. stamens from inside, F4. pistil (A PNH 78168, B Carr 12072, CI—C2 PHENG-NaREN, C, C3—C4
Put 4307, D S 23342, E KEP 76619).

A4

1982]

prominent beneath, elevated above; petiole 8-10 mm
long, slender. Panicle unknown; flowers unknown.
Fruit pedicel c. 2 mm long, stout; 2 longer calyx lobes
to 5 by | cm, spatulate, obtuse, c. 2 mm broad above
the to 6 by 3 mm narrowly ovate saccate thickened
base; 3 shorter lobes to 5 by 4 mm, ovate, obtuse to
acute, similarly saccate. Nut to 15 by 6 cm, narrowly
ovoid, the stylopodium showing as a prominent
medially thickened apiculus.

Distr. Malesia: W. New Guinea (R. Digoel, Muku
Subdistr.); S. New Guinea; Strickland R., E. New
Guinea.

Ecol. Local in lowland forest.

Vern. Et, teh, keilmoen, katiau, jeruieh.

41. Hopea dasyrrhachis SLoor. Bull. Bot. Gard. Btzg
III, 17 (1941) 130, f. 18; AsHton, Gard. Bull. Sing. 31
(1978) 32.—TIsoptera dasyrrhachis SLOOT. ex DEN
BERGER & ENDERT, Med. Proefst. Boschw. 9 (1925)
117, nomen.— Shorea dasyrrhachis SLooT. ex
ENDERT, Tectona 28 (1935) 282, 295, nomen.

Large, dark flaky-barked tree. Twig apices, ovary
apex and calyx caducous buff puberulent, panicles
and parts of corolla exposed in bud persistently
densely so. Twig c. 2 mm @, terete; stipule scars short,
obscure. Bud 1 by | mm, minute. Stipule fugaceous.
Leaves 7-15 by 2.5-8 cm, ovate-falcate, somewhat
coriaceous, base cuneate; acumen to 1.5 cm long;
nerves 12—14 pairs, slender, arched, at 50°-60°; typi-
cally with prominent glabrous pore-like axillary
domatia; tertiary nerves slender, scalariform; midrib
raised on both surfaces; petiole 5-15 mm long. Panicle
to 8.5 cm long, terete, singly branched, branchlets
bearing to 10 secund flowers. Flower bud to 2 mm
long, ovoid. 3 outer calyx lobes ovate, subacute, 2
inner suborbicular, villous on parts exposed in bud.
Petals pubescent on parts exposed in bud, cream.
Stamens 10, in a single verticil; filaments compressed
at base, tapering abruptly medially, filiform at the
base of the oblong anthers; appendage to connective
filiform, as long as anther. Ovary and stylopodium
subcylindrical, sericeous in the distal half, terminating
in a short truncate style. Fruit pedicel to 3 mm long,
slender. 2 longer fruit calyx lobes to 4 by 3.5 cm,
suborbicular, chartaceous, frequently subauriculate
with to 4 by 3 mm small thickened area at base; 3
shorter lobes to 6 by 7 mm, suborbicular, saccate. Nut
to 15 by 15 mm, ovoid, shortly apiculate.

Distr. Malesia: Borneo (Kapuas valley, Lower
Dyak, C. Dusun, Marabahan).

Ecol. On river banks in W. and S. Borneo.

Vern. Damar puteh (S. Borneo), tékam, t. lampung,
t. rayap, t. ayér (W. Borneo).

42. Hopea similis SLoot. Reinwardtia 2 (1952) 30; v.
RoyeEN, Man. Forest Trees Papua New Guinea 8
(1965) 41.

Medium-sized, buttressed tree with flaky bark. Leaf
buds, twigs, petioles and panicles + persistently
somewhat scabrid buff pubescent, parts of petals
exposed in bud evenly so; calyx and leaf nervation

DIPTEROCARPACEAE (Ashton)

417

below caducously so, glabrescent. Twig c. 2 mm g
apically, stout, rugose, becoming ribbed; stipule scars
distinct, short, + horizontal. Bud small, ovoid;
stipules unknown, caducous. Leaves 10-22 by 3.5-8
cm, elliptic-oblong to lanceolate, coriaceous; margin
revolute (less so in young trees); base obtuse to
subcordate; acumen to 8 mm long, short, broad:
nerves 12-16 pairs, prominent beneath, evident and
elevated above; petiole 11-14 mm long, c. 2 mm g,
rather stout. Panicle to 6 cm long, axillary to rami-
florous, slender, to 2-axillary; singly branched,
branchlets to 1.5 cm long, bearing up to 6 flowers.
Flower buds to 4 by 2 mm, lanceolate; 2 outer sepals
broadly lanceolate-acuminate, 3 inner suborbicular,
obtuse, fimbriate. Stamens 15, in 3 subequal verticils;
filaments broad and compressed at base, tapering and
filiform beneath the subglobose anthers; appendages
25-3 times length of anther, slender. Ovary and
stylopodium narrowly pyriform, slightly constricted
medially; stylopodium punctate distally, surmounted
by a short columnar style. Fruit pedicel to 3 mm long,
stout. 2 longer calyx lobes to 7.5 by 1.5 cm, obtuse, c. 2
mm wide above the to 8 by 4 mm narrowly ovate
prominently saccate thickened base; 3 shorter lobes to
8 by 6 mm, ovate, mucronate or obtuse, similarly
saccate; nut to 15 by 7 mm, narrowly ovoid, tapering,
prominently slender-apiculate.

Distr. Malesia: New Guinea (Rouffaer river, W.
Irian; Milne Bay Distr., Woitaki, Paiawa, and North-
ern Distr. Papua).

Ecol. Local, especially on ridges, in lowland (in-
cluding Anisoptera and oak Castanopsis) forests; also
recorded from freshwater swamp.

Vern. Lomas (Waigani), koperitoma (Upper
Waria).

43. Hopea forbesii (BRANDIS) SLOoT. Reinwardtia 5
(1961) 477; v. RoYEN, Man. Forest Trees Papua New
Guinea 8 (1965) 34.— Shorea forbesii BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 92; BAKER f. J. Bot. 61,
Suppl. (1923) 5; SLoor. Reinwardtia (1952) 61, f.
20. — Fig. 72 B-B4.

Tall, flaky-barked, hard-wooded tree. Young parts
densely buff pubescent, leaf buds, stipules, twig
apices, petioles, and parts of corolla expanded in bud
persistently so, becoming sparse on leaf undersurface
and calyx, caducous elsewhere. Twig c. 1 mm @
apically, becoming smooth, terete; internodes short,
branchlets hence densely leaved. Buds minute, ovoid;
stipules to 4 by 1 mm, linear-lanceolate, caducous.
Leaves 5.5—11 (to 13 in young trees) by 1.8—4 cm,
lanceolate, thinly coriaceous; margin subrevolute;
base obtuse, + equal; acumen to 13 mm, slender;
nerves (11—)13 to 15 pairs, dense, slender but promi-
nent beneath, + obscure and depressed above as also
the midrib, without domatia; tertiary nerves scalari-
form, + evident but unraised beneath; petiole 5-8 mm
long. Panicle to 3.5 cm long, slender, to 2-axillary,
singly branched; branchlets to 12 mm long, bearing to
5 secund flowers; bracteoles to 2 mm long, deltoid, not
at first caducous. Flower bud to 3 by 2 mm, lanceolate.

418

2 outer sepals narrowly deltoid-acuminate, 3 inner
suborbicular, mucronate. Stamens (15—)16—19,
shorter than style; filaments long, compressed but
slender, tapering to the oblong anthers; appendages c.
3 times length of anthers, slender. Ovary small,
tapering into a somewhat longer stoutly columnar
stylopodium tapering into short style. Fruit pedicel
short, stout. 2 longer calyx lobes to 6.5 by 1.0 cm,
spatulate, obtuse, c. 2 mm broad above the to 8 by 6
mm ovate saccate thickened base; 3 shorter lobes 12
by 8 mm, ovate, similarly saccate, mucronate, shorter
than the nut; or one sometimes to 23 mm long,
spatulate. Nut to 18 by 8 mm, narrowly ovoid,
resinous, tapering, prominently apiculate.

Distr. Malesia: East New Guinea (Central and
Milne Bay Distr.).

Ecol. Locally common in lowland seasonal semi-
evergreen forest, sometimes dominant, especially on
ridges, below 1000 m.

Vern. Tatami, uvami, yala yala, emi sapu, walei.

44. Hopea helferi (DYER) BRANDIS, J. Linn. Soc. Bot.
31 (1895) 62: Ind. Trees (1906) 67; FISCHER, Kew Bull.
(1927) 206; Sym. Gard. Bull. S. S. 8 (1934) 22; ibid. 8
(1935) 274; Mal. For. Rec. 16 (1943) 128, f. 69, 72, pl.
20; SMITINAND, Thai For. Bull. 1 (1954) 18. — Vatica
helferi Dyer, Fl. Br. Ind. 1 (1874) 302. — Shorea
helferi KURZ, Fl. Burma 1| (1877) 119. — H. dealbata
Hance, J. Bot. 15 (1877) 329; LANgEssAN, Pl. Util.
Colon. Fr. 1 (1886) 300; PrerRE, For. Fl. Coch. 4
(1891) t. 246; Heim, Rech. Dipt. (1892) 61; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 62; GugrRIN, FI. Gén. I-C. 1
(1910) 374, fig.; LECOMTE, Bois Indochine (1926) 110;
Foxw. Mal. For. Rec. 10 (1932) 128; Burk. Dict.
(1935) 1189.

Large dark flaky-barked tree with hard wood. Twig
apices, buds, petiole, leaf nervation beneath, stipules,
panicles and perianth outside densely buff puberulent,
+ caducous on twigs, leaf beneath, and calyx. Twig c.
3mm g apically, blackish, becoming ribbed along the
leaf traces; stipule scars obscure. Bud to 3 by 2 mm,
ovoid, acute; stipules to 9 mm long, linear, caducous.
Leaves (S—)10—24 by (2—)4.5—8 cm, oblong-lanceolate
or occasionally oblanceolate, coriaceous, silvery lepi-
dote beneath; base cuneate to occasionally cordate
(especially sucker-shoots and young trees), subequal;
apex shortly broadly acuminate or obtuse; nerves
(12-)14-16 pairs, ascending, rather straight, slender
but prominent beneath, obscure above; tertiary nerves
densely scalariform, evident beneath; midrib promi-
nent beneath, evident but shallowly channelled above;
petiole (S—)7-11 mm, c. 3 mm @, stout. Panicle to 11
cm long, terminal or subterminal axillary, becoming
lax; twice-branched, with to 2 cm long branchlets
bearing to 10 + secund cream flowers. Flower bud to 3
by 2 mm, lanceolate. Outer 2 sepals lanceolate,
subacute; inner 3 shorter, ovate, subacuminate.
Stamens 15; filaments broadly compressed at base,
tapering and filiform distally; appendages slender, c.
15 x length of the narrowly ellipsoid anthers. Ovary
and stylopodium cylindrical, subtruncate, equal in

FLORA MALESIANA

[ser. 1, vole

height but the ovary the broader; style shorter than
either, columnar. Fruit pedicel to 2 mm long, slender. 2
longer fruit calyx lobes to 6.5 by 1.8 cm, spatulate,
obtuse, c. 4 mm wide above the c. 5 by 3 mm small
elliptic saccate base; 3 shorter lobes to 5 by 3 mm,
ovate (occasionally to 18 mm long and distally lorate).
Nut to 10 by 7 mm, ovoid-apiculate.

Distr. Andamans, Burma, Thailand, Cambodia,
and in Malesia: Malaya (Langkawi, S. Kedah, Upper
Perak).

Ecol. Deep soils, especially on sedimentary rocks on
hill slopes and undulating land in semi-evergreen forest.

Uses. A valuable construction timber.

Vern. Lintah bukit (Langkawi), damar mata kuch-
ing, d. siput (Perak).

45. Hopea aptera AsHTON, Gard. Bull. Sing. 31 (1978)
32)

Small, smooth, barked stilt-rooted tree. Twigs,
petioles and panicles + persistently greyish sericeous,
outside of calyx and leaf nervation below caducously
so, parts of petals exposed in bud densely pubescent.
Twig c.2 mm @ apically, becoming terete, + rugulose.
Buds minute, ovoid; stipules fugaceous, not seen.
Leaves 10.5—25 by 4-8 cm, oblong-lanceolate, coria-
ceous; margin subrevolute, base obtuse or shallowly
cordate; acumen to 3 cm long, prominent, slender;
nerves 15-21 pairs, slender but prominent beneath, +
obscurely depressed above, arched towards the mar-
gin, at 55°-70°, without secondaries; tertiary nerves
scalariform, very slender but elevated beneath; midrib
prominent beneath, elevated above; petiole 8-12 mm
long. Panicle to 7 cm long, |-axillary to ramiflorous,
lax, slender; singly branched, branchlets to 1.5 cm
long, bearing to 3 secund flowers. Flower buds to S by 3
mm, ellipsoid, rather long. Sepals subequal, suborbi-
cular, pubescent, fimbriate, patent. Stamens 15, in 3
unequal verticils; filaments compressed and broad at
base, tapering and filiform in the distal 5; anthers
small, subglobose; appendage c. 35 times length of
anthers, very long and slender. Ovary and stylopodium
narrowly hour-glass shaped, with short but distinct
columnar style. Fruit pedicel very short. Calyx lobes to
8 by 6 mm, subequal, ovate, acute, saccate, + thinly
incrassate. Nut to 10 by 6 mm, ovoid, crowned by a
prominent medially swollen stylopodium.

Distr. Malesia: N.W. New Guinea (Vogelkop
Peninsula).

Ecol. Locally common, primary and secondary
forest below 300 m, clay soil including limestone.

Vern. Gamur (Maibrat).

Note. This is one of the several species in New
Guinea which apparently belong to sect. & subsect.
Hopea, yet which share with subsect. Pierrea a large
oblong leaf with more or less unequal base and narrow
tapering, though not spindle-shaped, stylopodium. H.
aptera is distinguished by its short subequal fruit
sepals.

46. Hopea ultima AsHTON, Gard. Bull. Sing. 31 (1978)
S35

1982]

Medium-sized, unbuttressed tree with flaky bark.
Twigs, petioles, panicles and calyx densely pale tawny
puberulent, midrib beneath sparsely so; parts of petals
exposed in bud densely pubescent. Twigs c. 2mm @
apically, terete, + rugose, pale brown, becoming
smooth. Leaves 6-17 by 3-6 cm, oblong, subsericeous;
base subcordate, + equal, margin narrowly revolute;
acumen to 5 mm long, stout; nerves 16-18 pairs,
slender but prominent beneath, at 50°-65°; tertiary
nerves slender, barely elevated, scalariform; midrib
slender but prominent beneath, elevated above;
petiole 9-14 mm long. Panicles to 5(—8) cm long,
3-axillary to ramiflorous or sometimes terminal,
short, slender; |—2 branched, the branchlets bearing to
6 secund flowers. Flower buds to 5 by 2 mm, ellipsoid.
Sepals unequal, the outer 2 lanceolate, inner 3 ovate,
erose. Stamens 15; filaments dilated at base, tapering
medially; anthers subglobose; appendage 23-3 times
length of anthers, aristate. Ovary and stylopodium
narrowly fusiform, tapering into a shorter style. Fruit
unknown.

Distr. Malesia: E. New Guinea (Milne Bay area,
Normanby I.).

47. Hopea novoguineensis SLOoT. Nova Guinea 14
(1924) 224, t. 19; Reinwardtia 2 (1952) 31; v. ROYEN,
Man. Forest Trees Papua New Guinea 8 (1965)
38. — H. celebica (non BURCK) DIELS, Bot. Jahrb. 57
(1922) 462.

Medium-sized, flaky-barked tree with tall but-
tresses. Parts of petals exposed in bud, panicles, bracts
and stipules densely persistently buff pubescent; calyx,
twigs and petiole caducously so. Twig c. 2 mm @
apically, becoming smooth, terete, blackish. Buds
minute, ovoid; stipules to 4 by 2 mm, lanceolate,
acute, caducous. Leaves 10-28 by 3.5—10 cm, variable
in size and shape, chartaceous, undersurface pale and
dull, margin + revolute; base + prominently un-
equal, cuneate abaxially, the adaxial side cuneate to
deeply cordate; acumen to | cm long, slender; nerves
(11-)14-16 pairs, slender but prominent beneath,
narrowly elevated above, arched, at 50°—65°, usually
with + prominent glabrous porous canaliculate
domatia; tertiary nerves densely scalariform, slender,
evident beneath; petiole 9-15 mm long. Panicle to 9 cm
long, to 3-axillary or terminal, singly branched;
branchlets to 2.5 cm long, bearing to 4 flowers;
bracteoles to 2 by 2mm, ovate, acute. Flower buds to 3
by 2 mm, ovoid. Sepals subequal, broadly ovate,
acute. Stamens 15, shorter than style, in 3 subequal
verticils; filaments compressed and broad at base,
tapering to the subglobose anthers; appendages +
equal to length of anthers, slender. Ovary small,
tapering into a distinct equally long subcylindrical
stylopodium and short style. Fruit pedicel to 3 mm
long, stout. 2 longer calyx lobes to 8 by 2 cm,
spatulate, obtuse, c. 5 mm broad above the to 5S by 4
mm ovate saccate thickened base; 3 shorter lobes to 7
by 4 mm, ovate, acuminate, shorter than nut. Nut to
17 by 8 mm, narrowly ovoid, tapering, prominently
apiculate.

DIPTEROCARPACEAE (Ashton)

419

Distr. Malesia: New Guinea (Sorong Distr., and
Munju Subdiv., South New Guinea, W. Irian; Wes-
tern Distr. T.P.N.G.), Moluccas (Halmahera).

Ecol. Common in lowland forest.

Vern. Puwokigih, woigik, wokidjih (Mooi), arid
(Mandobo), kielmun (Muju), tanjung (Halmahera).

Note. Very variable, especially in the depth of the
cordate leaf base and in the lustrousness of the
laminar surface; it is possible that further collections
will show there to be more than one species.

48. Hopeascabra AsHTON, Gard. Bull. Sing. 31 (1978)
33, non BucH.-HAM. Mem. Wern. Soc. 6 (1832) 300,
nom. inval., in syn.

Medium-sized buttressed tree. Twigs, petioles, leaf
buds and stipules + persistently pale rufous scabrid
pubescent, leaf nervation below and midrib above
sparsely but distinctly so; calyx outside fugaceous
puberulent. Twigs c. 2mm g apically, much branched,
becoming terete, smooth, pale brown. Leaf buds
minute, ovoid; stipules to 7 by 3 mm, lanceolate,
caducous. Leaves 6.5-17 by 2.3-6.5 cm, oblong to
lanceolate-falcate, coriaceous; margin subrevolute;
base unequal, cordate; acumen to 1.5 cm long, +
caudate, slender; nerves 15-24 pairs; slender but
prominent beneath, obscurely + shallowly depressed
above, arched, at 70°-80°, with many short but
distinct secondary nerves; tertiary nerves scalariform,
distinctly elevated beneath; midrib slender but promi-
nent beneath, elevated above; petiole 4-6 mm long.
Panicle to 4.5 cm long, short, slender, 1-axillary,
singly branched, branchlets to 13 mm long, bearing to
4 flowers; bracts to 1 mm long, deltoid, acute. Flower
buds at anthesis unknown. Very young fruit with 2
narrowly deltoid-acuminate outer sepals, 3 suborbi-
cular fimbriate obtuse inner sepals, and ovoid ovary
surmounted by a slightly narrower, equally long,
prominent stylopodium and short but distinct style.
Fruit pedicel short. 2 longer calyx lobes to 8 by 1.8 cm,
spatulate, obtuse, 2 mm wide above the to 5 by 4mm
ovate deeply saccate thickened base; 3 shorter lobes to
8 by 6mm, ovate, acuminate, similarly saccate. Nut to
8 by 6 mm, ovoid; stylopodium prominent, tapering.

Distr. Malesia: W. New Guinea (Hollandia Div.,
Madang).

Ecol. Clay soils on undulating land, locally fre-
quent.

49. Hopea papuana Diets, Bot. Jahrb. 57 (1922) 461;
SLoorT. Reinwardtia 2 (1952) 33, f. 11; v. ROYEN, Man.
Forest Trees Papua New Guinea 8 (1965) 40.
Medium-sized tree with dark flaky bark. Twigs,
petioles, leaf buds and stipules persistently rufous
scabrid tomentose; nerves below and midrib on both
surfaces thus at first, + glabrescent. Twigs c. 3mm @
apically, stout, + persistently ribbed, pale brown;
stipule scars prominent, descending. Buds to 2 by 2
mm, ellipsoid, obtuse; stipules to 12 by 5 mm,
lanceolate, subpersistent. Leaves 11—28 by 4.2-10 cm,
large, oblong, coriaceous; base + unequal, cordate;
acumen to 2.5 cm long, tapering; nerves 16-24 pairs,

420

arched towards the margin, at 70°—80°, slender but
prominent beneath, applanate above, without doma-
tia or secondaries; tertiary nerves scalariform, barely
evident beneath; midrib prominent beneath, elevated
above; petiole 5-7 mm long, stout. Complete panicle
unknown, short, l-axillary, glabrous. Flowers un-
known. Fruit pedicel short, stout. 2 longer calyx lobes
to 5.5 by 1.8 cm, broadly spatulate, obtuse, c. 2 mm
broad above the to 6 by 5 mm ovate saccate thickened
base; 3 shorter lobes to 9 by 6 mm, ovate, subacute,
similarly saccate. Nut to 16 by 9 mm, narrowly ovoid,
tapering to a prominent stout apiculate stylopod-
ium.

Distr. Malesia: New Guinea (Hollandia Div.; R.
Digul, Muju, in S. New Guinea; Idenburg R.; Papua
New Guinea: Central Distr., Amau hinterland, and
Western Highlands).

Ecol. Locally abundant; alluvium, low hills, river
banks (W. Irian); once on ridges with Araucaria
klinkti in Papua New Guinea.

Vern. Gointa (Western Highlands), riheu (Nemo),
pasang kesereep (Djair), keilmun, ogerie (Muju), linak-
iong (Manikiong), matre (Berak).

50. Hopea acuminata Merk. Philip. Gov. Lab. Bur.
Bull. 29 (1905) 30; Philip. J. Sc. 1 (1906) Suppl. 98; En.
Philip. 3 (1923) 93; WHITFORD, Philip. J. Sc. 4 (1910)
Bot. 703; Bull. Bur. For. Philip. 10 (1911) 75; Foxw.
Philip. J. Sc. 2 (1907) Bot. 389; ibid. 4 (1909) Bot. 514;
ibid. 6 (1911) Bot. 264; ibid. 13 (1918) Bot. 183; REYES,
Philip. J. Sc. 22 (1923) 339; Sym. Gard. Bull. S. S. 8
(1934) 22; GUTIERREZ, Act. Manil. 4, A, 2 (1968) 47, f.
9, pl. 5. — H. maquilingensis Foxw. Philip. J. Sc. 13
(1918) Bot. 184. — Fig. 64a.

Medium-sized tree with flaky bark. Young parts
fugaceous puberulent, panicle persistently so or gla-
brous, domatia and parts of petals exposed in bud
persistently so. Twig c. 1mm @ apically, slender, much
branched, terete, rugulose, dark brown. Buds minute;
stipules fugaceous, not seen. Leaves 4.5—12 by 2-4.5
cm, elliptic-falcate to ovate-lanceolate, thinly coria-
ceous; base unequal, cuneate; acumen to | cm long,
slender, tapering; nerves 9-11 pairs, slender but
prominent beneath, + applanate above, arched, at
45°-65°, with (young trees) or without small axillary
pubescent domatia; tertiary nerves densely scalari-
form, very slender, barely elevated beneath; petiole
6-8 mm long. Panicle to 3.5 cm long, slender, terminal
or to 2-axillary, singly branched; branchlets bearing to
4 cream flowers; bracteoles fugaceous. Flower buds to
3 by 2 mm, ellipsoid. 2 outer sepals lanceolate-acu-
minate; 3 inner sepals suborbicular, mucronate.
Stamens 10, equal; filaments compressed, tapering,
rather broad; appendages aristate, 14 times as long as
the oblong anthers. Ovary and stylopodium broadly
cylindrical, truncate, slightly tapering, densely puber-
ulent; style shorter than ovary but prominent. Fruit
pedicel c. 2 mm long, slender. 2 longer calyx lobes to
5.5 by 1 cm, spatulate, obtuse, c. 2 mm broad above
the to 4 by 4 mm elliptic saccate thickened base; 3
shorter lobes to 3 by 2 mm, elliptic, similarly saccate.

FLORA MALESIANA

[ser. I, vol. 9?

Nut to 6 by 6mm, broadly ovoid, with minute slender
apical style remnant.

Distr. Malesia: Philippines.

Ecol. Widespread and common in semi-evergreen
and evergreen forests, especially at 300-800 m but
down to 100 m.

Vern. Baniakau (Ibn.), barosingsing (Ilk.), daling-
ding (Tag.), dalingdingan (Tag., S.L.Bis.), ka/or (I1k.),
manggachapui (official and general name), mangga-
chinoro (Tag.), siayu (S.L.Bis.), yakal (Tag.).

Note. Allied to the widespread H. sangal which it
replaces in the Philippines.

51. Hopea depressinerva ASHTON, Gard. Bull. Sing. 22
(1967) 275, pl. 20; Man. Dipt. Brun. Suppl. (1968) 50,
teenie

Medium-sized tree with slightly cracked bark.
Young parts glabrescent. Twig 1-2 mm @ apically,
terete, smooth; stipule scars short, obscure. Bud to |
by 1 mm, minute. Stipule unknown. Leaves 5—13 by
2-5 cm, lanceolate to narrowly elliptic, coriaceous,
base cuneate; acumen to 1.5 cm long, slender; nerves
6-8 pairs, slender but raised beneath, depressed
above, at 40°-S0°; tertiary nerves slender, densely
scalariform; midrib prominently terete beneath,
slender, somewhat depressed, above; petiole 11-13
mm long, terete. Panicle to 7 cm long, terete, terminal
or axillary, densely persistently buff pubescent; singly
branched, branchlets to 1.5 cm long, bearing to 5
secund flowers; bracteoles fugaceous, unknown.
Flower bud to 3 by 2 mm, ellipsoid. Calyx densely buff
sericeous, lobes ovate, acute, the outer 2 somewhat
longer, relatively narrower, than the inner 3. Petals
elliptic, obtuse, densely pubescent on parts exposed in
bud, pink outside, pale orange within. Stamens 10,
equal, forming a ring round the ovary; filaments
broad, compressed at base, tapering and filiform
below the subglobose anthers; appendages to connec-
tives c. 2 times length of anthers, filiform, slender.
Ovary and stylopodium glabrous, cylindrical, trun-
cate, surmounted by a short style. Mature fruit
unknown; calyx lobes unequal, the 2 longer aliform,
considerably larger than the other 3.

Distr. Malesia: Borneo (W. Sarawak).

Ecol. Rare, granodiorite hill slopes below 500 m.

52. Hopea sangal KorTHu. Kruidk. (1841) 75; WALP.
Rep. 5 (1845) 128; Ann. 4 (1857) 339; BL. Mus. Bot.
Lugd.-Bat. 2 (1852) 34; Mia. Fl. Ind. Bat. 1, 2 (1859)
504; DC. Prod. 16, 2 (1868) 635; SLooT. ex HEYNE,
Nutt. Pl. ed. 2 (1927) 1111; Sym. Gard. Bull. S. S. 8
(1934) 18, pl. 5; Mal. For. Rec. 16 (1943) 141. f. 68C,
69, 77; BROWNE, For. Trees Sarawak & Brunei (1955)
121; BACKER & Baku. f. FI. Java 1 (1963) 331;
ASHTON, Gard. Bull. Sing. 20 (1963) 260; Man. Dipt.
Brun. (1964) 108, f. 12, pl. 25 (bark); ibid. Suppl.
(1968) 56; MeER & Woop, Sabah For. Rec. 5 (1964)
56, pl. 21b (stem), f. 31.— Dryobalanops sericea
Kort. Kruidk. (1841) 72; DC. Prod. 16, 2 (1868)
606; WALP. Rep. 5 (1845) 125. — H. sericea BL. Mus.
Bot. Lugd.-Bat. 2 (1852) 35; WALP. Ann. 4 (1857) 339;

1982]

DIPTEROCARPACEAE (Ashton)

421

Mia. FI. Ind. Bat. 1, 2 (1859) 504; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 85; DC. Prod. 16, 2 (1868) 635:
Burck, Ann. Jard. Bot. Btzg 6 (1887) 238: BRANDIs, J.
Linn. Soc. Bot. 31 (1895) 64; MerR. En. Born. (1921)
403. — Petalandra micrantha Hassk. Hort. Bog.
Desc. (1852) 105; Mig. Fl. Ind. Bat. 1, 2 (1859) 505;
DC. Prod. 16, 2 (1868) 636; KING, J. R. As. Soc. Beng.
Sc. 62, 2 (1893) 126. — H. fagifolia Mig. Sum. (1862)
490; DC. Prod. 16, 2 (1868) 635; WALP. Ann. 7 (1868)
379; SCHEFF. Nat. Tijd. N. I. 31 (1870) 351; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 65; BoerRL. Cat. Hort. Bog. 2
(1899) 103; K. & V. Bijdr. 5 (1900) 124: MoLL &
JANSSONIUS, Mikogr. Holz. (1906) 369; HEYNE, Nutt.
Pl. 1, 3 (1917) 290; ed. 2 (1927) 1111; Sym. Gard. Bull.
S. S. 7 (1933) 151, 154. — H. diversifolia (non MiQ.)
ScHEFF. Nat. Tijd. N. I. 31 (1870) 351; Burck, Ann.
Jard. Bot. Btzg 6 (1887) 239; BrRaANpis, J. Linn. Soc.
Bot. 31 (1895) 64; Foxw. Mal. For. Rec. 10 (1932)
123. — H. odorata (non Roxs.) HANCE, J. Bot. 5
(1876) 308, et auct. (1876-1927) p.p. quoad syn. H.
sangal. — Doona odorata (ROxB.) BURCK, Ann. Jard.
Bot. Btzg 6 (1887) 233, p. min. p., quoad syn.
H. sangal.— Doona’~ micrantha Burck, L.c.
234. — Doona javanica BurRcK, I.c. 235, t. 29, f.
7. — Dryobalanops neglectus KORTH. ex BURCK, /.c.
243. nomen pro syn.— H. micrantha {non (HAssK.)
Hook. f.] Hem, Rech. Dipt. (1892) 64. — H. hasskar-
liana Hem, I.c. 64.— H. javanica HEM, l.c. 64;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 64. — H. curtisii
KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 124;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 65; BRUHL &
KING, Ann. Bot. Gard. Calc. 5, 2 (1896) 155, t. 187;
Burk. J. Str. Br. R. As. Soc. 81 (1920) 66, fig.; RIDL.
Fl. Mal. Pen. 1 (1922) 212, 236, fig.; Foxw. Mal. For.
Rec. 10 (1932) 130; Burk. Dict. (1935) 1189. — H.
globosa BRANDIS, J. Linn. Soc. Bot. 31 (1895) 61:
RIDL. J. Str. Br. R. As. Soc. 54(1910) 26; Fl. Mal. Pen.
1 (1922) 236; HEYNE, Nutt. Pl. ed. 2 (1927) 1104;
Foxw. Mal. For. Rec. 10 (1932) 121; Burk. Dict.
(1935) 1190. — H. lowii DYER ex BRaNpis, J. Linn.
Soc. Bot. 31 (1895) 63; RipL, J. Str. Br. R. As. Soc. 73
(1916) 143; Fl. Mal. Pen. 1 (1922) 237; Merr. En.
Born. (1921) 402: Foxw. Mal. For. Rec. 10 (1932)
123. — H. minutiflora C.E.C.FISCHER, Kew Bull. 1
(1927) 207; SMITINAND, Nat. Hist. Bull. Siam Soc. 19
(1958) 74. — H. multiflora (non BRANDIS) Foxw. Mal.
For. Rec. 10 (1932) 110, p.p. — H. albescens (non
RIDL.) Foxw. Mal. For. Rec. 10 (1932) 122. — Fig.
15, 72 E1-E4.

Medium-sized to large, dark flaky-barked, but-
tressed tree with prominent opaque white resin exu-
dates on bole. Young twig, leaf bud, stipule, petiole,
leaf undersurface, midrib above and panicle + spar-
sely pale grey-brown pubescent, leaf sometimes gla-
brescent. Twig to 1 mm @ apically, slender, much
branched, smooth. Bud to 2 mm long, ovoid. Stipule
to 3 mm long, linear, fugaceous. Leaves 5.5—10 by
3.5-5 cm, ovate, thin, base + broadly cuneate;
acumen to 1.2 cm long, caudate; nerves 10-12 pairs,
slender, prominent beneath, slightly curved, at
50°—60°, with or without tomentose axillary domatia;

tertiary nerves very slender, scalariform, indistinct,
oblique to nerves; midrib slender, prominent beneath,
+ applanate above; petiole 0.5—1 cm long. Panicle to 7
cm long, terminal or axillary, lax, terete; singly or
doubly branched, branchlets bearing to 8 secund
flowers; bracteoles to 1 mm long, small, deltoid, buff
pubescent. Flower buds very small, ellipsoid-ovoid.
Calyx densely pubescent on both surfaces; 2 outer
lobes deltoid, subacute, 3 inner lobes suborbicular,
obtuse. Petals oblong-lanceolate, densely tomentose
on parts exposed in bud, cream. Stamens 10; filaments
slender, tapering; anthers oblong; appendage as long
as anther, slender. Ovary and stylopodium short.
broadly cylindrical, truncate, broader towards base,
puberulent at apex; style 3 length of ovary and
stylopodium, filiform, glabrous. Fruit calyx glabres-
cent; 2 longer lobes to 7 by 1.5 cm, oblong-spatulate,
obtuse, tapering to 4mm broad above the 5 by 4mm
somewhat saccate thickened base; 3 shorter lobes to 7
by 4mm, obtuse. Nut to 7 by 4mm, ovoid, persistently
subtruncate at the short style remnant, sparsely
pubescent.

Distr. Peninsular Thailand, and in Malesia:
Malaya, Sumatra, Banka, Billiton, W. Java (rare),
Lesser Sunda Is. (Bali, rare), Borneo.

Ecol. Locally common on clay rich soils on river
banks; scattered on fertile clay hillsides to 500 m.

Vern. Mérsiput, chengal pasir, c. mata kuching,
damar siput, mérawan hitam (Mal.), gagil (Sabah), m.
télor, m. batu, tongon banwah, chéngal, c. hitam, c.
bunga, damar bintang, d. géndiran, timbalun (Suma-
tra), awang awang, d. lampung, d. item, d. puteh, d.
kunyit, lampong gunung, lantang arong, tékam, t. paya
(Borneo).

53. Hopea ferrea LANEss. Pl. Util. Colon. Fr. 1 (1886)
300; PIERRE, For. Fl. Coch. 4 (1891) t. 249: Hem,
Rech. Dipt. (1892) 62; BRANDIS & GiLG in E. & P. Pfl.
Fam. ed. 1, 3, 6 (1894) 262; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 61; GuErtn, Fl. Gén. I.-C. 1 (1910) 371, 377,
fig.; Foxw. Mal. For. Rec. 10 (1932) 126; BurK. Dict.
(1935) 1189; Sym. Mal. For. Rec. 16 (1943) 124, f.
68H, 69. — Balanocarpus anomalus KiNG, J. R. As.
Soc. Beng. Sc. 62, 2 (1893) 132; BRANDis, J. Linn. Soc.
Bot. 31 (1895) 109; RipL. Fl. Mal. Pen. 1 (1922)
247. — H. anomala Foxw. J. Mal. Br. R. As. Soc. 5
(1927) 340; Mal. For. Rec. 10 (1932) 126. — Fig. 72
C-C4.

Small, occasionally stout and large, shaggy-barked
trees, often twisted and gnarled. Panicle and outside
of petals and sepals densely buff puberulent, otherwise
glabrous. Twig c. 1 mm g, slender, slightly zig-zag,
drying dark brown, minutely rugulose; stipule scars
obscure. Leaf bud minute; stipules fugaceous. Leaves
4-10 by 2-5.5 cm, ovate; base + broadly cuneate or
rarely obtuse; acumen to 2 cm long, slender; nerves
(6—-)8(-9) pairs, arched, slender but distinctly raised
beneath, evident above as also the midrib, frequently
with minute glabrous axillary domatia; tertiary nerves
densely scalariform, slender, evident beneath, obscure
above: petiole 9-13 mm long, slender. Panicle to 8 cm

422

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 73. Hopea odorata Roxs. a. Flowering branch, with domatia on leaves visible, b. leaf with infructescence, c.
fruit, d. nut, all x 2/3 (a BKF 40492, the domatia from BKF 51769, 6 Cult. Hort. Bog. VIII-A-26, c Cult. Hort.
Bog., sheet 902.146—395 in L).

long, slender, with to 3 cm long branchlets bearing
many (to 12) secund cream flowers. Flower bud to 3 by
2 mm, small, ovoid. Sepals broadly ovate, subacu-
minate, subequal. Stamens 15, the filaments lorate but
becoming filiform beneath the subglobose anthers;
appendages somewhat shorter than anther. Ovary and
stylopodium pyriform, glabrous; style short, obscure.
Fruit pedicel to 2 by | mm, broadening into receptacle.
2 longer calyx lobes to 4 by 1 cm, broadly spatulate,
obtuse, c. 2mm broad above the to 3 by 1 mm minute
thickened saccate base; 3 shorter lobes to 5 by 1 mm,
lanceolate-acicular, thickened, saccate. Nut to 13 by 4
mm, cylindrical, tapering abruptly to a terminal
apiculus.

Distr. Indochina, Thailand, and in Malesia: N. W.
Malaya (Kedah, Perlis, Langkawi).

Ecol. Locally abundant on rocky ridges and slopes,
especially on limestone.

Vern. Malut (Langkawi), chéngal laki (Kedah,
Perlis).

54. Hopea odorata Roxs. [Hort. Beng. (1814) 42,
nomen] PI. Corom. 3 (1819) 7, t. 210; Fl. Ind. ed. Carey
2 (1832) 609; WALP. Rep. 5 (1845) 128; Dyer, FI. Br.
Ind. | (1874) 308; PreRRE, For. Fl. Coch. 4 (1891) t.
244; BRANDis, J. Linn. Soc. Bot. 31 (1895) 59; Ind.
Trees (1906) 67; GUERIN, FI. Gen. I.-C. 1 (1910) 373;
HEYNE, Nutt. Pl. ed. 1, 3 (1917) 292; Troup, Silv. Ind.
Trees 1 (1921) 47; Cras, Fl. Siam. Enum. | (1925)
147; LECOMTE, Bois Indochine (1926) 11; Foxw. Mal.
For. Rec. 2 (1927) 75; ibid. 10 (1932) 118; Burk. Dict.
(1935) 1192; CoRNER, Wayside Trees 1 (1940) 212;

1982]

Sym. Mal. For. Rec. 16 (1943) 137, f. 69; SMITINAND,
Thai For. Bull. 1 (1954) 18. — H. vasta WALL. [Cat.
(1828) 962, nomen] ex DC. Prod. 16, 2 (1868)
633. — H. decandra BucH.-HaM. ex WIGHT, Ill. Ind.
Bot. 1 (1840) 88. — Fig. 73.

Tall buttressed tree with dark brown flaky bark.
Twig apices sparsely fugaceous pale buff puberulent;
panicle, leaf buds, flower calyx and petals outside
densely persistently so; fruit calyx lobes and fre-
quently nut glabrescent. Twig c. 2 mm @ apically,
terete, dark brown, smooth to rugulose; stipule scars
pale, minute. Buds minute, ovoid; stipules minute,
fugaceous. Leaves 7-14 by 3—7 cm, ovate-lanceolate,
falcate; base broadly cuneate, unequal; acumen to 1.5
cm long, broad; nerves 9-12 pairs, prominent
beneath, narrowly depressed above, arched, with
prominent porous saccate axillary domatia; tertiary
nerves densely sinuate-scalariform, very slender but
clearly evident on both surfaces; midrib prominent
beneath, applanate to somewhat channelled above;
petiole 10-16 mm long, slender. Panicle to 12 cm long,
terminal or axillary, twice-branched, with many to 2.5
cm long branchlets bearing to 9 secund flowers.
Flower buds to 3 by 2 mm, broadly ovoid. Outer 2
sepals lanceolate, obtuse or subacuminate; inner 3
broadly ovate, acute. Petals pale cream-yellow.
Stamens 15; filaments slender, compressed at base,
tapering; appendage slender, as long as narrowly
ellipsoid anthers. Ovary ovoid, punctate or glabrous,
surmounted by an equally tall columnar style. Fruit
pedicel to 1 mm long, short, slender. 2 longer calyx
lobes to 5.5 by 2cm, + broadly spatulate, obtuse, c. 3
mm broad above the to 4 by 4 mm small elliptic
saccate base; 3 shorter lobes to 4 by 4 mm, ovate,
subacuminate. Nut to 6 by 5 mm, small, ovoid,
subobtuse.

Distr. Andamans, Burma, Thailand, Indochina
and in Malesia: Malaya (N. Perak & Trengganu
northwards).

Ecol. Common, often gregarious, on river banks in
Malaya.

Uses. Cultivated as a shade tree.

Vern. Chéngal pasir, c. kampong, c. pulau, c. mas.

Note. Allied to H. parviflora BEDD. of the Western
Ghats, India.

55. Hopea centipeda AsHTON, Gard. Bull. Sing. 22
(1967) 274, pl. 19; Man. Dipt. Brun. Suppl. (1968) 48,
pl. 11 (stem-base). — H. acuminata (non MERR.) ASH-
TON, Man. Dipt. Brun. (1964) 94. — Fig. 72 D1-D4.

Medium-sized, prominently stilt-rooted tree with
smooth bark. Young twig, leaf bud, stipule (both
surfaces), petiole, and nervation beneath densely
shortly persistently brown pubescent. Twig to 1 mm @
apically, branching horizontally, smooth, glabrous.
Bud to 1.5mm !ong, subglobose. Stipule to 3mm long,
linear. Leaves 5.5—9 by 1.5—3.5 cm, lanceolate; base
unequal, cuneate; acumen to 1.5 cm long, caudate;
nerves 7—9 pairs, prominent beneath, slightly curved,
at 30°-40°, with prominent pilose axillary domatia;
tertiary nerves slender, scalariform, with slight reticu-

DIPTEROCARPACEAE (Ashton)

423

lations, diagonal to nerves; midrib prominent,
rounded beneath, narrow, depressed above; petiole
4-7 mm long, slender. Panicle to 2 cm long. Flowers
cream; bud to 6 by 2 mm, lanceolate. Calyx sparsely
pubescent; sepals ovate, acute, the outer 2 somewhat
longer than the inner 3. Stamens 15, in unequal
verticils; anthers subglobose; appendage to connec-
tive c. 3 times length of anthers, slender. Ovary and
stylopodium pyriform, glabrous, surmounted by a
prominent style equalling the stylopodium in length.
Fruit calyx puberulent, glabrescent; 2 longer lobes to
2.8 by 0.6 cm, spatulate, obtuse, to 1.5 mm broad
above the to 2.5mm by 2 mm elliptic shallowly saccate
thickened base; 3 shorter lobes to 4 mm long, obtuse,
similar at base. Nut to 4 by 3 mm, ovoid, glabrous.

Distr. Malesia: Borneo (Rejang hinterland to
Lawas, S. E. Sabah).

Ecol. Local, on banks of fast flowing inland rivers.

Vern. Mérawan daun bérbulu.

56. Hopea plagata (BLCO) VIDAL, Sinopsis (1883) t.
15A; Rev. Pl. Vasc. Filip. (1886) 62; BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 64; MeRR. Publ. Govt. Lab. Philip.
27 (1905) 22; MERRITT, Bull. Bur. For. Philip. 8 (1908)
48: WHITFORD, Philip. J. Sc. 4 (1910) 715; Bull. Bur.
For. Philip. 10 (1911) 73; Merr. Sp. Blanc. (1918)
2691; En. Philip. 3 (1923) 94; Reyes, Philip. J. Sc. 22
(1923) Bot. 447; GUTIERREZ, Act. Manil. 4, A, 2 (1968)
55, f. 11, pl. 7; ASHTON, Gard. Bull. Sing. 31 (1978)
32. — Mocanera plagata Buco, FI. Filip. ed. 1 (1837)
447. — Dipterocarpus plagatus Buco, FI. Filip. ed. 2
(1845) 311; ibid. ed. 3, 2 (1878) 212; DC. Prod. 16, 2
(1868) 614.— Anisoptera plagata Bi. Mus. Bot.
Lugd.-Bat. 2 (1856) 42; WaLP. Ann. 4 (1867) 336; DC.
Prod. 16, 2 (1868) 616. — Shorea reticulata (non THW.
ex Dyer) F.-VILL. Nov. App. (1880) 21. — H. odorata
(non Roxs.) Foxw. Philip. J. Sc. 13 (1918) Bot.
183. — H. sp. ‘““Gyam’’ Foxw. Philip. J. Sc. 6 (1911)
Bot. 263. — H. dasyrrachis (non SLooT.) ASHTON,
Man. Dipt. Brun. Suppl. (1968) 49, f. 7. — Fig. 72
A-A4.

Large flaky barked buttressed tree. Parts entirely
glabrous or ovary pubescent. Twig c. | mm @ apically,
much branched, terete, becoming smooth, dark
brown. Bud c. 2 by 1 mm, ovoid, stipules fugaceous,
not seen. Leaves 6-12 by 2.5—7 cm, elliptic-lanceolate
to ovate, + falcate, coriaceous; base cuneate to
obtuse, + markedly unequal; acumen to 1.5 cm long,
tapering; nerves 8-11(—12) pairs, slender, elevated
beneath, + applanate above, ascending at 35°-65",
sometimes pale stellate lepidote beneath, usually with
glabrous pore-like domatia; tertiary nerves densely
scalariform, hardly elevated beneath; midrib +
prominent on both surfaces; petiole 6-16 mm long.
Panicle to 3 cm long, slender, 1-axillary or terminal,
singly branched; branchlets to 2 cm long, bearing to 6
secund flowers; bracteoles fugaceous. Flower bud to 3
by 2 mm, ellipsoid, 2 outer sepals broadly ovate,
subacute; 3 inner suborbicular, obtuse. Stamens c. 35,
subequal, slightly shorter than style; filaments long,
slender, narrowly compressed at base, tapering and

424

filiform in the distal 4; anther elongate, tapering;
appendages + equal in length to anthers; acicular.
Ovary ovoid, with obscure stylopodium and short
broad style. Fruit pedicel to 2 mm long; 2 longer calyx
lobes to 45 by 20 mm, broadly oblong-spatulate,
sometimes suborbicular; 3 shorter lobes to 7 by 4mm
ovate. Nut to 10 by 7 mm, narrowly ovoid, +
pubescent or glabrous.

Distr. Malesia: Philippines (Luzon, Mindoro,
Tablas, Basilan, Tawitawi; rare in Mindanao); N. E.
Borneo.

Ecol. Widespread and common in semi-evergreen
seasonal forests, more local in evergreen non-seasonal
forests, on limestone in Sarawak.

Uses. The most widespread heavy construction
timber in the Philippines.

Vern. Banutan (Gad.), batik (Ilk.), gisok gisok
(Bik.), haras (P. Bis.), nutik (Buk.), paina (Tag.),
panggian (Ilk.), saplingan, siakal (Tag.), siggai, s. na
nalabaga, s. mulata, s. apuran (Ik.), tagai (Ik. Neg.),
yakal (Ilk., Pang., Sbl., Tag., Sul.), yakal saplingan
(official name).

Note. Two flowering collections, LOHER 12914 and
14901 from Luzon, differ in possessing flowers with
pubescent ovary and 15 stamens. They could, when
more material is available, merit separation as a
separate taxon, and serve to underline the close
affinity of this species to H. dasyrrhachis SLOOT. (q.v.)
whose distinct suborbicular fruit sepals occur some-
times in the present species; indeed sterile and fruiting
collections from Sarawak attributed here to H. dasyrr-
hachis may in fact belong to H. plagata.

57. Hopea nutans RipL. Fl. Mal. Pen. 1 (1922) 235;
Foxw. Mal. For. Rec. 10 (1932) 123; BurK. Dict.
(1935) 1191; Sym. Mal. For. Rec. 16 (1943) 136, f. 69,
76: BROWNE, For. Trees Sarawak & Brunei (1955) 125;
ASHTON, Man. Dipt. Brun. (1964) 104, f. 12; ibid.
Suppl. (1968) 54; MEVER & Woop, Sabah For. Rec. 5
(1964) 217, f. 29. — H. lowii (non DyER) Foxw. Mal.
For. Rec. 1 (1922) 67.

Medium-sized to large flaky-barked buttressed tree.
Young twig, leaf bud, stipule, panicle and petiole very
shortly sparsely pale brown pubescent. Twig to 2mm
@ apically, much branched, smooth, becoming gla-
brous. Bud to 1.5 mm long, ovoid. Stipule to 2.5 mm
long, linear, fugaceous. Leaves 8-13 by 4.5-8.5 cm,
broadly ovate, coriaceous, greyish lepidote below in
mature tree; base obtuse; acumen to | cm long; margin
frequently slightly revolute; nerves 7-10 pairs, dis-
tinct, hardly raised beneath, curved, at 50°—60°,
usually with large glabrous swollen pore-like domatia;
tertiary nerves slender but distinct, scalariform with
slight reticulations, diagonal to nerves; midrib broad,
slightly raised on both surfaces; petiole 1-1.5 cm long.
Panicle to 7 cm long, terminal or axillary, terete or
angular; singly branched, branchlets to 1.5 cm long,
bearing to 5 secund pale yellow flowers; bracteoles to 3
mm long, hastate, subacute. Flower bud to 4 by 2.5
mm, broadly ellipsoid, obtuse. Calyx densely pubes-
cent outside, glabrous within; 2 outer lobes hastate,

FLORA MALESIANA

[ser. I, volage

subacute; 3 inner lobes broadly ovate, thinner, taper-
ing at base. Petals linear, puberulent on parts exposed
in bud. Stamens 15, in 3 subequal whorls; filaments
broad at base, tapering and filiform below anthers;
anthers subglobose, tapering apically; appendage to
connective c. 2 times length of anther, slender,
scabrous towards apex. Ovary and stylopodium subcy-
lindrical, glabrescent; style short, glabrous. Fruit
calyx sparsely pale brown pubescent or glabrescent; 2
longer lobes to 8 by 1.5 cm, oblong, thinly coriaceous,
broad, obtuse, tapering to the to 2.5 by 3 mm ovate
saccate thickened base; 3 shorter lobes to 10 mm long,
acute, similar at base. Nut to 15 by 8 mm, ovoid,
glabrous; style remnant, short, acute.

Distr. Malesia: Malaya (E. coast from Trengganu
southwards), Borneo (Sarawak, Sabah, Nunukan).

Ecol. Sandy soils, often periodically swampy, near
and on coastal hills.

Vern. Giam, chéngal, c. batu, c. kéras, c. pélandok,
téngkawang (Mal.), garang buaya daun kéchil (Brun.).

58. Hopea bancana (BoERL.) SLOOT. Reinwardtia 3
(1956) 317. — Balanocarpus bancanus BOERL. Cat.
Hort. Bog. (1901) 111.

Medium-sized flaky-barked hard-wooded tree.
Twigs, petiole, midrib above and calyx outside cadu-
cous buff pubescent, persistent in young trees and on
parts of petals exposed in bud and leaf buds. Twig c. 2
mm @ apically, terete, rugulose, blackish. Bud minute;
stipules not seen. Leaves 8-15 by 3.5—7.5 cm, ovate-
falcate, thinly coriaceous, lustrous; base + broadly
cuneate to obtuse, frequently shortly decurrent; un-
equal; acumen to 1.5 cm long, tapering; nerves 6-8
pairs, slender but distinctly elevated on both surfaces
(more so below) as also the midrib, arched, at 45°—60°,
with prominent glabrous porous canaliculate domatia
(absent in immature trees); tertiary nerves densely
scalariform, very slender, + elevated on both sur-
faces; petiole 11-14 mm long (shorter in young trees),
slender. Panicle to 8 cm long, slender, pendant, to
2-axillary or terminal, singly branched; branchlets to 2
cm long, bearing to 5 flowers; bracts fugaceous.
Flower bud to 3 by 2 mm, ellipsoid. 2 outer sepals
ovate, acute, 3 inner ovate, acuminate. Stamens 15, in
3 unequal verticils; filaments compressed but rather
narrow at base, tapering to the narrowly ellipsoid
subacute anthers; appendage as long as anthers,
relatively stout. Ovary and stylopodium stoutly pyri-
form, subtruncate, with short columnar style. Fruit
pedicel c.2mm long, stout. 2 longer calyx lobes to 9 by
2.5m, lorate, obtuse, c. 5mm broad and subrevolute
above the c. 5 by 5 mm broadly ovate saccate
thickened base; 3 shorter lobes to 6 by 6 mm, ovate,
acute, similarty saccate. Nut to 9 by 6 mm, ovoid,
shortly apiculate, resinous.

Distr. Malesia: Central W. Sumatra (P. Musala).

Ecol. Very rare, low hills.

Note. Most collections come from a tree in Kebun
Raya Indonesia, the provenance of which was said to
be Banka, though no collections exist from that
island.

1982]

59. Hopea pentanervia SYM. ex Woop, Gard. Bull.
Sing. 17 (1960) 495; BRowneg, For. Trees Sarawak &
Brunei (1955) 126; ANDERSON, Gard. Bull. Sing. 20
(1963) 157; ASHTON, Man. Dipt. Brun. (1964) 106, f.
12, pl. 31 (bark); ibid. Suppl. (1968) 55; MEER &
Woop, Sabah For. Rec. 5 (1964) 219, f. 30.

Medium-sized, flaky-barked buttressed _ tree.
Young parts puberulent, glabrescent. Twig to 1.5 mm
@ apically, terete, much branched, smooth. Bud to 1.5
mm long, small. Stipule to 2 mm long, linear,
fugaceous. Leaves 5S—10 by 3.2-5 cm ovate, coria-
ceous; base obtuse or broadly cuneate; acumen taper-
ing, to 1.5 cm long, narrow, margin slightly revolute;
nerves c. 5 pairs, prominent beneath, slender, at
45°-S5° but strongly curved, with small glabrous
pore-like domatia; tertiary nerves slender, distinct,
densely scalariform, at 90° to midrib; midrib slender,
raised beneath, + applanate above; petiole 6-11 mm
long. Panicle to 8 cm long, terminal or to 2-axillary,
terete, lax; regularly singly or doubly branched,
branchlets to 1.5 cm long, bearing to 6 secund flowers;
bracteoles small, narrowly deltoid, glabrous, cadu-
cous. Flower bud small, ellipsoid, on c. 1.5 mm long
pedicel. Calyx glabrous but for fimbriate margin; 2
outer lobes oblong, acute; 3 inner lobes short, ovate,
acuminate. Petals narrowly lanceolate, shortly pubes-
cent on parts exposed in bud. Stamens 15, small, well
spaced round the ovary; filaments broad at base,
tapering abruptly and filiform distally; anthers sub-
globose; appendage to connective c. 3 times length of
anthers, very slender. Ovary and stylopodium cylindri-
cal, truncate, glabrous; style short, abrupt, slender.
Fruit calyx glabrous; 2 longer lobes to 5 by 1.2 cm,
spatulate, chartaceous, obtuse, to 3 mm broad above
the to 3 by 3 mm small ovate saccate somewhat
thickened base; 3 shorter lobes to 4 by 3 mm, ovate,
acute, saccate. Nut to 4 by 3.5 mm, ovate, glabrous;
style remnant short.

Distr. Malesia: Borneo (Sarawak to W. Sabah,
Sandakan).

Ecol. Mixed peat swamp forest over sand, podsols
on cuestas, plateaux and terraces, near present or
Pleistocene coastlines; on ultrabasic rocks in E.
Sabah.

Vern. Sélangan lima urat (Sabah), mang, m. bési
(Sar.), chéngai paya (Iban).

60. Hopea basilanica Foxw. Philip J. Sc. 6 (1911) Bot.
260, pl. 42; ibid. 13 (1918) 183; Merr. En. Philip. 3
(1923) 93; GUTIERREZ, Act. Manil. 4, A, 2 (1968) 53, f.
10, pl. 6.

Tall, flaky-barked, hard-wooded tree. Young twig,
petiole, midrib above, panicle, calyx and corolla
outside and ovary greyish cinereous, persisting only on
corolla outside. Twig 1-2 mm @ apically, terete, much
branched, dark brown. Leaves (7.5—)10-14 by
(2.5—)3.5—5 cm, oblanceolate or elliptic, thickly coria-
ceous; base cuneate, unequal; acumen to 15 mm long,
prominent, slender; nerves (9—)10—13 pairs, slender
but prominent beneath, applanate above, with promi-
nent glabrous pore-like domatia in the axils of the

DIPTEROCARPACEAE (Ashton)

425

basal 2-3 pairs; tertiary nerves scalariform, evident
but hardly elevated on either surface; petiole 6-10 mm
long, short. Panicles to 7 cm long, to 3 axillary or
ramiflorous, slender, singly branched; branchlets to
17 mm long, bearing to 7 secund flowers. Flower bud
to 3 by 2 mm, very small. Sepals ovate, the outer 2
subacute, the inner 3 shortly slender-acuminate.
Stamens 15, in 3 unequal verticils; filaments com-
pressed, broad at base, tapering in the distal + and
filiform beneath the subglobose anthers; appendage
slender, filiform, c. 25 times length of anthers. Ovary
and stylopodium equal in length, together broadly
hour-glass shaped with a distinct median constriction;
style as long as ovary, columnar, slightly tapering.
Fruit pedicel c. 1 mm long, short, slender. 2 longer
calyx lobes to 4.5 by 1.5 cm, broadly spatulate, obtuse,
chartaceous, c. 4mm broad above the 2-6 by 5 mm
elliptic saccate base; 3 shorter lobes to 7 by 5 mm,
ovate, acute. Nut to 6 by 5 mm, small, ovoid, shortly
apiculate.

Distr. Malesia: Philippines (Basilan, Mindanao).

Ecol. Undulating land and hills below 70 m.

Uses. Locally used for construction.

61. Hopea andersonii ASHTON, Gard. Bull. Sing. 22
(1967) 272, pl. 17; Man. Dipt. Brun. Suppl. (1968) 46,
f. 6.

a. ssp. andersonii.

Medium-sized tree to 40 m tall, with chocolate-
brown thickly flaky bark. Vegetative parts glabrous.
Twigs c. 1mm @ apically, terete, smooth; stipule scars
short, obscure. Bud to 1 by | mm, subglobose. Stipule
unknown. Leaves 5—14 by 2-6 cm, lanceolate-falcate
to elliptic, coriaceous, drying tawny with the ner-
vation beneath dark red-brown; base obtuse on
adaxial side, cuneate on the other, unequal; acumen to
2 cm long, slender, frequently falcate; nerves 9-12
pairs, slender, not prominently raised, arched, at
65°-75°, some with prominent axillary pustular pored
domatia; tertiary nerves scalariform; midrib evident
but unraised or hardly so above, prominent beneath;
petiole 5-10 mm long. Panicle to 12 cm long, terminal
or axillary, terete, frequently fascicled, densely evenly
+ persistently pale grey puberulent; singly branched,
branchlets bearing to 9 secund flowers; bracteoles to 2
mm long, linear, puberulent, fugaceous. Flower bud to
3 by 2 mm, ellipsoid. Sepals ovate, acute, subequal,
densely pubescent outside, more sparsely so within.
Petals lanceolate, densely pubescent on parts exposed
in bud. Stamens 15, in 3 unequal verticils; filaments
compressed at base, tapering and filiform below the
broadly ellipsoid anthers; appendage to connective
slender, 2-24 times as long as anther. Ovary and
stylopodium cylindrical, very slightly constricted
medially, the stylopodium slightly narrower than the
ovary, both densely papillose and puberulent; style
short, columnar. Fruit pedicel to 3 mm long, slender. 2
longer calyx lobes to 6 by 2 cm, oblong, obtuse,
puberulent at base but otherwise glabrous, to c. 3mm
broad above the to 4 by 3 mm elliptic somewhat

426

FLORA MALESIANA

[ser. I, vol. 9?

thickened saccate base; 3 shorter lobes to 4 by 3 mm,
ovate, obtuse, saccate. Nut to 8 by 5 mm, ovoid,
glabrous, with a small apical truncate stylopodium
and c. 1 mm long mucronate style remnant.

Distr. Malesia: Borneo (W. and N.E. Sarawak).

Ecol. Common on the lower slopes of limestone
hills, to 400 m.

Vern. Luis somit.

b. ssp. basalticola ASHTON, Gard. Bull. Sing. 22 (1967)
DADs

Bark surface coppery-brown, thinly flaked. Leaf
drying pale grey-brown.

Distr. Malesia: Borneo (Central and N.E. Sar-
awak, E. Sabah, S.E. Borneo to Pleihari, Ulu
Kapuas).

Ecol. Mixed Dipterocarp forest on clay rich soils,
especially on basic volcanic rocks; to 400 m.

62. Hopea ovoidea ASHTON, Gard. Bull. Sing. 31
(1978) 34. — H. plagata [non (BLCO) VIDAL] Foxw.
Philip. J. Sc. 3 (1907) Bot. 396, p.p.; ibid. 4 (1909) Bot.
515: ibid. 6 (1911) Bot. 262, 285; ibid. 13 (1918) Bot.
183; ibid. 67 (1938) 278, p.p.

Large buttressed flaky-barked tree. Leaf bud, pani-
cle, parts of perianth exposed in bud, and ovary
densely persistently pale buff pubescent, parts other-
wise glabrescent. Twigs c. | mm @ apically, slender,
becoming terete, smooth or rugulose. Buds minute;
stipule unknown, fugaceous. Leaves 9-13 by 3-6.5
cm, elliptic to narrowly ovate, chartaceous and undu-
late on drying; base + equal, cuneate, + shortly
decurrent; acumen to 2 cm long, slender, tapering;
nerves 7-8 (to 10 in young trees) pairs, slender but
distinctly elevated beneath, arched, ascending at
65°—55°, without or with a few porous canaliculate
domatia; tertiary nerves scalariform, + distinctly
elevated beneath; petiole 10-15 mm long, slender.
Panicle to 13 mm long, erect, slender, to 2-axillary or
terminal, singly branched; branchlets to 3.5 cm long,
bearing to 7 secund flowers; bracteoles fugaceous.
Flower buds to 3 by 2 mm, ovoid; 2 outer sepals
narrowly deltoid, subacute; 3 inner sepals broadly
ovate, acute; stamens 15, in 3 subequal verticils;
filaments compressed at base, tapering to the nar-
rowly elliptic subacute anthers; appendage 13—2 times
length of anther, acicular, relatively stout; ovary and
stylopodium ovoid, surmounted by a short glabrous
columnar style c. 4 their length. Fruit unknown.

Distr. Malesia: N.E. Borneo (Sandakan to
Tawau).

Ecol. Low hills near coast. Rare.

Notes. LOHER 12914 & 14901, from Luzon, con-
fused by FoxwortTny with H. plagata VIDAL, appear
to belong to this species, though the leaves are
narrower and the nerves less prominent beneath.

H. ovoidea closely resembles H. semicuneata SYM.
when sterile, but can nevertheless be distinguished by
its brown, rather than grey-brown, drying leaves and
small, canaliculate rather than pustular domatia
which are at times absent; the pubescent ovoid ovary

and rather stout connectival appendages (slender and
thrice as long as the anthers in H. semicuneata) define
this species.

63. Hopea semicuneata Sym. Gard. Bull. S.S.8 (1934)
24, pl. 6; Mal. For. Rec. 16 (1943) 143, f. 69; MEIER &
Woop, Sabah For. Rec. 5 (1964) 224, f. 32; ASHTON,
Man. Dipt. Brun. Suppl. (1968) 56, f. 8. — H. sp.
SLoor. ex MerR. Pl. Elm. Born. (1929) 202. — H.
diversifolia M1Q. Sum. (1860) 491, p.p.; DC. Prod. 16,
2 (1868) 635; WALP. Ann. 7 (1868) 379. — ‘Sama rupa
chengal’ Foxw. Mal. For. Rec. 10 (1932) 71,
p.p. — H. multiflora (non BRANDIS) Foxw. Mal. For.
Rec. 10 (1932) 119, p.p.— H. plagata (non VIDAL)
Sym. Gard. Bull. S. S. 7 (1933) 154.

Tall flaky-barked and buttressed tree. Young twigs
and petioles fugaceous puberulent, vegetative parts
otherwise glabrous. Twig c. | mm @ apically, terete,
becoming striated. Bud to 2 by | mm, conical. Stipule
to 2mm long, linear, fugaceous. Leaves 6.5—14 by 2-7
cm, elliptic to ovate-lanceolate, chartaceous, fre-
quently undulate; base narrowly or broadly cuneate;
acumen to 2.5 cm long, slender, tapering; nerves 6—9
pairs, scalariform, depressed above, slender but
prominent beneath, set obliquely at 45-65”, fre-
quently with prominent, pustular domatia; tertiary
nerves densely scalariform, at 90° to the midrib;
midrib slightly raised to slightly depressed above, +
prominent beneath; petiole 6-12 mm long, slender.
Panicle to 7 cm long, terminal or axillary, terete,
densely shortly evenly persistently pale cream-buff
pubescent; singly branched, the branchlets bearing to
7 cream flowers; bracteoles to 1 mm, short, deltoid,
fugaceous. Flower bud to 2 by | mm, ellipsoid. Sepals
broadly ovate, acute, subequal, glabrous or pubes-
cent. Petals lanceolate, puberulent on parts exposed in
bud. Stamens 15, in 3 unequal verticils; filaments
compressed at base, tapering filiform below the
ellipsoid anthers; appendage to connective slender, c.
3 times length of anther, reaching almost to style apex.
Ovary and stylopodium cylindrical, truncate, some-
times constricted medially, glabrous but for the
papillose apical platform; style shortly columnar,
glabrous. Fruit calyx at first pale buff puberulent,
caducous except at base. Pedicel to 2 mm long. 2
longer calyx lobes to 9.5 by 2.2 mm, broadly lorate,
obtuse, c. 4 mm wide above the 8 by 6 mm elliptic
thickened saccate base; 3 shorter lobes to 4 by 6 mm,
broadly ovate, subacute, saccate. Nut to 6 by 5 mm,
subglobose, shortly apiculate.

Distr. Malesia: Malaya, Sumatra (Atjeh, Lam-
pong), Borneo.

Ecol. Local, clay rich alluvium, undulating land,
and hillsides below 500 m.

Vern. Sama rupa chéngal, chéngal, c. batu, c. mas,
pénak, p. batu (Mal.), giam kulit mérah (Sabah).

64. Hopea megacarpa AsHTON, Gard. Bull. Sing. 22
(1967) 278, pl. 24; Man. Dipt. Brun. Suppl. (1968) 53,
iio: Ue

Small or medium-sized smooth-barked tree. Young

1982]

twig, petiole and leaf nervation beneath sparsely
caducous puberulent, leaf bud persistently so. Twig c.
1 mm @ apically, slender, terete, smooth; stipule scars
small, obscure. Bud to | by | mm, minute. Stipule to 2
mm long, linear-falcate, caducous. Leaf 6—12 by 1.5—S
cm, + narrowly elliptic, thinly coriaceous, undulate,
with cuneate base and to 2 cm long prominent caudate
acumen; nerves 6—7 pairs, slender, raised beneath, at
25°—40°; tertiary nerves slender, densely scalariform;
midrib prominent, terete beneath, applanate to
slightly raised, slender, above; petiole c. 6 mm long,
short, grooved on upper side. Panicle to 3 cm long,
axillary, terete, glabrous, lax; singly branched,
branchlets bearing to 3 flowers; bracts and bracteoles
to 2 mm long, linear. Flower bud 4 by 3 mm,
subglobose. Calyx lobes fimbriate; 2 outer lobes ovate,
acute; 3 inner lobes suborbicular, submucronate.
Petals elliptic-oblong, obtuse, sparsely puberulent on
parts exposed in bud, pale pink. Stamens 15; filaments
broad at base, tapering and filiform distally; anthers
oblong; appendage to connective c. 3 times length of
anthers, slender, filiform. Ovary and stylopodium
cylindrical, subtruncate, glabrous, surmounted by a
short style. Fruit entirely glabrous. Pedice/ to 3 mm
long, broadening into fruit. 2 longer calyx lobes to 10
by 1.3 cm, narrowly spatulate, subacute, c. 5 mm wide
above the to 9 by 15 mm subauriculate saccate base; 3
shorter lobes to 20 by 9 mm, ovate, similarly subauricu-
late, enclosing and obscuring the nut. Nut to 12 by 10
mm, ovoid, with minutely truncate mucronate apex.

Distr. Malesia: N.W. Borneo (Central Sarawak;
W. and S.E. Kalimantan).

Ecol. Locally frequent in Mixed Dipterocarp for-
ests below 600 m in the Rejang hinterland.

65. Hopea samarensis GUTIERREZ, Kalikasan 4 (1975)
AO Je Sle

Small smooth-barked tree. Parts glabrous but for
the pubescent domatia and parts of petals exposed in
bud, and glabrescent sepals. Twigs c. 2mm @ apically,
slender. Leaves (4—)8-15 by (1.5—)3—5.5 cm, oblong-
elliptic, thinly coriaceous; acumen to | cm long,
subcaudate; base subequal, cuneate; nerves 9—12(—13)
pairs, slender but prominent beneath, evident above,
with small axillary domatia; tertiaries densely scalari-
form, very slender, evident beneath; petiole 8—12 mm
long, slender. Panicles to 4 cm long, singly branched,
to 3-ramiflorous; branchlets to 2 cm long, bearing to 4
secund flowers; bracteoles minute, deltoid, fugaceous.
Flower bud to 8 by 4mm, ovoid; sepals broadly ovate,
the inner 3 acute, the outer 1 acuminate, becoming
rotate before anthesis; petals lorate; stamens 15 in 3
unequal verticils; filaments broad at base, tapering
and filiform beneath the broadly ellipsoid anthers;
appendage aristate, c. 2-3 times length of anthers;
ovary ovoid, surmounted by a slightly shorter slender
cylindric tapering stylopodium with short terminal
style c. 4 its length. Fruit unknown.

Distr. Malesia: Philippines (Samar I.).

Ecol. Locally common in moist lowland valleys in
Mixed Dipterocarp forest.

DIPTEROCARPACEAE (Ashton)

427

66. Hopea nodosa SLooT. Reinwardtia 2 (1952) 25, f.
8; v. RoYEN, Man. Forest Trees Papua New Guinea 8
(1965) 38.

Medium-sized tree with flaky bark and steep but-
tresses. Parts glabrous but for the persistently pubes-
cent parts of the petals exposed in bud. Twigs 2-3 mm
@, rather stout, terete; buds and stipules not seen.
Leaves (6—)8—-20 by (2.5—)3.8-7 cm, elliptic, coria-
ceous, lustrous, minutely punctate above; margin
narrowly revolute; base broadly cuneate; acumen to |
cm long, tapering; nerves 8-11 pairs, prominent
beneath, elevated but frequently set in a groove above,
arched, ascending at 50°—70°; domatia minute,
porous, or absent; without secondaries; tertiary
nerves scalariform, obscure; midrib prominent
beneath, distinctly elevated above; petiole 7-15 mm
long, rather stout. Panicle to 4 cm long, slender,
terminal or to 2-axillary or ramiflorous; singly
branched, branchlets to 12 mm long, bearing to 6
flowers; bracteoles fugaceous, not seen. Flower buds to
3 by 2 mn, ellipsoid. Sepals subequal, broadly
ovate-deltoid, obtuse. Stamens 15, shorter than style,
in 3 unequal verticils; filaments broad and compressed
at base, tapering and filiform in the distal 4; anthers
subglobose; appendages c. twice as long as anthers,
slender. Ovary and stylopodium narrowly pyriform
with distinct medial constriction, tapering to the short
but distinct columnar style. Fruit subsessile; calyx
lobes to 7 by 8 mm, subequal, shorter than nut,
broadly ovate, subacute, saccate, thickened. Nut to 10
by 6 mm, ovoid, shortly apiculate.

Distr. Malesia: N.W. New Guinea (Beriat,
Teminabuan; Sorong).

Ecol. Locally common in secondary and primary
lowland forest.

Vern. Megun gun (Telid).

67. Hopea celebica BURCK, Ann. Jard. Bot. Btzg 6
(1887) 237; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 64;
HEyYnegE, Nutt. Pl. ed. 2 (1927) 1110; SLoor. Reinward-
tia 2 (1952) 15, f. 5; BACKER & BAKH. f. Fl. Java 1
(1963) 331; AsHTON, Gard. Bull. Sing. 31 (1978)
35. — H. dolosa SLooT. Reinwardtia 2 (1952) 18, f. 6.

Medium-sized, hard-wooded, scaly-barked tree.
Twig apices, petiole, panicle and calyx outside +
caducous buff pubescent, parts of petals exposed in
bud and sometimes panicle persistently so. Twigs c. 2
mm @ much branched, terete, smooth. Buds small,
ovoid: stipules not seen. Leaves (5.5—)8—22 by
(2.2-)2.5-8 cm, ovate-lanceolate, coriaceous, + lus-
trous; margin + revolute; base subequal, obtuse or
sometimes broadly cuneate; acumen to 1.5 cm long,
usually short, tapering; nerves 8-11 pairs, slender but
prominent beneath, slightly elevated above, arched, at
45°_55° except at base, often with prominent axillary
porous canaliculate domatia; tertiary nerves subsca-
lariform, evident on both surfaces though more
distinct beneath; midrib prominent beneath, distinctly
elevated above; petiole 10-16 cm long. Panicle to 9 cm
long, terminal, to 3-axillary or ramiflorous, slender,
pendant, singly branched; branchlets to 13 mm long,

428

FLORA MALESIANA

(ser. I, vol. 9?

short, bearing to 4 flowers; bracteoles minute, deltoid,
fugaceous. Flower buds to 2 mm long, ellipsoid. 2
outer sepals long, narrowly deltoid-lanceolate; 3 inner
broadly ovate, + distinctly acuminate. Stamens 15, in
3 subequal verticils, short; filaments broad and com-
pressed at base, tapering and filiform beneath the
oblong anthers; appendages c. 15 times length of
anther, slender. Ovary and stylopodium stoutly pyri-
form, stylopodium punctate; style short but distinct,
columnar. Mature fruit unknown; pedicel at least 3
mm long; sepals unequal, 2 aliform, spatulate, obtuse;
3 shorter lobes exceeded by the ovoid apiculate nut.

Distr. Malesia: Celebes (S.W. Peninsula: Maros;
Central: Malili).

Ecol. Locally common in semi-evergreen forest
below 500 m.

Uses. Construction.

Vern. Hulo dereh, kérih (Maros), hulodere, damar
derehitém, d.d. lotang, bisik bisik, rinni rinni, saréh
paréh, torinih (Malili).

68. Hopea iriana SLOoT. Reinwardtia 2 (1952) 28, f.
10; v. ROYEN, Man. Forest Trees Papua New Guinea 8
(1965) 35; ASHTON, Gard. Bull. Sing. 31 (1978)
35. — H. nabirensis SLOOT. Reinwardtia 2 (1952) 27, f.
9: v. ROYEN, Man. Forest Trees Papua New Guinea 8
(1965) 38.

Tall, hard-wooded tree with blackish flaky bark
and flying buttresses. Young parts buff puberulent,
becoming sparse but + persistent on calyx, persistent
on panicle and parts of petals exposed in bud. Twig c.
2 mm @ apically, much branched, terete, becoming
smooth, blackish. Leaf bud minute, ovoid; stipules
fugaceous, not seen. Leaves S—13 by 2-4.5 cm, lanceo-
late-falcate, coriaceous, dull greyish minutely stellate
beneath; margin frequently subrevolute; base un-
equal, cuneate; acumen to 1.5 cm long, slender,
tapering; nerves 7—11 pairs, slender but distinctly
elevated beneath, slightly elevated above, usually with
prominent porous glabrous canaliculate domatia;
tertiary nerves scalariform, evident on both surfaces,
+ distinctly elevated beneath; petiole 7-9 mm long,
slender. Panicle to 6 cm long, slender, terminal or
l-axillary; singly branched, branchlets to 1.5 cm long,
bearing to 5 dense secund flowers; bracteoles fuga-
ceous, not seen. Flower buds to 3 by 2 mm, ellipsoid. 2
outer sepals narrowly lanceolate, subacute; 3 inner
suborbicular, mucronate. Stamens 15, shorter than
style, in 3 subequal verticils; filaments broad and
compressed at base, tapering and filiform in the distal
3; anthers broadly oblong; appendage c. 3 times length
of anthers, long, slender. Ovary and stylopodium
conical-cylindric, sericeous, tapering somewhat
abruptly beneath the short but distinct columnar
style. Fruit pedicel to 2 mm long. 2 longer calyx lobes
to 7 by 1.8 cm, broadly spatulate, obtuse, c. 4 mm
broad above the 6 by 4 mm ovate saccate thickened
base; 3 shorter lobes to 6 by 4 mm, ovate, acute,
similarly saccate. Nut to 13 by 7 mm, ovoid, with
tapering apiculus.

Distr. Malesia: New Guinea (West: Manokwari

area, Hollandia; Papua New Guinea: Morobe Distr.),
Japen I.; Aru Is. (sterile collections only).

Ecol. Evergreen forests; widespread hills below 600
m, especially on ridges.

Vern. Sian, saindorih (Japen), lilipga (Manikiong).

Notes. A variable species which, as H. novo-
guineensis, appears to possess regional diversification;
collections from western New Guinea have notably
larger more lustrous leaves.

69. Hopea glabrifolia C.T. Wuite, Proc. R. Soc.
Queens}. 43 (1932) 49; SLooT. Reinwardtia 2 (1952)
35, f. 13; v. RoYEN, Man. Forest Trees Papua New
Guinea 8 (1965) 35.

Tall, buttressed tree with hard wood and flaky bark.
Young parts and panicle greyish puberulent, glabres-
cent; petals persistently puberulent on parts exposed
in bud. Twig c. 2 mm @ apically, mbbed, much
branched, becoming terete, rugose, dark brown. Leaf
buds minute, ellipsoid; stipules not seen, fugaceous.
Leaves 18-19 by 2-5.5 cm, lanceolate, falcate, coria-
ceous, lustrous; margin narrowly subrevolute; base
prominently unequal, cuneate abaxially, cordate
adaxially; acumen to 1.5 cm long, broad, tapering;
nerves 9-12 pairs, slender but prominent beneath, +
narrowly depressed above, arched, ascending at
45°—50° except at base; without secondaries; tertiaries
densely scalariform, very slender but distinctly ele-
vated beneath; midrib prominent beneath, slender but
prominent above; petiole S-8 mm long, short. Panicles
to Scm long, terminal or to 3-axillary, slender; singly
branched, branchlets to 1.5 cm long, bearing to 5
flowers. Flower buds to 3 by 2 mm, ellipsoid. Sepals
ovate to suborbicular, subacute. Stamens 15; appen-
dage c. 4 times the length of the anther cells.
Gynoecium glabrous; ovary ovoid, surmounted by a
cylindrical stylopodium twice its length and very short
style. Fruit pedicel c. 1 mm long, short. 2 longer calyx
lobes to 7 by 1.3 cm, spatulate, obtuse, c. 3 mm broad
above the to 8 by 6mm ovate saccate thickened base; 3
shorter lobes to 9 by 7 mm, ovate, acute. Nut to 14 by8
mm, ovoid, stoutly apiculate.

Distr. Malesia: Papua New Guinea (Milne Bay
area) and Louisiades (Sudest I., Misima I.).

Ecol. Locally abundant in semi-evergreen seasonal
forest below 350 m.

Vern. Kapilatana, matapo, malabia.

70. Hopea gregaria SLOoT. Reinwardtia 2 (1952) 21, f.
7; ASHTON, Gard. Bull. Sing. 31 (1978) 32.
Medium-sized or large flaky-barked tree with hard
wood. Young parts buff puberulent, becoming sparse
but + persistent on calyx, persistent on panicle and
parts of petals exposed in bud. Twig 1-2 mm @
apically, much branched, terete, becoming smooth,
blackish. Leaf bud minute, ovoid; stipules fugaceous,
not observed. Leaf 6-13 by 2.5—6.5 cm, lanceolate-fal-
cate to ovate, thinly coriaceous, with dull minutely
stellate undersurface; base cuneate, unequal; acumen
to 1.5 cm long, slender, tapering; nerves 7-10 pairs,
slender but distinctly elevated beneath, + applanate

1982]

above, rarely with glabrous porous canaliculate axil-
lary domatia; tertiary nerves scalariform, + evident
on both surfaces; midrib slender but prominent
beneath, evident and + elevated above; petiole 6-10
mm long, slender. Panicle to 6 mm long, slender,
terminal or |-axillary, singly branched, branchlets to
12 mm long, bearing to 4 flowers. Flower buds to 3 by 2
mm, ellipsoid. Sepals subequal, ovate, acute. Stamens
15, shorter than style, in 3 subequal verticils; filaments
broad, compressed, tapering and becoming filiform
below the broadly oblong anthers; appendages c. 15

DIPTEROCARPACEAE (Ashton)

429

times length of anthers. Ovary and stylopodium
broadly pyriform, punctate distally, somewhat
abruptly tapering to the short columnar style. Mature

fruit unknown. Pedicels c. 3 mm long; sepals unequal

but the 2 longer relatively short, broad, becoming
reflexed; nut ovoid, shortly apiculate.
Distr. Malesia: S.E. Celebes (Kendari), S.E.
Moluccas (Aru Is. ?) and Japen I. (sterile collections)?
Vern. Pooti (Celebes), koereh, mandonor (Biak),
kamoera (Aru).

2h. Subsection Pierrea

(Herm) AsHTon, Gard. Bull. Sing. 20 (1963) 259; Man. Dipt. Brun. (1964) 91;
GUTIERREZ, Act. Manil. 4, A, 2 (1968) 25, 26.— Pierrea HEIM, non HAN-
cE. — Pierreocarpus RIDL. ex. SYM. in syn. — Hopea, Pierrea group Sym. Gard.
Bull. S. S. 9 (1934) 32; Mal. For. Rec. 16 (1943) 108. — Fig. 74.

Panicles glabrescent, fascicled; ovary and stylopodium hour-glass-shaped,
elongate; style short, obscure; bark surface generally smooth, or shallowly papery

flaked, usually stilt-rooted.

Distr. Malesia: Malaya, Borneo, Philippines.

Note. The least well defined of the four subsections. The New Guinea species, with their elongate stylopodia
and in several cases large leaves with unequal bases are almost intermediate between subsections Hopea and
Pierrea, as also to a large extent are some Indochinese species in the type subsection (e.g. H. oblongifolia DYER,
H. reticulata Tarpieu, H. hongayanensis TARDIEU), while H. glaucescens and H. wyatt-smithii share the flower

colour and leaf-shape of the H. nervosa group within subsect. Sphaerocarpae.

71. Hopea glaucescens Sym. J. Mal. Br. R. As. Soc. 19
(1941) 142, pl. 2; Mal. For. Rec. 16 (1943) 126, f.
69. — Hopea sp. nov. SyM. J. Mal. Br. R. As. Soc. 14
(1936) 348.

Medium-sized, smooth-barked tree, often stilt-
rooted. Panicles sparsely persistently puberulent,
twigs and petioles fugaceously so, petals outside
persistently densely gold pubescent. Twig 2-3 mm @
apically, becoming dark brown, terete; with promi-
nent ribs along the leaf traces; stipule scars minute.
Bud to 2 by 2 mm, ovoid; stipules fugaceous. Leaves
9-18 by 3.5-9 cm, + elliptic, coriaceous, glaucescent
beneath in mature trees; base cuneate; acumen to | cm
long, slender, prominent; nerves 12-15 pairs, slender
but relatively prominent beneath, obscure above, with
many distinct short secondary nerves; tertiary nerves
slender, subscalariform, evident to obscure in mature
trees; midrib distinctly elevated on both surfaces;
petioles 10-15 mm long. Panicle to 4cm long, terminal
or axillary to ramiflorous, with to 2 cm long branch-
lets bearing to 8 congested secund flowers. Flower bud
to 4 by 3 mm, ovoid. Sepals subequal, broadly ovate,
the outer 2 subacuminate, the inner 3 mucronate.
Stamens 15; filaments broadly compressed at base,
tapering and filiform distally; appendages aristate, c.
3 x length of the small subglobose anthers; style and
stylopodium hour-glass-shaped, equal in height but
the ovary the broader; style short, tapering. Fruit
pedicel to 1 mm long, slender. 2 longer calyx lobes to7
by 1.5 cm, spatulate, obtuse, c. 5 mm broad above the

to 8 by 5 mm ovate saccate thickened base; 3 shorter
lobes to 20 by 5 mm, linear-lorate, similar at base,
completely enclosing nut. Nut to 9 by 7 mm, ovoid,
apiculate.

Distr. Malesia: Malaya.

Ecol. Rare, below 500 m, in Mixed Dipterocarp
forest.

Vern. Mérawan kélabu, m. galor, m. jangkang, m.
téngkok biawak.

72. Hopea wyatt-smithii Woop ex ASHTON, Gard.
Bull. Sing. 19 (1962) 260, pl. 4; Man. Dipt. Brun.
(1964) 113, f. 12; ibid. Suppl. (1968) 58; MEUER &
Woop, Sabah For. Rec. 5 (1964) 227, f. 33. — Fig.
74B-B4.

Small to medium-sized smooth-barked promi-
nently stilt-rooted tree. All parts glabrous but for
petals and ovary. Twig c. 1 mm @ apically, smooth.
Bud to 1.5 mm long, ovoid. Stipule to 2 mm long,
linear, fugaceous. Leaves 9-14 by 5.5—9 cm, broadly
ovate to elliptic, undersurface sparsely greyish lepi-
dote; base broadly cuneate, occasionally obtuse:
acumen caudate to 2 cm long; nerves 4-8 pairs, well
spaced, irregularly disposed owing to frequent pre-
sence of short but prominent secondaries; slender,
slightly raised beneath, basal 2-3 pairs straight, at first
decurrent with midrib, distal pairs curved at 40°—50°;
tertiary nerves well spaced, scalariform, at 90° or
ascending; midrib slender, applanate beneath acute
above; petiole 1.2-1.7 cm long. Panicle to 6 cm long,

430 FLORA MALESIANA [ser. I, vol. 9?

J Ai

Fig. 74. Flower details in Hopea sect. Hopea subsect. Pierrea (HEIM) ASHTON. All x 10. Sepals drawn from

inside. — Hopea philippinensis DyER. A. Bud, A/. outer sepal, 42. inner sepal, 43. stamens from inside, A4.

pistil. — H. wyatt-smithii Woop ex ASHTON. B. Bud, B/. outer sepal, B2. inner sepal, B3. stamens from inside,

B4. pistil. — H. bilitonensis ASHTON. C. Bud, C/. outer sepal, C2. inner sepal, C3. stamens from inside, C4.

pistil. — H. apiculata Sym. D3. Stamens from inside, D4. pistil (A RAMos & EDANO BS 31296, B Woop 15061, C
bb. 23087, D KEP 98175).

1982]

DIPTEROCARPACEAE (Ashton)

43]

terminal or to 2-axillary, glabrous, terete or +
compressed; singly branched, branchlets to 1.5 cm
long, bearing to 6 secund flowers; bracteoles small,
deltoid, glabrous. Flower bud to 3 mm long, subglo-
bose. Calyx glabrescent, with a shortly fimbriate
margin, spreading; lobes subequal, broadly ovate to
suborbicular, obtuse. Petals dark red, oblong-lanceo-
late, obtuse, densely pubescent on parts exposed in
bud. Stamens 15; filaments broad at base, tapering
abruptly; anther broadly oblong, cells subequal;
appendage to connective about twice length of anther,
slender. Ovary and stylopodium narrowly hour-glass
shaped; ovary glabrous, stylopodium somewhat
longer than ovary, puberulent, crowned by a short
glabrous style. Fruit calyx entirely glabrous; 5 lobes
equal, to 10 by 8 mm, ovate, subacute, shallowly
saccate, thickened, closely imbricate and adpressed to
the nut. Nut to 1.2 cm long and g, ovoid, acute,
enclosed by calyx but for an up to 3 mm g apical gap.

Distr. Malesia: Borneo (Sarawak N.E. of the
Lupar, S.W. and S.E. Sabah).

Ecol. Local, on clay rich soils in Mixed Diptero-
carp forest on low hills.

Vern. Mérawan puteh (Brun.), sélangan daun bulat
(Sabah).

73. Hopea polyalthioides Sym. J. Mal. Br. R. As. Soc.
19, 2 (1941) 146, pl. 4; Mal. For. Rec. 16 (1943) 140, f.
69.

Small monopodial smooth-barked tree. Twig and
petiole densely persistently fulvous tawny pubescent,
leaf undersurface and midrib above sparsely so;
panicle glabrous. Twig c. 3 mm @ apically, stout,
terete, becoming dark brown, with a prominent short
rib at first following the leaf trace; internodes 1-5 cm
long; stipule scars obscure; stipules unknown. Leaves
(8—)14-27 by (2.5-)4-6 cm, narrowly oblong-lanceo-
late, coriaceous; base subcordate; acumen short,
broad; nerves 10—13 pairs, arched and becoming wavy
at the margin, prominent beneath, + obscure above,
sometimes with a few short indistinct secondary
nerves; midrib stout and prominent beneath, slender,
elevated to shallowly channelled, above; petiole 5-8
mm long, c. 3 mm @, short, stout. Panicles to 11 cm
long, very slender, laxly shortly branched; flowers and
fruit unknown.

Distr. Malesia: Malaya (S. Johore).

Ecol. Rare, in well-drained forest.

Vern. Giam rambai, resak, r. rambai, sélumbar.

74. Hopea cagayanensis (Foxw.) SLooT. Reinwardtia
3 (1956) 318. — Balanocarpus cagayanensis Foxw.
Philip. Journ. Sc. 13 (1918) Bot. 194, pl. 2; ibid. 67
(1938) 285, pl. 3; Merr. En. Philip. 3 (1923) 101;
Reyes, Philip. J. Sc. 22 (1923) 335; Sym. Gard. Bull. S.
S. 8 (1934) 17, 32.

Large, flaky-barked tree. Twigs, leaf bud, stipules,
petioles, domatia and parts of petals exposed in bud
densely persistently tawny pubescent; nerves and
midrib above caducously so; panicle and nerves and
midrib beneath sparsely caducously so. Twigs c. 1 mm

@ apically, slender, much branched, becoming
smooth, terete. Buds c. 2 by 1 mm, small, ovoid;
stipules to 4 mm long, linear, not at first caducous.
Leaves 8-10 by 2.54 cm, lanceolate, thinly coria-
ceous; base unequal, broadly cuneate or obtuse on the
adaxial side, subcordate on the abaxial; acumen to 1.5
cm long, slender, subcaudate; nerves 9-12 (to 14 in
young trees) pairs, slender but prominent beneath,
applanate above, arched, at 55°-65°, with domatia;
secondary nerves short, obscure; tertiary nerves den-
sely scalariform, evident and + elevated beneath;
midrib slender but evident and distinctly elevated on
both surfaces; petiole 5-7 mm long, short. Panicle to 5
cm long, to 2-axillary, slender: singly branched,
branchlets bearing to 3 flowers; bracts and bracteoles
to 1 mm long, minute, deltoid, not at first caducous.
Flower bud to 2.5 by 1.5 mm, ellipsoid. Sepals
subequal, broadly ovate, shortly acuminate. Stamens
15, shorter than style, in 3 + unequal verticils;
filaments broad and compressed at base, tapering and
filiform in the distal 3; anthers broadly oblong:
appendages c. twice length of anther, slender. Ovary
small, ovoid, tapering into an equally long somewhat
narrower cylindrical punctate stylopodium and
shorter columnar style. Fruit subsessile; sepals short,
subequal; adpressed to the nut; outer 2 to 9 by 7 mm,
ovate-acuminate, incrassate; inner 3 to 10 by 10 mm,
broadly ovate, obtuse, thin. Nut to 15 by 10 mm,
ovoid, apiculate, + thinly resin-coated.

Distr. Malesia: Philippines (N.E. Luzon: Cagayan
Prov.).

Ecol. Locally frequent, semi-evergreen forests.

Uses. House posts.

Vern. Narek.

75. Hopea paucinervis Paris in Fedde, Rep. 33 (1933)
243.

Medium-sized tree. Young parts fugaceous buff
puberulent. Twigs slender. Stipules fugaceous, un-
known. Leaf 4.5—-8 by 2.5—4.5 cm, ovate-lanceolate,
coriaceous; base subequal, obtuse or broadly cuneate;
apex shortly acuminate; nerves 7-10 pairs, ascending,
curved, prominent beneath; tertiary nerves subscalari-
form, elevated beneath. Petiole 8-12 mm long. Panicle
to 4 cm long, to 2-axillary, axillary to ramiflorous.
Buds and opened flowers and fruit are unknown.
Sepals of old flowers unequal, ovate-lanceolate.
Stamens 15,in 3 + unequal verticils: filaments lorate,
tapering in distal 4; anthers oblong; appendages equal
to anther, short. Ovary and stylopodium equal, overall
pyriform with prominent intermediate construction.

Distr. Malesia: S.E. Sumatra (Djambi).

76. Hopea apiculata Sym. Gard. Bull. S. S. 8 (1935)
277, pl. 21; Mal. For. Rec. 16 (1943) 120, f. 69,
70. — Fig. 74 D3-D4.

Small smooth-barked tree with sharp, often stilted
buttresses. Twigs caducous tawny puberulent,
petioles and outside of petals densely persistently
pubescent, otherwise glabrous. Twigs c. 2 mm @
apically, straight, infrequently branched, pendant,

432

MALESIANA

[ser. I, vole?

Fig. 75. Hopea pachycarpa (HEIM) SYM. a. Node with leaf and inflorescence, b. branch with young fruits, c. ripe
fruit, d. ditto with sepals removed, all x 5 (aS 22406, with flowers of ANDERSON S 15408, c-d bb. 35260).

terete, pale brown; stipule scars minute, horizontal;
leaves distant. Buds minute; stipule small, linear,
caducous. Leaves alternate, 12-26 by 4-8 cm, nar-
rowly oblong-lanceolate, subcoriaceous; base cor-
date, equal; acumen short; nerves 12-15 pairs, the first
2-3 arising from the base, slender but prominent
beneath, shallowly depressed above, arched; midrib
prominent beneath, obscure and depressed or some-
times evident and elevated above; tertiary nerves
densely subreticulate, evident beneath, obscure
above; petiole 6-12 mm long, c. 3 mm Qg, stout.
Panicles to 20 cm long, terminal or ramiflorous, borne
densely along the twigs, slender, laxly branched;
branchlets to 2 cm long, unbranched, bearing to 6
secund flowers. Flower bud to 5 by 3mm, ovoid; sepals
ovate-lanceolate, fimbriate, acuminate, the outer 2
somewhat larger, sometimes obtuse; petals pale yel-
low. Stamens 15, filaments dilated at base, tapering;
appendages very slender, 4—5 times as long as subglo-
bose anthers. Ovary and stylopodium hour-glass-
shaped, the latter somewhat the larger, with interven-
ing frequently puberulent constriction; style colum-
nar, shorter than ovary. Fruit pedicel to 1 mm long,
short. Sepals to 2.5 by 0.5 cm but usually shorter than

nut (variable, even on one tree), unequal to subequal,
spatulate to ovate-acuminate, thickened and saccate
at base; nut to 20 by 12 mm, ovoid, prominently
apiculate.

Distr. Malesia: Malaya (Perak). SMITINAND (Thai
For. Bull., Bot. 12, 1980, 45) records this also from
Peninsular Thailand and S. Burma (Kemas); I have
not seen this material.

Ecol. Very local, common in two valleys east of the
Keledong Saiong range.

Vern. Mélukut, résak mélukut.

77. Hopea pachycarpa (HEIM) Sym. Gard. Bull. S.S.8
(1934) 30, pl. 8; BRowNne, For. Trees Sarawak &
Brunei (1955) 125; ASHTON, Man. Dipt. Brun. (1964)
105, f. 12; ibid. Suppl. (1968) 54; Gard. Bull. Sing. 22
(1967) 271; ibid. 31 (1978) 35. — Pierrea pachycarpa
Her, Bull. Mens. Soc. Linn. Paris 2 (1891) 958; Rech.
Dipt. (1892) 78, pl. 7; BRANDIS & GILG in E. & P. Pf.
Fam. ed. 1, 3, 6(1895) 268; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 113; Merr. En. Born. (1921) 408; GILG in E.
& P. Pfl. Fam. ed. 2, 21 (1925) 263. — Balanocarpus
pubescens RiDL. Fl. Mal. Pen. 1 (1922) 247; Foxw.
Mal. For. Rec. 10 (1932) 147; Sym. Gard. Bull. S.S.8

1982]

(1934) 28, 32; Burk. Dict. (1935) 288. — Pierreo-
carpus pachycarpa RIDL. ex SyM. Gard. Bull. S. S. 8
(1934) 30, nomen in syn. — H. laxa Sym. Gard. Bull. S.
S. 8 (1934) 33, pl. 9; BRowne, For. Trees Sarawak &
Brunei (1955) 125; cf. ASHTON, Gard. Bull. Sing. 22
(1967) 271; ASHTON, Man. Dipt. Brun. Suppl. (1968)
54. — H. resinosa Sym. Gard. Bull. S. S. 8 (1935) 278,
pl. 23; Mal. For. Rec. 16 (1943) 141, f. 68B, 69. — Fig.
75, 76.

Young twig, leaf bud, stipule outside (glabrous
within) and petiole shortly persistently pale fawn
tomentose; leaf glabrous. Twig to 2 mm @g apically,
terete, becoming smooth. Bud to 2.5 by 2mm, ovoid to
globose. Stipule to 4 mm long, linear, fugaceous.
Leaves 13-22 by 4~7 cm, thinly coriaceous, elliptic to
lanceolate, with or without silvery lepidote undersur-
face; base subequal, cuneate on one side, obtuse on the
other; acumen to 1 cm long, narrow; nerves
(10-)13-17 pairs, slender but distinctly raised
beneath, at 45°—55° but curving round to run parallel
to the margin before terminating at it, with small
tomentose domatia; tertiary nerves slender, densely
scalariform, sinuate, diagonal to nerves; midrib
rounded, raised on both surfaces; petiole 7-10 mm
long, short, stout. Panicle to 8 cm long, axillary to
ramiflorous, terete, puberulent or glabrescent, to
2-axillary, rarely branched; bracteoles to 2 mm long,
linear, not at first caducous. Flower bud to 4 by 2.5
mm, broadly ellipsoid. Calyx glabrous but for the
fimbriate margin; 2 outer lobes broadly ovate, sub-
acute; 3 inner lobes broader, slightly shorter, subacu-
minate. Petals broadly oblong, glabrescent, strongly
contorted in bud. Stamens 15, in 3 unequal verticils,
the 5 inner an anther’s length longer than the 10 outer;
filaments broad at base, tapering, filiform distally:
anthers subglobose: appendage to connective slender,
2-3 times length of anther, glabrous. Ovary small,
ovoid, glabrous; style and stylopodium spindle-
shaped, glandular-papillose towards apex, tapering
into short glabrous style. Fruit entirely glabrous. 5
calyx lobes subequal, to 2 by 1.5 cm, ovate, subacute,
thickened, deeply saccate, closely adpressed to nut;
apex of nut visible only at 5 mm @ apical gap. Nut to
1.5 cm g, subglobose; style remnant short, abrupt,
acute.

Distr. Malesia: Malaya (Pahang, E. Johore), Cen-
tral Sumatra, Ankola in W. Sumatra, Borneo (Ulu
Kapuas, Sarawak, Brunei, Berau).

Ecol. Locally abundant on moist soils on lower
hillsides and alluvium in Mixed Dipterocarp forest.

Vern. Bayan, mérawan mata kuching (Mal.), mélapi
bérjangkang (Kapuas), mérkoyong (Sar.).

78. Hopea bilitonensis ASHTON, Gard. Bull. Sing. 31
(1978) 35. — Fig. 74 C-C4.

Small smooth-barked tree with stilt roots. Leaf
buds and parts of petals exposed in bud densely tawny
pubescent, young twigs and panicles fugaceously so,
otherwise glabrous. Twigs c. 1 mm g apically, slender,
much branched, red-brown, terete, smooth. Leaf buds
c. 1 by 1 mm, ovoid, acute; stipules unknown,

DIPTEROCARPACEAE (Ashton)

433

fugaceous. Leaves 7.5—16 by 3.5—6 cm, ovate-lanceo-
late, + chartaceous, lustrous; base obtuse to subcor-
date, subequal; acumen to 2 cm long, attenuate;
nerves 6-8 pairs, slender but prominent beneath,
applanate above, arched, at 50°-60°; secondaries
absent; tertiary nerves remotely subscalariform, evi-
dent and distinctly elevated beneath; petiole 6-8 mm
long, short. Panicle to 18 cm long, slender, axillary,
solitary, lax, pendant; twice branched, branchlets to 4
cm long, bearing to 6 flowers; bracts and bracteoles
minute, deltoid, caducous. Flower bud to 3 by 2 mm,
lanceolate. Sepals fimbriate; 2 outer deltoid, subacute;
3 inner ovate, subacute. Stamens 15, in 3 unequal
verticils, shorter than style at anthesis; filaments
somewhat slender, compressed at base, tapering dis-
tally and filiform beneath the small subglobose
anthers; appendages c. 34 times as long as anthers,
very long and slender, + crisped. Ovary small, ovoid,
with somewhat longer oblanceolate stylopodium and
short terminal style. Fruit pedicel to 2 mm long, stout.
2 longer calyx lobes to 5 by 1.2 cm, broadly spatulate,
obtuse, c. 7 mm broad above the to 7 by 4 mm
subauriculate centrally thickened base; 3 shorter lobes
to 9 by 6 mm, ovate, acuminate, shorter than nut. Nut
to 10 by 6 mm, ovoid, prominently slender apiculate.

Distr. Malesia: E. Sumatra (Banka and Billiton):
Malaya (Perak, once).

Ecol. Locally common in lowland forest, once
recorded from limestone in N.W. Malaya.

Vern. Pélepak.

Notes. An interesting species, locally common on
the sandy islands of Banka and Billiton and now
recorded from limestone in a distinctly disjunct
position in relatively seasonal N.W. Malaya, a range
that has apparently become disjunct since the Pleisto-
cene.

Though an isolated species on account of its leaves,
solitary axillary inflorescences and small fruit, the
leaves nonetheless recall those of the anomalous H.
polyalthioides SyM. of East Johore, still unknown in
flower or fruit, though they differ in size and shape.

79. Hopea bullatifolia ASHTON, Gard. Bull. Sing. 22
(1967) 274, pl. 18; Man. Dipt. Brun. Suppl. (1968) 48.
i265

Small smooth-barked stilt-rooted tree. Leaf bud,
stipule outside (glabrous within), petiole and midrib
beneath densely persistently evenly tawny pubescent,
nervation and lamina beneath and midrib above
sparsely so. Twig c. 2 mm @, terete, becoming smooth.
Bud to 1 by 1 mm, small, subglobose. Stipule to 4mm
long, linear, caducous. Leaves 16-34 by 4.5—9 cm,
oblong, prominently bullate between the tertiary
nerves; base cordate, subequal; acumen to | cm long,
slender; nerves 17-26 pairs, slender but prominent
beneath, depressed above, at 35—45°, frequently
linked to form a looped intramarginal nerve distally;
tertiary nerves scalariform, prominent beneath,
depressed above; midrib prominent on both surfaces;
petiole 3-6 mm long, short, stout. Flowers unknown.

434

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 76. Hopea pachycarpa (HEM) SyM. Leaf from
sterile branch, x 4(S 22035).

Inflorescences unknown. Fruit entirely glabrous; 2
longer calyx lobes to 8 by 1.5 cm, spatulate, obtuse, c.
4 mm broad above the to 8 by 3 mm ovate thickened
saccate base; 3 shorter lobes to 15 mm long, lanceo-
late, acute, slender, similar at base, enclosing the nut.
Nut c. 10 by 7 mm, ovoid, apiculate, enclosed in the
sepals.

Distr. Malesia: Borneo (Central Sarawak; S.E.
Kalimantan, Pulau Laut).

Ecol. Rare, in Mixed Dipterocarp forest on shale
knolls.

80. Hopea pterygota ASHTON, Gard. Bull. Sing. 22
(1967) 280, pl. 26; Man. Dipt. Brun. Suppl. (1968) 55,
fae?

Small tree. Vegetative parts at first densely pale
tawny pubescent, caducous first on lamina, nervation
beneath, petiole and then twig, persistent on buds and

stipules. Twig c. 2 mm @ apically, becoming smooth,
glabrous. Bud to 2 by 1 mm, ovoid, acute. Stipule to 4
mm long, linear, caducous. Leaves 12-28 by 5-9 cm,
oblong-lanceolate to oblanceolate; base obtuse to
subcordate, unequal, the larger side adjacent to the
twig; acumen to 2 cm long, subcaudate; margin
somewhat revolute; nerves 12-21 pairs, slender, sca-
lariform, diagonal to midrib and nerves; midrib
prominent on both surfaces, more so beneath than
above; petiole 3-8 mm long, short, stout. Panicle to 8
cm long, terete, glabrous, slender, frequently fascicu-
late, frequently borne up the branches behind the leafy
twigs; singly branched, branchlets bearing to 8 secund
flowers. Flower bud to 3 by 2 mm, ellipsoid. Calyx
glabrous, 2 outer sepals lanceolate, acuminate, 3 inner
suborbicular, somewhat shorter, shortly mucronate.
Petals linear, pubescent on parts exposed in bud.
Stamens 15, in 3 unequal verticils; filaments com-
pressed at base, tapering and filiform below the
subglobose anthers; appendage to connective slender,
3-4 times as long as anthers. Ovary narrowly ovoid,
glabrous, surmounted by a spindle-shaped glabrous
stylopodium and style somewhat shorter than the
ovary in length. Fruit glabrous. Pedicel to 1 mm long,
short. 2 longer calyx lobes to 10 by 1.5 cm, spatulate,
chartaceous, c. 4mm broad above the to 2 cm long, to
7 mm wide paired basal auricles, with a to 5 by 4mm
central basal ovate saccate thickened area; 3 shorter
lobes to 3 cm long, with up to 12 by 4 mm tapering
subacute lobe above a similar base. Nut to 7 by 5mm,
ovoid, acute, entirely concealed by auricles.

Distr. Malesia: Borneo (Sarawak, Brunei).

Ecol. Locally common in Heath forest and ridge
tops; podsols and skeletal soils, to 1000 m.

Note. This species is variable in leaf size; some
collections of H. bullatifolia from Kalimantan
approach it, and the two species may eventually prove
conspecific.

81. Hopea philippinensis Dyer, J. Bot. 16 (1878) 100;
VIDAL, Phan. Cuming. (1885) 97; Rev. Pl. Vasc. Filip.
(1886) 62; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 64;
EVERETT & WHITFORD, Bull. Bur. For. Philip. 5 (1906)
16, 28, 53; WHITFORD, Bull. Bur. For. Philip. 10, 2
(1911) 75; Foxw. Philip J. Sc. 4 (1909) Bot. 515; ibid. 6
(1911) Bot. 261; ibid. 13 (1918) Bot. 183; ibid. 67 (1938)
276; MerR. En. Philip. 3 (1923) 94; Reyes, Philip. J.
Sc. 22 (1923) 338; Sym. Gard. Bull. S. S. 8 (1934) 32;
ibid. 8 (1935) 279; GUTIERREZ, Act. Manil. 4, A, 2
(1968) 65, f. 13, pl. 9.— H. odorata (non ROxB.)
VIDAL, Sinopsis (1883) t. 15, f. A. — Fig. 74 A—-A4.
Small, smooth-barked buttressed tree. Twigs,
petioles, domatia, leaf bud, stipules and parts of petals
exposed in bud densely persistently pale tawny pubes-
cent; midrib above (densely puberulent in young trees)
and panicle glabrescent. Twig c. 2 mm @ apically,
ribbed, becoming terete, red-brown. Buds small,
ovoid; stipules to 5mm long, linear, fugaceous. Leaves
(7-)12-25 by (2.5-)4-7 cm, narrowly elliptic-oblong
to lanceolate, thinly coriaceous; base unequal,
cuneate abaxially, cordate adaxially; acumen to 2 cm

1982]

long, slender, subcaudate; nerves 17—22 pairs, slender
but prominent beneath, obscurely depressed above,
arched, at S0°—60°, with axillary tomentose domatia,
without secondaries; tertiary nerves densely scalari-
form, very slender but evident and distinctly elevated
beneath; midrib slender but prominent beneath, ele-
vated above; petiole S—7 mm long, short. Panicle to 5
cm long, l-axillary to ramiflorous, slender, rather lax,
singly branched; branchlets to 1.5 cm long, bearing to
4 flowers; bracteoles minute, deltoid, fugaceous.
Flower bud to 3 by 2 mm, ellipsoid. 2 outer sepals
lanceolate-acuminate; 3 inner broadly ovate, shortly
acuminate. Stamens 15, slightly shorter than style at
anthesis; filaments compressed at base, tapering and
filiform in the distal 4; anthers subglobose; appen-
dages c. 2} times length of anthers, slender. Ovary and
stylopodium hour-glass shaped, the ovary slightly the
broader; style short but distinct. Fruit pedicel short,
base of fruit impressed. 2 longer calyx lobes to 12 by 3
cm, spatulate, obtuse, c. 3 mm broad above the to 9 by
8 mm ovate deeply saccate thickened base; 3 shorter
lobes to 7 by 8 mm, broadly ovate, mucronate, shorter
than nut. Nut to 11 by 8 mm, broadly ovoid, shortly
apiculate.

Distr. Malesia: Philippines.

Ecol. Widespread and common in evergreen non-
seasonal forest on hills to 500 m.

Vern. Bagitarim (Tag.), baguatsan, bagupsan
(Bik.), bantaya (S. L. Bis.), gisok (P. Bis., Mbo., Lan.,
C. Bis.), gisok gisok (P. Bis., Sul., Sh. Bis.), gisok
nga-salngan (S. L. Bis.), Auli lisian, maka tayring
(Tag.), malatamban (Bik.), malibato (Mbo.), mang-
laum (Tag.), malalamba, malatagum, pagak son (Bik.),
paina (Tag.), pongo, subigan (S. L. Bis.).

82. Hopea mindanensis Foxw. Philip. J. Sc. 6 (1911)
Bot. 261, pl. 43; ibid. 13 (1918) Bot. 183; ibid. 67 (1938)
277; Merr. En. Philip. 3 (1923) 93; Reyes, Philip. J.
Sc. 22 (1923) 334; Sym. Gard. Bull. S. S. 8 (1934) 32;
GUTIERREZ, Act. Manil. 4, A, 2 (1968) 63, f. 12, pl. 8.

Medium-sized hard-wooded tree, with blackish
flaky bark. Twigs, petioles, leaf buds and domatia +
persistently pale tawny pubescent, parts elsewhere
glabrescent. Twig c. 5mm @g apically, stout, becoming
terete, pale brown. Leaves 30-60 by 9.5—18 cm, large,
narrowly oblong, thinly coriaceous; base unequal,
cordate; acumen to 3 cm long, prominent; nerves
22-28 pairs, prominent beneath, + shallowly
depressed above ascending at 50°—60°, straight but
arched at the margin and running along parallel to it
before terminating; with small pubescent pore-like
axillary domatia; tertiary nerves + densely scalari-
form, very slender but distinctly elevated beneath;
petiole 15-18 mm long, c. 5 mm @g, short, relatively
stout, panicle to 6 cm long, short, to 2-axillary or
ramiflorous, singly branched. Flower buds to 3 by 2
mm, ovoid-lanceolate. 2 outer sepals ovate, acu-
minate; 2 inner suborbicular, acute. Stamens 15,
subequal; filaments compressed, rather broad, taper-
ing; appendages very slender, c. 14 as the ellipsoid
anthers. Ovary ovoid, surmounted by a prominent

DIPTEROCARPACEAE (Ashton)

435

spindle shaped stylopodium of equal length, with a
prominent intervening constriction; style shorter than
stylopodium but prominent, tapering. Fruit pedicel
short; 2 longer calyx lobes to 11 by 15 cm, narrowly
spatulate, obtuse, c. 3 mm broad above the to 9 by 7
mm ovate saccate thickened base; 3 shorter lobes to 12
by 8 mm, ovate, thin, saccate, appressed to nut; nut to
12 by 8 mm, ovoid, prominently apiculate, resinous.

Distr. Malesia: Philippines (Mindanao).

Ecol. Frequent in lowland evergreen forest.

Vern. Bagasusu (Zamboanga), magasusu (Sulu),
ganon (Sub.), yakal magasusu (official name).

83. Hopea tenuineryula ASHTON, Gard. Bull. Sing. 22
(1967) 281, pl. 27, 349 (phot. habit); Man. Dipt. Brun.
Suppl. (1968) 57.— H. philippinensis (non Dyer)
ASHTON, Man. Dipt. Brun. (1964) 107, f. 12.

Small to medium-sized stilt-rooted tree with papery
flaky bark. Young vegetative parts densely pale tawny
tomentose, persistent on buds and stipules, + persis-
tent on nervation beneath and midrib above; else-
where caducous. Twig to 2 mm @ apically, becoming
smooth, glabrous. Bud to 3 by 1.5 mm, lanceolate.
Stipule to 5 mm long, linear, subpersistent. Leaves
alternate, 10-27 by 3-S.5 cm, narrowly ovate to
lanceolate; base obtuse, pronouncedly unequal, the
larger side adjacent to the twig, acumen to | cm long,
narrow; margin revolute; nerves 12-21 pairs, slender,
prominent beneath, at 50°-60°; curving and running
parallel to margin before terminating at it; tertiary
nerves slender, densely scalariform, at 90°; midrib
rounded, raised on both surfaces; petiole 3-7 mm
long, short, stout. Panicle to 8 cm long, to 2-axillary,
rarely terminal, terete, lax, glabrous; singly branched,
branchlets to 3 cm long, bearing to 4 secund pale
yellow flowers; bracteoles to 1 mm long, deltoid,
glabrous, subpersistent. Bud to 4 mm long, ellipsoid,
subsessile. Ca/yx glabrous but for fimbriate margin; 2
outer lobes narrowly ovate, prominently acuminate; 3
inner lobes suborbicular to broadly ovate, mucronate.
Petals oblong, lanceolate, acute, densely pubescent on
parts exposed in bud, otherwise glabrous. Stamens 15;
filaments broad at base, tapering; anthers oblong-
ellipsoid; appendage to connective slender, about
twice length of anther. Ovary ovoid, glabrous; stylo-
podium as long as ovary, spindle-shaped, glabrous,
indistinct from style. Fruit entirely glabrous; 2 longer
calyx lobes to 10 by 1.7 cm, spatulate, narrowly
obtuse, to 2 mm broad above the to 6 by 4 mm
narrowly ovate prominently saccate thickened base; 3
shorter lobes to 3 cm long, subequal, acute, slightly
auriculate at the to 7 mm broad base but otherwise
similar. Nut to 12 by 8 mm, ovoid, almost completely
enveloped by shorter calyx lobes; style remnant short,
acute.

Distr. Malesia: Borneo (Sarawak, Brunei, S.E.
Borneo).

Ecol. Locally abundant on leached sandy soils in
Mixed Dipterocarp forest on low hills near Pleisto-
cene coastlines.

Vern. Mérawan daun serong.

436

84. Hopea enicosanthoides ASHTON, Gard. Bull. Sing.
22 (1967) 276, pl. 21; Man. Dipt. Brun. Suppl. (1968)
50, f. 6.

Small smooth-barked stilt-rooted tree. Young
twigs. buds, stipules, petioles and base of midrib
above caducous pale tawny pubescent or glabrous.
Twig c. 3 mm g, terete to somewhat compressed,
smooth, prominently ribbed below the petiole inser-
tion; stipule scars short, obscure. Bud to 2 by 1 mm,
conical, acute. Stipule to 8 mm long, linear, subpersis-
tent. Leaf (16-)27-46 by (5—)8-15 cm, very large,
oblong, coriaceous, prominently convex between the
depressed nerves on the upper face; base cordate,
unequal; acumen to 2.5 cm, long, slender; nerves
16-30 pairs, slender but prominent beneath; tertiary
nerves densely scalariform, slender but evident
beneath; petioles 5-8 mm long, short, stout. Flowers
unknown. Panicle and fruit entirely glabrous. Panicle
to 12 cm long, axillary to ramiflorous, frequently to
2-axillary, terete, lax, singly branched; branchlets to 3
cm long, ascending, bearing to 5 flowers; bracteoles c.
2mm long, linear, subpersistent. Pedicel c. 1 mm long,
short. 2 longer calyx lobes to 13 by 3 cm, + broadly
spatulate, obtuse, c. 6mm broad above the c. 12 by 10
mm ovate thickened saccate base; 3 shorter lobes to 20
mm long, lanceolate, acute, similarly saccate but
hardly thickened at base, enfolding the nut. Nut to 10
by 6mm, ovoid; style remnant to 2 mm long, filiform.

Distr. Malesia: Borneo (Sarawak, from the Rejang
R. to Miri).

Ecol. On low, damp hillsides and, most frequently,
banks of sluggish or tidal, but not brackish rivers, but

FLORA MALESIANA

[ser. I, vol. 9?

apparently intolerant of prolonged root immersion.
Locally frequent.

Dubious

Hopea parvifolia (WARB.) SLooT. Reinwardtia 2
(1952) 37. — Anisoptera parvifolia WARB. Bot. Jahrb.
13 (1891) 382; BRANDis, J. Linn. Soc. Bot. 31 (1895)
45; Diecs, Bot. Jahrb. 57 (1922) 461; SLoor. Bull.
Jard. Bot. Btzg III, 8 (1926) 5 (misspelt A. parviflora);
Nova Guinea 14 (1926) 225.

Nothing is known of this species apart from the
inadequate original description; the type (WARBURG
20034) has not been found by myself and was not seen
by BRANDIS or VAN SLOOTEN either. It was apparently
in fruit, though it is not even possible from the
description to be certain of the genus.

Excluded

Hopea gracilis MiQ. Sum. (1860) 490; DC. Prod. 16, 2
(1868) 635. BURCK (Ann. Jard. Bot. Btzg 6, 1887, 237)
correctly excluded this species, based on a sterile
TEYSMANN collection from Padang, Sumatra at
Utrecht, from the family. Its correct identity remains
obscure.

Hopea siranda Miq. Sum. (1860) 489 = Annonaceae.

Hopea sumatrana KiNG ex GILG in E. & P. Pfl. Fam.
ed. 2, 21 (1925) 249, nomen. Quoted as producing resin
of high quality. There is no other record of this name,
which was certainly never published by KING.

10. SHOREA

ROxB. ex GAERTN. f. Fruct. 3 (1805) 48; Roxs. Pl. Corom. 3 (1815) t. 212; B. & H.
Gen. Pl. 1 (1862) 193; DC. Prod. 16, 2 (1868) 628; Dyer, FI. Br. Ind. 1 (1874) 303;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 204; Herm, Rech. Dipt. (1892) 36, incl. sect.
Anthoshorea HEIM, /.c. 41, sect. Hopeoides HEM, /.c. 43, sect. Pachycarpae HEM,
l.c. 44, sect. Richetioides HEM, I/.c. 48; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 73,
incl. sect. Pinanga BRANDIS, /.c. 90, sect. Mutica BRANDIS, /.c. 100; Foxw. Philip.
J. Sc. 67 (1938) 290, incl. sect. Isoptera (SCHEFF. ex BURCK) Foxw. /.c. 291, 301;
SyM. Mal. For. Rec. 16 (1943) 1; MEER, Act. Bot. Neerl. 12 (1963) 322, incl. subg.
Richetia (HEM) MEER, nom. inval., subg. Rubroshorea MEIER, |.c. 322; ASHTON,
Gard. Bull. Sing. 20 (1963) 261, incl. sect. Neohopea ASHTON, I.c. 266, sect.
Rubella ASHTON, /.c. 267, sect. Ovalis ASHTON, /.c. 268; Man. Dipt. Brun. (1964)
115; ibid. Suppl. (1968) 60; MEER & Woop, Sabah For. Rec. 5 (1964) 48;
ASHTON, Blumea 20 (1972) 360; Gard. Bull. Sing. 31 (1978) 36, incl. sect.
Pentacme (A. DC.) ASHTON, /.c. 38; SMITINAND, Thai For. Bull. (Bot.) 12 (1980)
57. — Saul Roxs. ex W. & A. Prod. (1834) 84, nomen. — Doona THw. in Hook.
Kew J. 3 (1851) t. 12; ibid. 4 (1852) 7; Dyer, FI. Br. Ind. 1 (1874) 311; cf. ASHTON,
Blumea 20 (1972) 361. — Pentacme A. DC. Prod. 16, 2 (1868) 626; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 72; Sym. Mal. For. Rec. 16 (1943) 104, f. 63
(map). — Isoptera SCHEFF. ex BURCK, Med. Lands PI. Tuin 3 (1886) 27; BRANDIS,

1982] DIPTEROCARPACEAE (Ashton) 437

J. Linn. Soc. Bot. 31 (1895) 105. — Ridleyinda O. K. Rev. Gen. Pl. 1 (1891)
65. — Richetia HeEIM, Bull. Mens. Soc. Linn. Paris 2 (1891) 975. — Anthoshorea
PIERRE ex HEIM, Rech. Dipt. (1892) 41, nomen in syn. — Parahopea Heim, L.c.
66. — Pachychlamys (DYER ex KING) DYER ex RIDL. Fl. Mal. Pen. 1 (1922)
233. — Fig. 77-118.

Medium-sized or large buttressed trees without stilt-roots; mature crown large,
hemispherical or dome-shaped, sympodial. Jnflorescence paniculate. Calyx lobes
free to receptacle; 3 outer lobes thicker, somewhat longer, narrower, than 2 inner
lobes in flower. Petals usually connate at base on falling, sometimes free. Stamens
10—co; filaments variable, lorate to filiform; anthers subglobose to narrowly
oblong; appendage to connective vestigial or prominent. Ovary tomentose, rarely
glabrous; style with or without a distinct stylopodium. 3 outer fruit calyx lobes
usually much longer than 2 inner lobes, thin, spatulate; or all lobes subequal; base
of lobes + thickened, expanded, saccate. Nut free from calyx, pericarp splitting
irregularly at germination. Unicellular scales, if present, broad-lobed.

Distr. About 194 spp. in Ceylon, India, Burma, Thailand, Indochina, and 163 spp. in Malesia: Malaya,
Sumatra, Borneo and intervening islands, Java, Philippines and Moluccas. Fig. 79.

Fossil wood has been identified as Shoreoxylon from Timor in the Lesser Sunda Islands by SCHWEITZER
(Palaeontographica 105B, 1959, 1-66) in which islands Shorea does not occur at present.

Ecol. The dominant emergent tree genus of the lowland forests of West Malesia; others occur and often are
gregarious on river banks, on podsols and peat swamps; a few are montane, to 1750 m; some do not emerge from
the main canopy.

Uses. The most important timber genus in the Asian humid tropics. Sections Shorea, Pentacme (Asian species
only) and Neohopea yield a hard heavy timber suitable for construction and decking (balau, Malaya, sélangan
batu, Borneo, yakal, Philippines); sect. Richetioides a yellowish brown light hardwood (yellow méranti, Malaya,
yellow séraya, Sabah, lun, Sarawak); sect. Anthoshorea a white siliceous timber veneers (white méranti, Malaya,
raruk, Sarawak, mélapi, Sabah), and the remaining sections pink, red or occasionally white or brown light
medium or sometimes heavy hardwoods favoured for furniture as well as light construction (red méranti,
Malaya, /up, pérawan, Sarawak, red séraya, Sabah, red lauan, Philippines).

Note. The subdivision of the genus accepted here has been more fully discussed by me in my paper of 1963.

SUBDIVISION OF SHOREA IN MALESIA

1. Appendage to connective barbate, stout, anthers 4-locular, with apices of anterior loculi barbate. Fig. 78, 82
1. Sect. Shorea
2. Flower buds elongate, petals narrow ae separately; appendage to connective sparsely ciliate. Fig. 78.

Spel —Otaes. \ -. . .la. Subsect. Shorea
2. Flower buds globose before anthesis: “petals ‘short, ‘oblong, connate at base on falling; appendages to
connective and apices of anther cells densely barbate. Fig. 82. Spp. 29-35 . . 1b. Subsect. Barbata

1. Appendages, if barbate surmounting a 2-locular anther, otherwise scabrous or glabrous; anthers glabrous.
3. Anthers linear, prolonged apically into prominent horn-like processes. Fig. 84. Spp. 36-37
2. Sect. Pentacme
3. Anthers not as above.
4. Appendage to connective not exceeding anther-apex, stout; stamens 15; anthers subglobose. Fruit calyx
lobes subequal, elongated, much longer than the nut. Fig. 85. Sp.38 . . . . 3. Sect. Neohopea
4. Appendage exceeding anther apex or, of short, either slender, recurved or with more than 30 stamens. Fruit
calyx lobes subequal but shorter than nut, or unequal, 3 eases and spatulate.
5. Anthers with 2 pollen sacs. Fig. 87,89... . . . . . . 4, Sect. Richetioides
6. Stamens more than 100; anthers lorate. Fig. 87. Sp. 39... ; . 4a. Subsect. Polyandrae
6. Stamens at most 17; anthers subglobose or oblong. Fig. 89. Spp. 40-71 . 4b. Subsect. Richetioides
5. Anthers with 4 pollen sacs.
7. Anthers linear to oblong; filaments lorate or tapering gradually (not abruptly medially); ovary without
distinct stylopodium; style at least as long as ovary.
8. Appendage to connective at least as long as anther, frequently scabrous. Wood yellow or white, with
reticulate vessel arrangement in TS. Female silica present. Fig. 91 B—C, 92. Spp. 72-92
5. Sect. Anthoshorea

438 FLORA MALESIANA [ser. I, vol. 9?

Fig. 77. Shorea falciferoides Foxw. a. Habit, b. leaf of seedling 1.2 m high, c. fruit, d. nut, all x $(aSAN 37512,5
S 5718, c-d S 2125).

8. Appendage to connective less than } the length of the anther; short, glabrous. Wood pink, with loose
oblique vessel-arrangement in TS. Silica absent. Fig. 94, 95. Spp. 93-97. . . .6. Sect. Rubella
7. Anthers broadly oblong to subglobose; appendage to connective vestigial, or shorter than the anther
and reflexed, or, if long and unreflexed filaments tapering + medially or united in a tube round the
ovary; stylopodium usually distinct. Wood pink, without silica; pore arrangement oblique, clustered or
solitary, variable though never reticulate.

9. Stamens less than 30; filaments compressed and broad at base.

10. Appendage to connective filiform, slender, at least twice as long as the anther, not reflexed; filaments
broad at base, tapering abruptly and filiform below the anthers.

11. Filaments broad at base, tapering abruptly medially, filiform, not fixed into a distinct tube round
the ovary though frequently connate at base; ovary either with a distinct stylopodium and +
pyriform, or without and small and ovoid; style filiform. Fig. 91D, 98-100

7. Sect. Brachypterae
12. Stamens 24-28. Fig. 98. Sp.98 . . so) MP ko aon eo 7a. SubseceSmithiana
12. Stamens 15. Fig. 91D, 99-100. Spp. 99- 122 i935 2) $2 Tb. Subsecit Brachypterae

1982] DIPTEROCARPACEAE (Ashton) 439

11. Filaments united into a tube around the base of the ovary, tapering abruptly near apex;
stylopodium merging into the style; style and stylopodium spindle-shaped, narrow and tapering at
base and apex; or stylopodium indistinct, style at least twice as long as ovary, filiform. Fig. 103, 104.

Spp., 123-32: 3 . . . 8. Sect. Pachycarpae
10. Appendage to connective shorter than o oras long as s the anther, becoming reflexed; filaments tapering
gradually from base to anther. Fig. 106,107. . . sy Nae RaoSect. Mutea
13. Fruit calyx lobes auriculate at base. Fig. 107. Spp. 133- 138. ae, fea Oat Subsect. Auriculatae
13. Fruit calyx lobes not auriculate. Fig. 106. Spp. 139-159. . . . . .9b. Subsect. Mutica

9. Stamens 50-70, appendage to connective vestigial; filaments filiform. ‘Fig. 115. Sp. 160
10. Sect. Ovalis

KEY TO THE SPECIES*

1. Appendage to connective barbate, stout, anthers 4-locular with apices of anterior loculi barbate. Fig. 78, 82.
Spp. 1-35.1. Sect. Shorea
2. Flower buds elongate, petals narrow, falling separately; appendage to connective sparsely ciliate. Fig. 78.
Spp. 1-28. 1a. Subsect. Shorea
3. Appendage to connective typically with 1-4 (a few appendages with up to 8) bristles, stamens otherwise
glabrous.
4. Stamens more than 50; fruit calyx lobes vestigial though aliform and slightly exceeding ripe nut
1. S. collina
4. Stamens at most 30; fruit calyx not as above.

5. Leaf undersurface persistently scabrid pubescent . . . . . . . +. +. 2. S. ochrophloia
5. Leaf undersurface glabrescent.
meres aticase 1S pairs 2 854. ee peay >: Late ee ome! ea ee eS ee
6. Nerves at most 14 pairs.
7. Fruit calyx lobes short, subequal . . . . -. =. - -.. - ~- =. ~. +. 4 S.bavilandi
7. Fruit calyx lobes unequal, 3 aliform.
Seesiamens aeleast 40 2 Se ee re ee eS. - ORECSCCMS
8. Stamens less than 35.
9. Appendage mostly with a single apical bristle . . . . . . . . . 6. S.scrobiculata
9. Appendage with 2-4 bristles.
10. Petiole up to 14mm long; stamensc.30 . . . . . . . . . +. ‘7. S. leptoderma
10. Petiole at least 15 mm long; stamens less than 28.
RiteStamensic. 26sleamlancedlate 2 6) -) 2 8 = 9. 3) 2 oe 8. S. ciliata
11. Stamens 20; leaf ovate. . eA =. submontana

3. Appendage to connective with at least 5 bristles, or shoulder of filaments and usually anther apices setose.
12. Fruit calyx lobes subequal, shorter than nut.

13. Stamens c. 55, nutinfruituptoSbyScm. .. . . . . . . . .- 10. S. geniculata
13. Stamens 25—40, nut in fruit c. 1 by 1 cm.
Pastamens 50-408) oo ee ee ST ce a, te ee Sey eS
14-Stamens 25° = . . . . . 12. S. sumatrana

12. Fruit calyx lobes unequal, 5 longer lobes greatly exceeding nut.
15. Filaments distinctly barbate along the distal lateral margins.

16. Leaf of mature tree not cream or white lepidote beneath. . . . . . . 13. S. foxworthyi
16. Leaf of mature tree distinctly white or cream lepidote beneath.
{7 ‘Stamens ‘at most 24. so we Oe ey 2 es AS ess

17. Stamens at least 25.
18. Young twigs compressed.
19. Tomentum on twig and petiole yellow-brown, scabrid; petiole less than 1.5 cm long
15. S. exelliptica
19. Tomentum on twig and petiole buff, even; petiole 1.5-2.6cmlong . . 16. S. inappendiculata
18. Twigs entirely terete.
20. Leaf falcate, base subequal, nerves beneath slender.
21 Stamens 33-44 2 5S OS 2 OR 8 8 ee i a ee
Fi Stamens 25-92". 9 POS. ee a A er ee L

De eee eee

* Flowers are unknown from the spp. 161-163 which can, for that reason, also not be placed in a section, and
are omitted from the key.

440 FLORA MALESIANA [ser. I, vol. 9?

20. Leaf subequal, base equal, nerves beneath stout.

224sStamens:25=33i5 slash ale: boot e hap ln ies we Oe Sigal S satrinenvosa
22. Stamens 35-46.

23 Petioles S-sicmilong,.. : 5 2 = «= «1 .5: 0 ¢5 «5 <> =o nfl en RoMZO MBS acrassal
232eRetiolesl2—vemilonye silky ©.) oc GIR MS ME) Ee O71 eI Sobscara’

15. Filaments glabrous.
24. Stamens more than 40.

25. Leaf subequal, not lepidote beneath; nerves 11-15 pairs, stout beneath . . 22. S. lunduensis

25. Leaf falcate, cream lepidote beneath; nerves less than 12 pairs, slender beneath

23. S. falciferoides

24. Stamens 37 or less.

26. Nerves of leaf 16-24 pairs, pilose beneath . . . . . =. =. =. =. +. +. 24. S. superba

26. Nerves of leaf less than 16 pairs, epilose beneath.

27. Nerves 11-16 pairs; domatia if present very small.

28. Stamens c. 33; nerves applanate above. . . . 4 + + ge ee op eZee Sally polencal
28. Stamens 35-37; nerves narrowly depressed above . Jaak: . . . . 26. S. malibato
27. Nerves at most 12 pairs, with prominent pore-like axillary domatia.

29. Ovary with prominent stylopodium; stamensc.32. . . . . . . +. +. 27. S. astylosa
29. Ovary without stylopodium; stamens 25-30... 28. S. domatiosa

. Flower buds globose before anthesis; petals short, oblong, connate at base on 1 falling; appendages to
connective and apices of anther cells densely barbate. Fig. 82. Spp. 29-35. 1b. Subsect. Barbata
30. Stamens at least 45.
315, Meat 6-15 by 2:3-9 cm;merves 7-l0)pairs. . - « = = = 35 5 = =) ceo ES selanca
3) Leaf 6:5—10'by 2:54 cm; nerves 11-14 pairs. . . . . =. = | «= 4» = SO liSalaevis
30. Stamens at most 35.
32. Nerves 6-7 pairs, not sunken above; leaf less than 8 cm long.
33. Fruit calyx lobes subequal, shorter than nut. ws a te pS eas wasabi
33. Fruit calyx lobes unequal: 3 spatulate, thin, much longer than nut . « 8 4 ps 932s Samicans
32. Nerves more than 8 pairs, depressed above or, if not depressed, leaf exceeding 10 cm long.
34. Fruit calyx lobes subequal, shorter than nut.

35. Leaf coriaceous, nerves 5—6 pairs, not depressed above . . »; . «. «= « 2 33a Saladiana
35. Leaf thin, nerves more than 8 pairs, depressed above. . . > = « = 2 S45 Soblawals
34. Fruit calyx lobes unequal, 3 spatulate, thin, much longer than nut . . . . 3§. S. maxwelliana

1. Appendages, if barbate, surmounting a 2-locular anther, otherwise scabrous or glabrous; anthers glabrous.
36. Anthers linear, prolonged apically into prominent horn-like processes. Fig. 84. Spp. 36-37.
2. Sect. Pentacme
37. Deciduous hard-wooded trees; leaf base equal, nerves 13-16 pairs . . . . . .36. S. siamensis
37. Evergreen soft-wooded trees; leaf base unequal, nerves 7-9 pairs . . . . . . 37. S. contorta
36. Anthers not as above.
38. Appendage to connective not exceeding anther apex, stout; stamens 15; anthers subglobose. Fruit calyx
lobes subequal, elongated, much longer than the nut. Fig. 85. Sp. 38. 3. Sect. Neohopea
38. S. isoptera
38. Appendage exceeding anther apex or, if short, either slender, recurved or with more than 30 stamens.
Fruit calyx lobes subequal but shorter than nut or unequal, 3 prolonged and spatulate.
39. Anthers with 2 pollen sacs. Fig. 87, 89. Spp. 39-71. 4. Sect. Richetioides
40. Stamens more than 100; anthers lorate. Fig. 87. Sp. 39. 4a. Subsect. Polyandrae
39. S. polyandra
40. Stamens at most 17; anthers subglobose or oblong. Fig. 89. Spp. 40-71. 4b. Subsect. Richetioides
41. Fruit calyx shorter than nut.
42. Fruit calyx lobes unequal, spatulate . . . . . . . . +. +. +. + +. 40. S. kuantanensis
42. Fruit calyx lobes subequal, + ovate.
43. Nut glabrous.

44. Petiole 1-1.2 cm long; floral ovary glabrous; stamens 15. . . . . .~ «Al. S. longiflora

44. Petiole 1.8—3.8 cm long; floral ovary pubescent; stamens 10. . . . 42. S. macrobalanos
43. Nut pubescent.

AS ulecatuotamaturestreejpeltates = 1) 2.) 26. 1s | elenn =) des Soa Pelli

45. Mature tree leaf not peltate.

46. Leaf less than twice as long as broad.
Aj<Stamenus 6-17... s . « = = © + © -5 © we 6 os 2) Oo nines
47. Stamens 10-15.
48. Leaf nervation persistently cream pubescent beneath. . . . . . . . 45. S.laxa

1982] | DIPTEROCARPACEAE (Ashton) 44]

48. Leaf entirely glabrous.
49. Leaf 7-12 cm long, nerves 5-7 pairs; nut to 3 by 1.3 cm, drying mauve tomentose
46. S. balanocarpoides
49. Leaf less than 8 cm long, nerves 7 or more pairs; nut to 2 by 1.2 cm, cream-buff tomentose or
glabrous.
50. Nerves 8-10 pairs, hardly raised beneath; leaf base cuneate but not decurrent at petiole
insertion; petiole drying rugulose, black; style without distinct stylopodium

: A. : 47. S. multiflora
50. Nerves 7-9 pairs, slender but distinctly raised beneath; leaf base cuneate, somewhat

decurrent; petiole drying smooth, black; style with distinct stylopodium
48. S. patoiensis
46. Leaf at least twice as long as broad.

51. Leaf grey-brown scabrid tomentose beneath; base cordate . . . . 49. S. induplicata
51. Leaf not as above.
52. Nerves and midrib sunken in shallow furrows above. . . . . .50. S. subcylindrica

52. Leaf nervation not as above.
53. Leaf nervation pubescent beneath.
54. Leaf base equal; midrib prominently depressed, obscure, above . . 51. S. obovoidea
>4. Leaf base unequal; midrib not as above. = . .. . . =. = . “$2. 'S. chaiana
53. Leaf nervation glabrous beneath.
55. Petiole persistently pale cream puberulent throughout.
56. Petiole San 19 mm long, leaf undersurface matte, pets stamens 10; fruit calyx +
patentee = Ewe 2753. +S: collaris
56. Petiole to 15 mm long: leaf undersurface and fruit calyx not : as above.
57. Petiole 6-8 mm long; flower buds to 2.5mm long; stamens 15; panicle to 10 cm long
54. S. angustifolia
57. Petiole exceeding 10 mm long; flower buds exceeding 12 mm long; stamens 10; panicles to

7mmlong. . . RING (Ove, Aiea SSS smaxmia
55. Petiole epilose, exceeding 8 mm long.

58. Leaf undersurface matte, distal end of petiolecream. . . . 56. S. tenuiramulosa

58. Leaf undersurface lustrous; petiole drying entirely black.
59 eNerves:at most: G pairs sees HST Be eg OS ee Se S75 Sy conica

59. Nerves at least 9 pairs.

60. Leaf coriaceous, midrib applanate above . . . . . . . .58. S. bakoensis
60. Leaf thin, midrib elevated above. . . . . . .59, S. xanthophylla

. Fruit calyx lobes unequal, 3 prolonged, thin, spatulate, sreatly Sree the nut.
oe Leaf large, thickly coriaceous, with 14-16 pairs of slender nerves and tertiary nerves equally elevated
‘onjedch SUDfACE ae =e) eis re ee) Pe ee ee 00 SS blunmtensis
61. Leaf not as above.
62. Petiole at least 19 mm long, nerves at least 10 pairs.

63. Petiole 3-4 mm @, stout; leaf thickly coriaceous . . . . . . . . +. =. 61. S. iliasii

63. Petiole to 2mm g, slender; leaf chartaceous . . . . . 62. S. faguetioides
62. Petiole less than 18 mm long or, if longer, than nerves less than 9 pairs.

64suNervesr14=19 sparse 2) 710 YA §C-Sie tee OLE toe 2 eS He ee, PEGS) Ss alutacea

64. Nerves at most 12 pairs.
65. Leaf nervation beneath persistently tomentose.

66. Leaf undersurface densely pale grey-green pubescent . . . . . . 64. S. longisperma
66. Leaf undersurface glabrous but for nervation.
67. Leaf margin revolute, nerves 9-12 pairs, tomentum scabrid . . . 65. S. acuminatissima
67. Leaf margin not revolute, nerves 7-9 pairs, tomentumeven. . . . . 66. S. gibbosa

65. Leaf nervation beneath fugaceous puberulent or glabrous.
68. Leaf nerves very slender, hardly raised beneath, with distinct secondaries; midrib drying reddish

or blackish, frequently with a pair of glabrous domatia at base . . . 67. S. hopeifolia
68. Leaf not as above.
69. Petiole 17-22 mm long, nerves 7-9 pairs . . . . . . . +. +. 68. S. kudatensis
69. Petiole at most 16 mm long.
70: Nerves:5—7 paitss <0 2 4.9. 4 Oke 6 Spel se ep ee EE BS oeeaia
70. Nerves 8 or more pairs.
71. Nerves c. 8 pairs, prominently raised beneath . . . . . . . 70. S. mujongensis
71. Nerves 9-12 pairs, hardly raised beneath . . . . . . +. +. # Ti. S. faguetiana

39. Anthers with 4 pollen sacs.

442 FLORA MALESIANA [ser. I, vol. 9?

72. Anthers linear to oblong: filaments lorate or tapering gradually (not tapering abruptly medially); ovary
without distinct stylopodium; style at least as long as ovary.
73. Appendage to connective at least as long as anther, frequently scabrous; wood yellow or white, with
reticulate vessel arrangement in TS; silica present. Fig. 91B—C, 92. Spp. 72-92.
5. Sect. Anthoshorea
74. Stamens at least 17.
75. Stamens 17; nerves 20-24 pairs; leaf undersurface pale pink lepidote. . . . 72. S. dealbata
75. Stamens at least 20.
76. Stamens less than 40.
77. Leaves elliptic-oblong; petiole 2-3 cm long; semi-deciduous tree . . . . 73. S. farinosa
77. Leaves eliptic lanceolate or broadly elliptic-ovate; petiole less than 2 cm long; evergreen trees.
78. Nerves at most 14 pairs; leaves broadly elliptic-ovate.

79. Stamens 21-25; tertiary nerves scalariform. . . . . . . . . . .74.S. polita
79. Stamens 25; tertiary nerves subreticulate . . . . . . . . . 75. S. gratissima
78. Nerves 17-20 pairs; leaves elliptic-lanceolate . . . . . . . +. .76. S. henryana
76s Stamensimore than'45" 2). = = 29) Avil ay Eel toe) ee ee eS Onegers

74. Stamens 15.
80. Stipules ovate, auriculate, not at first caducous; leaves not usually exceeding 9 by 4.5 cm (in mature
trees), with at least 13 pairsof nerves... » 1:03) S785.SFassamica
80. Stipules not as above, caducous; leaves at least l0c cm long or with less than 13 pairs of nerves.
81. Leaf nervation beneath persistently tomentose.
82. Twigs, petioles, and nerves and midrib beneath red-brown to cream-brown pubescent; leaf
undersurface bright yellow lepidote . . , lie a7 Ssochracea
82. Tomentum dull grey to tawny brown, leaf undersurface dull grey.
83. Leaves typically obovate, with at most 17 pairs of nerves; twigs prominently compressed
80. S. virescens
83. Leaves typically ovate-oblong; nerves 17—26 pairs; twigs terete.

84. Tomentum short, even; anther thrice aslongas broad . . . . . . 81. S. javanica
84. Tomentum uneven, scabrid; anther twice aslongas broad . . . ._ . 82. S. lamellata
81. Leaf nervation beneath early glabrescent.
85. Midrib evident, applanate above; deciduous tree. . . . . . . . 83. S. roxburghii
85. Midrib obscure, depressed above; evergreen trees.
86. Fruit calyx lobes vestigial, about twice length of nut . . . . . . 84. S. bentongensis

86. Fruit calyx lobes at least thrice as long as nut.
87. Nerves at least 15 pairs; twig apices compressed.

88. Petiole at least 2 cm long; leaf undersurface cream lepidote in mature trees

85. S. hypochra
88. Petiole less than 2 cm long: leaf undersurface not cream lepidote

89. Fruit pedicel c. 1 cm long, c. 5mm g, very stout and prominent; 3 longer fruit calyx lobes to
18cmlong. . . . . 86. S. symingtonii
89. Fruit pedicel at most 2n mm long, not prominent: fruit calyx lobes not exceeding 13 cm long.
90. Leaf to 12 by 4.8 cm, narrowly elliptic to lanceolate; base obtuse or broadly cuneate;

undersurface coppery lepidote . . . . . . 87. S. retinodes
90. Leaf at least 8 by 5.5 cm, oblong to obovate; base cordate: undersurface not lepidote
88. S. cordata

87. Nerves at most 15 pairs or, if more, then twigs terete.
91. Twig apices + compressed; leaf subchartaceous, drying chocolate brown
89. S. bracteolata
91. Twigs terete; leaf coriaceous, drying tawny to pale yellow-brown.

92. Leaf base cuneate . . ste by Cp Ee 2ciicemn hav St Sees
92. Leaf base obtuse to cordate.
93. Base of fruit impressed; flower bud to5by2.5mm. . . . . . . 91. S. agamii
93. Base of fruit tapering; flower budto9by5mm . . . 92. S. confusa

73. Appendage to connective less than 3 length of anther, short. glabrous. Wood pink, with loose oblique
vessel arrangement (in TS); silica absent. Fig. 94, 95. Spp. 93-97. 6. Sect. Rubella
94. Midrib obscure above; twigs compressed; stamens 20-24. . . . . . . 93. S. albida
94. Midrib evident above; twigs terete.
95. Stamens at most 20.
Dorstamens15) 4c a) Soc dieeted geen guruoo. 4. OR See
GeeSiamenst9-20) 0. 4 2 ys fs Sekeeed toning. oly Seer CESS alee
95. Stamens at least 25.

1982] DIPTEROCARPACEAE (Ashton) 443

OFMESTAMENSIC W255 hee rae se het LRN) © ve Pe es ee eee) 1. 96:2 Sudispar
Offers tamensica48 a . . . 97. S. negrosensis
72. Anthers broadly oblong to subglobose; appendage to connective vestigial, or shorter than anther and
reflexed, or, if long and unreflexed, filaments abruptly tapering + medially or united in a tube round
the ovary; stylopodium usually distinct. Wood pink, without silica; pore arrangement oblique,
clustered or solitary, variable though never reticulate.
98. Stamens less than 30; filaments compressed and broad at base.
99. Appendage to connective filiform, slender, at least twice as long as anther, not reflexed; filaments
broad at base, tapering abruptly and filiform below the anthers.

100. Filaments broad at base, tapering abruptly medially, filiform, not fixed into a distinct tube round
the ovary though frequently connate at base; ovary either with a distinct stylopodium and +
pyriform, or without and small and ovoid; style filiform. Fig. 91D, 98-100. Spp. 98-122.

7. Sect. Brachypterae
01. Stamens 24-28. Fig. 98. Sp. 98. 7a. Subsect. Smithiana
Only species. 98. S. smithiana
101. Stamens 15. Fig. 91D, 99-100. Spp. 99-122. 7b. Subsect. Brachypterae
102. Ovary ovoid, stylopodium indistinct or absent; style as long as ovary and stylopodium or
longer, frequently + pubescent in the basal half.
103. Leaf base distinctly unequal, midrib curved to one side.
104. Leaf broadly ovate; nerves at most 13 pairs . . . . . . . 99. S. inaequilateralis
104. Leaf oblong-ovate; nerves at least 19 pairs . . . . . . . . . 100. S. selanica
103. Leaf base equal.
105. Leaf undersurface, petiole and twig scabrid tomentose.
106. Leaf coriaceous, margin revolute, cream lepidote beneath; longer fruit calyx lobes to 6.5 by
IS Gon 4 « . . . . 101. S. flemmichii
106. Leaf + chartaceous, not lepidote beneath; margin not revolute; longer fruit calyx lobes
exceeding 10 by 2 cm.
107! Weaticoncavenw AR) irae ep Re a ae Ft eee 0G1025°S. almon
107. Leafapplanate . . . . . 103. S. parvistipulata
105. Leaf undersurface, petiole, and twig densely shortly evenly pubescent to glabrous.
108. Nerves at least 13 pairs, very slender, hardly elevated beneath; tertiary nerves densely
scalariform beneath, unraised.
109. Leaf beneath prominently cream lepidote . . . . . . . . 104. S. balangeran
109. Leaf entirely glabrous.
110. Leaf 10-15 by 5—8 cm; base subpeltate, discernible as a rib over the base of the midrib in
mature trees; stipule scars cuneate. . . des 105. S. coriacea
110. Leaf 6-10 by 3—S cm; base not subpeltate; stipule scars amplexicaul 106. S. venulosa
108. Nerves, if more than 13 pairs, prominent beneath and tertiary nerves not as above.

111. Nerves at least 16 pairs, leaf pinkish cream lepidote beneath . . . 107. S. waltoni
111. Nerves most 16 pairs and then leaf not pale lepidote beneath.
112 Petiole4=6:em long) +2 “ue Se). eee: eae ey Se SOSS> pachyphylla
112. Petiole at most 3.5 cm long.
113. Appendage to connective setose . . . . . . . . . . 109. S. paucifiora

113. Appendage glabrous.
114. Leaves with to 3 pairs of white scale-like domatia at base; style glabrous except at base .
110. S. johorensis
114. Leaves without white scale-like domatia.
115. Fruit calyx lobes short, subequal . . . . . . . ~~~ IMI. S. palembanica
115. Fruit calyx lobes long, aliform, unequal.
116. Leaf undersurface cream lepidote (mature trees); corolla mauve; nut to 14 by 7 mm
112. S. andulensis
116. Leaf undersurface glabrous; corolla cream; nut to 18 by 14mm 113. S. polysperma
102. Ovary and stylopodium distinctly pyriform, the stylopodium distinct from the style; style not
longer than ovary and stylopodium, glabrous.
117. Twigs compressed, nerves hardly raised beneath; midrib sharply acute beneath
114. S. platyclados
117. Twigs terete, nerves distinctly raised beneath; midrib prominent beneath but not sharp.
118. Fruit calyx lobes relatively short, less than twice as long as ripe nut.
119. Style and stylopodium densely pubescent but for the apex, hardly distinguishable from one
another, filiform at the apex, stouter beneath, frequently further swollen in the basal half;
fipemutto’s by 25cm); A eee el eS Seabird

444 FLORA MALESIANA [ser. I, vol. 97

119. Style and stylopodium well defined; ovary and stylopodium ovoid, crowned by an equally

long filiform glabrous style; ripe nutto3.5by1.5cem. . . . . . 4116. S. fallax
118. Fruit calyx lobes more than 3 times as long as nut.
120. Style and stylopodium densely pubescent but forthe apex . . . 117. S. pubistyla
120. Ovary and stylopodium crowned by an equally long filiform glabrous style.
121. Stipules large, subpersistent, leaving amplexicaul scars . . . . 118. S. palosapis

121. Stipules fugaceous, small, leaving short scars.
122. Appendages exceeding style at anthesis, very slender, crisped; leaf bullate between tertiary
nerves; nervation beneath, petiole and twig scabrid tomentose. . . 119. S. bullata
122. Appendages not exceeding style apex, awn-like; leaf applanate, evenly pubescent or
glabrous.
123. Style c. + length of ovary and stylopodium; ovary and stylopodium glabrescent; anthers
large, fused, strongly tapering towards apex. . -» = . 920. S: flavifiora
123. Style + equal in length to ovary and stylopodium; ovary and stylopodium shortly
pubescent; anthers not large, not fused, hardly tapering.
124. Nerves 13-16 pairs; leaf undersurface gold lepidote, without domatia
121. S. monticola
124. Nerves 6-8 pairs; leaf glabrous, with pore-like domatia . . . 122. S. kunstleri
100. Filaments united in a tube around the base of the ovary, tapering abruptly near apex; stylopodium
merging into style; style and stylopodium spindle-shaped, narrow and tapering at base and apex;
or stylopodium indistinct, style at least twice as long as ovary, filiform. Fig. 103, 104. Spp. 123-132.
8. Sect. Pachycarpae
125. Stipule scars amplexicaul.

126. Leaf densely gold-brown tomentose beneath . af) Suntec ai eetek! ABBIZ3 SS 3pilosa
126. Leaf shortly sparsely tomentose beneath or glabrous.
127. Stipule with cordate subequal base. . . . . . . =. . +. +. 124. S. splendida

127. Stipule not cordate.
128. Nerves 11-20 pairs; nut to 6 by 4cm, relatively large; fruit calyx lobes to 11 cm long; relatively
short.

129. Panicles to 35 cm long, axillary on modified parts of twigs with short internodes, the
subtending leaves mainly abortive; stipules to 2 cm long, subrevolute 125. S. stenoptera
129. Panicles to 20 cm long, borne in axils of normal leaves on normal twig; stipule to 5 cm long,
applanate .. . . 126. S. macrophylla
128. Nerves 9-12 pairs; nut to 3) 7 by 2 5 cm, relatively small; fruit calyx lobes exceeding 15 cm

long, relatively long.

1BOfLeat25-35.cmlone.<. .< = )acte;cuswen name ete col ZS a peaesines
130. Leaf 11-21 cm long.
131. Leaf broadly ovate to suborbicular, 10-13 cm wide . . . ._ . 128. S. rotundifolia
131. Leaf elliptic, S-8.com wide. . 2 «ew is) ee)» 129.8, amplexicaniis

125. Stipule scars not amplexicaul.
132. Leaf base cordate, undersurface densely persistently golden pubescent; nerves 16—20 pairs
130. S. mecistopteryx
132. Leaf base not cordate; tomentum not as above or leaf glabrous.
133. Stipule scars short, + straight or slightly up-pointing; nerves 11—14 pairs
131. S. beccariana
133. Stipule scars falcate, down-curved; nerves normally 14-19 pairs. . . . 132. S. pinanga
99. Appendage to connective shorter than, or as long as, anther, becoming reflexed; filaments tapering
gradually from base to anther. Fig. 106, 107. Spp. 133-159. 9. Sect. Mutica
134. Fruit calyx lobes auriculate at base. Fig. 107. Spp. 133-138. 9a. Subsect. Auriculatae
135. Leaf narrowly oblong; nerves 19-37 pairs.
136. Leaf concave, nerves depressed above . . . . . . . . . . .133. S. slootenii
136. Leaf applanate.
137. Nerves 24-28 pairs; leaf beneath sparsely tufted tomentose or glabrescent
134. S. myrionerva
137. Nerves 19-25 pairs; leaf beneath densely shortly persistently pink-brown scabrid tomentose
135. S. sagittata
135. Leaf elliptic-ovate to lanceolate; nerves 10-15 pairs.
138. Leaf 7-16 by 2.2-6 cm.
139. Leaf undersurface lustrous, nerves prominent. . . . . . . . 136. S. macroptera
139. Leaf undersurface matte, nerves hardly raised. . . . . . . . 137. S. ferruginea
138. Leaf 14-26 by 6.5-12 cm, thickly coriaceous . . . . . . . +. +. #4138. S. acuta

1982} DIPTEROCARPACEAE (Ashton) 445

134. Fruit calyx lobes not auriculate. Fig. 106. Spp. 139-159. 9b. Subsect. Mutica
140. Stipules exceeding 20 mm long, broad, boat-shaped, coriaceous, lustrous, not at first caducous;
leaves unequal, ovate, glabrescent beneath.
BtieeNerves|4—S: pairs" Boaoe ay pe) are! Ge dsr go ( eh edie g? @ CIS Siineryis
141. Nerves 7-10 pairs . . . - = - o@§ + €s = SAE SZ acmmata
140. Stipules less than 20 mm long, thin, 35 - applanate.
142. Flower buds at least 14 mm long, large.

143. Petiole to 6 mm long; leaf base cordate, unequal. . . . . . . 141. S. macrantha

143. Petiole 6-12 mm long; leaf base obtuse,equal. . . . . . . . 142. S. hemsleyana
142. Flower buds at most 10 mm long.

144. Fruit calyx lobes less than twice aslongasnut . . . . . 143. S. singkawang

144. Longer fruit calyx lobes more than three times as long as nut.
145. Leaf undersurface entirely glabrous.

146 sea apex retuse. 2. = 3 ps is us) so SR ae 2D pee reins
146. Leaf apex acuminate.
147. Nerves 14-16 pairs; stipule to 20 mm long, subpersistent. . . . . 145. S. lepidota
147. Nerves at most 12 pairs; stipules to 8 mm long, fugaceous.
148. Nerves 8-9 pairs, with prominent axillary pore-like domatia . 146. S. foraminifera

148. Nerves 1 1—12 pairs, with only a few pairs of small domatia at the base of the midrib
147. S. teysmanniana
145. Leaf undersurface sparsely or densely tomentose, or cream lepidote.
149. Leaf with at least 14 pairs of nerves or, if 12-13, with cream pubescent undersurface or
with pale scale-like domatia up either side of the midrib.
150. Leaf evenly pinkish velutinate beneath; nerves 20—25 pairs, with prominent secondaries
148. S. argentifolia
150. Leaf tomentum not as above; nerves 14~21 pairs, without secondaries.
151. Leaf beneath, petiole and twigs densely fulvous scabrid tufted tomentose.
152. Leaf prominently concave, chartaceous; petiole 22-32 mm long 149. S. uliginosa

152. Leaf applanate, coriaceous; petiole 13-23 mmlong. . . . . 150. S. rugosa

151. Tomentum not as above, young tree leaf with prominent pale domatia up each side of
the midrib.

153. Mature tree leaf cream pubescent beneath; nerves 12-15 pairs. 151. S. leprosula

153. Mature tree leaf sparsely scabrid beneath; nerves 16-20 pairs . 152. S. platycarpa
149. Leaf with at most 13 pairs of nerves; domatia and indumentum not as above.
154. Leaf narrowly ovate to lanceolate, + cream to pink ee beneath; nerves slender,
hardly raised beneath; midrib acute beneath . =»: (. 153.S: cartisn
154. Leaf elliptic or broadly ovate, undersurface not as above; nerves prominent beneath.
155. Leaf lustrous beneath.
156. Leaf 5—9 by 3-5 cm, elliptic, frequently retuse; margin hardly or not revolute
154. S. scabrida
156. Leaf 10-15 by 5.5-10 cm, broadly ovate, acuminate, with revolute margin
155. S. revoluta
155. Leaf not lustrous beneath.
157. Leaf broadly ovate; nerves 8-10 pairs. . . . . . . . . 4156. S. ovata
157. Leaf ovate to elliptic; nerves c. 11 pairs.
158. Leaf beneath, petiole and twigs densely evenly vinous tomentose. 157. S. rubra
158. Leaf beneath, petiole and twigs sparsely pale brown pubescent 159. S. parvifolia
98. Stamens 50-70, appendage to connective vestigial, filaments filiform. Fig. 115. Sp. 160.
10. Sect. Ovalis, 160. S. ovalis

1. Section Shorea

ASHTON, Gard. Bull. Sing. 20 (1963) 265; Man. Dipt. Brun. (1964) 116. — Shorea
sect. Eushorea BRANDIS, J. Linn. Soc. Bot. 31 (1895) 79. — Shorea, Balau group
Sym. Mal. For. Rec. 16 (1943) 14. — Fig. 78, 82.

Flowers cream, often pink at base. Stamens 20-60, in several verticils; filaments
broad at base, gradually tapering; anthers with 4 pollen sacs, + broadly oblong,

446 FLORA MALESIANA [ser. 1, voR@2

6a =<

eee

<<a

——-

a

Fig. 78. Flower details in Shorea sect. Shorea subsect. Shorea. All x 10. — S. atrinervosa SYM. A. Bud, A/. outer

sepal, 42. inner sepal, both from inside, 43. stamens and pistil. — S. brunnescens ASHTON. B. Stamens and pistil,

B1. stamen from inside. — S. scrobiculata BuRCK. C. Bud, C1]. stamens and pistil, C2. stamen from inside. — S.

hypoleuca Meuer. D. Bud, D1. stamens from outside, D2. stamen from inside, D3. pistil (A SAN 27274, B
SMYTHIES 15218, C BRUN 740, D BaKarR 36229).

cells attenuate at base and apex; appendages shorter than anthers, with 1—co
bristles. Ovary tomentose, with stylopodium. Stipules and bracts fugaceous,
small. Leaf with scalariform tertiary nerves; midrib raised or depressed above,
always evident. Trees with narrow prominent buttresses. Bark surface flaky or
dippled (S. biawak excepted); radially oblique stone-cell fingers often present;
phelloderm pale, thick, conspicuous; expansion tissue in short fingers, more

numerous towards outer surface.
Vern. Balau (Mal., Sum.), sélangan batu (Borneo, Mal.), tékam (Dayak), yakal (Philippines).
Note. The species are, with adequate material well defined and most vary little geographically.

1982] DIPTEROCARPACEAE (Ashton) 447

la. Subsection Shorea

Saul Roxs. ex W. & A. Prod. (1834) 84, nomen. — Isoptera SCHEFF. ex
BurcK. — Ridleyinda O. K.— Shorea sect. Isoptera (SCHEFF. ex BURCK) Foxw.
Philip. J. Sc. 67 (1938) 291, 301; SLoor. Bull. Bot. Gard. Btzg 17 (1941)
116. — Shorea ciliata subgroup Sym. Mal. For. Rec. 16 (1943) 5. — Fig. 78.
Flower buds elongate. Petals linear, falling separately. Appendage to connec-

tive with few bristles.

Distr. Southern India and Ceylon to Indochina and through Malesia to the Moluccas.

Ecol. Evergreen, semi-evergreen and savanna (S. robusta of India, S. obtusa of S.E. Asia) forests, especially
below 1000 m and always below 1500 m. Only S. robusta, the sal, can be strictly gregarious, most being scattered
on well drained soil and river banks.

Note. S. sumatrana has been observed to be thrips pollinated and this is likely to be so with other species.

1. Shorea collina Ript. Agr. Bull. Str. & F.M.S. 9
(1910) 182; Fl. Mal. Pen. 1 (1922) 231; BURN-Mur-
DOCH, Trees and Timbers Mal. Pen. 1 (1911) 13;
HEyNneE, Nutt. Pl. ed. 1, 3 (1917) 299; ibid. ed. 2 (1927)
1116; Foxw. Mal. For. Rec. 3 (1927) 63, p.p.; ibid. 10
(1932) 173, p.p.; SyM. Gard. Bull. S.S. 9 (1935) 270, pl.
18; Mal. For. Rec. 16 (1943) 12, f. 6, 7.—S.
angustiloba Foxw. Mal. For. Rec. 10 (1932) 168, pl.
11; SyM. apud Descu, Mal. For. 3 (1934) 195; Sym.
Mal. For. 4 (1935) 26.

Medium-sized or large tree. Stipules outside, leaf
bud, parts of petals exposed in bud, ovary and
panicles persistently densely shortly evenly buff
pubescent, twig and sepals caducously so, panicle
sparsely caducously so. Twigs c. 3 mm @ apically,
stout, ribbed at first, becoming terete, smooth, black-
ish; stipule scars obscure, falcate, downcurved. Buds
to 3 by 2 mm, ovoid, acute. Stipules to 10 by 5 mm,
lanceolate, acute, caducous. Leaves 9-21 by 5-13 cm,
broadly elliptic-oblong, coriaceous; base obtuse to
cordate; apex obtuse, acute or with short broad
acumen; nerves 11—15 pairs, slender but prominent
beneath, arched, at to 90° at the base, down to 30°
near the apex; tertiary nerves very slender, densely
scalariform; midrib applanate above, prominent and
terete beneath; petiole 2-4 cm long, stout. Panicle to 4
cm long, terminal or axillary, slender; singly
branched, branchlets to 1 cm long, bearing to 3 secund
flowers. Flower buds to 8 by 4 mm, fusiform; sepals
broadly ovate, subequal, submucronate; stamens c.
55, subequal; filaments broadly dilated and connate at
base, tapering; anthers narrowly oblong, glabrous;
appendages somewhat exceeding anthers, with 2-5
prominent apical bristles; ovary broadly ovoid; style
short, columnar. Fruit pedicel by 2 by 3 mm, stout:
calyx lobes vestigial; 3 longer lobes to 5 by 1.3 cm,
obtuse, spatulate, c. 6 mm wide above the to 1.7 by 1.4
cm ovate somewhat thickened saccate base; 2 shorter
lobes to 2.5 by 0.5 cm apically, otherwise similar; nut
to 3.5 by 2.5 cm, large, crowned by a to 8 mm long
tapering style remnant.

Distr. Malesia: Malaya (E. coast from Trengganu
southwards).

Ecol. Low lying land; local.

Vern. Balau meérah, b. bukit, b. tiong, sélimbar,
térbak, téngkawang batu.

2. Shorea ochrophloia [SyM. apud Descu, Mal. For.
Rec. 4 (1935) 28, nomen] STRUGNELL ex SyM. Gard.
Bull. S. S. 8 (1935) 268, pl. 17; Mal. For. Rec. 16(1943)
23flS:

Large buttressed tree. Twigs, buds, stipules,
petioles, panicles, sepals, part of petals exposed in
bud, ovary, midrib above and leaves beneath persis-
tently densely shortly dark golden-brown scabrid
pubescent. Twig. c. 2mm g apically, terete, becoming
blackish, rugulose to smooth. Bud to 4 by 3 mm,
ellipsoid, subacute. Stipules to 10 by 5 mm, oblong-
elliptic, obtuse, early caducous. Leaves (4-)6—-12 by
3.5-6.5 cm, ovate to elliptic-oblong. coriaceous; mar-
gin frequently revolute; base obtuse to subcordate:
apex obtuse, acute or with a short broad acumen;
nerves 13-18 pairs, prominent beneath, + depressed
above, set at 80° at the base but down to 25° near the
apex; tertiary nerves slender, scalariform; midnb
prominent and terete beneath, somewhat depressed
above; petiole 7-17 mm long, short, geniculate. Pani-
cle to 6 cm long, terminal or axillary, short, terete;
singly branched, branchlets bearing to 6 flowers;
bracteoles fugaceous, unknown. Flower buds to 7 by 2
mm, fusiform. Sepals ovate, the 3 outer subacute, the
2 inner subacuminate. Petals cream with a deep pink
patch at the base within. Stamens 30, subequal,
filaments tapering, compressed; anthers oblong, gla-
brous; appendage exceeding anther apex. with to 5 +
prominent bristles; ovary with distinct stylopodium,
both pubescent; style short but prominent. Fruit
pedicel to 2 by 2 mm. 3 longer calyx lobes to 7 by 1.5
cm, spatulate, obtuse, c. 5mm broad above the to 7 by
5 mm ovate saccate thickened base; 2 shorter lobes to
6 by 0.7 cm, otherwise similar. Nut to 2 by 1 cm,
narrowly ovoid, tapering to an up to 3 mm long
columnar style.

Distr. Malesia: Malaya, W. Sumatra (Painan).

Ecol. Scattered, local, on well drained undulating
land and old alluvium below 350 m.

Vern. Séraya batu.

Note. A local segregate of S. guiso, with which it
now nevertheless grows in mixture without apparent
hybridisation.

3. Shorea guiso (BLCO) BL. Mus. Bot. Lugd.-Bat. 2
(1852) 34; WaLp. Ann. 4 (1857) 338; DC. Prod. 16, 2

448

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 79. Density map of Shorea RoxB. ex GAERTN. f. in
Malesia; number of endemics above the hyphen,
number of non-endemics below it.

(1868) 632; VIDAL, Sinopsis (1883) t. 1SC; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 89; PERK. Fragm. FI. Philip.
(1904) 23; Merr. Philip. J. Sc. 1 (1906) Suppl. 98; Sp.
Blanc. (1918) 270; En. Philip. 3 (1923) 97; Foxw.
Philip. J. Sc. 2 (1907) Bot. 384; ibid. 4 (1909) Bot. 509;
ibid. 6 (1911) Bot. 272; ibid. 13 (1918) Bot. 191; Mal.
For. Rec. 10 (1932) 175; Philip. J. Sc. 67 (1938) 292;
EVERETT & WHITFORD, Bull. Bur. For. Philip. 5 (1906)
16; MERRITT & WHITFORD, ibid. 5 (1906) 36; MERRITT,
ibid. 8 (1908) 48; WHITFORD, Philip. J. Sc. 4(1910) Bot.
703; Bull. Bur. For. Philip. 10 (1911) 71; Reyes, Philip.
J. Sc. 22 (1923) 337; SLoot. in Merr. Pl. Elm. Born.
(1929) 203; Descu, Mal. For. 4 (1935) 29; Mal. For.
Rec. 12 (1936) 20; Sym. Gard. Bull. S. S. 8 (1935) 266,
pl. 16; Mal. For. Rec. 16 (1943) 16, f. 11; BROWNE,
For. Trees Sarawak & Brunei (1955) 153; MEUER &
Woop, Sabah For. Rec. 5 (1964) 171, f. 18; ASHTON,
Man. Dipt. Brun. Suppl. (1968) 71. — Euphoria
malaanonan Buco, FI. Filip. ed. 1 (1837) 286. — Mo-
canera guiso BLCO, FI. Filip. ed. 1 (1837) 449. — Dip-
terocarpus guiso BLCO, FI. Filip. ed. 2 (1845) 313; DC.
Prod. 16, 2 (1868) 614; BLco, FI. Filip. ed. 3, 2 (1878)
215. — Euphoria vel Nephelium Buco, FI. Filip. ed. 2
(1845) 200; ibid. ed. 3, 2 (1878) 9. — Anisoptera guiso
DC. Prod. 16, 2 (1868) 616. — S. pierrei HANcE, J.
Bot. 16 (1878) 302; BRANpbis, J. Linn. Soc. Bot. 31
(1895) 81.— S. robusta (non GAERTN. f.) F.-VILL.
Nov. App. (1880) 21. — S. vulgaris PrERRE ex LANES-
SAN, Pl. Util. Colon. Fr. (1886) 301; Pierre, For. FI.
Coch. 3 (1889) t. 232; BRANDis, J. Linn. Soc. Bot. 31
(1895) 81, t. 2, f. 15-16; BRANDIS & GitG in E. & P. Pf.
Fam. ed. 1, 3, 6 (1895) 264, 265, fig.; GUERIN, Fl. Gén.
I.-C. 1 (1910) 300; Gite in E. & P. Pfl. Fam. ed. 2, 21
(1925) 261. — S. vidaliana BRANDIS, J. Linn. Soc. Bot.
31 (1895) 83. — Isoptera burckii BoERL. Cat. Hort.
Bog. 2 (1901) 111; SLoor. Bull. Bot. Gard. Btzg III, 17
(1941) 117. — S. obtusa var. kohchangensis HEI™, Bot.

Tidsskr. (1902) 263. — S. robusta var. schmidtii HE™,
l.c. 263. — S. warburgii (non GILG) Perk. Fragm. FI.
Philip. (1904) 23. — S. longipetala Foxw. Mal. For.
Rec. 10 (1932) 174, t. 13.—S. scrobiculata (non
BuRCK) Foxw. Mal. For. Rec. 10 (1932) 174; Sym.
apud Descu, Gard. Bull. S. S. 8 (1935) 28, 29.

Large buttressed tree. Young parts at first greyish
buff pubescent, early caducous except on inflores-
cence, calyx, corolla outside, ovary and nut; twig
apices and stipule outside densely so at first, becoming
sparse as parts expand, remaining dense on inflores-
cence and nut. Twig. c. 1 mm @ apically, slender,
smooth, with minute inconspicuous stipule scars. Bud
to 5 by 3 mm, compressed, ovate-falcate. Stipule to 7
by 5 mm, oblong, subacute, caducous. Leaves 5.5—-14
by 2.5-6 cm, oblong-lanceolate, thinly coriaceous,
glabrescent; base obtuse to broadly cuneate, acumen
to 1 cm long, prominent, broad; nerves (11—)15—19
pairs, slender, at 45°—55° to the midrib but up to 90° at
the base; tertiary nerves very slender, densely scalari-
form, frequently obscure; midrib slender, raised
beneath, evident but somewhat depressed above;
petiole \-1.8 cm long, slender. Panicle to 10 cm long,
slender, lax, pendent, the branchlets bearing to 5
secund flowers; bracts and bracteoles minute, linear,
fugaceous. Flower bud to 1 cm long, slender. Sepals
broadly ovate; outer 3 acute, inner 2 acuminate, with
thin margins. Petals bright yellow-red at base within,
narrowly lanceolate, densely pubescent on parts
exposed in bud. Stamens 20-28: filaments lorate,
tapering, glabrous; anthers narrowly oblong, gla-
brous, the two outer cells somewhat the larger:
appendage c. 5 as long as anther, with 14(-8) long
bristles. Ovary ovoid-conical, densely pubescent, sur-
mounted by a columnar glabrous style c. its length. 3
longer fruit calyx lobes to 5.5 by 1 cm, spatulate,
obtuse, c. 3 mm broad above the to 8 by 5 mm saccate
thickened base; 2 shorter lobes to 3 cm long, linear but
similar at base. Nut to 8 by 5 mm, ovoid, apiculate,
hidden by base of calyx lobes.

Distr. Cochinchina, S.E. and Peninsular Thailand;
Malesia: Malaya, Sumatra (Atjeh, Tapanuli, Palem-
bang), Borneo (mainly in east), Philippines.

Ecol. Scattered in lowland forest on red soils, most
common in slightly seasonal climates; rare and con-
fined to limestone hills in W. and Central Borneo.

Uses. The hard red timber is valuable for light
construction in the Philippines.

Vern. Mémbatu, mérbatu, lemesa, |. kulor, |. ner-
ang, chéngal pasir, méranti bulu, m. pahang, résak
samak, sélimbar (Mal.), béraja, damar kénuar batu,
meéranti hitam (Sumatra), sélangan batu merah
(Sabah), pélapak, lauan, raru, pakulin batu, keping
burong, ménkabang, ambam, bérukan baju (S.E.
Borneo), guijo (Philippines).

4. Shorea havilandii BRANDIS, J. Linn. Soc. Bot. 3
(1895) 82; Merr. En. Born. (1921) 405; BROWNE, For.
Trees Sarawak & Brunei (1955) 168; ASHTON, Man.
Dipt. Brun. (1964) 136, f. 13, pl. 34 (stem); ibid. Suppl.
(1968) 72, f. 9; MEWER & Woop, Sabah For. Rec. 5

1982]

(1964) 173.— Hopea ovalifolia BoeRL. Cat. Hort.
Bog. 2 (1901) 102.

Small or medium-sized tree. Twig, petiole, midrib
above, buds and stipule on both surfaces + persis-
tently shortly evenly greyish tawny pubescent. Twig c.
1 mm @g apically, terete, slender; stipule scars c. 1 mm
long, short, cuneate, horizontal or slightly descend-
ing. Bud c. 2 by 1.5 mm, ovoid, obtuse. Stipule c. 6 by
2.5 mm, oblong, subacute, fugaceous. Leaves 8—16 by
3.+6 cm, thinly coriaceous, ovate-elliptic; base
obtuse or broadly cuneate; acumen c. 0.7 cm long,
narrow; nerves 9—12 pairs, dense, parallel, slender but
prominent beneath, at c. 30°-45° along the lamina, to
80° at the base; with small tomentose domatia; tertiary
nerves slender, densely scalariform; midrib grooved:
petiole 8-12 mm long. Panicle to 12 cm long, terminal
or axillary, straight, terete or slightly compressed,
shortly sparsely persistently pale brown pubescent;
singly branched, the branchlets to 1.5 cm long, short,
bearing to 7 secund flowers; bracteoles to 4 mm long,
elliptic, subacute, shortly pubescent, fugaceous.
Flower bud to 8 by 2.5 mm, lanceolate. Calyx shortly
pubescent outside, glabrous within; lobes ovate, sub-
acute, the inner 2 somewhat smaller, relatively
broader and thinner than the outer 3. Petals cream
pink at base. Stamens 30—S0; filaments broad at base.
tapering abruptly and filiform distally, glabrous:
anthers oblong, the cells tapering, outer cells longer
than inner cells; appendages to connective short.
glabrous but for a single long apical bristle. Ovary
broadly ovoid, pubescent. Fruit calyx to 10 by 8 mm;
lobes subequal, shortly buff-pubescent on both sur-
faces. thin, ovate, prominently narrowly acuminate,
shallowly saccate. Nut to 1.5 by 1 cm, globose, obtuse;
style remnant c. 2 mm long, slender.

Distr. Malesia: Borneo (Sarawak, E. Sabah).

Ecol. Locally abundant in Heath forest and fresh
water swamp forest on white and yellow sand. and on
peat overlying limestone.

Vern. Sélangan batu pinang.

5. Shorea brunnescens AsHTON, Gard. Bull. Sing. 22
(1967) 283, pl. 28; Man. Dipt. Brun. Suppl. (1968) 68,
f. 9. — Fig. 78 B-B1.

Medium-sized tree. Leaf bud densely shortly persis-
tently buff pubescent, young twigs and petiole spar-
sely caducously so; lamina glabrous. Twig c. 1mm @
towards apex, terete, smooth; stipule scars short,
horizontal, obscure. Bud to 2.5 by 2mm, small, ovoid,
acute. Stipules unknown. Leaves 6-12 by 2.5-6 cm,
broadly ovate to lanceolate, coriaceous; base broadly
cuneate; acumen to | cm long, narrow; nerves 9-11
pairs, very slender, hardly raised beneath, curved, at
45~—60°; tertiary nerves dense, subreticulate, evident
but hardly raised on either surface; midrib obscure.
depressed above, prominent, furrowed, beneath:
petiole \-1.5cm long. Panicle to 9 cm long, terminal or
axillary, angular, sparsely shortly buff pubescent;
singly branched, branchlets to 1.5 cm long; bracteoles
unknown. Flowers secund; buds to 4 by 2 mm,
narrowly ellipsoid. Calyx densely pubescent outside,

DIPTEROCARPACEAE (Ashton)

449

glabrous within; lobes ovate, acute, the inner two
somewhat shorter and relatively broader than the
outer 3. Petals lanceolate, shortly pubescent on parts
exposed in bud. Stamens 40-62; filaments glabrous,
compressed at base, tapering abruptly and filiform
distally, anthers elliptic-oblong, glabrous, the inner 2
sacs somewhat smaller than the outer 2; appendage to
connective shorter than anther, glabrous but for up to
2 long apical setae. Ovary and stylopodium pyriform,
densely pubescent, crowned by a short columnar
glabrous style. Mature fruit unknown: Fruit calyx at
first greyish pubescent; calyx lobes aliform, unequal, 3
much longer than the other 2.

Distr. Malesia: Borneo (W. Sarawak eastwards to
Kinabalu, S.E. Borneo northwards to Belajan R.).

Ecol. Locally frequent on skeletal soils on high
ridges to 1500 m, also sometimes on leached clay soil
on undulating land.

Vern. Sélangan batu tinteng.

6. Shorea scrobiculata Burck, Med. Lands P|. Tuin 3
(1886) 223; Ann. Jard. Bot. Btzg 6 (1887) 207:
BRANpIs, J. Linn. Soc. Bot. 31 (1895) 83: MERR. En.
Born. (1921) 406; BRowne, For. Trees Sarawak &
Brunei (1955) 169; AsHton, Gard. Bull. Sing. 20
(1963) 272; Man. Dipt. Brun. (1964) 144, f. 13; ibid.
Suppl. (1968) 74. — S. pierreana Hem, Rech. Dipt.
(1892) 48. — S. sp. nov. SyM. ex Descu, Mal. For.
Rec. 12(1936) 9. — S. meadiana Sym. Gard. Bull. S.S.
10 (1939) 366, pl. 23; Mal. For. Rec. 16 (1943) 22. f. 6;
cf. ASHTON, Gard. Bull. Sing. 22 (1967) 282. — Fig. 78
C-C2.

Medium-sized tree. Twig, panicle, leaf bud, stipule
and petiole densely shortly buff pubescent: midrib on
both surfaces and nerves beneath sparsely dotted with
minute hair tufts. Twig c. 1.5 mm g apically, terete,
much branched, slender, smooth, becoming finely
rugulose; stipule scars short, pale, horizontal. Bud 2-3
by 1-2 mm. ovoid, frequently slightly compressed.
Stipule c. 7 by 3 mm, hastate, acute, fugaceous. Leaves
5.5-11 by 2.5-4cm, narrowly ovate to oblong-lanceo-
late; base broadly cuneate; acumen to | cm long,
narrow; nerves 10-12 pairs, dense, prominent,
straight, at c. 35°—-55°; midrib slightly depressed
above; tertiary nerves densely scalariform, at 90° to
the nerves: petiole c. 8 mm long. Panicle to 7 cm long,
terminal or axillary, terete; singly branched, branch-
lets to 1.5 cm long, short, bearing to 9 secund flowers:
bracteoles to 3 mm long, elliptic, subacute, shortly
pubescent. Flower bud to 6 by 2 mm, narrowly
lanceolate. Calyx shortly tomentose outside, glabrous
within: lobes ovate, subacute: inner 2 somewhat
smaller, relatively broader, thinner, than outer 3.
Petals pink, cream at margin, linear, pubescent on
both surfaces. strongly contorted in bud. Stamens
20-30: filaments broad at base, tapering abruptly and
filiform distally, glabrous; anthers oblong, glabrous.
tapering, the outer cells larger than the inner cells;
appendage to connective short, glabrous but for a
single (—2) apical bristle. Ovary broadly ovoid, pubes-
cent; stylopodium narrowly cylindrical, shorter than

450

the ovary, pubescent; style as long as stylopodium,
tapering, glabrous. Fruit calyx shortly buff pubescent;
3 longer lobes to 5 by 1.2 cm, spatulate, chartaceous,
obtuse, c. 2.5 mm broad above the c. 5 by 4 mm
saccate thickened base; 2 shorter lobes to 35 by 4mm,
subequal, spatulate, similar at base. Nut to 10 by 7
mm, ovoid, densely shortly buff tomentose; style
remnant c. 1.5 mm long.

Distr. Malesia: Malaya (Perak, Trengganu,
Pahang), Borneo (Ulu Kapuas, Sarawak, E. Sabah,
W. Kutei, Muaratewe).

Ecol. Undulating land and hills to 700 m, Mixed
Dipterocarp forest.

Vern. Sélangan batu zang (Sar.), palepek gunong
(S.E. Borneo), balau séngkawang, damar laut kuning
(Mal.).

7. Shorea leptoderma MEER, Act. Bot. Neerl. 12
(1963) 331, pl. 5; MEIER & Woop, Sabah For. Rec. 5
(1964) 180, pl. 14b (stem).

Small to medium-sized tree. Young twigs, leaf buds,
parts of petals exposed in bud, ovary, panicles,
petioles, stipules outside (sparsely so within) and
nervation above (glabrescent beneath) densely evenly
shortly persistently pale ocherous pubescent; sepals
thus at first, becoming sparse in fruit. Twig c.2mm @
apically, ribbed at first, becoming terete, chocolate-
brown, minutely pale lenticellate. Buds to 3 by 2mm,
ellipsoid, subacute. Stipules to 8 by 5 mm, elliptic,
obtuse, fugaceous. Leaves 4-18 by 1.5—6.5 cm, elliptic,
chartaceous; base cuneate; apex acute or with to | cm
long slender acumen; nerves 7—12 pairs, slender but
prominent beneath, arched, at 35°—45°; tertiary nerves
slender, scalariform; midrib applanate above, promi-
nent beneath; petiole 8-10 mm long. Panicle to 10 cm
long, ribbed; singly branched, branchlets short, bear-
ing to 8 congested secund flowers; bracteoles un-
known. Flower buds to 5 by 2 mm, lanceolate; sepals
broadly ovate, subequal, the outer 3 acute, the inner 2
subacuminate; stamens c. 30; filaments compressed,
tapering; anthers ellipsoid, glabrous; appendages
hardly exceeding anthers, with to 5 long bristles; ovary
ovoid, with short columnar style. Fruit pedicel to 2
mm long, slender; 3 longer calyx lobes to 8 by 1.5 cm,
spatulate, obtuse, c. 4mm wide above the to 5 by Smm
saccate thickened base; 2 shorter lobes to 4.5 by 0.4
mm, linear, acute, otherwise similar. Nutto 12 by6mm,
ovoid, tapering to the 2 mm columnar style remnant.

Distr. Malesia: Borneo (N.E. Borneo, Sandakan
Distr. and Tawau).

Ecol. Lowland forests.

Vern. Sélangan batu biabas, s. b. jambu.

Note. Closely allied to S. scrobiculata from which it
is doubtfully distinct.

8. Shorea ciliata KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 118; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 82;
BRUHL & KING, Ann. R. Bot. Gard. Calc. 5, 2 (1896)
154, t. 187; Rip. Fl. Mal. Pen. 1 (1922) 229; Foxw.
Mal. For. Rec. 10 (1932) 171; Sym. Mal. For. Rec. 16
(1943) 10, f. 6.

FLORA MALESIANA

[ser. I, vol#92

Medium-sized tree. Sepals, ovary and parts of
petals exposed in bud densely shortly evenly buff
pubescent; young innovations fugaceous cream lepi-
dote, leaf undersurface sometimes sparsely persis-
tently so, otherwise glabrous. Twigs c. 2 mm @
apically, slender, much branched, terete, dark brown,
smooth. Buds to 3 by 2 mm, ellipsoid, subacute.
Stipules to 6 mm long, linear, fugaceous. Leaves
(4—-)5—13(-17) by (1.5—)2.5-5(-8) cm, ovate-lanceo-
late, coriaceous; base cuneate; acumen to | cm long,
slender; nerves 9-11 pairs, slender, hardly elevated
beneath, arched, at 25°—65°; tertiary nerves slender,
subreticulate; midrib elevated on both surfaces
though more prominent below than above, terete;
petiole |.5—2.5 cm long, slender, geniculate. Panicle to
10 cm long, terminal or axillary, terete; singly or
doubly (if terminal) branched, branchlets to 3 cm
long, bearing to 6 flowers; bracteoles unknown,
fugaceous. Flower bud to 8 by 3mm, lanceolate. Petals
cream. Sepals ovate, subacute, unequal. Stamens c.
26-28, subequal; filaments compressed, tapering;
anthers oblong, glabrous; appendages short, with to 5
prominent bristles; ovary with distinct stylopodium;
style short but prominent, glabrous. Fruit pedicel to |
mm, short. 3 longer fruit calyx lobes to 6 by 1.5 cm,
spatulate, obtuse, to 5 mm wide above the to 8 by 5
mm ovate saccate thickened base; 2 shorter lobes to 3
by 0.4.cm, lorate, obtuse, otherwise similar. Nut to 12
by 8 mm, ovoid, tapering to a c. 3 mm long style
remnant.

Distr. Malesia: Malaya.

Ecol. Widespread in Upper Dipterocarp Forests
on ridges of the main range, at 800-1200 m, and down
to 300 m on Penang Hill.

Vern. Balau gunong.

9. Shorea submontana Sym. Gard. Bull. S.S. 10(1939)
368: Mal. For. Rec. 16 (1943) 24, f. 6. — S. costata
(non (CORREA) PRESL) KING, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 119; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
81; BurK. J. Str. Br. R. As. Soc. 81 (1920) 67, 86, fig.;
ibid. 86 (1922) 281; Ripv. Fl. Mal. Pen. 1 (1922) 231,
p.p.; Foxw. Mal. For. Rec. 3 (1927) 67; ibid. 10 (1932)
176, p.p.; DescH, Mal. For. Rec. 12 (1936) 9.

Large buttressed tree. Buds, stipules outside, pani-
cles, parts of petals exposed in bud and ovary shortly
densely evenly persistently buff pubescent; sepals thus
at first, becoming sparsely so; twigs, petioles, midribs
above and nervation beneath caducously so. Twigs c.
3 mm @g apically, ribbed, becoming terete, smooth,
blackish. Buds to 5 by 3mm, ellipsoid, obtuse. Stipules
to 14 by 6mm, oblong, obtuse, caducous. Leaves 7-20
by 4-10 cm, ovate to obovate, thinly coriaceous; base
broadly cuneate to cordate; apex acute or with up to |
cm long, slender acumen; nerves 9-14 pairs, slender
but prominent beneath, at 90° at the base, down to 45°
towards the apex; tertiary nerves densely scalariform,
slender, obscure; midrib prominent, terete, beneath,
evident but + applanate above; petiole 15-33 mm
long, geniculate. Panicle to 5 cm long, terete, terminal
or axillary, singly branched; bracts and bracteoles and

1982]

mature flowers unknown. Sepals broadly ovate, sub-
equal; stamens c. 20; filaments compressed, tapering;
anthers oblong; appendages exceeding anthers,
setose; ovary narrowly ovoid; style short, glabrous.
Fruit pedicel to 1 by 1 mm, short. 3 longer fruit calyx
lobes to 9 by 1.8 cm, spatulate, obtuse, c. 6 mm wide
above the to 13 by 10 mm ovate saccate thickened
base; 2 shorter lobes to 6 by 0.4 mm, lorate, obtuse,
otherwise similar. Nut to 1.8 by 1.4 cm, ovoid,
crowned by an up to 3 mm tapering style remnant.

Distr. Malesia: Malaya (Selangor and W. Pahang
to Penang and Trengganu).

Ecol. Locally very common on high hills of the
main range, usually 800-1000 m, but down to 350 m
near coast.

Vern. Balau gajah.

10. Shorea geniculata SYM. ex [BROWNE, For. Trees
Sarawak & Brunei (1955) 167, nomen] ASHTON, Gard.
Bull. Sing. 19 (1962) 291, pl. 19; Man. Dipt. Brun.
(1964) 133, f. 13, pl. 32 (habit, stem); ibid. Suppl.
(1968) 71.

Large tree with stout buttresses. All vegetative parts
glabrous. Twig c. 3 mm @ apically, stout, much
branched, terete, smooth; nodes prominent, with
raised round petiole scar and c. 2-3 mm long straight
pale linear ascending stipule scars (only visible on
young twigs). Bud 4~7 by 1—1.5 mm, linear to falcate.
Stipule to 10 by 3 mm, narrowly oblong, acute,
caducous. Leaves 11-17 by 7-13 cm, broadly ovate to
suborbicular, cream lepidote beneath, coriaceous;
base obtuse or subcordate; acumen to 8 mm long;
nerves 9-11 pairs, distant, arched, at c. 40°—S0°;
tertiary nerves sinuate, densely scalariform, slender;
petiole 4-6 cm long. Panicle to 12 cm long, terminal or
axillary, + terete, densely shortly persistently pale
buff pubescent, glabrescent; regularly singly
branched, branchlets to 1.5 cm, short, bearing to 4
secund flowers; bracteoles to 4 mm long, oblong,
puberulent, fugaceous. Flower buds to 20 by 4 mm,
large, narrowly lanceolate, acute. Calyx shortly
pubescent outside, glabrous within; lobes ovate,
acute, the outer 3 slightly longer and narrower than
inner 2. Petals cream, to 2.5 cm long, linear, pubescent
on both surfaces. Stamens c. 55; filaments broad at
base, tapering and filiform distally; anthers oblong,
glabrous, the posterior lobes slightly smaller; appen-
dage to connective somewhat shorter than anther but
prominent, stout and setose. Ovary broadly ovoid,
densely tomentose but for the glabrous base, abruptly
tapering to a short broad trifurcate glabrous style;
stigma minute. Fruit calyx lobes c. 1.5 cm long and
broad, equal, deltoid, subacute, incrassate, sparsely
pale grey-buff pubescent, the apices adpressed to the
base of the nut. Nut to 5 by 5 cm, globose, very large,
densely shortly grey-buff tomentose, ridged longitu-
dinally and transversely rugose when dry, shortly
mucronate.

Distr. Malesia: Borneo (Sarawak and Brunei).

Ecol. Very local, on deep leached yellow soils in
Mixed Dipterocarp Forest on subcoastal hills.

DIPTEROCARPACEAE (Ashton)

45]

Vern. Upun pényau (Brun.).

il. Shorea seminis (DE VRIESE) SLOOT. in Merr. Pl.
Elm. Born. (1929) 204; Bull. Bot. Gard. Btzg III, 17
(1941) 117; Dakkus, Bull. Jard. Bot. Btzg III, Suppl. 1
(1930) 268; Foxw. Mal. For. Rec. 10 (1932) 237:
Philip. J. Sc. 67 (1938) 301; BaL, Landbouw 9 (1934)
275; BURK. Kew Bull. (1935) 317; RowaAANn, Land-
bouw 13 (1937) 310; Bericht n. 13 Afd. Handelsmus.
Kol. Inst. (1937) 5; BROWNE, For. Trees Sarawak &
Brunei (1955) 170; AsHTON, Gard. Bull. Sing. 20
(1963) 272; Man. Dipt. Brun. (1964) 145, f. 13; ibid.
Suppl. (1968) 75; MEER & Woop, Sabah For. Rec. 5
(1964) 185, f. 2d, pl. 17 (habit). — Hopea seminis DE
VriEsE, Minyak Tengkawang (1861) 32; T. & B. Cat.
Hort. Bog. (1866) 202. — Hopea lanceolata DE VRIESE,
Minyak Tengkawang (1861) 32.—S. schefferiana
HAnceE, J. Bot. 16 (1878) 303; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 82; Merr. En. Born. (1921)
406. — Isoptera borneensis SCHEFF. ex BURCK, Med.
Lands Pl. Tuin 3 (1886) 27; Ann. Jard. Bot. Btzg 6
(1887) 222, t. 25; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
106, p.p.; BRANDIS & GILG in E. & P. Pfi. Fam. ed. 1, 3,
6 (1895) 263; BoeRL. Cat. Hort. Bog. 2 (1901) 110;
Becc. For. Born. (1902) 155, 192, 571; Wand. (1904)
92, 124, 390; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 310;
Foxw. Philip. J. Sc. 13 (1918) Bot. 194, pl. 1; MERR.
En. Born. (1921) 407; En. Philip. 3 (1923) 101; REYEs,
Philip. J. Sc. 22 (1923) 301; Gite in E. & P. Pfl. Fam.
ed. 2, 21 (1925) 259; DEN BERGER & ENDERT, Med.
Proefst. Boschw. 11 (1925) 117; ENDERT, M.O. Born.
Exp. (1927) 204, 289, 311; HeyNne, Nutt. Pl. ed. 2
(1927) 1113, 1114, 1127; Watson, Mal. For. Rec. 5
(1928) 45, 205, p.p.; SLooT. Bull. Bot. Gard. Btzg III,
17 (1941) 117. — S. borneensis PIERRE, For. Fl. Coch.
3 (1889) t. 234. — Ridleyinda borneensis O. K. Rev.
Gen. Pl. 1 (1891) 65. — Hopea ovalifolia (non BOERL.)
Foxw. Philip. J. Sc. 6 (1911) Bot. 263; ibid 13 (1918)
Bot. 183. — Isoptera seminis BURK. Kew Bull. (1935)
317: Dict. (1935) 1254; BACKER & BAKH. f. FI. Java 1
(1963) 332.

Medium-sized to large tree. Twig, panicle, leaf bud,
stipule, petiole, midrib on both surfaces and nerves
beneath shortly evenly pale grey-brown pubescent.
Twig c. 1.5 mm @g apically, straight, slender, fre-
quently ribbed when young; stipule scars to 1 mm
long, short, descending. Bud 1.5-2.5 by 1-2 mm,
slightly compressed, ovoid to falcate, acute. Stipule to
7 by 3.5 mm, oblong, obtuse, caducous. Leaves
variable in size, 9-18 by 2.5-8 cm, oblong-ovate to
lanceolate, glabrous or greyish lepidote beneath,
thinly coriaceous; base obtuse or cuneate; acumen
0.8-2 cm long, narrow; nerves 9-15 pairs, slender,
rather straight, prominent beneath, at c. 40°-55°;
tertiary nerves slender, densely scalariform, sinuate;
petiole 1-1.5 cm long. Panicle to 10 cm long, terminal
or axillary to ramiflorous, terete, drying angular;
regularly singly branched, branchlets to 1.7 cm long,
bearing to 5 secund flowers; bracteoles linear, shortly
pubescent, fugaceous. Flower bud to 8 by 2 mm,
narrowly lanceolate. Calyx shortly pubescent outside,

452 FLORA MALESIANA [ser. I, vol. 9?

Let

re
BSS
pane ths

us

Fig. 80. Shorea sumatrana (SLOOT. ex THORENAAR) SYM. ex DESCH; tree c. 30 m high. Palembang (Photogr.
THORENAAR, 1924).

1982]

glabrous within; lobes subequal, ovate, obtuse, the
inner 2 somewhat narrower and thinner than the outer
3. Petals cream, pink towards base, linear, hardly
contorted, shortly tomentose outside, glabrous
within; stamens 30—40; filaments compressed at base,
tapering, with a few long bristles; anthers oblong,
glabrous, the outer cells larger than the inner cells;
appendage to connective short, setose. Ovary and
stylopodium conical to hour-glass shaped, densely
pubescent; style short, glabrous. Fruit calyx shortly
sparsely greyish buff pubescent, lobes subequal, to 2
by 1.8 cm (usually c. 1.5 by 1.4 cm), incrassate,
orbicular, rotate. Nut c. 1 cm long and @, ovoid or
globose; style remnant to 12 mm long, stout.

Distr. Malesia: Borneo, Philippines.

Ecol. Alluvium banks of sluggish river, often
gregarious.

Uses. The fruit are prepared in Borneo as a source
of illipe butter, but are too small to be valued for
export.

Vern. Engkabang chengai, e. tégelam, e. térindak, e.
mayoh, e. pélépak, tekam tégelam (Sar.), sangkawang
(Brun.), téngkawang ayer, t. batu, t. pélepak (kéle-
pak), t. tanggoi, t. térendak, t. chépak (Indon. Bor-
neo).

12. Shorea sumatrana (SLOOT. ex THORENAAR) SYM.
ex Descu, Mal. For. 3 (1934) 195; ibid. 12 (1936) 9; J.
Str. Br. R. As. Soc. 19 (1941) 161; Mal. For. Rec. 16
(1943) 25, f. 6, 16; SLoor. Bull. Bot. Gard. Btzg III, 17
(1941) 124, f. 17. — Isoptera borneensis (non SCHEFF.
ex BURCK) KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893)
129, p.p.; RipL. Trans. Linn. Soc. 3 (1893) 284;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 106, p.p.; MERR.
En. Born. (1921) 407, p.p.; Foxw. Mal. For. Rec. 1
(1921) 71; ibid. 2 (1922) 170; ibid. 3 (1927) 70; BurK. J.
Str. Br. R. As. Soc. 86 (1922) 281, 287; Bot. Gard.
Sing. Guide (1925) 22; Rip. Fl. Mal. Pen. 1 (1922)
245; Merr. En. Philip. 3 (1923) 101, p.p.; WATSON,
Mal. For. Rec. 8 (1930) 30; BUNTING & MILSUM,
Guide Govt. Exp. Plant. Serdang (1931) 122. — Isop-
tera sumatrana SLOOT. ex [ENDERT, Tectona 18 (1925)
73, 78, 135; DEN BERGER & ENDERT, Med. Proefst.
Boschw. 11 (1925) 117, t. 15, f. 58, nomen] THOR-
ENAAR, Med. Proefst. Boschw. 16 (1926) 115, f. 19;
HEYNE, Nutt. Pl. ed. 2 (1927) 1127; Foxw. Mal. For.
Rec. 10 (1932) 238; Burk. Dict. (1935) 1257. — Fig.
80, 81.

Description as in S. seminis, but stamens 25.

Distr. Pattani in S.E. Peninsular Thailand and in
Malesia: Malaya (mainly East Coast Res.), Sumatra.

Ecol. Alluvium banks of sluggish but not brackish
rivers, sometimes semi-gregarious.

Vern. Séngkawang (Mal., Sum.), kédawang, séng-
kawang bésak (Palembang), téngkawang, t. ijok, t.
batu (Mal.).

Note. Of the characters by which VAN SLOOTEN
distinguished this taxon from S. seminis only the
number of stamens is consistently reliable, and this
species is hence of dubious specific status.

DIPTEROCARPACEAE (Ashton)

453

13. Shorea foxworthyi Sym. Gard. Bull. S. S. 8 (1935)
272, pl. 19; Mal. For. Rec. 16 (1943) 14, f. 6,9; MEUER
& Woop, Sabah For. Rec. 5 (1964) 169; AsHTON,
Man. Dipt. Brun. Suppl. (1968) 70, f. 9. — S. collina
(non RIDL.) Foxw. Mal. For. Rec. 3 (1927) 63; ibid. 10
(1932) 173; Sym. apud Descu, Mal. For. 3 (1934) 195.

Tall buttressed tree. Twigs, petioles, buds and
stipules outside (sparsely within) densely shortly
evenly persistently golden-tawny pubescent, leaf ner-
vation beneath sparsely caducously so. Twig c. 2 by 3
mm @ apically, somewhat compressed, rugose when
dry, becoming smooth, terete; stipule scars short,
dark, slender, horizontal. Bud to 3 by 2 mm, ellipsoid,
obtuse. Stipule to 15 by 4 mm, narrowly elliptic to
falcate, subacute, caducous. Leaves 8-13 by 3—6.5 cm,
elliptic, coriaceous; base broadly cuneate; acumen to |
cm long, broad; nerves 10-14 pairs, very prominent
beneath, applanate or depressed above with the
lamina bullate between them, at 85° at the base,
40°-5S° towards the apex; tertiary nerves slender,
densely scalariform; midrib prominent beneath,
somewhat depressed above; petiole 11-20 mm long,
drying rugose. Panicle to 5 cm long, terminal or
axillary, short, rugose when dry, persistently evenly
shortly golden-tawny pubescent, branchlets bearing
to 3 flowers; bracteoles fugaceous. Flower bud to 10 by
3 mm, fusiform. Calyx shortly densely pubescent on
parts exposed in bud, the inner 2 lobes fimbriate
distally; lobes ovate, acute, the inner 2 somewhat
smaller than the outer 3. Petals cream, pale carmine at
base, linear, shortly densely pubescent outside, spar-
sely sericeous within. Stamens 32-41; filaments com-
paratively densely setose, compressed at base, taper-
ing distally; anthers narrowly oblong, sparsely setose
on the distal margin, the 2 outer cells somewhat the
larger; appendage to connective hardly exceeding
anther apex, densely setose. Ovary ovoid, densely
pubescent, surmounted by a similarly tomentose
narrowly conical stylopodium and glabrous filiform
style; style as long as ovary and stylopodium. Fruit
pedicel and base of calyx lobes outside sparsely
pubescent, nut densely evenly shortly persistently buff
pubescent, fruit otherwise glabrous. 3 longer calyx
lobes to 10 by 2.5 cm, broadly spatulate, obtuse, to 7
mm broad above the to 15 by 12 mm ovate thickened
saccate base; 2 shorter lobes to 8 by | cm, linear, acute,
similar at base. Nut to 2.5 by 1.4 cm ovoid; stylopo-
dium to 6 mm long, prominent.

Distr. Malesia: Peninsular Thailand, in Malaya
(W. coast from Kedah to Selangor; E. coast), Sumatra
(N.E. at Langkat), Borneo (Central & N.E. Sarawak,
E. Sabah, S.E. Borneo to Puruktjau).

Ecol. Scattered, sometimes common, undulating
land or hills below 700 m; sandy clay soils.

Vern. Balau bukit, damar laut kuning (Mal.),
sélangan batu kuning, s. b. bérsisek, tékam (Borneo).

14. Shorea lumutensis Sym. Gard. Bull. S.S. 10 (1939)
364, pl. 22; Mal. For. Rec. 16 (1943) 19, f. 6. — S. ?
inappendiculata (non BURCK) SYM. ex DeEscH, Mal.

454 FLORA MALESIANA [ser. I, vol. 9?

Fig. 81. Trunk-base of Shorea sumatrana (SLOOT. ex THORENAAR) SYM. ex DESCH with some root-climbers
ascending, the soil in front carpeted with seedlings. Measuring staff 2 m; same tree as in fig. 80. Palembang
(Photogr. THORENAAR, 1924).

1982]

DIPTEROCARPACEAE (Ashton)

455

For. Rec. 12 (1936) 9, p.p.; Govt. Gaz. F. M.S. 29, 26
notice n. 5884 (1937), p.p.

Medium-sized to large tree. Young twigs and fruit
calyx caducous cream puberulent, panicles, parts of
perianth exposed in bud, ovary and base of style
persistently grey-brown, rough pubescent. Twigs c. 3
mm @ apically, terete, drying black. Stipules not seen.
Leaves 7-29 by 2.5—7 cm, oblong or elliptic, coria-
ceous, persistently cream lepidote beneath, fuga-
ceously so above; margin + sinuate; base cuneate or
obtuse; acumen short, broad; nerves 12-16 pairs,
obscure above, prominent and drying black beneath;
tertiary nerves densely scalariform, sinuate, evident
beneath; midrib prominent beneath, evident but +
applanate above; petiole 2—3.5 cm long, geniculate, +
persistently cream lepidote. Panicle to 10 cm long,
terminal or axillary; singly or (if terminal) doubly
branched, branchlets to 1.5 cm long, bearing to 5
secund flowers. Flower buds c. 9 mm long at anthesis; 3
outer sepals ovate-deltoid, obtuse; 2 inner ovate, acute
or acuminate; stamens 20-24, of several heights;
filaments rather short, broad at base, tapering and
filiform beneath the ellipsoid anthers; outer anther
cells + sparsely barbate at apices; appendages short,
with 1-2 terminal bristles; ovary and stylopodium
ovoid-conical, tapering into the short stout style. Fruit
+ sessile; 3 longer calyx lobes to 7 by 1.5 cm,
spatulate, obtuse, tapering to c. 5mm above the to 10
by 8 mm elliptic saccate thickened base; 3 shorter
lobes to 4.0 by 0.4mm, + linear, similar at base; nut to
20 by 12 mm, ovoid, prominently beaked.

Distr. Malesia: W. coastal Malaya (Dindings).

Ecol. Local but common on coastal hills above 100
m.
Vern. Balau puteh, b. bukit, damar laut (Mal.).
Note. Without flowers this species is indistinguish-
able from the Bornean S. inappendiculata BURCK.
Sterile specimens from N.W. Johore, Sumatra’s E.
coast, Karimun and Lingga could belong to either
species.

15. Shorea exelliptica MEIER, Act. Bot. Neerl. 12
(1963) 323, pl. 1; MEIER & Woop, Sabah For. Rec. 5
(1964) 167; ASHTON, Man. Dipt. Brun. (1964) 132, f.
13; ibid. Suppl. (1968) 69. — S. ? elliptica (non BURCK)
Sym. Mal. For. Rec. 16 (1943) 13, f. 6, 8. — Shorea sp.
Browne, For. Trees Sarawak & Brunei (1955) 170.
Tall buttressed tree. Twig, panicle, leaf bud, stipule
(short on inner, long on outer surfaces), petiole,
midrib on both surfaces and nerves on undersurface
densely persistently purplish brown to gold-brown
scabrid tomentose; sometimes glabrous on midrib
upper surface in mature trees. Twigs c. 2 mm @
apically, ridged at first, becoming terete or somewhat
compressed, smooth, with c. 2 mm long pale linear
descending stipule scars. Bud to 3.5 by 3mm, ovoid, +
compressed. Stipule to 8 by 4 mm, not at first
caducous, broadly ovate, acute. Leaves 9-15 by 3.5—7
cm, coriaceous, oblong to broadly ovate, golden to
silver lepidote beneath; base broadly cuneate; acumen
to 1 cm long; nerves 12-18 pairs, prominent beneath,

at c. 50°—60°; tertiary nerves slender, sinuate, scalari-
form; petiole 1.2-1.7 cm long, rugose. Panicle to 12cm
long, terminal or axillary, ribbed and slightly com-
pressed; singly branched, branchlets to 2.5 cm long,
bearing to 8 secund flowers; bracteoles to 4mm long,
elliptic, acute, shortly pubescent, fugaceous. Flower
bud to 10 by 3mm, narrowly lanceolate. Calyx densely
pubescent outside, puberulent within; lobes deltoid,
subequal. Petals cream, pink at base, linear, densely
tomentose outside, shortly pubescent within. Stamens
30-40; filaments compressed at base, slender, taper-
ing, slightly hispid; anthers oblong, the outer cells
somewhat larger than the inner cells; appendage to
connective as long as anther, stout, setose. Ovary and
stylopodium shortly ovoid, densely tomentose except
at the base; style as long as ovary and stylopodium,
slender, glabrous. Fruit calyx puberulent to glabrous;
3 longer lobes to 8 by 2.4 cm, spatulate, obtuse, to 4
mm broad above the to 12 by 7 mm ovate thickened
saccate base; 2 shorter lobes to 4.5 by 1.7 cm,
otherwise similar. Nut to 1.5 by 1 cm, ovoid, densely
pale grey-buff pubescent; style remnant to 3 mm long.

Distr. Malesia: Malaya (Kedah, Perak, E. coast),
Borneo.

Ecol. Widespread, leached clay soils on undulating
land, ridges and plateaux to 600 m.

Vern. Balau témbaga (Mal.), sélangan batu tém-
baga (N. Borneo).

16. Shorea inappendiculata Burck, Ann. Jard. Bot.
Btzg 6 (1877) 206; BRANDis, J. Linn. Soc. Bot. 31
(1895) 102, t. 2, f. 21, 30; Merr. En. Born. (1921) 405;
BROWNE, For. Trees Sarawak & Brunei (1955) 171;
ASHTON, Man. Dipt. Brun. Suppl. (1968) 73.

Large buttressed tree with flaky bark. Twigs, buds,
stipules, petioles, midrib above and panicles densely
persistently scabrid rufous pubescent, leaf nervation
beneath sparsely so (denser, longer in young trees);
leaf undersurface frequently + densely silvery lepi-
dote. Twig 2-4 mm @ apically, blackish, prominently
ribbed or compressed at first; stipule scars pale,
horizontal. Bud to 6 by 4mm, ovoid, conical; stipules
to 10 by 4 mm, broadly lanceolate, acute. Leaves
(7-)10-16 by (2-)3-8 cm (to 30 by 15 cm in young
trees), oblong-elliptic, coriaceous; base + unequal,
obtuse or cordate; apex shortly acuminate to obtuse;
nerves 13-24 pairs, prominent beneath, + distinctly
depressed above with the lamina bullate between;
tertiary nerves densely scalariform, slightly elevated
beneath; midrib prominent beneath, depressed above;
petiole 15-30 mm long, + stout. Panicle to 10 cm
long, axillary, ribbed when dry, rather unevenly
persistently buff pubescent; singly branched, branch-
lets to 2 cm long, bearing to 7 secund flowers;
bracteoles to 3 by 2 mm, elliptic, obtuse, shortly
evenly pubescent, fugaceous. Flower bud to 7 by 3mm,
fusiform. Calyx lobes ovate, densely pubescent on
parts exposed in buds, inner sepals fimbriate; 3 outer
sepals subacute, larger than the 2 acuminate inner
sepals. Petals linear, densely pubescent on parts
exposed in bud. Stamens 28-34; filaments setose

456

distally abaxially, compressed, tapering; anthers nar-
rowly oblong-elliptic, sparsely setose distally, the 2
outer sacs larger than the 2 inner; appendage to
connective as long as anthers on outer stamens,
shorter on inner, with a few terminal setae. Ovary and
stylopodium ovoid, densely pubescent, tapering into
the short glabrous style. Fruit pedicel to 4 by 3 mm,
broadening into the receptacle; fruit calyx glabres-
cent; 3 longer lobes to 13 by 3 cm, broadly spatulate,
obtuse, c. 8 mm broad above to 15 by 12 mm ovate
saccate thickened base; 2 shorter lobes to 10 by 0.9 cm,
lorate, subacute, similar at base; nut to 33 by 16 mm,
ovoid, prominently apiculate.

Distr. Malesia: Malaya (N.W. Johore), Sumatra
(Langkat), ?Lingga, northern Borneo (Sarawak to E.
Sabah and south to Tiding and Muara Tewe).

Ecol. Lowland Mixed Dipterocarp forest on coas-
tal hills and immediately behind the peat swamps;
rather rare.

Vern. Pelepak, p. gunong, damar pangin.

Note. See under S. /umutensis, with which it is
vicarious.

17. Shorea falcifera DyER ex BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 86; BEcc. For. Born. (1902) 571; MERR.
En. Born. (1921) 571; ASHTON, Gard. Bull. Sing. 31
(1978) 36. — Hopea linggensis BOERL. Cat. Hort. Bog.
2 (1901) 103. — S. glauca (non KING) BROWNE, For.
Trees Sarawak & Brunei (1955) 168, p.p. — S. flava
MEueR, Act. Bot. Neerl. 12 (1963) 325, pl. 2; ASHTON,
Man. Dipt. Brun. Suppl. (1968) 69, f. 9. — Fig. 10.
Medium-sized tree. Twig, bud, stipule (sparsely so
within), nervation and lamina beneath densely persis-
tently pale yellow lepidote. Twig c. 1 mm @ apically,
slender, terete, smooth; stipule scars short, horizontal,
obscure. Bud to 2 by 1 mm, small, ellipsoid, subacute.
Stipule to 6 by 4mm, elliptic, obtuse, fugaceous. Leaf
6.5-12 by 2.5—-5 cm, narrowly ovate to lanceolate,
coriaceous; base broadly cuneate, subequal; acumen
to 2 cm long, slender; nerves c. 10 pairs, slender,
somewhat elevated beneath, applanate above, at
40°—60°; tertiary nerves obscure, scalariform; midrib
slender, prominent beneath, shallowly depressed
above; petiole 11-18 mm long, somewhat geniculate.
Panicle to 11 cm long, terminal or axillary, terete or
ribbed, shortly evenly persistently buff pubescent;
singly branched, branchlets to 3 cm long, short,
bearing to 5 distichous flowers. Flower buds to 8 by 2
mm, fusiform. Calyx pubescent on parts exposed in
bud; lobes ovate, the two inner subacuminate, some-
what narrower than the outer 3; outer 3 subacute.
Petals cream, linear, densely pubescent on parts
exposed in bud, sparsely so elsewhere. Stamens 33-34;
filaments setose, compressed at base, tapering and
filiform distally; anthers oblong, the two outer cells
slightly the larger, sparsely setose on the distal margin;
appendage to connective shorter than anther, shortly
setose. Ovary and stylopodium pyriform, densely
sericeous, tapering into the short glabrous style. Fruit
pedicel, base of calyx and nut shortly densely evenly
cream pubescent, calyx elsewhere sparsely so. 3 longer

FLORA MALESIANA

(ser. I, vol. 9?

calyx lobes to 9.5—1.8 cm, broadly spatulate, obtuse,
to 7 mm broad above the to 2 by 1.3 cm ovate
thickened saccate base; 2 shorter lobes to 7 by 0.8 cm,
narrowly spatulate, obtuse, similar at base. Nut to 2.5
by 1.5 cm, ovoid; stylopodium to 7 mm, prominent,
filiform.

Distr. Malesia: Malaya (E. coast: Trengganu,
Pahang), N.E. Sumatra (Idi in Atjeh; Langkat;
Lingga), Borneo (Sarawak west of K. Lupar).

Ecol. Locally abundant on dry hillslopes near coast.

Vern. Sélangan batu kéring (Sar.).

18. Shorea materialis Ripv. Agr. Bull. Str. & F. M.S.
9 (1910) 183; Fl. Mal. Pen. 1 (1922) 227; BURN-Mur-
DOCH, Tr. Timb. Mal. Pen. | (1911) 11; HEYNE, Nutt.
Pl. ed. 1, 3 (1917) 306; ibid. ed. 2 (1927) 1123; BurK. J.
Str. Br. R. As. Soc. 81 (1920) 68; Foxw. Mal. For.
Rec. | (1921) 69; ibid. 3 (1927) 62; ibid. 10 (1932) 172;
Sym. Mal. For. Rec. 16 (1943) 29, f. 6, 13; ASHTON,
Man. Dipt. Brun. (1964) 140, f. 13; ibid. Suppl. (1968)
74.— S. glauca (non KING) BROWNE, For. Trees
Sarawak & Brunei (1955) 168, p.p.

Medium-sized tree. Young twig, panicle, leaf bud
and petiole caducous cream lepidote. Twig 1.5mm @
apically, terete, smooth, dark chocolate-brown; sti-
pule scars c. | mm long, pale, falcate. Leaf bud to 3 by
1 mm, linear, acute. Stipule to 12 mm long, linear,
fugaceous. Leaf 8-15 by 3.5—8.5 cm, broadly ovate,
glabrous above, cream lepidote beneath; base cuneate
to subcordate, unequal; apex with to 1.5 cm long
slender acumen; nerves 9-12 pairs, slender but promi-
nent beneath, set at 30°—45° to the midrib at base,
glabrous; tertiary nerves slender, densely scalariform.
Petiole 1.2-2 cm long, to 2 mm @g, cream lepidote.
Panicle to 15 cm long, terminal or axillary, terete or
ribbed; singly branched, branchlets to 12 mm long,
bearing to 11 secund flowers; bracteoles to 2 mm long,
ovate, acute, lepidote. Flower bud to 10 by 4 mm,
lanceolate. Calyx sparsely evenly tomentose outside,
glabrous within; lobes ovate, subacute, subequal.
Corolla cream; petals linear, acute, shortly pubescent
on parts exposed in bud. Stamens c. 30; filaments
glabrous, flat at base, tapering and filiform below the
anther; anthers ellipsoid, the 2 outer cells somewhat
the larger; appendage to connective short, setose.
Ovary and stylopodium ovoid-conical, densely evenly
cream pubescent, tapering into a short glabrous style.
Fruit pedicel 2 mm long, short. Fruit calyx sparsely
pubescent towards base; outer lobes to 9 by 3 cm,
spatulate, obtuse, tapering to 8 mm broad above the c.
8 by 8 mm saccate thickened base; 2 shorter lobes to 6
by 0.8 cm, linear, subacute, similar at base. Nut to 12
by 12 mm, broadly ovoid, densely shortly cream
tomentose, tapering to the 4 mm acute style remnant.

Distr. Malesia: Malaya (E. coast: S. Trengganu,
Pahang, Johore), N.W. Borneo (Brunei, N.E. Sar-
awak), E. coast Sumatra (sterile collections)?

Ecol. Heath forest on white sand giant podsols of
Quaternary marine and estuarine terraces, and sand-
stone cuestas to 800 m. Locally semi-gregarious.

Vern. Balau pasir, b. laut, b. betul (Mal.).

1982]

19. Shorea atrinervosa Sym. Gard. Bull. S. S. 10
(1939) 363, pl. 21; Mal. For. Rec. 16 (1943) 9, f. 5A, 6;
BROWNE, For. Trees Sarawak & Brunei (1955) 167:
MEER & Woop, Sabah For. Rec. 5 (1964) 163, f. 17,
pl. 12b; ASHTON, Man. Dipt. Brun. Suppl. (1968) 68, f.
9. — S. collina (non Ripi.) Foxw. Mal. For. Rec. 10
(1932) 173, p.p. — S. ? inappendiculata (non BURCK)
Sym. ex Descu, Mal. For. Rec. 12 (1936) 9, p.p.; Govt.
Gaz. F.M.S. 29, 26, notice n. 5994 (1937) 2,
p.p. — Fig. 78 A-A3.

Large tree. Bud, twig and petiole at first caducous
buff pubescent, outside of stipule persistently sparsely
so, vegetative parts otherwise glabrous. Twig c. 2mm
@ apically, terete, drying ribbed; stipule scars short,
pale horizontal. Bud to 4 by 2 mm, narrowly ovoid,
acute. Stipule to 8 by 3 mm, lanceolate, acute,
caducous. Leaves 8-16 by 3.5—9 cm, elliptic to ovate,
frequently irregular in shape, coriaceous, undulate, +
persistently white lepidote beneath; base broadly
cuneate to subcordate; acumen to | cm long; nerves
10-12 pairs, prominent beneath, glabrous, frequently
somewhat sinuous, at 45°-55° except at base, some-
times with small glabrous axillary domatia; tertiary
nerves sinuous, slender, scalariform; midrib raised
above, prominently so beneath; petiole 12-22 mm
long, terete. Panicle to 11 cm long, terminal or
axillary, straight, terete, densely somewhat unevenly
shortly pale grey pubescent; singly branched, branch-
lets to 2 cm long, bearing to 6 secund flowers;
bracteoles to 3 by 2 mm, elliptic, acute, sparsely
pubescent, caducous. Flower bud to 8 by 3 mm,
fusiform. Calyx shortly pubescent on parts exposed in
bud; lobes ovate, acute, the inner 2 somewhat shorter,
thinner, than the outer 3; petals crimson at centre,
cream at margins, linear, sericeous on both surfaces.
Stamens 25-33; filaments compressed at base, taper-
ing, sparsely setose; anthers oblong-elliptic, glabrous,
the inner pair of pollen sacs shorter, setose. Ovary and
stylopodium ovoid, pubescent, surmounted by a short
glabrous style. Fruit pedicel to 4 by 2 mm, broadening
into calyx. Fruit calyx sparsely pubescent, tomentum
caducous except at base; 3 longer lobes to 11 by 2.5
cm, spatulate, obtuse, c. 1 mm broad above the to 10
by 8 mm ovate thickened prominently saccate base; 2
shorter lobes to 8.5 by 0.9 cm, lorate, acute, similar at
base. Nut to 2 by 1.3 cm, ovoid, densely shortly evenly
buff pubescent; stylopodium to 2 mm long, short,
conical.

Distr. Malesia: Malaya (E. coast: Trengganu to
Johore), Sumatra (Atjeh, West coast south to Bang-
kahulu, Tapanuli; Riouw Distr. and Langkat in E.),
northern Borneo (Ulu Kapuas, Sarawak, Sabah,
Bulungan to W. Kutei).

Ecol. Undulating land in valleys, and on hillsides
on clay rich soil, in Mixed Dipterocarp forest; locally
common.

Vern. Balau hitam (Mal.), résak bamban, résak
bunga, laru bétina, méranti hursik, rihir minyak kuyung
(Sumatra), /émbasung (Tarakan), mélapi bukit
(Kapuas), sélangan batu hitam (Sabah).

DIPTEROCARPACEAE (Ashton)

457

20. Shorea crassa AsHTON, Gard. Bull. Sing. 20
(1963) 271; Man. Dipt. Brun. (1964) 130, f. 13; ibid.
Suppl. (1968) 69, pl. 13 (bark).

Medium-sized to large tree. Twig, panicle, bud,
stipule (outside only; glabrescent within) and petiole
densely shortly tomentose, with minute adpressed
persistent hair tufts; leaf nervation beneath sparsely
so. Twig to 5 by 2.5 mm @g apically, compressed,
becoming terete and glabrous, sometimes narrowly
evenly cracked; stipule scars c. 2 mm long, cuneate,
pale, ascending. Bud to 6 mm long and broad, ovoid,
compressed, subacute. Stipule to 8 by 4 mm, ovate,
cupped, subacute, fugaceous. Leaves 10-18 by 5-10
cm, elliptic to ovate, pale cream to golden lepidote
beneath; base equal or subequal, cuneate or narrowly
obtuse, occasionally subcordate; acumen to | cm
long; nerves 7-11 pairs, prominent, well spaced, at c.
40°—55°; tertiary nerves slender, sinuate, densely sca-
lariform; midrib applanate at base, depressed towards
apex, above; petiole 3.5—5 cm long, stout. Panicle to 13
cm long, terminal or axillary, terete or somewhat
compressed, ribbed on drying, stout; regularly singly
branched, branchlets to 4.5 cm long, bearing to 12
distichous flowers; bracteoles to 3 mm long, suborbi-
cular, shortly pubescent, caducous. Flower bud to 15
by 3.5 mm, narrowly lanceolate. Calyx pubescent
outside, glabrescent within; lobes broadly ovate,
subacute; outer lobes slightly longer, more obtuse
than inner. Petals cream, pink at base, linear, shortly
pubescent, glabrescent within. Stamens 38-46; fila-
ments broad at base; tapering, hispid; anthers nar-
rowly oblong, cells tapering, sacs subequal or the
outer slightly larger; appendage to connective promi-
nent but always shorter than the anther, setose. Ovary
ovoid, glabrous at base, otherwise tomentose; stylo-
podium longer than ovary, cylindrical, tomentose;
style very short, glabrous. Fruit calyx sparsely puberu-
lent towards apex, more densely so towards base; 3
longer lobes to 9 by 2.5 cm, broadly spatulate,
coriaceous, obtuse, c. 8 mm broad above the c. 1.5 by
1.3 cm elliptic shallowly saccate thickened base; 2
shorter lobes to 7 by 0.7 cm, narrowly oblong, similar at
base. Nut to 2.5 by 2cm, ellipsoid, densely shortly cream
tomentose; style remnant stout, tapering, to 8 mm.

Distr. Malesia: Borneo (Sarawak, Brunei, Ulu
Kapuas, ? Puruktjau, sterile collections).

Ecol. Common on deep leached yellow sandy and
sometimes clay soils, on low hills, terraces and
occasionally dry sandstone ridges to 1000 m; mainly
near the coast.

Vern. Sélangan batu daun tébal (Brun.).

21. Shorea obscura MEtsER, Act. Bot. Neerl. 12
(1963) 333, pl. 6; MEER & Woop, Sabah For. Rec. 5
(1964) 184, pl. 16 (bark); AsHToN, Man. Dipt. Brun.
(1964) 142, f. 13; ibid. Suppl. (1968) 74.

Large tree. Leaf nervation beneath, petiole, twig,
panicle, bud and stipule shortly evenly pale brown
pubescent, almost entirely caducous except on pani-
cle, bud and stipule. Twig c. 1.5-2 mm @ apically,
terete, slender, becoming glabrous; stipule scars short,

458

FLORA MALESIANA

[ser. I, vol. 9?

pale, obscure. Bud 2-4 by 1.5-2.5 mm, ovoid to
falcate. Stipule to 7 by 2.5 mm, oblong, acute,
caducous. Leaves 7-12 by 2.5—5 cm, ovate-lanceolate,
coriaceous, cream lepidote beneath in mature trees,
base cuneate; acumen to | cm long, narrow; with or
without small pore-like axillary domatia; nerves 7-9
pairs, curved, rather prominent, at 30°45’, tertiary
nerves slender, scalariform at 90°; midrib applanate or
slightly raised above; petiole 1.2—2 cm long, slender.
Panicle to 12 cm long, terminal or axillary, + terete;
singly branched, branchlets to 1.5 cm long bearing to 5
secund flowers; bracteoles to 3 mm long, ovate-ellip-
tic, acute, puberulent, fugaceous. Flower bud to 6 by 2
mm, lanceolate. Calyx shortly pubescent outside,
glabrous within; lobes ovate, acute, the inner 2
somewhat thinner than the outer 3. Petals bright pink,
cream at margin, linear, pubescent outside, puberu-
lent within. Stamens 35-45; filaments compressed at
base, tapering, glabrous; anthers oblong-elliptic, the
inner sacs shorter than the outer; appendage to
connective almost as long as anther, stout, setose.
Ovary and stylopodium ovoid, pubescent but for the
glabrous base, tapering into the short glabrous style.
Fruit calyx shortly sparsely caducously cream pubes-
cent; 3 longer lobes to 10 by 2cm, spatulate, obtuse, to
5 mm broad above the c. 7 mm broad saccate base; 2
shorter lobes to 5 by 0.5 cm, subacute, narrowly
oblong, similar at base. Nut to 12 by 9 mm, globose to
ellipsoid, shortly buff pubescent, abruptly apiculate.

Distr. Malesia: Borneo (Sarawak N.E. of Rejang
valley, Sabah, S-E. Borneo to Sampit).

Ecol. Local, skeletal shale soils on ridges and steep
hillsides, typically at 600-800 m, but rarely below and
to 1400 m.

Vern. Mélapi bukit (Kapuas), buntok, bénuas (S.E.
Borneo), sélangan batu tandok (Sabah), s.b. padi
(Brun.).

22. Shorea lunduensis ASHTON, Gard. Bull. Sing. 22
(1967) 284, pl. 29; Man. Dipt. Brun. Suppl. (1968) 73.

Large tree. Twigs and buds shortly evenly fuga-
ceous buff pubescent; leaves glabrous. Twig c. 3 by 2
mm @ apically, at first compressed, somewhat lus-
trous; stipule scars c. 3 mm long, pale, prominent,
ascending. Bud to 5 by 3 mm, ovoid, acute. Stipules
unknown. Leaves 14-24 by 6-15 cm, large, + broadly
ovate to elliptic; base broadly cuneate to subcordate;
acumen to | cm long; nerves 11-15 pairs, obscure
above, prominent beneath, at 40°—SS° to the midrib;
tertiary nerves sinuate, densely scalariform; midrib
hardly elevated above, prominent beneath; petiole
2-3.5 cm long, stout. Panicle to 12 cm long, terminal
or axillary, subterete, densely shortly tufted buff
pubescent, singly or doubly branched, branchlets to 3
cm long; bracteoles to 4 by 2 mm, ovate, densely
shortly pubescent, fugaceous. Flowers secund; bud to
10 by 3 mm, narrowly ovoid, with the calyx somewhat
spreading. Calyx sericeous in parts exposed in bud;
sepals narrowly ovate, subacute, the inner 2 smaller,
relatively broader, than the outer 3. Petals cream,
linear, densely pubescent on parts exposed in bud.

Stamens 47-52; filaments compressed, tapering, gla-
brous; anthers oblong, tapering distally, glabrous;
appendage to connective exceeding length of anther,
densely setose. Ovary and stylopodium pyriform,
densely pubescent, crowned by a glabrous columnar
style. Mature fruit unknown; sparsely shortly pubes-
cent at first. Calyx lobes unequal, 3 long and two
short, spatulate, subacute. Nut crowned by a short, c.
1 mm long, apiculus.

Distr. Malesia: Borneo (W. Sarawak).

Ecol. Local, lower slopes of granodiorite moun-
tains to 650 m, and along porphyry dikes in sedimen-
tary areas.

Note. Possibly vicarious with S. collina of E.
coastal Malaya.

23. Shorea falciferoides Foxw. Philip. J. Sc. 13 (1918)
Bot. 189; ibid. 67 (1938) 296; AsHToNn, Gard. Bull.
Sing. 31 (1978) 37, incl. ssp. glaucescens (MEIJER)
ASHTON. — S. balangeran (non BURCK) VIDAL, Phan.
Cuming. (1885) 97; Rev. Pl. Vasc. Filip. (1886) 61;
BRANDIS, J. Linn. Soc. Bot. 31 (1885) 86; Foxw.
Philip. J. Sc. 4 (1909) Bot. 508, 516; ibid. 6 (1911) Bot.
269; WHITFORD, Bull. For. Bur. Philip. 10 (1911) 73;
Foxw. Philip. J. Sc. 13 (1918) Bot. 187; MerR. En.
Philip. 3 (1923) 96; Reyes, Philip. J. Sc. 22 (1923) 336;
Sym. Gard. Bull. S. S. 8 (1935) 273; ASHTON, Gard.
Bull. Sing. 31 (1978) 36. — S. gisok Foxw. Philip. J.
Sc. 67 (1938) 294, pl. 4 — Fig. 77.

a. ssp. falciferoides.

Large tree. Young twig, panicle, petiole, leaf bud
and stipule shortly densely evenly cream pubescent.
Twig c. 2-3.5 mm g apically, terete or + compressed
and ribbed, stout, smooth; stipule scars c. 1.5 mm
long, pale, cuneate, horizontal. Bud to 5 by 3.5 mm,
globose to ovoid, subacute, slightly compressed.
Stipules c. 10 by 4 mm, obtuse, fugaceous. Leaves
10-18 by 4.5-8 cm, broadly ovate, chartaceous, pale
cream-brown lepidote beneath; base obtuse to
cuneate, subequal; acumen to c. 8 mm long; nerves
8-11 pairs, slender, well spaced, raised but not
prominent beneath, at c. 40°—-S0°; tertiary nerves
slender, densely scalariform, at c. 90° to the nerves;
petiole 1.5-2 cm long, stout. Panicle to 4 cm long,
terminal or axillary, ribbed and somewhat com-
pressed; singly branched, branchlets to 1.2 cm long,
bearing to 6 close secund flowers; bracteoles to 3 mm
long, elliptic, shortly pubescent, fugaceous. Flower
bud to 5 by 2.5 mm, small, lanceolate. Calyx densely
pubescent outside, glabrous within; lobes suborbicu-
lar, obtuse, subequal. Petals cream, narrowly elliptic,
acute, shortly pubescent on both surfaces. Stamens c.
45: filaments glabrous, applanate at base, tapering
and filiform distally, somewhat gibbous; anthers
subglobose, the 2 outer cells slightly larger; appendage
to connective very short on inner anthers, + 4 length
of anthers on outer anthers, sparsely shortly setose
but for a single long apical bristle. Ovary and
stylopodium ovoid, densely pubescent, tapering into
the short glabrous style. Fruit calyx sparsely puberu-

1982]

lent, more densely so at the base; 3 longer lobes to 9.5
by 2.2cm, broadly spatulate, obtuse, with c. 8 by 8mm
thickened saccate base closely adpressed to the base of
the nut; 2 shorter lobes to 7 by | cm, subequal, broad,
tapering to Smm broad above the saccate base. Nut c.
15 by 15 mm, broadly ovoid, shortly densely cream-
buff tomentose; style remnant c. 4 mm long, tapering.

Distr. Malesia: throughout the Philippine islands
to Prov. Zambales and Bulacan (Central Luzon) in
the moderately seasonal northwest.

Ecol. In Mixed Dipterocarp forest to 1000 m, more
or less confined to ridge tops in the everwet areas.

Note. Specimens from the more seasonal N.E.
Luzon (S. falciferoides) have more or less smaller
leaves than other Philippine collections (formerly
named S. gisok).

b. ssp. glaucescens (MEIJER) ASHTON, Gard. Bull. Sing.
31 (1978) 37.—S. glaucescens Meer, Act. Bot.
Neerl. 12 (1963) 327, pl. 3; MEER & Woop, Sabah
For. Rec. 5 (1964) 170; ASHTON, Man. Dipt. Brun.
(1964) 134, f. 13; ibid. Suppl. (1968) 71.

Leaves broadly ovate-falcate, chartaceous, base
subequal; nerves 8-12 pairs, well spaced, raised but
not prominent beneath; petiole rather stout.

Distr. Malesia: Borneo, except the west and south-
west.

Ecol. Clay rich soils in Mixed Dipterocarp forest to
600 m.

Vern. Sélangan batu daun nipis (Brun.), s. b. laut
(Sabah).

Note. Differs from the type subspecies only in the
leaf characters.

24. Shorea superba Sym. Gard. Bull. Sing. 17 (1960)
491; ASHTON, Man. Dipt. Brun. (1964) 146, f. 13; ibid.
Suppl. (1968) 146; MEER & Woop, Sabah For. Rec. 5
(1964) 187, f. 22, pl. 18-19 (habit). — Fig. 9.

Vast, prominently buttressed tree. Twig, panicle,
leaf bud, stipule, petiole and nerves beneath densely
shortly evenly pink-brown pubescent. Twig c. 2.5 by 1
mm g@ apically, compressed and ridged at first,
becoming terete, much branched; stipule scars c. 1.5
mm long, pale, linear, horizontal. Leaf bud to 4 by 3
mm, ovoid, compressed, slightly asymmetrical, sub-
acute. Stipule to 12 by 5 mm, ovate-lanceolate,
fugaceous. Leaves 7-12 by 4-7 cm, oblong, thinly
coriaceous, silver to cream lepidote beneath; base
broadly cuneate; acumen 5-10 mm long, broad;
nerves 16—24 pairs, dense, straight, at 50°—60°; midrib
slender, prominent beneath, depressed above; tertiary
nerves densely scalariform, slender, + obscure;
petioles \-1.5 cm long, slender. Panicle to 8 cm long,
terminal or axillary, somewhat compressed and
ribbed; regularly singly branched, branchlets to 2 cm
long, bearing to 8 close secund flowers; bracteoles to 5
mm long, lanceolate, acute, caducous. Flower bud to 7
by 2 mm, lanceolate. Calyx shortly pubescent outside,
glabrous within; 3 outer lobes deltoid-ovate, acute; 2
inner lobes smaller, ovate, acuminate, thinner. Petals
cream, linear, shortly tomentose outside, glabrous

DIPTEROCARPACEAE (Ashton)

459

within, hardly contorted. Stamens c. 30, subequal;
filaments compressed and broad at base, tapering;
anthers oblong, the outer sacs larger than the inner,
glabrous; appendage to connective short, setose.
Ovary and stylopodium cylindrical to conical, tomen-
tose; style short, glabrous. Fruit calyx puberulent; 3
longer lobes to 6 by 1.2 cm, spatulate, acute or obtuse,
tapering to a c. 8 by 8 mm elliptic saccate thinly
incrassate base; 2 shorter lobes to 30 by 4 mm linear,
acute, similar at base. Nut to 12 by 7 mm, ovoid,
shortly pubescent; style remnant short.

Distr. Malesia: Borneo (Sarawak N.E. of Bintulu,
Sabah, Tidung, Berau, Sampit; sterile coll.).

Ecol. Fertile clay soils at low altitudes in Mixed
Dipterocarp forest to 600 m.

Vern. Sélangan batu daun halus (Sabah), s.b. tulang
ikan (Brun.).

25. Shorea hypoleuca MEER, Act. Bot. Neerl. 12
(1963) 329, pl. 4; MEWER & Woop, Sabah For. Rec. 5
(1964) 174, pl. 13; AsHToN, Man. Dipt. Brun. Suppl.
(1968) 72, f. 9. — Fig. 78 D-D3.

Large tree. Midrib above, bud and stipule outside
shortly evenly persistently ocherous pubescent, twig
caducously so; twig, midrib above, petiole and leaf
beneath yellowish lepidote. Twig 1.5-2 mm @ api-
cally, ribbed, becoming terete, smooth; stipule scars
short, falcate, descending. Stipule to 7 by 4 mm,
oblong-ovate, subacute, fugaceous. Leaves 8.5—17 by
3.5-8 cm, ovate to narrowly elliptic, coriaceous,
yellowish lepidote beneath; margin undulate distally;
base cuneate to obtuse; acumen to 1.5 cm long,
slender; nerves 11-16 pairs, prominent beneath,
straight, oblique at 30°—50°, with small subglabrous
axillary domatia; tertiary nerves slender, scalariform;
midrib depressed above, prominent beneath; petiole
10-16 mm long, rather short, terete. Panicle to 14 cm
long, terminal or axillary, ribbed, shortly sparsely
puberulent; singly branched, branchlets to 2.5 cm
long, bearing to 5 secund flowers; bracteoles fuga-
ceous. Flower buds to 10 by 2 mm, fusiform. Calyx
pubescent on parts exposed in bud; lobes ovate; inner
2 subacuminate, smaller than outer 3; outer 3 sub-
acute. Petals pale yellow, linear, pubescent on parts
exposed in bud. Stamens c. 33; filaments glabrous,
broad and compressed at base, tapering abruptly and
filiform distally; anthers elliptic-oblong, glabrous;
appendage to connective as long as anther, setose, the
apical bristles longer and exceeding length of anther;
ovary and stylopodium pyriform, densely pubescent,
crowned by a short glabrous filiform style. Fruit calyx
lobes sparsely puberulent basally, glabrous distally; 3
longer lobes to 8.5 by 1.7 cm, narrowly spatulate,
obtuse, to c. 3 mm broad above the to 10 by 6 mm
narrowly elliptic thickened saccate base; 2 shorter
lobes to 6 by 0.4 cm, subacute, similar at base. Nut to
15 by 10 mm, ovoid, densely evenly pale buff pubes-
cent; stylopodium to 3 mm long, slender.

Distr. Malesia: Northern Borneo (Sarawak,
Sabah, Tidung).

Ecol. Alluvium, undulating land, hillsides below

460

300 m, on fertile deep soils in Mixed Dipterocarp
forest.
Vern. Sélangan batu kélabu.

26. Shorea malibato Foxw. in Elmer, Leafl. Philip.
Bot. 6 (1913) 1955; Philip. J. Sc. 13 (1918) Bot. 189, pl.
6; ibid. 67 (1938) 298; Merr. En. Philip. 3 (1923) 97;
SLOOT. in Merr. Pl. Elm. Born. (1929) 203.

Large buttressed tree. Buds, twigs and parts of
perianth exposed in bud caducously pale cream
puberulent, petioles more persistently so; panicles and
ovary persistently so. Twigs c. 1 mm @ apically, much
branched, terete, becoming smooth to rugulose,
blackish. Buds to 4 by 2 mm, lanceolate-falcate, acute.
Stipules fugaceous, not seen. Leaves 7-12 by 2-5 cm,
elliptic to lanceolate, + thinly coriaceous; base
broadly cuneate; acumen to 1.5 cm long, tapering,
slender; nerves 11-14 pairs, slender but distinctly
elevated beneath, distinctly depressed above except in
young trees, at 35°—50°; tertiary nerves densely scala-
riform, obscure; midrib prominent beneath, obscure
and depressed above; petiole 9-20 mm long, very
slender. Panicles to 8 cm long, slender, terminal or
axillary, pendant, singly or doubly branched. Flower
bud to 5 by 3 mm; sepals broadly ovate, acute.
Stamens 35-37; filaments glabrous; anthers ellipsoid;
appendages prominent, with 1—3 long terminal, and
2-3 shorter lateral setae. Ovary ovoid-conical, style
short. Mature fruit unknown. Fruit subsessile; calyx
lobes unequal, 3 longer lobes c. 5 by 1.3 cm, spatulate,
obtuse, c. 3 mm broad above the c. 5 by 4 mm elliptic
saccate thickened base; 2 shorter lobes c. 25 by 3 mm,
linear, similar at base; nut small, ovoid, apiculate.

Distr. Malesia: Philippines (Mindanao, Leyte and
Luzon).

Ecol. Local in non-seasonal evergreen forests of
lowlands.

Vern. Malibato, yakal (Tayabas, Zamboanga),
guisok, g. amarillo (Camarines), g. madlao (Leyte).

Note. Replacing S. hypoleuca in the Philippines,
from which it differs notably in the number of stamens
and the smaller, less coriaceous leaf with narrowly
channelled nerves above.

27. Shorea astylosa Foxw. Philip. J. Sc. 13 (1918)
Bot. 188, pl. 5; ibid. 67 (1938) 297; MerR. En. Philip. 3
(1923) 96; AsHToN, Gard. Bull. Sing. 31 (1978)
37. — S. ciliata (non KING) Foxw. Philip. J. Sc. 13
(1918) Bot. 188; ibid. 67 (1938) 300.

Large tree. Young twigs and petioles caducous buff
puberulent, panicles and buds persistently so, calyx
thus at first, becoming sparsely so, parts of petals
exposed in bud and ovary densely persistently cream
pubescent. Twigs c. | mm @ apically, slender, much
branched, terete, smooth. Buds small, ovoid; stipules
fugaceous, not seen. Leaves 6.5—12 by 2.5-6.5 cm,
ovate, thinly coriaceous, + lustrous above, glabrous
or + densely cream lepidote beneath the nerves
excepted; base cuneate to obtuse, subequal; acumen to
1.5 cm long, slender, tapering; nerves 8-9 pairs,
slender, somewhat elevated beneath, evident but +

FLORA MALESIANA

[ser. I, vol. 9?

applanate above as also the midrib, arched and
somewhat sinuate, set at 45°—75°, with + prominent
glabrous pore-like domatia; tertiary nerves densely
scalariform, obscure; petiole 11-25 mm long, long,
slender, geniculate. Panicle to 9 cm long, terminal or
axillary, singly branched; branchlets short, few-flow-
ered. Flower buds to 5 by 3 mm, fusiform; sepals
broadly ovate, acute, subequal; stamens 32, subequal;
filaments compressed, tapering; anthers oblong, gla-
brous, the outer cells tuberculate at base; appendages
exceeding anther apex, densely setose; ovary ovoid,
tapering into a prominent stylopodium; style short,
glabrous. Fruit (? immature): Pedicel c. 2 mm long,
slender; 3 longer calyx lobes to 6 by 1.5 cm, spatulate,
obtuse, c. 3 mm wide above the 7 by 4 mm ovate
elliptic saccate thickened base; 2 shorter lobes to 25 by
2mm, linear, similar at base; nut to 15 by 8 mm, ovoid,
tapering into a long slender stylopodium.

Distr. Malesia: Philippines (Luzon,
Samar, Negros, Mindanao).

Ecol. Local, in Lowland Evergreen Dipterocarp
forest.

Vern. Yacal.

Notes. This and the following species differ princi-
pally in the number of stamens as in the pair S. seminis
and S. sumatrana, and like them occupy geographi-
cally separate ranges.

FoxworTHY maintained that BS 18575 from
Biliran I. and FB 22788 from Quezon Prov., Luzon
represented a different taxon from the type of S.
astylosa (FB 13271, from Zamboanga, Mindanao); he
compared them with S. ciliata KiNG of Malaya.
MERRILL was in disagreement and I concur with
MERRILL that they belong to S. astylosa.

The species differs from the Bornean S. domatiosa
ASHTON, with which it is vicarious, principally in
having a prominent stylopodium and c. 32, in com-
parison with 25-30, stamens.

Biliran,

28. Shorea domatiosa ASHTON, Gard. Bull. Sing. 19
(1962) 285, pl. 16; Man. Dipt. Brun. (1964) 131, f. 13;
ibid. Suppl. (1968) 69; MEER & Woop, Sabah For.
Rec. 5 (1964) 166.

Large tree. Panicle, flowers and stipule outside pale
cream pubescent, domatia fimbriate, parts otherwise
+ glabrous. Twig c. 1.5 mm @, much branched
slender, terete, frequently wrinkled or compressed
when dried, smooth; stipule scars short, pale, horizon-
tal, obscure. Bud c. 4 by 2mm, ovoid, acute. Stipules c.
6 by 2.5 mm, ovoid, narrowly obtuse, caducous.
Leaves 6.5—10 by 3-7 cm, broadly ovate to obovate, +
chartaceous, cream lepidote on undersurface only in
fully mature and old tree; base obtuse or subcordate;
acumen to 8 mm long; nerves 8-12 pairs, slender,
curved, set at 45°-65°, with prominent axillary pore-
like fimbriate domatia; tertiary nerves slender, densely
scalariform, sinuate; midrib prominent beneath;
petiole 1.5—2.5 cm long, slender, long, geniculate with
the distal half swollen. Panicle, bracts and bracteoles
unknown. Bud to 1.3 by 4 mm, fusiform. Calyx
shortly pubescent outside, glabrous within; 3 outer

1982]

DIPTEROCARPACEAE (Ashton)

46]

lobes broadly ovate, subacute, 2 inner lobes narrowly
ovate, acute. Petals linear, obtuse, hardly contorted in
bud, sparsely pubescent on parts exposed in bud,
elsewhere glabrous. Stamens c. 25-30: filaments
applanate towards base, tapering; anthers broadly
oblong, the inner 2 pollen sacs shorter than the outer
2, with sparsely setose apices; appendage as long as
anther, stout, densely setose. Ovary ovoid, densely
tomentose except at base; stylopodium 14 times as
long as ovary, very long, pubescent towards base,
elsewhere glabrous. Fruit calyx glabrous; 3 longer

lobes to 13 by 3 cm, broadly spatulate, obtuse, to 8
mm broad above the to 1.8 by 1.6 cm elliptic saccate
thickened base; 2 shorter lobes to 7 by 1.2 cm,
narrowly spatulate, with similar base. Nut 8.5 by 2.5
cm, ovoid, densely shortly pale cream pubescent: style
remnant to | cm long, filiform.

Distr. Malesia: N.E. Borneo (Rejang valley north-
eastwards to S.W. and S.E. Sabah and Nunukan).

Ecol. Scattered on clay soils in Mixed Dipterocarp
forest below 600 m.

Vern. Sélangan batu lobang idong.

lb. Subsection Barbata

SyM. ex ASHTON, Gard. Bull. Sing. 20 (1963) 266; Man. Dipt. Brun. (1964)
166. — Barbata group Sy. J. Mal. Br. R. As. Soc. 79 (1941) 162. — Fig. 82.
Flower buds subglobose. Petals cream, short, elliptic-oblong, obtuse, connate

at base on falling. Appendages and apices of outer anther cells densely setose.
Distr. Peninsular Burma and Thailand to Malesia: Sumatra, Borneo.

Ecol. Scattered in lowland forests below 1500 m.

29. Shorea glauca KING, J. R. As. Soc. Beng. 62, 2
(1893) 117; BRANpis, J. Linn. Soc. Bot. 31 (1895) 102:
RIDL. Fl. Mal. Pen. 1 (1922) 223; HEYNE, Nutt. Pl. ed.
1, 3 (1917) 300; ibid. ed. 2 (1927) 1118, 1121; Foxw.
Mal. For. Rec. 3 (1927) 64; ibid. 10 (1932) 232; BurRK.
Dict. (1935) 2010; Sym. Mal. For. Rec. 16 (1943) 15, f.
6, 10.

Medium-sized or large tree. Parts of petals exposed
in bud, ovary and panicles densely persistently evenly
ocherous pubescent, sepals thus at first, becoming
sparsely so in fruit; twigs, stipules outside, petiole,
midrib above and leaves beneath persistently cream
lepidote. Twig c. 1 mm @ apically, terete, becoming
smooth, pale brown. Leaves 6-15 by 2.3—9 cm, ovate
to lanceolate, thin; margin undulate; base cuneate:
acumen to 2 cm long, slender, tapering, nerves 7-10
pairs, slender, hardly raised beneath, arched, at
50°-70° to the midrib; secondary nerves slender,
scalariform, obscure; midrib applanate above, slightly
elevated beneath; petiole 10-20 mm long, slender,
terete. Panicles to 7 cm long, slender, lax, terminal or
axillary, terete or compressed; singly or doubly
branched, branchlets bearing to 8 secund flowers;
bracteoles short, linear, fugaceous. Flower bud to 3 by
3 mm, globose; sepals broadly ovate, the 3 outer
somewhat larger, acute, the inner 2 subacuminate;
stamens c. 60, subequal; filaments broadly compressed
at base, tapering; anthers lorate-oblong, barbate at
base and apex, appendages not exceeding anthers,
very short, villous; ovary ovoid, style short, broadly
cylindrical. Fruit pedicel to 1 by 1 mm. 3 outer calyx
lobes to 7 by 1.8 cm, spatulate, obtuse, c. 5 mm wide
above the to 6 by 5 mm ovate, saccate thickened base:
2 shorter lobes to 5 by | cm, otherwise similar. Nut to
1.5 by 1.5 cm, broadly ovoid, shortly apiculate.

Distr. Peninsular Thailand, and in Malesia:
Malaya, west coast Sumatra (from Painan to Atjeh),
Simalur I.

Ecol. Local, but often semi-gregarious, on hills,
especially rocky slopes and ridges, generally near the
coast, to 600 m.

Vern. Damar laut daun bésar, résak, r. rémenia,
terbak, sélimbar, damar laut kuning, tengkawang.

Note. Flowering specimens, without which the
species cannot always be distinguished with certainty
from S. materialis and S. falcifera in subsect. Shorea,
are unknown from east coastal Malaya, east coastal
Sumatra and Riouw, and it seems unlikely that it
occurs there in spite of earlier reports (e.g. SYMING-
TON).

30. Shorea laevis RIDL. Fl. Mal. Pen. 1 (1922) 232:
HEYNE, Nutt. Pl. ed. 2 (1927) 1121; Foxw. Mal. For.
Rec. 10 (1932) 179; Sym. Mal. For. Rec. 16 (1943) 18,
f. 6, 12; BROWNE, For. Trees Sarawak & Brunei (1955)
169; ASHTON, Man. Dipt. Brun. (1964) 139, f. 13, pl.
33 (habit), 35 (stem-base); ibid. Suppl. (1968) 73, f. 9:
MEUER & Woop, Sabah For. Rec. 5 (1964) 178, pl. 12a
(habit), pl. 14a (stem), f. 20; ASHTON, Gard. Bull. Sing.
31 (1978) 38. — S. ciliata (non KING) RIDL. Agr. Bull.
Str. & F.M.S. 4 (1905) 63; Foxw. Mal. For. Rec. 1
(1921) 69; ibid. 3 (1927) 66; ibid. 8 (1930) 19;
Epwarps, Mal. For. Rec. 9 (1931) 142. — Hopea
laevifolia Parus in Fedde, Rep. 33 (1933) 244. — S.
laevifolia ENDERT, Tectona 28 (1935) 292; BROWNE,
For. Trees Sarawak & Brunei (1955) 171. — S. roger-
siana RAIZADA & SMITINAND, Thai For. Bull. Bot. |
(1954) 7.

Vast, prominently buttressed tree. All vegetative
parts epilose. Twig c. 1 mm @ apically, much
branched, slender, terete, smooth; stipule scars small,
obscure. Bud c. 3.5 by 1.5 mm, narrowly ovoid. Stipule
to 8 by 2 mm, narrowly lanceolate, acute, fugaceous.
Leaves 6.5-10 by 2.5-4 cm, narrowly ovate-lanceo-
late; falcate, thinly coriaceous, cream lepidote
beneath in mature trees; base subequal, broadly

462 FLORA MALESIANA [ser. I, vol. 92

Fig. 82. Flower details in Shorea sect. Shorea subsect. Barbata SyM. ex ASHTON. All x 10. Sepals drawn from

inside. — S. ladiana Asuton. A. Bud, A/. outer sepal, A2. inner sepal, 43. androecium, 44. three stamens and

pistil, 45. stamen from inside. — S. asahii ASHTON. B. Stamens and pistil, B/. stamen from inside. — S.

maxwelliana KiNG. C. Bud, Cl. outer sepal, C2. inner sepal, C3. stamens and pistil, C4. stamen from

inside. — S. biawak AsuTon. D. Bud, D1. outer sepal, D2. inner sepal, D3. stamens and pistil (A Rost 15025, B
bb. 35219, C KEP 69917, D S 28778)

1982]

cuneate; acumen to 2 cm long, slender; nerves 11-14
pairs, slender, coriaceous, dense, curved, at c. 50°—60°,
unraised on either surface as also the midrib; tertiary
nerves slender, densely scalariform, diagonal to
nerves; petiole 1-1.5 cm long, slender, geniculate.
Panicle to 12 cm long, terminal or axillary, slender,
terete, shortly persistently pale cream grey pubescent,
+ glabrescent; singly or doubly branched, branchlets
to 3 cm long, bearing to 9 distichous flowers; brac-
teoles to 2 mm long, linear, pubescent, fugaceous.
Flower bud to 2 mm long, globose. Calyx shortly pale
cream-brown tomentose outside, glabrous within; 3
outer lobes ovate-deltoid, subacute; 2 inner lobes
smaller, suborbicular. Petals small, oblong, shortly
pubescent on parts exposed in bud. Stamens c. 50;
filaments slender, tapering, sparsely barbate towards
the apex; anther oblong, the longer, outer, sacs
barbate at base and apex; appendage to connective
small, barbate. Ovary and stylopodium small, conical,
densely buff tomentose, crowned by a short glabrous
style. Fruit calyx shortly buff pubescent on both
surface; 3 longer lobes to 6.5 by 1 cm, oblong,
chartaceous, slightly broader at the obtuse apex,
untapering but saccate and adpressed to the nut at the
base; 2 shorter lobes to 4 by 0.5 cm, subacute,
otherwise similar. Nut to 1.5 by 0.9 cm, ovoid, shortly
densely evenly buff pubescent; style remnant to 4mm
long, tapering, frequently bent over.

Distr. Peninsular Burma and Thailand (Pattani); in
Malesia: Malaya, N. Sumatra (Atjeh), Borneo.

Ecol. Widespread, often common and even gregar-
ious on skeletal or dry soils on ridges in Hill Diptero-
carp forest, typically at 200-1000 m, but sometimes
lower.

Vern. Kumas, k. merah, k. hitam, sélimbar, damar
laut kuning (Mal.), mikai (Sar.), sélangan batu kumus
(Sabah), pényau (W. Borneo), bangkirai tanduk, b.
lampong, mérenting, tenggelan ménpelam, gélam (S.E.
Borneo), bénuas, b. layang (S. Borneo).

31. Shorea asahii ASHTON, Gard. Bull. Sing. 19 (1962)
279, pl. 13; Man. Dipt. Brun. (1964) 128, f. 13; ibid.
Suppl. (1968) 68; MEER & Woop, Sabah For. Rec. 5
(1964) 163. — Fig. 82 B-B1.

Medium-sized tree. Young parts at first shortly
pubescent, otherwise glabrous but for panicles and
flowers. Twig c. 0.7 mm @ apically, slender, terete;
stipule scars small, horizontal, obscure. Bud c. 1.5 by
1.0 mm, falcate, small. Stipule to 5mm long, narrowly
deltoid, fugaceous. Leaves 6-10 by 3-5 cm, ovate,
coriaceous, lustrous; base broadly cuneate, subequal:
acumen c. | cm long, narrow; nerves 6—7 pairs, arched,
slender, at c. 60°: midrib unraised on either surface,
indistinct; tertiary nerves scalariform, dense, very
slender and obscure, at 90° to midrib or slightly
ascending though at 90° to the nerves nearer the
margin; petiole c. | cm long, slender. Panicle to 2 cm
long, terminal or axillary, slender, terete, straight,
densely shortly cream tomentose; singly branched,
branchlets to 7 mm long, short, zig-zag, bearing to 4
close secund flowers; bracteoles to 2 mm long,

DIPTEROCARPACEAE (Ashton)

463

elliptic, acute, puberulent, fugaceous. Flower bud to 2
mm long, globose. Ca/yx shortly pubescent outside,
glabrous within; 3 outer lobes ovate-deltoid, sub-
acute; 2 inner lobes smaller, relatively broader, ovate,
less acute. Petals elliptic-oblong, acute, shortly pubes-
cent outside, puberulent within. Stamens c. 30; fila-
ments broad at base, abruptly tapering, filiform and
sparsely barbate distally; anther narrowly oblong, 2
outer sacs larger, sparsely barbate apically; appen-
dage to connective up to 2 times length of anther.
stout, barbate towards apex. Ovary and stylopodium
ovoid, densely pubescent, tapering into the short
slender glabrous style. Fruit calyx lobes to 4 by 4mm,
subequal, ovate, subacute, sparsely buff pubescent,
adpressed to nut. Nut to 11 by 6 mm, oblong-ovoid,
acute, densely buff pubescent.

Distr. Malesia: N.W. Borneo (Kapuas valley;
Rejang valley to Brunei).

Ecol. Local, on shale spurs below 850 m in Mixed
Dipterocarp forest.

Vern. Tékam padi (Iban), kumus bukit (Mal.).

32. Shorea micans ASHTON, Gard. Bull. Sing. 31
(1978) 38.

Medium-sized tree. Panicles and nut densely
greyish puberulent, fruit calyx sparsely so, other
known parts glabrous. Twig c. 1 mm @ apically,
slender, much branched, smooth, terete. Buds minute;
stipules not seen. Leaves 5-10 by 1.8-4.7 cm, ovate-
lanceolate, thinly coriaceous, lustrous on both sur-
faces; margin narrowly subrevolute; base broadly
cuneate, + unequal; apex to 1.3 cm caudate; nerves
7-8 pairs, very slender, slightly elevated beneath, +
applanate above, at 50°—55°; tertiary nerves obscure,
scalariform; midrib slender, evident and slightly ele-
vated on both surfaces; petiole 7-12 mm long, rather
short, very slender. Panicle to 7 cm long, terminal or
subterminal axillary, slender, shortly branched.
Flowers unknown. Fruit pedicel c. 1 mm long, slender:
3 longer calyx lobes to 5 by 1.5 cm, spatulate, obtuse,
c.4mm broad above the to 8 by 7 mm elliptic saccate
thickened base; 2 shorter lobes to 25 by 4 mm,
narrowly spatulate, acute, similar at base; nut to 19 by
7 mm including the prominent slender apiculus,
ovoid.

Distr. Malesia: N.E. Borneo (once collected north
of Sandakan).

Ecol. On ultrabasic rock in lowlands.

Note. Differing (in the absence of flowering collec-
tion) from S. asahii only in the fruit sepals.

33. Shorea ladiana AsHTON, Gard. Bull. Sing. 19
(1962) 295, pl. 21; Man. Dipt. Brun. (1964) 138, f. 13;
ibid. Suppl. (1968) 73. — Fig. 82 A—AS5.

Small to medium-sized tree. Young twig and
petioles shortly puberulent, bud and panicle grey
tomentose, vegetative parts otherwise glabrous. Twig
1.5-2 mm g apically, terete, much branched, smooth
to rugulose; stipule scars small, slightly descending,
obscure. Bud c. 1 by 0.5 mm, small, conical. Stipule
unknown. Leaves 10-14 by 4.5-7.5 cm, coriaceous,

464

FLORA MALESIANA

[ser. I, vol. 97

Fig. 83. Shorea maxwelliana KiNG. a. Habit, b. fruit, both x 5, c. venation, enlarged (aS 29235, b-c SAN 16988).

lustrous, ovate; base subequal! to equal, obtuse or
broadly cuneate; acumen 1—1.5 cm long; nerves 5—6
pairs, prominent beneath, well spaced, arched, set at c.
40°—50°; tertiary nerves slender, indistinct, densely
scalariform, at 90°; margin usually narrowly revolute:
petiole \-2.2 cm long. Panicle to 15 cm long, terminal
or to 2-axillary, somewhat compressed, straight, lax,
shortly persistently pale cream-grey pubescent; singly
or doubly branched; branchlets to 8 mm long, short,
zigzag, bearing to 9 close secund flowers; bracteoles to
4.5 mm long, ovate-deltoid, acute, shortly pubescent,
fugaceous. Flower bud to 1.5 mm g, globose (only
young bud known). Calyx densely puberulent outside,
glabrous within; sepals ovate, subequal. Petals
broadly elliptic. Stamens 30-35; filaments short,
tapering; anthers oblong, the outer sacs the larger,
shortly barbate apically; appendage to connective
longer than anther, long barbate. Ovary and stylopo-
dium ovoid, densely pubescent, with short glabrous
style. Fruit pedicel to 3 mm long, slender, puberulent.
Calyx shorter than nut, the 5 equal lobes c. 8 mm long
and broad, broadly ovate, puberulent, subacute,
thickened, shallowly saccate, closely adpressed to the
nut. Nut to 1.7 by 1.4 cm, obovoid-globose, densely

shortly buff pubescent, abruptly tapering to the c. |
mm long narrow acute style remnant.

Distr. Malesia: Borneo (Sarawak, Brunei).

Ecol. Local, deep yellow sandy soils in Mixed
Dipterocarp forest on low hills to 300 m.

Vern. Sélangan batu kilat.

34. Shorea biawak AsHTON, Gard. Bull. Sing. 19
(1962) 281, pl. 14; Man. Dipt. Brun. (1964) 129, f. 13,
pl. 37 (bark); ibid. Suppl. (1968) 68; MEveER, & Woop,
Sabah For. Rec. 5 (1964) 165. — Fig. 82 D—D3.
Small tree. Young twig, panicle, leaf bud and
petiole shortly pale buff pubescent, persistent on
panicle and leaf bud. Twig to 1 mm @ apically,
slender, terete, much branched, smooth or striated.
Bud to 1 mm long, small. Stipule unknown. Leaves
6-10 by 2.5-4.5 cm, obovate, thinly coriaceous; base
obtuse or broadly cuneate; acumen to 1.5 cm long,
narrow to caudate; nerves 5—6 pairs, slender, slightly
raised beneath, slightly depressed above, arched, at
50°-60°, with small puberulent axillary domatia;
tertiary nerves obscure, scalariform at c. 90°; midribs
slightly raised beneath, slightly raised or applanate
above; petiole 7-10 mm long, slender. Panicle to 14cm

1982]

long, terminal or 1-axillary, terete; singly branched,
branchlets to 6mm long, short, + secund, bearing to 6
close secund flowers; bracteoles to 2 mm long, ovate,
acute, puberulent, fugaceous. Flower bud to 1.5 mm
long, globose. Calyx shortly puberulent outside,
glabrous within; lobes broadly ovate, acute, subequal,
the inner 2 slightly shorter and relatively wider,
thinner. Petals elliptic, obtuse, puberulent outside,
glabrous within. Stamens c. 35; filaments slender,
tapering, glabrous; anthers narrowly oblong, sparsely
barbate apically, tapering, the 2 outer sacs the larger;
appendage to connective somewhat shorter than
anther, densely barbate. Ovary and stylopodium ovoid
to pyriform, densely pubescent, crowned by a short
glabrous style. Fruit calyx lobes to 7 by 9 mm,
subequal, shorter than nut, suborbicular, obtuse, thin,
closely adpressed to nut, shortly persistently buff
puberulent. Nut to 1.3 by 1 cm, subglobose, obtuse,
densely pale buff pubescent; style remnant to 1.5 mm
long.

Distr. Malesia: Borneo (Rejang valley to S.
Sabah).

Ecol. Local, clay ridges below 600 m, in Mixed
Dipterocarp forest.

Vern. Resak biawak (Brun.), sélangan batu buaya
(Mal.).

Note. Forming, with S. maxwelliana, a pair similar
to S. asahii and S. micans, and differing from the
following species principally in the fruit, though also
in the androecium.

35. Shorea maxwelliana KING, J. R. As. Soc. Beng.
Sc. 62, 2 (1893) 114; BRANbis, J. Linn. Soc. Bot. 31
(1895) 86; RipL. Fl. Mal. Pen. 1 (1922) 227; Foxw.
Mal. For. Rec. 3 (1927) 35; ibid. 10 (1932) 194; Sym.
Gard. Bull. S. S. 7 (1933) 146, pl. 44; ibid. 8 (1934) 28;
ibid. 9 (1938) 325, 326; Mal. For. Rec. 16 (1943) 21, f.
5B, 6, 14; BRowneE, For. Trees Sarawak & Brunei
(1955) 169; ASHTON, Man. Dipt. Brun. (1964) 141, f.
13, pl. 36 (bark); ibid. Suppl. (1968) 74; MEER &
Woop, Sabah For. Rec. 5 (1964) 182, t. 15, f.21. — S.
utilis KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 119;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 81; RIpDL. Agr.
Bull. Str. & F.M.S. 1 (1901) 58; Fl. Mal. Pen. 1 (1922)
230; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 310; ibid. ed. 2
(1927) 1126; Foxw. Mal. For. Rec. 3 (1927) 65; ibid.
10 (1932) 177; Burk. J. Str. Br. R. As. Soc. 81 (1920)
69, fig.; ibid. 86 (1922) 281. — S. barbata BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 81; Rupv. Agr. Bull. Str. &
F.M.S. 1 (1901) 59; J. Str. Br. As. Soc. 54 (1909) 23; FI.
Mal. Pen. 1 (1922) 230; Foxw. Mal. For. Rec. 1 (1921)

DIPTEROCARPACEAE (Ashton)

465

68; ibid. 3 (1927) 68; ibid. 10 (1932) 178. — S. ciliata
(non KING) RipL. Agr. Bull. Str. & F.M.S. 4 (1905)
63. — S. alba Riwt. J. Str. Br. R. As. Soc. 82 (1920)
171; Fl. Mal. Pen. 1 (1922) 230. — Balanocarpus
ovalifolius Raw. J. F.M.S. Mus. 10 (1920) 130, p.p.;
Fl. Mal. Pen. 1 (1922) 247, p.p. — Fig. 82 C-—C4,
83.

Tall, prominently buttressed tree. Vegetative parts
glabrous but for the shortly pubescent to glabrescent
buds and stipules. Twig c. 0.7 mm @ apically, slender,
much branched, smooth; stipule scars minute, short,
obscure. Bud c. | mm long and broad, minute, globose
to conical. Stipule to 4 mm long, puberulent on both
surfaces, linear, fugaceous. Leaves 6-10 by 2.5—4 cm,
coriaceous, ovate-lanceolate, sometimes sparsely
cream lepidote beneath; base obtuse or broadly
cuneate; apex to 2 cm long caudate; nerves 8—10 pairs,
slender; midrib elevated beneath, depressed above, at
c. 40°-50°, well spaced, curved, with or without
minute axillary domatia; tertiary nerves very slender,
scalariform, at 90°; petiole 0.7—1 cm long, slender.
Panicle to 5 cm long, terminal or to 3-axillary, terete,
straight, shortly evenly buff pubescent; singly
branched, branchlets to 1.3 cm long, short, bearing to
6 close secund flowers; bracteoles to 1.5 mm long,
elliptic, puberulent, fugaceous. Flower bud to 2.5 mm
long, globose. Calyx shortly pubescent outside, gla-
brous within. Stamens c. 30; filaments broad at base,
tapering gradually to anther, glabrous; anther nar-
rowly oblong, outer cells larger, barbate apically;
appendage to connective as long as anther, barbate at
apex. Ovary and stylopodium ovoid to pyriform,
densely pubescent; style glabrous, short. Fruit calyx
+ puberulent, more densely so towards base; 3 longer
lobes to 10 by 1.5 cm, spatulate, obtuse, to 4 mm
broad above the to 1.5 by 1 cm elliptic somewhat
thickened prominently saccate base; 3 shorter lobes to
6 by 0.6 cm, otherwise similar. Nut to 2 by 1.5 cm,
ovoid, tapering, apiculate, densely buff pubescent.

Distr. Malesia: Malaya (Penang and Trengganu
southwards), Sumatra (Atjeh, West Coast, Lam-
pong), Borneo.

Ecol. Widespread, sometimes common, in Mixed
Dipterocarp forest in low hills on well drained
clay-rich soils to 700 m.

Vern. Damar laut daun kéchil, kumus hitam, chén-
deras, chéngal batu, résak, r. hitam, balau, séngkaw-
ang, damar laut kuning (Mal.), sélangan batu asam
(Sabah), pakit (W. Borneo), tékam tégelam (Iban,
Tidung), rikir, r. minyak, r. séga, damar bintang, résak
tanduk (Sumatra).

2. Section Pentacme

(DC.) AsHton, Gard. Bull. Sing. 31 (1978) 38. — Pentacme A. DC. Prod. 16, 2
(1868) 626; Kina, Ann. R. Bot. Gard. Cale. 5, 2 (1896) 151; BRANDIs, J. Linn. Soc.
Bot. 31 (1895) 72; Foxw. Mal. For. Rec. 10 (1932) 154; Sym. Mal. For. Rec. 16
(1943) 104, f. 63 (map), 64-66. — Fig. 84.

466 FLORA MALESIANA [ser. I, voles?

Flowers large, cream, ovoid, on lax spreading racemes. Petals broadly elliptic,
ovate, hardly contorted, falling separately. Stamens 15, in 3 verticils; filaments
short, applanate, tapering; anthers linear, glabrous, with 4 pollen sacs each
prolonged and tapering apically into a prominent awn at least as long as the
stoutly acicular + recurved appendage. Ovary ovoid, style filiform. Stipules and
bracts fugaceous, small. Leaf with scalariform tertiary nerves; midrib raised,

evident, above. Bark surface V-section fissured. Wood anatomy and properties

widely divergent between S.E. Asia and Philippine species.

Distr. One species in seasonal S.E. Asia and Malesia: Malaya, and one in the Philippines.

Ecol. See under the species.

Note. S. siamensis differs from the Philippine species in having a hard heavy durable wood anatomically
similar to that in sect. Shorea; in the Philippine species the wood is soft, light and unsuitable for exterior work.

36. Shorea siamensis Miq. Ann. Mus. Bot. Lugd.-Bat.
1 (1864) 214; DC. Prod. 16, 2 (1868) 631; WaALP. Ann.
7 (1868) 379: Dyer, FI. Br. Ind. 1 (1874) 304; RYAN &
Kerr, J. Siam Soc. 8 (1911) 7, 15, 35, t. 1-4; ASHTON,
Gard. Bull. Sing. 31 (1978) 39. — Hopea suava WALL.
[Cat. (1828) n. 959, nomen] ex A. DC. Prod. 16, 2
(1868) 635. — Pentacme suavis A. DC. Prod. 16, 2
(1868) 626; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 72;
Indian Trees (1906) 68; GAMBLE, Man. Ind. Timb.
(1922) 77; Troup, Silv. Ind. Trees 1 (1921) 145; Crap,
Fl. Siam. Enum. | (1925) 145; Gite in E. & P. Pfi.
Fam. ed. 2, 21 (1925) 259; SMITINAND, Thai For. Bull.
1 (1954) 9, 10, 24.— Pentacme siamensis (MuIQ.)

Kurz, J. R. As. Soc. Beng. Sc. 39, 2 (1870) 66; FI.

Burma | (1877) 119; PrerreE, For. Fl. Coch. 3 (1889) t.
225-227, incl. var. laevis PIERRE et var. suavis (DC.)
PIERRE; HEIM, Rech. Dipt. (1892) 56; Bot. Tidsskr. 25
(1902) 46; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1,
3, 6 (1895) 263, fig.; GUERIN, Fl. Gén. L.-C. 1 (1910)
379, 385, fig.; CRAIB, Fl. Siam. Enum. | (1925) 145;
GILG in E. & P. Pfl. Fam. ed. 2, 21 (1925) 238; Foxw.
Mal. For. Rec. 8 (1930) 15, 35; ibid. 10 (1932) 155, pl. 2
(seedlings); Burk. Dict. (1935) 1690; Sym. Mal. For.
Rec. 16 (1943) 105, f. 64-66. — S. bracteata PIERRE ex
LANESSAN, Pl. Util. Colon. Fr. (1886) 301.—S.
mekongensis PIERRE ex LANESSAN, /.c. — S. suavis
PIERRE ex LANESSAN, /.c. — S. tomentosa (non Miq.)
PIERRE, For. Fl. Coch. 3 (1889) sub t. 225 in
syn. — Pentacme malayana KING, J. R. As. Soc. Beng.
Sc. 62, 2 (1893) 107; BRUHL & KING, Ann. R. Bot.
Gard. Calc. 5, 2 (1896) 151, t. 184; Rip. Agr. Bull.
Str. & F.M.S. 1 (1901) 62; Fl. Mal. Pen. 1 (1922) 220;
Crais, Fl. Siam. Enum. 1 (1925) 145. — Pentacme
tomentosa CRAIB, Kew Bull. (1915) 423; Fl. Siam.
Enum. | (1925) 145.

Small gnarled, + shortly deciduous tree (in Male-
sia). Young calyx, twigs and panicle caducous puberu-
lent, otherwise glabrous (in Mal.). Twigs 3-S mm @
apically, terete, smooth; stipule scars short, pale. Bud
small, ovoid, acute; stipule to 18 by 7 mm, ovate-fal-
cate, fugaceous. Leaves 9-12 by 6-13 cm (smaller if
subtending panicles), broadly ovate-oblong, charta-
ceous; base deeply cordate to cuneate (if subtending
panicles); acumen to | cm long, short, broad; nerves
13-16 pairs, slender but prominent beneath, barely
elevated above as also the midrib, arched, the basal

pair with prominent lateral branchlets; tertiary nerves
remotely scalariform, sinuate, slender but typically
prominently elevated beneath; petiole 3-5 mm long, c.
2mm g, straight. Panicle to 14 mm long, terminal or
axillary, lax, peduncle stout at base; irregularly
branched, branchlets to 7 cm long, bearing a few +
secund flowers. Anthesis directly following leaf fall;
bud to 15 by 6 mm, large, ellipsoid; sepals narrowly
ovate, prominently slender, acuminate, subequal;
petals broadly elliptic, glabrous (in Mal.):; stamens 15,
subequal; filaments lorate, slightly tapering; anther
cells linear, extended apically beyond the connective
into prominent tapering horns c. 4 their length;
appendages acicular, glabrous, c. 5 length of anthers;
ovary narrowly ovoid, tapering into a stoutly colum-
nar style c. twice its length and exceeding the stamens
at anthesis. Fruit pedicel to 5 by 3 mm, broadening
into the receptacle; 3 longer calyx lobes to 12 by 1.3
cm, narrowly spatulate, narrowly obtuse, c. 4 mm
broad above the to 8 by 7 mm elliptic saccate
thickened base; 2 shorter lobes to 7 by 0.5 cm, lorate,
subacute, similar at base. Nut to 20 by 12 mm, ovoid,
tapering into an up to 8 mm long prominent acicular
style remnant.

Distr. Burma, Indochina and Thailand south to
Malesia: N.W. Malaya (Langkawi and once in Perlis).

Ecol. A tree of Dry Dipterocarp forests, especially
on skeletal soils and overlying granite; occurring
scattered on rocky headlands in Malaya.

Vern. Témak batu, témak, méranti témak.

Note. Varying greatly in the distribution and
density of the tomentum, reduction of which is
roughly correlated with increasing humidity of cli-
mate or soil. The species Pentacme malayana (leaf
glabrescent) and P. tomentosa (both surfaces of leaf
tomentose) were distinguished from P. siamensis
(=suavis) with tomentose leaf undersurface, but the
continuous variation which exists in nature suggests
merely ecotypic differentiation in panmictic popula-
tions.

37. Shorea contorta VIDAL, Sinopsis (1883) 15, t. 15E;
Rev. Pl. Vasc. Filip. (1886) 61; BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 88; MerR. Philip. J. Sc. 1 (1906) Suppl.
98: AsHTON, Gard. Bull. Sing. 31 (1978) 40. — Pen-
tacme contorta (VIDAL) MERR. & ROLFE, Philip. J. Sc.

DIPTEROCARPACEAE (Ashton)

467

Fig. 84. Flower details in Shorea sect. Pentacme (DC.)
ASHTON. — S. contorta VIDAL. A. Bud, BI. outer
sepal, B2. inner sepal, both from inside, C. petal, D.
stamens from outside, E. pistil, all x 10 (FB 10721).

3 (1908) Bot. 115; Merritt, Bull. Bur. For. Philip. 8
(1908) 48; WuitForD, Philip. J. Sc. 4 (1910) Bot. 703;
Bull. Bur. For. Philip. 10 (1911) 61; Foxw. Philip. J.
Sc. 4 (1910) Bot. 511; ibid. 6 (1911) Bot. 266; ibid. 13
(1918) Bot. 186; ibid. 67 (1938) 287; Merr. En. Philip.
3 (1923) 95; Reyes, Philip. J. Sc. 22 (1923) 332. — Pen-
tacme paucinervis BRANDIS, J. Linn. Soc. Bot. 31
(1895) 73. — Pentacme mindanensis Foxw. Philip. J.
Sc. 13 (1918) Bot. 185; ibid. 67 (1938) 289; Merr. En.
Philip. 3 (1923) 95; Reyes, Philip. J. Sc. 22 (1923)
332. — Fig. 84.

Medium-sized, sometimes large evergreen V-fis-
sured buttressed tree. Panicle, parts of petals exposed
in bud, ovary and leaf buds densely persistently pale
brown puberulent, twigs, petioles and calyx outside
caducously so. Twig c. 2 mm @ apically, terete or +
ribbed; stipule scars short, descending. Leaf buds to 6
by 3 mm, lanceolate; stipules fugaceous. Leaves 9-29
by 5.5—11 cm, ovate to lanceolate, thinly coriaceous;
base subequal, obtuse or rarely cordate (subpeltate in
young trees); apex broadly to 1 cm long acuminate;
nerves 5—8(—9) pairs, slender but prominent beneath,
distant, arched, + applanate above as also the midrib,
set at 45°—70°; tertiary nerves densely scalariform,
slender, hardly elevated on either surface; petioles
20-33 mm long, slender. Panicles to 22 cm long (if
terminal), to 14 cm long (if axillary), singly or doubly
branched; branchlets to 4cm long. Flower buds to 8 by
4 mm, ovoid, lanceolate; sepals ovate, obtuse, the
outer 3 somewhat the larger; petals broadly oblong-
elliptic, acute; stamens 15, subequal; filaments short,
broad, applanate; anther cells linear, subequal, pro-
longed into prominent distal horns c. 5 their length
and as long as the stout appendage; sty/e columnar, c.
thrice length of ovary. Fruit shortly pedicellate; 3
longer calyx lobes to 12 by 3 cm, spatulate, obtuse,
tapering toc. 8 mm wide at the incrassate saccate base;
2 shorter lobes to 9 by 1.5 cm, otherwise similar; nut to
35 by 15 mm, narrowly ovoid, apiculate.

Distr. Malesia: throughout the Philippines.

Ecol. Common, often semi-gregarious in more or
less seasonal Semi-evergreen Dipterocarp forests in
the lowlands, uncommon in everwet areas.

Vern. Malaanonan, lauan.

Note. Leaves of Mindanao and some other speci-
mens in the non-seasonal areas are usually longer
leaved, and formerly named Pentasacme mindanensis
(cf. S. falciferoides, S. polysperma).

3. Section Neohopea

ASHTON, Gard. Bull. Sing. 20 (1963) 266; Man. Dipt. Brun. (1964) 116. — Fig. 85.

Flower small, cream, bud globose. Peta/s broadly elliptic, falling separately.
Stamens 15, in 3 verticils; filaments stout, short, compressed, hardly tapering;
anthers with 4 pollen sacs, subglobose; appendage to connective hardly exceeding
anther apex, short, stout, glabrous. Ovary and stylopodium conical, both
puberulent; style very short. Stipules and bracts minute, fugaceous. Leaf with

468

FLORA MALESIANA

[ser. I, vol. 9?

scalariform tertiary nerves; midrib broad, evident above. Bark and wood as in

sect. Shorea.
Distr. & Ecol. See under the species.

Fig. 85. Flower details in Shorea sect. Neohopea

ASHTON. — S. isoptera ASHTON. A. Bud, B/. outer

sepal, B2. inner sepal, both from inside, C. stamens

abaxial view, C/. outside view on staminal group, C2.

inside view on staminal group, D. pistil, all x 10
(BRUN 3018).

38. Shorea isoptera ASHTON, Gard. Bull. Sing. 19
(1962) 293, pl. 20; Man. Dipt. Brun. (1964) 137, f. 13:
ibid. Suppl. (1968) 73; MEER & Woop, Sabah For.
Rec. 5 (1964) 189, f. 19. — Fig. 85, 86.

Tall, stoutly buttressed tree. All vegetative parts
glabrous. Twig to 1.5mm g apically, slender, smooth,
terete; stipule scars short, obscure. Bud to 1.5 mm
long, small, globose. Stipule caducous, unknown.
Leaves 9-16 by 5.8-8 cm, ovate; base broadly cuneate
to obtuse, decurrent to 2 mm down petiole (peltate in
saplings); acumen to 1.5 cm long, narrow; margin
undulate; nerves 9-11 pairs, curved, slightly raised
beneath, depressed above, at 45°—65”; tertiary nerves
scalariform, sinuate, slender; midrib terete, slightly

elevated, beneath, + applanate above; petiole |.3—2
cm long. Panicle to 11 cm long, terminal or to
3-axillary, terete or somewhat compressed, lax,
shortly persistently greyish puberulent; regularly
alternately doubly branched, branches to 5 cm long,
branchlets to 1 cm long, short, bearing to 6 close
secund flowers; bracts unknown; bracteoles to 1 mm
long, linear, pubescent, fugaceous. Flower bud to 1.5
mm long, globose. Calyx shortly pubescent outside,
glabrous within; lobes subequal, imbricate, acute; 3
outer lobes ovate, 2 inner lobes deltoid. Petals densely
pubescent outside, shortly sparsely pubescent within,
oblong, obtuse, strongly contorted. Stamens 15, in 3
verticils, double alternating with single stamens; fila-
ments broad, compressed, hardly tapering, glabrous;
anthers oblong-globose, pollen sacs subequal or the
outer somewhat larger; appendage to connective
short, stout, glabrous, not exceeding the anther apex.
Ovary and stylopodium ovoid-conical, glabrous at
base, puberulent near apex, crowned abruptly by a
short glabrous style; stigma minute. Fruit calyx
glabrescent to puberulent at base; lobes to 5.5 by 1.5
cm, subequal, spatulate, obtuse, to 5mm broad above
the slightly broader saccate base, rotate when ripe,
narrowly imbricate at base. Nut to 9 by 10 mm, ovoid,
broader than long, shortly pale grey-brown pubes-
cent, on an up to 8 mm @ broad shallow receptacle;
style remnant to 4 mm long, tapering.

Distr. Malesia: Northern Borneo (Sarawak, S.W.
Sabah and Sandakan Distr.).

Ecol. Locally frequent on clay rich + calcareous
soils, and on limestone scarps, on low hills below 600
m.

Vern. Sélangan batu gélombang, s.b. main bulu
ayam.

Note. An isolated species whose fruit, with broad
shallow receptacle and subequal sepals, and androe-
cium are unique.

4. Section Richetioides

Herm, Rech. Dipt. (1892) 48; ASHTON, Gard. Bull. Sing. 20 (1963) 267; Man. Dipt.
Brun. (1964) 116; Gard. Bull. Sing. 22 (1967) 288. — Richetia HEI, Bull. Mens.
Soc. Linn. Paris 2 (1891) 975. — Shorea, Richetia group SyM. Gard. Bull. S.S.9
(1938) 330; Mal. For. Rec. 16 (1943) 44, f. 27 (map). — Shorea, Meranti Damar
Hitam group Sym. Gard. Bull. S. S. 9 (1938) 330; Mal. For. Rec. 16 (1943) 2,
44. — Shorea subg. Richetia (HEIM) MEER, Act. Bot. Neerl. 12 (1963) 322, nom.
inval. — Fig. 87, 89.

Flowers usually small, usually bright lemon yellow or pale yellow; petals
narrow, strongly twisted and forming a sharply defined though small cup at base
enclosing anthers at anthesis, falling as a rosette. Leaf with + reticulate pellucid

1982] DIPTEROCARPACEAE (Ashton) 469

n

Fig. 86. Shorea isoptera ASHTON. a. Flowering twig, b. fruit, c. nut, all x 4 (aS 17978, b-c SAN 15222).

tertiary nerves; midrib elevated or depressed above, evident. Young leaves often
deep violet or magenta. Large or small trees, the larger with stout prominent
buttresses. Bark usually appearing scaly; phelloderm thin, inconspicuous;

expansion tissue in long fingers, becoming wider outwards.
Distr. & Ecol. See under the species.
Vern. Damar hitam (Mal.), lun (Sarawak), mérakunyat (Sum., Dayak), séraya kuning (Sabah).

4a. Subsection Polyandrae

ASHTON, Gard. Bull. Sing. 22 (1967) 288. — Fig. 87.

Bud large, broadly ovoid. Stamens co; filaments compressed at base, tapering;
anthers narrowly oblong; appendages to connectives somewhat shorter than
anther apices, densely setose. Ovary ovoid, without stylopodium, pubescent; style
short, broad, prominently trifurcate.

470

Fig. 87. Flower details in Shorea sect. Richetioides

Heim subsect. Polyandrae ASHTON. — S. polyandra

AsHTON. A. Bud, B/. outer sepal, B2. inner sepal, both

from inside, CJ. staminal group from outside, C2.

staminal group from inside, D. pistil, all x 10 (Kos-
TERMANS 13302).

39. Shorea polyandra AsHTON, Gard. Bull. Sing. 22
(1967) 286, pl. 32; Man. Dipt. Brun. Suppl. (1968), f.
11. — Fig. 87, 88.

FLORA MALESIANA

[ser. I, vol. 97

Very large, buttressed tree. Twig, petiole and leaf
beneath persistently purplish rufous lepidote, fading
to grey; bud and stipule persistently shortly purplish
rufous puberulent. Twig c. 1 mm g apically, slender,
terete, smooth or minutely striated; stipule scars
short, horizontal, obscure. Bud to 2 by | mm, small,
ellipsoid, subacute. Stipule to 5 by 2 mm, lanceolate,
subacute, fugaceous. Leaves 8-13 by 3—5 cm, lanceo-
late, chartaceous, undulate; base cuneate; acumen to 2
cm long, slender; nerves 11—14 pairs, slender, elevated
beneath, at 40°-60°; tertiary nerves scalariform,
slender, at 90° to the midrib; midrib slender, evident,
applanate above, prominent beneath; petiole 14-20
mm long, slender. Panicle to 6 cm long, terminal or
axillary, terete, densely dark rufous pubescent; singly
branched, branchlets to 1 cm long, bearing to 3 more
or less distichous flowers; bracteoles to 4 by 3 mm,
elliptic, subacute, rufous pubescent, caducous. Flower
bud to 5 by 4mm, broadly ovoid. Calyx sericeous on
parts exposed in bud; lobes suborbicular, subequal.
Petals oblong, obtuse, densely pubescent on parts
exposed in bud. Stamens 102-107, very many, sub-
equal; filament compressed at base, tapering, c. }length
of anther, anther narrowly oblong, c. 4 times as long
as broad; appendage to connective filiform, tapering,
somewhat shorter than length of anther, densely
pubescent. Ovary ovoid, densely pubescent, tapering
into a short broadly columnar trifid style; style
pubescent except at apex. Fruit pedicel to 1 mm long,
short. Calyx sparsely purplish rufous pubescent to-
wards the base, elsewhere glabrescent; 3 longer lobes
to 8 by 1.4m, spatulate, obtuse or subacute, tapering
to 8 mm broad above the to 10 by 8 mm tuberculate
saccate thickened base; 2 shorter lobes to 5 by 0.5 cm,
linear, acute, similar at base. Nut to 30 by 13 mm,
narrowly ovoid, acute, densely rufous pubescent.

Distr. Malesia: Borneo (Ulu Kapuas, Sarawak, S.
E. Sabah, S. E. Borneo to Pulau Laut and Meratus
mountains).

Ecol. Locally abundant on fertile clay rich soils on
calcareous shales, igneous and volcanic rocks below
600 m.

Vern. Putang lénit, p. bési, mérakunyit, damar
hirang, d. kétuyang, d. kuning, d. jangkar, kélapih
pahit, lodan, duku mintola.

4h. Subsection Richetioides

ASHTON, Gard. Bull. Sing. 22 (1967) 288. — Fig. 89.
Flowers usually small, buds fusiform. Stamens (10—)15, in (2—)3 verticils;

filaments broad at base, frequently gibbous, tapering + abruptly medially,
filiform distally; anthers broadly oblong to subglobose; appendages to connec-
tives longer than anthers, + scabrous towards apex. Ovary with stylopodium,

shortly tomentose, with a slender columnar style.

Distr. Non-seasonal western Malesia including the Philippines.

Ecol. Lowland forests below 1500 m.

Notes. S. multiflora has been observed to be thrip-pollinated; in view of the uniformity of flower structure
other species are likely to be also. Sympatric species flower sequentially. The short-sepalled species do not forma

1982] DIPTEROCARPACEAE (Ashton) 47]

C Gg d = q

Fig. 88. Shorea polyandra ASHTON. a. Sterile twig, b. flowering twig, c. fruit, d. nut, all x 2/3 (a bb. 31167, 5
KOSTERMANS 13302, c—d bb. 12375).

MALESIANA

FLORA

472 [ser. I, vol. 97

Fig. 89. Flower details in Shorea sect. Richetioides HEIM subsect. Richetioides. All x 10, except BJ. — S. maxima

(KING) SyM. A. Young stamens from inside, 4/. mature stamens from outside, 42. pistil. — S. longiflora

(BRANDIS) SYM. B. Young stamens from inside, B/. anther of young stamen, x 20, B2. mature stamens from

outside, B3. pistil. — S. angustifolia ASHTON. C. Stamens from inside, C/. pistil. — S. richetia Sym. D. Stamens
from inside, D/. pistil (A KEP 80222, BS 19425, C BRUN 778, D RosLi s.n., tree 121).

natural group and some form species pairs with long-sepalled species, as in S. multiflora and S. hopeifolia.
Species with long fruit sepals are invariably emergent, whereas most with short sepalled are of the main canopy
and understorey. Endemism is heavily predominant among short sepalled species, while several of the
widespread species, e.g., S. gibbosa, S. faguetiana, and S. hopeifolia (long-sepalled) and S. multiflora
(short-sepalled) show complex geographical variation, some of which I recognize as local sibling species (S.
mujongensis, S. cuspidata, S. alutacea of S. gibbosa; S. iliasii, S. kuantanensis, S. kudatensis of S. faguetiana; S.
richetia of S. multiflora). Such patterns recall those of S. macroptera and its allies in sect. Mutica subsect.
Auriculatae where apomixis through adventive embryony is known to occur.

40. Shorea kuantanensis ASHTON, Gard. Bull. Sing. 31
(1978) 41.

Medium-sized tree. Twig apices and leaf buds
sparsely buff puberulent, ovary densely so. Twig c. 2
mm @ apically, terete, striated. Leaves 7.5—12 by 2.7-5
cm, lanceolate, thinly coriaceous; base cuneate, sub-
equal; acumen to | cm long, slender; margin subrevo-
lute; nerves 8-9 pairs, slender but elevated beneath,
obscurely depressed above; tertiary nerves reticulate;

midrib prominent beneath, hardly elevated above;
petiole 15-18 mm long, slender. Panicles and flowers
unknown. Fruit pedicel to 4 mm long, expanding into
the fruit base; calyx lobes + subequal, to 22 by 4mm,
linear except at the expanded incrassate saccate base;
nut to 25 by 11 mm, narrowly ellipsoid, acute,
exceeding fruit sepals.

Distr. Malesia; Malaya (Bukit Goh forest, Kuan-
tan).

1982]

—

DIPTEROCARPACEAE (Ashton)

473

Ecol. Apparently once, frequent on the basalt soils
in this one locality, now converted to plantation.
Vern. Damar hitam.

41. Shorea longiflora (BRANDIS) Sym. Gard. Bull. S.S.
9 (1938) 330; SLoor. Reinwardtia 3 (1956) 318;
BROWNE, For. Trees Sarawak & Brunei (1955) 163;
ANDERSON, Gard. Bull. Sing. 20 (1963) 158; ASHTON,
Man. Dipt. Brun. (1964) 156, f. 14; ibid. Suppl. (1968)
86. — Hopea longiflora BRANDIS, J. Linn. Soc. Bot. 31
(1885) 63. — Hopea longifolia (non DYER) MERR. En.
Born. (1921) 402. — Balanocarpus longiflorus Foxw.
ex SyM. Gard. Bull. S. S. 8 (1934) 29, pl. 7. — Balano-
carpus grandifolius RipL. ex Sym. Gard. Bull. S. S. 8
(1934) 29, nomen in syn. — Fig. 89 B—B3, 90.

Small, frequently crooked, tree, Young twig, pani-
cle, bud and stipule fulvous powdery pubescent,
caducous on twig and petiole. Twig to 2.5 mm @
apically, terete, stout, becoming striated and papery
flaked. Bud to 3 by 2mm, slightly compressed, globose
to ellipsoid. Stipule to 10 by 6 mm, oblong-elliptic,
obtuse, cupped. Leaves 10-24 by 4-6 cm, narrowly
ovate to lanceolate, coriaceous; base obtuse; acumen
narrowly acute, 1-2 cm long; margin prominently
revolute; nerves 12-15 pairs, distant, prominent
beneath, at 60°—70°; tertiary nerves broadly scalari-
form; petiole 1-1.2 cm long, short. Panicle to 11 cm
long, terminal or axillary, terete; unbranched or singly
branched; bracteoles minute, fugaceous. Flower bud
to 8 by 3.5 mm, lanceolate, relatively large. Calyx
glabrous but for the shortly fimbriate margin; lobes
broadly ovate, thickened, the inner 2 somewhat
smaller, thinner, more constricted basally, shortly
acuminate, the outer 3 obtuse. Petals brownish purple
to dark yellow, linear, strongly contorted, inner
margin shortly setose, puberulent on parts exposed in
bud. Stamens 15, in 3 unequal verticils; filaments
stout, gradually tapering; anther oblong, becoming +
reflexed at anthesis; appendage to connective as long
as anther, sparsely ciliate towards apex. Ovary small,
ovoid, glabrous; stylopodium glabrous, slender,
tapering; style glabrous; stylopodium and style about
twice length of ovary. Fruit subsessile; calyx glabrous,
lobes subequal, c. 7 mm long and broad, broadly
deltoid to ovate, rather thin, striated, not adpressed to
the base of the nut but somewhat spreading with the
apices turned slightly inwards. Nut to 4.5 by 1.7 cm,
lustrous, glabrous, finely striated, ellipsoid to sub-
ovoid or obovoid, often bent over to one side; style
remnant short, acute.

Distr. Malesia: Borneo (Sarawak, Brunei, Sangku-
lirang and Lower Mahakam region in south-east).

Ecol. Shallow peat swamps and rarely yellow
sandy soils, on low hills and plateaux to 1000 m.

Vern. Damar hitam paya (Brun.), /un paya (Sar.).

42. Shorea macrobalanos AsHTON, Gard. Bull. Sing.
22 (1967) 202, pl. 37; Man. Dipt. Brun. Suppl. (1968)
86, f. 11.

Tall buttressed tree. Leaf bud and stipule persis-
tently buff pubescent, otherwise glabrous. Twig c. 4

mm @ apically, terete, becoming prominently verru-
cose; stipule scars c. 3 mm long, obscure, ascending.
Leaf bud to 2 by 2mm, ovoid, acute. Stipule unknown.
Leaves 19-37 by 9-15 cm, oblong, coriaceous; margin
revolute; base cordate; apex shortly broadly acu-
minate or obtuse; nerves 12-16 pairs, prominent
beneath, set obliquely at 45°-65°; tertiary nerves
slender, remotely subreticulate; midrib applanate
above, prominent beneath; petiole 1.8-3.8 cm long,
stout. Panicle to 32 cm long, terminal or axillary,
terete to somewhat compressed, glabrous, pale
brown, becoming rugose, singly or doubly branched.
Flower to 9 by 3.5 mm, lanceolate, large. Calyx
glabrous; lobes broadly ovate, thickened; inner 2
somewhat smaller than outer 3. Perals linear, inner
margin shortly setose, puberulent on parts exposed in
bud. Stamens 10, in a single verticil; filaments slender,
tapering; anthers broadly oblong; appendages to
connective as long as anther. Ovary narrowly ovoid,
tapering, densely yellow-brown pubescent; style gla-
brous. Fruit sessile, entirely glabrous; calyx lobes to 8
by 8 mm, ovate, acute, + undulate and subrotate, not
closely adpressed to the nut, mounted on a to 1 cm g,
to 8 mm deep, receptacle. Nut to 5 by 2.5 cm, large,
oblong, shortly apiculate.

Distr. Malesia: Borneo (W. and Central Sarawak;
E. Kalimantan: W. Kutei).

Ecol. Rare, clay rich soils on undulating land and
ridges to 900 m.

Vern. Engkabang low (Sar.).

43. Shorea peltata Sym. J. Mal. Br. R. As. Soc. 19
(1941) 158, pl. 6; Mal. For. Rec. 16 (1943) 56, f. 29;
DescuH, Mal. For. Rec. 14 (1941) 27; SLoor. Rein-
wardtia 3 (1956) 337, f. 8.

Small tree. Panicles, petals outside and nut persis-
tently densely buff puberulent, calyx outside cadu-
cously so, parts of petals exposed in bud puberulent;
other parts glabrous. Twig c. 2mm @ apically, terete,
smooth. Buds minute, ovoid, acute. Stipules un-
known. Leaves 8-16 by 4-7.5 cm, oblong, charta-
ceous, prominently peltate; acumen to | cm long,
cuspidate; nerves 8—9 pairs the first 3 of which arise
from the petiole insertion, arched, slender but dis-
tinctly raised beneath, elevated above as also the
subreticulate tertiaries and terete midrib; petiole 1.8—3
cm long, slender. Panicle to 14 cm long, terminal or
subterminal axillary, with to 1 cm long branchlets
bearing to 5 secund flowers. Flower buds to 6 mm long,
lanceolate; sepals ovate, acute, subequal; petals yel-
low; stamens 15, in 3 unequal verticils; filaments broad
at base, tapering and filiform beneath the broadly
ellipsoid anthers; appendages c. 15 times length of
anther, scabrous towards the apex, slender; ovary
ovoid, sericeous distally, crowned by a glabrous
columnar style equal in length. Fruit subsessile; calyx
lobes to 8 by 5 mm, short, subequal, ovate, acute,
incrassate, tuberculate; nut to 30 by 9 mm, fusiform,
tapering, acute.

Distr. Malesia: S. E. Malaya (N. E. Johore), E.
Sumatra (Indragiri Uplands), West Borneo.

FLORA

Fig. 90. Shorea longiflora (BRANDIS) SYM. with nar-
row, thick leaves. Brunei (Photogr. G.H.S. Woop,
SAN 17535).

Ecol. Locally common or even gregarious, on
well-drained flat land or low hills in Mixed Diptero-
carp forest.

Vern. Méranti telepok,
(Malaya), manga (Sumatra).

sama rupa meranti

44. Shorea richetia SyM. Gard. Bull. S. S. 9 (1938)
330; SLOOT. Reinwardtia 3 (1956) 335, f. 6; ASHTON,
Man. Dipt. Brun. Suppl. (1968) 89, f. 11, pl. 15
(slash). — Richetia coriacea HEIM, Bull. Mens. Soc.
Linn. Paris 2 (1891) 975. — Balanocarpus coriaceus
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 112; Sym. Gard.
Bull. S. S. 8 (1934) 27; Merr. En. Born. (1921)
407. — Fig. 89 D-D1.

Medium-sized tree. Apices of twigs, buds and
stipules puberulent, other vegetative parts glabrous.
Twig 1-2 mm @ apically, terete, smooth, pale grey-
brown; stipule scars short, horizontal, obscure. Bud to
2 by 2 mm, ovoid, subacute. Stipule to 5 by 2 mm,
lanceolate, acute. Leaves 5—11 by 3-6.5 cm, broadly
elliptic, thickly coriaceous; base broadly cuneate;
acumen to | cm long; nerves S—7 pairs, comparatively
stout and prominent beneath, arched, at 45°—65°;
tertiary nerves subscalariform; midrib applanate to
slightly depressed above, prominent beneath; petiole
6-10 mm long, stout. Panicle to 8 cm long, terminal or
axillary, compressed or terete, lax, buff puberulent or

MALESIANA

[ser. I, volR?2

glabrous; singly branched, branchlets bearing to 5 +
distichous flowers. Buds to 7 by 2 mm, fusiform.
Sepals broadly ovate, fimbriate but otherwise gla-
brous; outer 3 acute, inner 2 relatively broader,
thinner at margin. Petals lime-yellow, lanceolate,
densely pubescent on parts exposed in bud, imbricate
at base after opening forming a cup, spreading and
contorted apically. Stamens 16-17 in 3 unequal
verticils; filaments compressed as base, tapering and
filiform below the broadly ellipsoid anthers; appen-
dage to connective slender, c. 14 times length of
anther, pubescent in the distal 5. Ovary ovoid, densely
pubescent; style columnar, tapering, 2—3 times as long
as ovary, pubescent in the basal 5. Fruit pedicel to 2
mm long, stout. Calyx glabrous; lobes to 1.7 by 1 cm,
subequal, narrowly ovate, acute, incrassate, saccate
and tuberculate at base. Nut to 2.5 by 1.5cm, obovoid,
acute, densely shortly persistently buff pubescent.

Distr. Malesia: N. W. Borneo (W. Sarawak and
lower Kapuas valley).

Ecol. Local, on leached soils on undulating land in
Mixed Dipterocarp and Heath forest.

Vern. Lun mélapi.

45. Shorea laxa SLooT. Reinwardtia 3 (1956) 345;
ASHTON, Man. Dipt. Brun. (1964) 155, f. 14, pl. 38
(habit, saplings); ibid. Suppl. (1968) 86, pl. 16 (bark);
MEER & Woop, Sabah For. Rec. 5 (1964) 75. — S.
peltata (non SYM.) BROWNE, For. Trees Sarawak &
Brunei (1955) 164.

Medium-sized to large tree. Bud, stipule outside,
twig, panicle, petiole and the basal half of the midrib
beneath persistently shortly + sparsely pale cream-
brown pubescent. Twig 2mm @ apically, smooth. Bud
c. 3 by 2mm, globose to ovoid, subacute. Stipule to 5
by 2mm, fugaceous, lanceolate, acute. Leaves 7-14 by
4-9 cm, ovate to elliptic, coriaceous; base obtuse or
broadly cuneate; acumen c. I—1.5 cm long, narrow;
margin slightly revolute; nerves 8-10 pairs, prominent
beneath, at 50°-85° towards the base and 30°-40° at
the apex, curved and following the margin for a short
distance distally; tertiary nerves well spaced, scalari-
form; petiole 1.5—2.2 cm long. Panicle to 18 cm long,
terminal or axillary, terete, slender, lax; irregularly
doubly branched, branchlets to 3 cm long bearing to 7
+ distichous flowers; bracteoles to 1.5 mm long,
minute, fugaceous. Flower bud to 8 by 2mm, relatively
large, lanceolate, acute. Calyx shortly cream pubes-
cent outside, glabrous within; lobes ovate, acute,
shortly acuminate, the inner 2 slightly smaller, thin-
ner, basally constricted, with a pronouncedly setose
margin. Petals cream, linear, pubescent on parts
exposed in bud, imbricate at base and forming a small
cup on opening, spreading distally but hardly twisted.
Stamens 15, in 3 unequal verticils; filaments slender,
tapering, hardly gibbous; anthers subglobose; appen-
dage to connective twice length of anther, slender,
reaching base of style, shortly ciliate towards apex.
Ovary and stylopodium ovoid-conical, densely shortly
pubescent, tapering distally; style short, densely setose
in basal half, otherwise glabrous. Fruit calyx glabrous,

1982]

DIPTEROCARPACEAE (Ashton)

475

lobes subequal, c. | cm long and broad, broadly ovate,
thickened at the base, tuberculate, closely adpressed
to the base of the nut, the obtuse thin apices recurved.
Nut c. 3.5 by 2.5 cm, large, broadly obovoid, shortly
buff pubescent, splitting open at germination un-
evenly to reveal brilliant red cotyledons.

Distr. Malesia: N. E. Borneo (N. E. Sarawak,
Brunei, S. E. Sabah).

Ecol. Local, deep yellow sandy soils, Mixed Dip-
terocarp forest on hills near coast; on ultrabasics in
Sabah.

Vern. Damar hitam timbul (Brun.).

46. Shorea balanocarpoides Sym. Gard. Bull. S. S. 9
(1938) 330; Mal. For. Rec. 16 (1943) 47, f. 29, 30;
Descu, Mal. For. Rec. 14 (1941) 27, 28; ibid. 15 (1941)
127: SLoot. Reinwardtia 3 (1956) 340, f. 9. — Balano-
carpus pahangensis Foxw. Mal. For. Rec. 10 (1932)
145: Burk. Dict. (1935) 287; Descn, Mal. For. Rec. 12
(1936) 37, 38. — S. dolichocarpa SLoot. Reinwardtia
3 (1956) 342: ASHTON, Man. Dipt. Brun. (1964) 151, f.
14; ibid. Suppl. (1968) 83.

Small to medium-sized tree. Young twig, leaf bud,
stipule outside, panicle and buds shortly sparsely grey
puberulent, otherwise glabrous. Twig c. 1.5 mm 2
apically, terete, much branched. Bud to 2 by 1.5 mm,
small, globose to ovoid, obtuse. Stipule to 4 mm long,
linear, fugaceous. Leaves 6-12 by 2—7 cm, coriaceous,
+ ovate; base obtuse or broadly cuneate, usually
unequal, decurrent to 1.5 mm along the petiole:
acumen to | cm long, narrow; nerves 5-7 pairs,
strongly curved, widely spaced, at 60°—70°, running
parallel to the margin for a short distance distally:
tertiary nerves distant, scalariform at the margin,
reticulate near the midrib; petiole 1.2-2 cm long.
Panicle to 8 cm long, terminal or axillary, slender,
terete; somewhat irregularly doubly branched,
branchlets to 1.5 cm long, short, frequently zigzag,
bearing to 8 distichous flowers; bracteoles minute,
fugaceous. Flower bud to 4.5 by 1.2 mm, lanceolate,
acute. Calyx shortly pubescent outside, glabrous
within; 3 outer lobes deltoid, acute; 2 inner lobes
subequal to them, suborbicular, mucronate, thin.
Petals yellow-brown with bright yellow margin,
linear, puberulent on parts exposed in bud, imbricate
in basal half, spreading and twisted distally when
opened. Stamens 15, in 3 slightly unequal verticils;
filaments broad and slightly gibbous at base, tapering
and filiform distally; anther broadly oblong to subglo-
bose; appendage to connective somewhat longer than
anther, shortly ciliate towards apex. Ovary and stylo-
podium conical, glabrescent, tapering; style short,
stout, glabrous. Fruit subsessile; calyx lobes c. 5 mm
long and broad, equal, short, deltoid, subacute,
glabrous, saccate and closely adpressed to the nut,
margin slightly revolute. Nut to 3 by 1.3 cm, ellipsoid
to obovoid, densely evenly pubescent, appearing
distinctly purplish; style remnant short, acute.

Distr. Malesia: Malaya (Kedah, Perak, E. coast),
Sumatra (Atjeh; Langkat), Borneo (Sarawak N.E. of
Rejang valley and Brunei).

Ecol. Common in Mixed Dipterocarp forest on
undulating ground and on ndges to 700 m.

Vern. Damar katup, mérawan, m. hijau, kala daun
bésar, damar hitam d.b. (Malaya), damar hitam gondol
(Brun.).

47. Shorea multiflora (BURCK) Sym. Gard. Bull. S. S.
9 (1938) 330; Mal. For. Rec. 16 (1943) 54, f. 28, 29, 33,
34: DescH, Mal. For. Rec. 14 (1941) 27, 28; ibid. 15
(1941) 127; SLoor. Reinwardtia 3 (1956) 320, f. 1;
Browne, For. Trees Sarawak & Brunei (1955) 63;
ASHTON, Man. Dipt. Brun. (1964) 157, f. 14, pl. 39
(bark); ibid. Suppl. (1968) 87; MEER & Woop, Sabah
For. Rec. 5 (1964) 76, f. 1b.— Doona multiflora
Burck, Ann. Jard. Bot. Btzg 6 (1887) 234. — Richetia
latifolia Hem, Bull. Mens. Soc. Linn. Paris 2 (1891)
976. — Richetia acuminata HEM, /.c. 979. — Richetia
oblongifolia Heim, I.c. 979. — Richetia penangiana
Hei, /.c. 980; BRANDIS & GiLG in E. & P. Pfi. Fam. ed.
1, 3. 6 (1895) 268. — Balanocarpus penangianus KING,
J. R. As. Soc. Beng. Sc. 62, 2 (1893) 131; BRANDIS &
Gitc in E. & P. Pfi. Fam. ed. 1, 3, 6 (1895) 268;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 109; BRUHL &
KING, Ann. R. Bot. Gard. Calc. 5, 2 (1896) 158, t. 191;
Burk. J. Str. Br. R. As. Soc. 81 (1920) 65, fig.; Dict.
(1935) 287; Rupv. Fl. Mal. Pen. 1 (1922) 246; Foxw.
Mal. For. Rec. 1 (1921) 66; ib. 3 (1927) 58; ib. 8 (1930)
11; ibid. 10 (1932) 143; Rip. Fl. Mal. Pen. 1 (1922)
246. — Hopea multiflora BRANDIS, J. Linn. Soc. Bot.
31 (1895) 60; BoERL. Cat. Hort. Bog. 2 (1901) 102, incl.
var. venosa BOERL.; RIDL. Fl. Mal. Pen. 1 (1922) 237;
Foxw. Mal. For. Rec. 10 (1932) 110; Burk. Dict.
(1935) 1191. — Balanocarpus latifolius BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 112; MerR. En. Born. (1921)
407: Foxw. Mal. For. Rec. 10 (1932) 145; Burk. Dict.
(1935) 286. — Balanocarpus acuminatus BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 113; MeRR. En. Born. (1921)
407. — Balanocarpus sibogae BOERL. Cat. Hort. Bog.
2 (1901) 112. — Balanocarpus sp. SLOOT. ex MERR. PI.
Elm. Born. (1929) 200. — Balanocarpus multiflorus
Sym. Gard. Bull. S.S. 7 (1933) 153, pl. 47; Descu, Mal.
For. Rec. 12 (1936) 37, 38. — Fig. 5.

Small to medium-sized, occasionally large tree.
Young twig, petiole, stipule outside and bud shortly
grey tomentose, caducous on twig and petiole. Twig c.
1 mm g apically, slender, terete, smooth, much
branched. Bud 1-2 by 1.5—2 mm, small, ovoid, obtuse
or acute. Stipule to 6 by 2 mm, narrowly lanceolate,
acute, fugaceous. Leaves 4.5-7.5 by 2-3.5 cm, small,
ovate-lanceolate; base cuneate, equal or subequal.
with or without paired domatia; acumen to 1.5 cm
long, narrow; nerves 8-10 pairs, very slender and
barely elevated beneath, distant, at 50°—60°; tertiary
nerves scalariform to reticulate; petiole 7-10 mm long,
slender. Panicle to 16 cm long, terminal or axillary,
terete, lax, shortly sparsely or densely evenly persis-
tently pale cream-buff pubescent; regularly doubly,
rarely trebly, branched, branchlets to 1 cm long,
zigzag, bearing to 9 distichous flowers; bracteoles
minute, fugaceous. Flower bud to 2.5 mm long, small,
lanceolate. Calyx shortly cream pubescent outside,

476

glabrous within; lobes ovate, acute, the inner 2 slightly
thinner, more constricted at the base. Petals pale
yellow, linear, pubescent on parts exposed in bud, the
basal 4 imbricate forming a cup, twisted and spreading
distally. Stamens (10—-)15, in (1—)3 slightly unequal
verticils; filaments short, broad at base, tapering,
filiform distally; anther subglobose; appendage to
connective 15—2 times length of anther, reaching base
of style, shortly ciliate distally. Ovary ovoid, tapering,
shortly pubescent; stylopodium pubescent, tapering:
style short, glabrous; the 2 latter as long as ovary.
Fruit calyx puberulent to glabrous but for fimbriate
margin; lobes to 5 by 4mm, equal, deltoid, thickened,
saccate, subacute. Nut to 2 by 1.2 cm, obovoid, finely
striated longitudinally, glabrous and lustrous or
shortly grey pubescent; style remnant mucronate.

Distr. Malesia: Malaya, Sumatra (Karimun, Asa-
han S.E. to Langsa, Tapanuli, and Palembang, S.W.
to Sibolga, Pariaman, Painan), Borneo.

Ecol. Widespread and common, on low hills near
coast and inland ridges to 700 m, in Heath and Mixed
Dipterocarp forest.

Vern. Damar hitam, d. katup, sénggai (Mal.), d.
tanduk, d. hitam, d. siput, riung, méranti képala rusa,
mandirawan, képala tupai, d. résak hitam manis
(Sumatra), tismantok (Murut), banjutan (Dusun), /oan
sane barit, bunbun puteh, bambéring, mandjin bukit,
képala pipit, puting délatit (Indon. Borneo).

48. Shorea patoiensis ASHTON, Gard. Bull. Sing. 19
(1962) 302, pl. 24; Man. Dipt. Brun. (1964) 159, f. 14;
ibid. Suppl. (1968) 88; MEER & Woop, Sabah For.
Rec. 5 (1964) 78.

Medium-sized tree. Vegetative parts glabrous but
for the fimbriate stipule and bud scales and frequently
a white waxy powder on young buds and twigs in
mature trees. Twig c. 1 mm @g apically, slender, much
branched in mature trees, terete, smooth or rugulose.
Leaf bud c. 3 by 1 mm (to 4 by 2 mm in young trees),
narrowly ovoid, obtuse with the bud scales conspi-
cuously patent. Stipule c. 8 by 2mm, glabrous but for
margin, oblong, acute. Leaves 5—8 by 2—3.5 cm, ovate,
chartaceous, frequently subfalcate; base + broadly
cuneate, decurrent for up to | mm at the petiole
insertion; acumen c. | cm long, narrow; nerves 7-9
pairs, curved, set obliquely (35°—45°), slightly elevated
above, slender but prominent beneath; tertiary nerves
densely scalariform; petiole 7-10 mm long. Panicle to
9 cm long, terminal or axillary, rather short and
compact, slender, terete, shortly persistently pale
greyish brown pubescent; singly or doubly branched,
branchlets to 1 cm long, bearing to 6 flowers;
bracteoles minute, fugaceous. Flower bud to 3.5 by 1.5
mm, lanceolate, acute. Ca/yx densely reddish brown
pubescent; lobes very small, suborbicular, obtuse, the
inner 2 somewhat thinner, constricted at the base.
Petals bright lemon-yellow, linear, imbricate and
forming a cup at the base, twisted and spreading
apically. Stamens 15, in inequal verticils; filaments
rather short, broad at base, abruptly tapering and
filiform distally; anther subglobose; appendage to

FLORA MALESIANA

[ser. I, vol. 9?

connective as long as anther, shortly ciliate towards
the apex, reaching almost to the style apex. Ovary
ovoid, shortly pubescent; stylopodium cylindrical,
pubescent; style short, glabrous. Fruit calyx shorter
than nut, caducous yellowish buff pubescent, margin
persistently fimbriate; lobes 3-S mm long and broad,
subequal, broadly ovate, + adpressed to the base of
the nut and united at the base to formac. 5mm @ cup.
Nut to 15 by 8 mm, small, oblong-ellipsoid, striated,
glabrescent at maturity, with a minute style remnant.

Distr. Malesia: Northern Borneo (Central and
N.E. Sarawak, S.E. Sabah, Tidung, E. Kutei).

Ecol. Local, on fertile clay rich soils on calcareous
shales and volcanic rocks, hills below 500 m.

Vern. Damar hitam padi (Brun.), séraya kuning
pinang (Sabah), njérakat (Berau).

49. Shorea induplicata SLoot. Reinwardtia 3 (1956)
327, f. 2; ASHTON, Man. Dipt. Brun. Suppl. (1968) 85,
f10:

Medium-sized tree. Innovations pale rufous scabrid
tomentose, turning dark grey-brown and persisting
on twig, buds, stipule, petiole and leaf beneath. Twig
c. 2 mm @ apically, terete, becoming verruculose;
stipule scars short, horizontal, obscure. Bud to 3 by 2
mm, ellipsoid, obtuse. Stipule c. 5 by 2 mm, linear, not
at first caducous. Leaves 8-17 by 2-5 cm, lanceolate,
coriaceous, margin revolute; base cordate; acumen to
2 cm long, slender; nerves 12-16 pairs, prominent
beneath, at 45°-55° except at base; tertiary nerves
subscalariform; midrib depressed above, prominent
beneath; petiole 6-13 mm long, rather short. Flowers
unknown. Fruit pedicel to 2 mm long. Calyx glabres-
cent; /obes to 8 by 6 mm, subequal, ovate, acute, the
apices becoming subpatent, thickly incrassate. Nut to
2.5 by 2.0 cm, broadly ellipsoid, obovoid, subacute,
densely shortly evenly fulvous pubescent.

Distr. Malesia: Borneo (North-west of the Kapuas
and Lupar).

Ecol. Local on podsols on sandstone hills; Mixed
Dipterocarp forest — Heath forest ecotone.

Vern. Lun puteh (Sar.), téenkuyung (Sanggau).

50. Shorea subcylindrica SLooT. Reinwardtia 3 (1956)
331, f. 4; ASHTON, Man. Dipt. Brun. Suppl. (1968) 90,
fealile

Small to medium-sized tree. Young twigs, bud,
stipules outside (subglabrous within) and petiole pale
brown frequently somewhat flocculent caducous
pubescent. Twig c. 2 mm @ apically, terete or
occasionally compressed, straight, minutely fissured;
stipule scars short, horizontal. Bud c. 3 by 2 mm,
ovoid, obtuse. Stipule to 10 by 4 mm, lanceolate,
subacute, fugaceous. Leaves 9-24 by 4-12 cm, rela-
tively large, narrowly elliptic or ovate to lanceolate,
coriaceous, lustrous; base cuneate, rarely obtuse;
acumen to 1.5 cm long, slender to caudate; nerves
8-11 pairs, prominent beneath, evident above and
sunken in shallow furrows as also the midrib, oblique,
at 35°-S0°; midrib prominent on both surfaces;
tertiary nerves subreticulate, the scalariform elements

1982]

set at 90° to the midrib; petiole 9-18 mm long. Panicle
to 12 cm long, terminal or axillary, terete, lax, +
densely somewhat flocculent buff pubescent, partially
caducous; doubly or trebly branched, branchlets
bearing to 5 + secund flowers; bracteoles c. 1 mm
long, minute, deltoid, acute, fugaceous. Flower bud to
3 by | mm, small, fusiform. Sepals ovate, pubescent
on parts exposed in bud; 3 outer acute, 2 inner
acuminate, thinner towards margin than outer 3.
Petals cream, narrowly lanceolate, sparsely pubescent
on parts exposed in bud, imbricate at base after
opening and thus forming a cup, spreading and
contorted apically. Stamens 15, in 3 unequal verticils;
filaments compressed at base, tapering and filiform
below the subglobose anthers; appendage to connec-
tive as long as or somewhat longer than anther,
slender, glabrous, exceeding style apex. Ovary ovoid,
sericeous; style columnar, stout, somewhat shorter
than ovary, sericeous in the basal 4. Fruit pedicel c. |
mm long, short. Calyx glabrescent; lobes to 12 by 9
mm, subequal, ovate, acute, relatively thin, hardly
saccate. Nut to 23 by 16 mm, ellipsoid, acute, densely
shortly pale buff pubescent.

Distr. Malesia: N.W. Borneo (Sarawak west of the
Lupar).

Ecol. Local, on leached clay soils on undulating
land in Mixed Dipterocarp forest.

51. Shorea obovoidea SLooT. Reinwardtia 3 (1956)
332, f. 5; ASHTON, Man. Dipt. Brun. Suppl. (1968) 88,
feedil:

Small to medium-sized tree. Leaf bud, twig and
petiole densely + unevenly shortly golden-brown
pubescent, nervation beneath sparsely so. Twig c. 2
mm g, terete, becoming pale red-brown, slightly
rough; stipule scars short, pale, cuneate, horizontal.
Bud to 2 by | mm, ovoid, acute. Stipule unknown.
Leaves 5—13 by 1.7-4.5 cm, elliptic, coriaceous; base
cuneate; acumen to 12 mm long, slender or caudate;
nerves 8-10 pairs, slender but prominent beneath,
arched, at 45°—55°; tertiary nerves subreticulate: mid-
rib depressed above, prominent beneath; petiole 5-8
mm long, slender. Panicle to 7 cm long, terminal or
axillary, terete, densely + unevenly shortly golden-
brown pubescent; singly branched, branchlets bearing
to 5 distichous flowers. Bud to 3 by 2mm, small. Calyx
lobes ovate, obtuse, pubescent on parts exposed in
bud, the outer 3 somewhat longer than the inner 2.
Petals pale yellow, pubescent on parts exposed in bud.
Stamens 15, in 3 unequal verticils; filaments com-
pressed at base, tapering medially and filiform below
the oblong anthers; appendage to connective slightly
longer than anther, filiform, villous in the distal 4.
Ovary narrowly ovoid, sericeous near the apex,
slender; style glabrous. Fruit pedice/ c. | mm long, c. 2
mm @, short. Calyx densely evenly persistently yel-
low-brown pubescent; lobes to 7 by 4 mm, subequal,
short, ovate, acute. Nut to 2.5 by 1.7 cm, obovoid
to ellipsoid, acute, densely evenly shortly buff
pubescent.

Distr. Malesia: N.W. Borneo (W. and Central

DIPTEROCARPACEAE (Ashton)

477

Sarawak; Kalimantan: Kapuas Valley; Upper Bar-
ito).

Ecol. Rare, Mixed Dipterocarp forest on shale hills
and undulating land to 500 m.

52. Shorea chaiana ASHTON, Gard. Bull. Sing. 31
(1978) 42.

Large buttressed tree. Petioles, panicles, perianth
outside and ovary persistently + densely cream-buff
puberulent; sepals, twigs and leaf nervation below
sparsely + caducously so; other parts glabrous. Twigs
c. 1mm @ apically, much branched, terete, becoming
smooth, dark brown. Buds minute. Leaves 6-11 by
2-4 cm, elliptic-lanceolate, + distinctly falcate, sub-
coriaceous, margin subrevolute; base cuneate or
obtuse, subequal; acumen to 15 mm long, slender,
caudate; nerves 8-11 pairs, slender but prominent
beneath, evident above, arched; tertiary nerves reticu-
late, distinctly elevated beneath; midrib prominent
and terete beneath, evident but applanate to shallowly
depressed above; petiole 5-8 mm long, short, slender.
Panicle to 6.5 cm long, terminal or axillary, slender,
singly branched; branchlets to 2 cm long. Flower buds
to 5 by 2 mm, lanceolate. Sepals broadly ovate,
subacuminate, subequal. Stamens 15; filaments
expanded and gibbous in the basal half, filiform
distally; appendages acicular, c. 25 times as long as the
narrowly ellipsoid 2-locular anthers. Ovary ovoid-
conical, surmounted by an equally tall columnar
puberulent stylopodium and shorter glabrous style.
Mature fruit unknown; sepals ovate, subequal; ovary
ovoid.

Distr. Malesia: Northern Borneo (Central and
N.E. Sarawak).

Ecol. Local, in Mixed Dipterocarp forest below
1000 m.

Note. The leaves somewhat resemble those of S.
longisperma Roxs. though the tomentum beneath is
more sparse, the base unequal, and besides, the fruit
calyx lobes are short and unequal. The leaf base and
tomentum also differentiates it from S. obovoidea
SLOOT.

53. Shorea collaris SLooT. Reinwardtia 3 (1956) 329;
f. 3; ASHTON, Man. Dipt. Brun. Suppl. (1968) 82, f. 10.

Large buttressed tree. Petiole and bud persistently
greyish buff sericeous, young twigs and midrib above
caducously so. Twig c. 2 mm @ apically, terete,
smooth; stipule scars short, horizontal, obscure. Bud
to 2 by 2 mm, ovoid, subacute. Stipule unknown.
Leaves 13-23 by 4.3-8.5 cm, oblong-lanceolate, some-
what chartaceous; margin frequently somewhat revo-
lute; base obtuse; acumen to | cm long, broad; nerves
13 pairs, slender but elevated beneath, at 45°—60°;

tertiary nerves slender, hardly raised, scalariform:
midrib applanate above, slender but prominent
beneath; petiole 19-30 mm long. Panicle to 9 cm long,
terminal or axillary, terete, densely pale greyish buff
pubescent or glabrous; doubly branched, branchlets
bearing to 8 + secund flowers; bracteoles fugaceous.
Flower bud to 5 by 3 mm, ellipsoid. Sepals ovate,

478

subacute, densely pubescent on parts exposed in bud;
2 inner sepals narrower at base, thinner at margins,
than outer 3. Petals lanceolate, densely pubescent on
parts exposed in bud or glabrous, strongly contorted.
Stamens 10, subequal; filaments compressed at base,
tapering and filiform below the narrowly ellipsoid
anthers; appendage to connective c. 24 times length of
anther, the inner 5 reaching to the style apex, slender,
sparsely pubescent in the distal 4. Ovary and stylopo-
dium pyriform, densely pubescent or glabrous,
crowned by a short glabrous trifid style. Fruit pedicel
to 3 mm long, stout. Calyx glabrescent; lobes to 10 by
9mm, subequal, ovate, acute, incrassate, becoming +
reflexed distally. Nut to 3 by 2 cm, broadly obovoid-
ellipsoid, large, apiculate, persistently pale fulvous
sericeous.

Distr. Malesia: Central Borneo (Central Sarawak,
Ulu Mahakam, Ulu Kapuas).

Ecol. Locally common, clay rich soils, often near
streams, in Mixed Dipterocarp forest on hills below
500 m.

Vern. Lun kélabu (Sar.), télingan (Kapuas), téglam
(Mahakam).

54. Shorea angustifolia ASHTON, Gard. Bull. Sing. 19
(1962) 277, pl. 12; Man. Dipt. Brun. (1964) 150, f. 14;
ibid. Suppl. (1968) 81; MevER & Woop, Sabah For.
Rec. 5 (1964) 67. — Fig. 89 C-Cl.

Small to medium-sized tree. Young twig and stipule
sparsely shortly caducous pale brown pubescent; bud,
panicle and petiole persistently so. Twig to 2 mm @
apically, straight, terete, superficially cracked, rugose
and coming away in small papery flakes. Leaf bud 0.5
by | mm, small, ovoid, obtuse. Stipu/e to 3 mm long,
linear, fugaceous. Leaves 8-14 by 2.5-4 cm, deep
violet when opening, ovate to lanceolate, coriaceous;
base cuneate; acumen to 1.5 cm long; nerves 8-10
pairs, hardly raised beneath, curved, well spaced, at c.
50°-60°; midrib beneath hardly elevated; tertiary
nerves densely scalariform; petiole 6-8 mm long,
finely cracked and drying distinct pale brown to
cream-grey. Panicle to 10 cm long, terminal or
axillary, slender, lax, terete; regularly singly or doubly
branched, branchlets to 2 cm long, bearing to 8
distichous flowers; bracteoles small, fugaceous.
Flower bud to 2.5 by 1.5 mm, lanceolate, acute. Calyx
shortly pubescent outside, glabrous within; lobes
ovate, acute, the 2 inner lobes thinner, more con-
stricted at base than 3 outer. Petals cream, linear,
shortly pubescent on parts exposed in bud, imbricate
and cupped at base on opening, twisted and spreading
distally. Stamens 15, in 3 unequal verticils; filaments
broad at base, tapering and filiform distally; anthers
oblong; appendage to connective 1-2 times length of
anther, ciliate towards apex, as long as style. Ovary
ovoid, shortly pubescent except at base; stylopodium
conical; style short, glabrous; style and stylopodium
as long as ovary. Fruit calyx lobes to 5 by 5 mm,
subequal, broadly ovate, incrassate, obtuse. Nut to 8
by 9 mm, obovoid, acute, shortly buff pubescent.

Distr. Malesia: Borneo (Sarawak N.E. of Rejang

FLORA MALESIANA

[ser. I, vol. 9?

valley, S.W. Sabah, Tidung, Upper Dyak in S.
Borneo).

Ecol. Local, on shale ridges at 500-1200 m (rarely
lower).

Vern. Damar hitam bukit (Brun.), séraya kuning
bukit (Sabah).

55. Shorea maxima (KING) Sym. Gard. Bull. S. S. 9
(1938) 330; Mal. For. Rec. 16 (1943) 53, f. 29,
32. — Balanocarpus maximus KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 133; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 110; BRUHL & KiNG, Ann. R. Bot. Gard.
Calc. 5 (1896) 159, t. 192; Ripv. Fl. Mal. Pen. 1 (1922)
248; Burk. J. Mal. Br. R. As. Soc. 1 (1923) 218;
HEyYNgE, Nutt. Pl. ed. 2 (1927) 1128; Foxw. Mal. For.
Rec. 3 (1927) 56; ibid. 10 (1932) 148; Burk. Dict.
(1935) 287. — Fig. 89 A—A2.

Small to medium-sized, often crooked tree. Petiole,
stipule, ovary and parts of petals exposed in bud
persistently buff puberulent, panicle and calyx outside
caducously so. Leaf bud to 3 by 2 mm, ovoid,
subacute. Stipule to 1 cm long, acicular, fugaceous.
Twig c. 3 mm g, pale red-brown, + prominently
ribbed and conspicuously papery flaky. Leaves 8.5-19
by 3.5-7 cm, dark violet when opening, elliptic to
lanceolate or rarely oblanceolate, coriaceous; base
broadly cuneate to obtuse; apex tapering into an up to
1 cm long short broad acumen; nerves 7—10 pairs, +
ascending, slender but distinctly elevated beneath,
obscure above; petiole 1-1.5 cm long, c. 3 mm @,
relatively short and stout. Panicle to 7 cm long, short,
axillary to ramiflorous or occasionally terminal, with
a few short branchlets bearing up to 4 secund flowers.
Flower buds to 15 by 4 mm, large, lanceolate. Sepals
broadly ovate, subequal; outer 3 acute, inner 2 shortly
acuminate. Corolla pale yellow; stamens 10, equal;
filaments long, lorate, slender, tapering; anthers sub-
globose, becoming reflexed; appendages as long as
anthers, relatively short. Ovary narrowly ovoid, taper-
ing into the prominent glabrous style. Fruit sessile;
sepals to 15 by 10 mm, subequal, ovate, acute, united
at the thickened tuberculate base. Nut to 6.5 by 2 cm,
very large, fusiform to narrowly obovoid, apiculate.

Distr. Malesia: Malaya (Perak and Pahang south-
wards).

Ecol. Local, on undulating land and hills to 1300 m.

Vern. Méranti séngkawang puteh, m. bahru, séng-
kawang puteh, m. kérbau, résak, damar katup, séraya.

56. Shores tenuiramulosa ASHTON, Gard. Bull. Sing.
31 (1978) 42.

Small to medium-sized tree. Panicles caducous
greyish puberulent; bracts persistently so, parts of
petals exposed in bud and ovary persistently densely
so. Twig 1-2 mm @ apically, terete, pale greyish
brown, rugulose. Buds and stipules not seen. Leaves
9-24 by 4-11 cm, elliptic to lanceolate, thinly coria-
ceous drying pale greyish brown; margin undulate,
somewhat revolute; base broadly cuneate to obtuse;
apex shortly broadly acuminate; nerves 8—9(—11)
pairs, arched, at 55°-60°, very slender but distinctly

1982]

elevated beneath, slightly so above, as also the laxly
reticulate tertiary nerves; midrib prominent on both
surfaces; petioles 11-20 mm long, drying cream-
brown at the ends, otherwise blackish. Panicles to 18
cm long, terminal or to 3-axillary or ramiflorous,
slender, many flowered; doubly branched, branchlets
to 2 cm long; bracts to 2 mm long, elliptic, fugaceous.
Flower buds to 5 by 2 mm. Sepals ovate-deltoid,
incrassate, subacute, glabrous, subequal. Stamens 15,
in 3 unequal verticils; filaments dilated at base,
tapering and filiform distally; appendages slender,
villous distally, c. 14 times as long as the narrowly
ellipsoid anthers. Ovary ovoid, tapering into the
somewhat shorter stout columnar style; style villous in
the basal 5. Fruit pedicel to 2 by 2 mm; sepals to 6 by 5
mm, equal, ovate, subacuminate, thickened; nut to 25
by 8 mm, fusiform-lanceolate.

Distr. Malesia: N.E. Borneo (E. Sabah, Sakar I.).

Ecol. Locally frequent on dry rocky ultrabasic
ridges near coast.

Note. Clearly allied to S. angustifolia ASHTON; the
rather broad, chartaceous leaf, curling over irregu-
larly at the margin and with the matte undersurface,
and the long and slender, epilose petiole cream
coloured only at the distal end, serve to distinguish
it.

57. Shorea conica SLOoOT. Reinwardtia 3 (1956) 336, f.
I:

Medium-sized tree. Young parts glabrescent; pani-
cles, parts of petals exposed in bud, calyx and ovary
densely persistently pale rufous puberulent. Twigs c. |
mm @ apically, much branched, terete, smooth,
blackish. Buds small, ovoid; stipule to 2 by 1 mm,
ovate, acute, fugaceous. Leaves 6-10 by 2-4.5 cm,
narrowly ovate, coriaceous; margin subrevolute; base
cuneate, subequal; acumen to 2 cm long, slender,
prominent; nerves 4-6 pairs, slender but prominent
beneath, hardly elevated above as also the midrib,
arched, ascending at 45°—5S0°; tertiary nerves densely
subreticulate, barely elevated beneath; petiole 8-12
mm long, slender. Panicles to 8 cm long, terminal or
axillary, slender; singly branched, branchlets to 2 cm
long; bracteoles and flowers unknown. Fruit pedicel to
3 mm long, stout; calyx lobes to 10 by 8 mm, short,
subequal, subacuminate, saccate, thickened,
appressed to the base of the nut. Nut to 32 by 10 mm,
narrowly ovoid-lanceolate, tapering, acute.

Distr. Malesia: E. Sumatra (Labuan Batu, Indra-
girl).

Ecol. Local, undulating land near coast.

Vern. Méranti pugil, m. kunyit, m. témpalo, m.
rambai, samarupa chéngal.

58. Shorea bakoensis ASHTON, Gard. Bull. Sing. 22
(1967) 289, pl. 34; Man. Dipt. Brun. Suppl. (1968) 81,
fe 10!

Small tree. All vegetative parts apparently gla-
brous. Twig c. 2mm @ apically, smooth; stipule scars
short, horizontal. Bud and stipule unknown. Leaves
13-18 by 5-6 cm, narrowly oblong to lanceolate,

DIPTEROCARPACEAE (Ashton)

479

coriaceous; base obtuse; acumen to | cm long, slender;
nerves 9-10 pairs, prominent beneath, at 55°-65°;
tertiary nerves slender, hardly raised beneath, subsca-
lariform; midrib applanate above, prominent
beneath; petiole 10-12 mm long, stout. Flower and
infloresence unknown. Fruit pedicel short, obscure.
Calyx lobes to 4 by 3 mm, subequal, oblong, obtuse,
incrassate, saccate, patent, sparsely persistently buff
sericeous on the outer surface. Nut to 25 by 14 mm,
ellipsoid-cylindric, densely evenly persistently buff
pubescent; style remnant to 1 mm long.

Distr. Malesia: Borneo (W. Sarawak).

Ecol. Rare (one collection); skeletal podsols near
coast.

59. Shorea xanthophylla Sym. Gard. Bull. S. S. 9
(1938) 342, pl. 24; Browne, For. Trees Sarawak &
Brunei (1955) 164; SLoor. Reinwardtia 3 (1956) 344;
ASHTON, Man. Dipt. Brun. (1964) 160, f. 14; ibid.
Suppl. (1968) 91; MEER & Woop, Sabah For. Rec. 5
(1964) 78.

Small tree. Young twig and petiole shortly buff
caducous pubescent; panicle, stipule (outside only),
and leaf bud persistently shortly evenly cream pubes-
cent. Twig 2-3 mm @ apically, straight, stout, terete;
petiole scars large, orbicular, swollen; stipule scars
short, indistinct. Bud to 3 by 1.5 mm, conical. Stipule
c. 8 by 3 mm, fugaceous, narrowly deltoid, cupped,
subacute. Leaves 12-25 by 4-7 cm, bright red at first,
oblong-lanceolate, thinly coriaceous, slightly bullate;
base obtuse or broadly cuneate; acumen to | cm long,
deltoid; margin sometimes slightly revolute; nerves
9-13 pairs, prominent beneath, at 40°—-50°, curving
and continuing parallel with the margin distally,
sometimes anastomosing to form an indistinct intra-
marginal nerve; tertiary nerves remotely scalariform;
petiole 8-15 mm long, c. 1.5mm g, short. Panicle to 10
cm long, terminal or 2-axillary to ramiflorous, angu-
lar; somewhat irregularly doubly branched, branch-
lets to 1.5 cm long, zigzag, bearing to 9 flowers;
bracteoles minute, fugaceous. Flower bud to 4.5 by 1.5
mm, lanceolate, acute. Calyx shortly puberulent
outside, glabrous within; lobes ovate, the inner 2 lobes
thinner, more constricted at the base. Petals cream-
yellow, linear, shortly pubescent on parts exposed in
bud, imbricate and cupped at the base, spreading and
twisted distally. Stamens 15, in 3 unequal verticils;
filaments broad at base, tapering and filiform distally;
anthers oblong; appendage to connective 1—2 times
length of anther, ciliate towards the apex, reaching the
base of the style. Ovary ovoid-conical, glabrous at
base, shortly pubescent apically, tapering into the
stylopodium; style short, glabrous. Fruit calyx lobes c.
7 mm long and broad, subequal, broadly ovate,
thickened, shortly pale fulvous caducous pubescent
outside, glabrous within, not adpressed to the nut but
prominently saccate. Nut c. 2.0 by 1.3 cm, obovoid,
subacute, slightly striated longitudinally, densely
shortly persistently pale fulvous pubescent.

Distr. Malesia: N.E. Borneo (Sabah, Brunei, Sar-
awak N.E. of Rejang valley).

480

Ecol. Mixed Dipterocarp forest below 1000 m, on
clay soil.

Vern. Séraya kuning barun (Sabah), mérabubok
(Iban).

60. Shorea blumutensis Foxw. Mal. For. Rec. 10
(1932) 236, pl. 20; Sym. Mal. For. Rec. 16 (1943) 49, f.
29.

Large tree. Leaf bud, twigs, petioles, midrib below,
stipules, panicles, bracteoles, calyx and parts of petals
exposed in bud densely tawny puberulent, caducous
except on bud, twig apices, petiole, panicle and petals;
ovary and nut densely pale buff puberulent. Leaf bud
to 4 by 2mm, ovoid, subacute. Stipules short, acicular,
fugaceous. Twig c. 2 mm @ apically, smooth, terete.
Leaves 8-17 by 2.8-6.5 cm; elliptic-lanceolate, coria-
ceous, lustrous beneath, dull above; base cuneate;
margin narrowly revolute; acumen to | cm long,
tapering; nerves 14-16 pairs, very slender, slightly
elevated on both surfaces as also the subreticulate
tertiary nerves, ascending at c. 50°; midrib prominent
beneath, evident and elevated above; petiole 1.2—2.2
cm long, c. 2mm @. Panicle to 16 cm long, terminal or
axillary, lax, singly branched; branchlets to | cm long,
short, bearing c. 4 secund flowers. Flower bud to 6 by 3
mm, fusiform; sepals ovate, subequal, the 2 inner
somewhat the narrower; stamens 15, in 3 subequal
verticils; filaments compressed, broad at base, taper-
ing and filiform beneath the subglobose anthers;
appendages slender, scabrous in the distal 3, c. 15
times length of anthers; ovary ovoid, with prominent
narrow stylopodium, both densely puberulent; style
short, obscurely trifid, glabrous. Fruit subsessile; 3
longer calyx lobes to 9 by 1.8 cm, spatulate, obtuse, c.
7 mm broad above the to 8 by 10 mm ovate saccate
thickened tuberculate base; 2 shorter lobes to 6.5 by
0.7 cm, lorate, similar at base. Nut to 3 by 1.3 cm,
obovoid, apiculate.

Distr. Malesia: Malaya (Johore), N.E. Sumatra
(Karimun).

Ecol. Rare, in Lowland Dipterocarp forest below
500 m.

Vern. Méranti kelim.

61. Shorea iliasii ASHTON, Gard. Bull. Sing. 22 (1967)
291, pl. 36; Man. Dipt. Brun. Suppl. (1968) 85, f. 10.

Medium-sized tree. Young parts greyish sericeous,
caducous. Twig c. 2 mm @ apically, terete, sometimes
patchily chartaceous flaky; stipule scars short, hori-
zontal. Bud c. 2 by 2 mm, conical, subacute. Leaf
19-25 by 9-12 cm, oblong-ovate, obtuse but shortly
decurrent and subequal at base; acumen to 8 mm long,
broad, short; nerves 12—14 pairs, prominent beneath,
arched, at 50°—75°; tertiary nerves remotely subscalar-
iform; midrib applanate or slightly elevated above,
prominent beneath; petiole 22-30 mm long, stout,
terete. Panicle to 10 cm long, terminal or axillary,
terete or somewhat compressed, caducous grey-buff
sericeous; singly or doubly branched; bracts and
bracteoles unknown. Buds to 4 by 2 mm, small,
fusiform. Calyx densely puberulent on parts exposed

FLORA MALESIANA

[ser. I, vol. 9?

in bud; 3 outer lobes ovate, acute; 2 inner lobes
somewhat smaller, narrower at base, shortly fim-
briate. Petals cream, linear, obtuse, strongly con-
torted, pubescent on parts exposed in bud. Stamens
15, in 2 unequal verticils, the inner 5 somewhat longer
than the outer 10; filaments broad at base, tapering
and filiform below the subglobose anthers; appendage
to connective c. 15 times length of anther, sericeous in
the apical 4. Ovary narrowly ovoid, pubescent, taper-
ing into the short glabrous style. Fruit calyx sparsely
greyish sericeous towards the base; 3 longer lobes to 8
by 2.2 cm, spatulate, obtuse to subacute, c. 4 mm
broad above the 6 by 5 mm thickened saccate
prominently tuberculate base; 2 shorter lobes to 5.5 by
1.2 cm, otherwise as longer lobes. Nut to 10 by 8 mm,
ovoid, acute, densely persistently greyish sericeous.

Distr. Malesia: Borneo (Central Sarawak).

Ecol. Local, clay soils on undulating land and hills
below 400 m.

Vern. Lun siput daun bésar.

62. Shorea faguetioides ASHTON, Gard. Bull. Sing. 19
(1962) 287, pl. 17; Man. Dipt. Brun. (1964) 154, f. 14;
ibid. Suppl. (1968) 83; MEWER & Woop, Sabah For.
Rec. 5 (1964) 80.

Large buttressed tree. Young twig sometimes
shortly buff caducous pubescent; other vegetative
parts entirely glabrous. Twig 3 mm @ apically, slightly
compressed at first, becoming terete, minutely striated
or rugulose. Bud 3-4 by 2.5 mm, ovoid, obtuse,
compressed. Stipule to 18 by 6 mm, narrowly deltoid,
saccate, obtuse, less caducous than in most species of
sect. Richetioides. Leaves 12-18 by 5-7 cm, brilliant
magenta when opening, narrowly ovate, chartaceous;
base broadly cuneate to subcordate, subequal;
acumen prominent, c. 2 cm long; nerves 10-15 pairs,
slender but prominent, at 60°—70°; tertiary nerves
scalariform, slightly oblique to the nerves; petioles
2.2-3 cm long, very long, slender. Panicle to 15 cm
long, terminal or axillary, terete, rugose when dry, lax,
sparsely persistently pale greyish brown pubescent;
regularly alternately doubly, rarely trebly, branched,
branchlets to 2 cm long, bearing to 7 flowers;
bracteoles small, fugaceous. Flower bud to 2.5 by 1.5
mm, small, elliptic, obtuse. Ca/yx densely shortly
pubescent outside, glabrous within; lobes ovate,
acute, the 2 inner slightly the smaller, thinner, more
constricted at base. Petals cream, linear, pubescent on
parts exposed in bud, imbricate in the basal half
forming a broad shallow cup, distally spreading and
twisted. Stamens 15, in 3 unequal verticils; filaments
slender, tapering, basally compressed and slightly
gibbous, distally filiform; anthers subglobose; appen-
dage to connective slender, 3 times length of anther,
the longest 5 exceeding the style apex, slightly ciliate
towards the apex. Ovary ovoid, sparsely pubescent;
stylopodium shorter than ovary, cylindrical, more
densely pubescent; style short, glabrous. Fruit calyx
puberulent at the base, otherwise glabrescent; 3 longer
lobes to 7 by 1.5 cm, chartaceous, broadly spatulate,
obtuse, tapering to 4 mm above the to 5 mm broad

1982]

prominently tuberculate saccate thickened base; 3
shorter lobes to 5 by 0.7 cm, spatulate, similar at base.
Nut c. 20 by 6 mm, ellipsoid to obovoid, pale buff
pubescent; style remnant c. 3 mm long, slender.

Distr. Malesia: Northern Borneo (Ulu Kapuas
and Sarawak to S.W. Sabah and W. Kutel).

Ecol. Local, on clay rich soils on hillsides below 700
m.
Vern. Damar hitam daun nipis (Brun.), bépisang
(Kapuas), pénuar (W. Kutei).

Note. One collection (bb 35287, Ng. Njaban, Ulu
Kapuas, W. Borneo) possesses fruit with short sub-
equal sepals; without further collections it is not
possible to assess whether it represents more than a
single aberrant tree.

63. Shorea alutacea ASHTON, Gard. Bull. Sing. 22
(1967) 288, pl. 33; Man. Dipt. Brun. Suppl. (1968) 81,
f. 10.

Medium-sized tree. Twigs, stipules. leaf buds,
petioles and midrib shortly densely persistently buff
pubescent, leaf nervation and surface beneath spar-
sely so. Twigs c.2 mm @ near the apices, terete; stipule
scars c. 1.5 mm long, horizontal, evident. Bud c. 2 by
1.5 mm, ovoid, acute, small. Stipule to 8 by 3 mm,
narrowly deltoid, acute, relatively large. Leaves 15—22
by 6-8 cm, ovate to lanceolate, undulate, chartaceous,
with cordate base; acumen to | cm long, narrow;
nerves 14-19 pairs, slender, narrowly depressed
above, slender and prominent beneath, at up to 100°
at base, 60°—-70° distally; tertiary nerves subscalari-
form, slender, indistinct; midrib depressed above,
prominent, terete beneath; petiole 6-10 mm long,
drying rugose. Panicle to 15 cm long, terete, straight,
densely shortly pale buff pubescent; singly or doubly
branched, branchlets to 2.5 cm long; bracteoles to 4 by
3 mm, ovate, acute, densely shortly buff fugaceous
pubescent. Flowers secund; buds to 6 by 3 mm,
ellipsoid, relatively large. Calyx densely pubescent on
parts exposed in bud; sepals broadly ovate, subequal,
the inner 2 shortly acuminate, thinner towards the
margin and narrower at the base than the acute outer
3. Petals lanceolate, pale yellow, pubescent on parts
exposed in bud; imbricate at base after opening
forming a cup; spreading and twisted distally.
Stamens 15, in 3 unequal verticils the appendages of
the inner 5 exceeding the style apex; filaments broad
and compressed at base, tapering and filiform distally;
anthers broadly oblong; appendage to connective
slender, 3-4 times as long as anther, glabrous. Ovary
and stylopodium pyriform, densely pubescent, taper-
ing into a glabrous columnar style equal in length to
the stylopodium. Fruit pedicel and calyx shortly
sparsely caducous pubescent. Pedicel c. | mm long,
short. 3 longer calyx lobes to 8 by 1.8 cm, spatulate, c.
4 mm broad above the to 7 by 7 mm ovate saccate
thickened base; 2 shorter lobes to 6 by 1.2 cm,
otherwise similar to longer lobes. Nut to 2.2 by 1 cm,
narrowly ovoid, shortly apiculate, densely shortly
evenly buff pubescent.

Distr. Malesia: Borneo (W. Sarawak).

DIPTEROCARPACEAE (Ashton)

48]

Ecol. Rare, lower slopes of granodiorite moun-
tains.

64. Shorea longisperma Roxs. (Hort. Beng. (1814) 93,
nomen] FI. Ind. ed. Carey 2 (1832) 618; DC. Prod. 16, 2
(1868) 632; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 103;
RIpDL. Fl. Mal. Pen. | (1922) 143; AsHton, Gard. Bull.
Sing. 31 (1978) 43. — Parashorea longisperma KurRzZ,
J. R. As. Soc. Beng. Sc. 39, 2 (1870) 66; Dyer, FI. Br.
Ind. 1 (1874) 441; SLoor. Bull. Jard. Bot. Btzg III, 8
(1927) 571; Foxw. Mal. For. Rec. 10 (1932) 226. — S.
resina-negra FOxW. Mal. For. Rec. 10 (1932) 205, pl.
16; BurK. Dict. (1935) 2021; Sym. Mal. For. Rec. 16
(1943) 56, f. 29, 35; ASHTON, Man. Dipt. Brun. Suppl.
(1968) 89, f. 11, pl. 18 (habit, bole, damar).

Very large, prominently buttressed tree. Leaf bud
and stipule persistently unevenly shortly fulvous
pubescent, twig caducously so. Leaf beneath and
petiole persistently shortly subscabrid pale grey-green
pubescent. Twig 1-2 mm @ apically, slender, terete,
becoming smooth: stipule scar small, horizontal to
descending, frequently obscure. Bud to 3 by 2 mm,
ovoid, acute. Stipule to 5 by 2 mm, ovate to lorate,
acute, caducous. Leaves 7—12 by 2.3-6 cm, elliptic to
ovate, somewhat chartaceous, pale mauve when fresh;
base cuneate to obtuse, acumen to 1.5 cm, long,
slender; nerves 10—13 pairs, at 45°—55°, slender, raised
beneath; tertiary nerves subscalariform; midrib
applanate to slightly depressed above, prominent
beneath; petiole 10-15 mm long. Panicie to 7 cm long,
terete or ribbed, slender, densely persistently pale
fulvous scabrid pubescent; shortly singly branched,
branchlets ascending, bearing to 4 flowers. Buds to 9
by 3 mm, lanceolate. Sepals broadly ovate, acute,
densely sericeous on parts exposed in bud, the inner 2
fimbriate, somewhat shorter than the outer 3. Petals
pale yellow, narrowly lanceolate, strongly contorted,
sericeous on parts exposed in bud. Stamens 15, in 3
unequal verticils; filaments broad and compressed at
base, tapering to the large broadly ellipsoid anthers;
appendage to connective slender, c. 2 times length of
anther at anthesis. Ovary and stylopodium pyriform,
shortly densely pubescent; style slender, columnar,
glabrous, as long as ovary and stylopodium. Fruit
sessile. Calyx sparsely pubescent; 3 longer lobes to 9
by 1.5 cm, spatulate, obtuse, c. 3 mm broad above the
to 6 by 4 mm elliptic saccate tuberculate thickened
base; 2 shorter lobes to 7 by 0.4 cm, linear, obtuse,
similar at base. Nut to 2.3 by 1.2 cm, ellipsoid, acute,
shortly evenly densely buff pubescent.

Distr. Malesia: Malaya (except seasonal areas), E.
Sumatra (Labuhan Batu), Borneo (Central and N.
Sarawak, S.E. and S. Borneo).

Ecol. Scattered on fertile clay rich soils in Mixed
Dipterocarp forest, especially on igneous and volcanic
rocks on undulating land and sheltered inland moun-
tain slopes to 1400 m.

Vern. Méranti damar hitam, kala, katup, mérawan,
sénggai (Mal.), lun méranti (Sar.), mérsiput, merak-
unyit, kérambukuh, mémukuh, awang sih (S.E. Bor-
neo).

482

65. Shorea acuminatissima Sym. Gard. Bull. S. S. 9
(1938) 340, pl. 23; BROWNE, For. Trees Sarawak &
Brunei (1955) 162; ASHTON, Man. Dipt. Brun. (1964)
149, f. 14; ibid. Suppl. (1968) 81; MEvER & Woop,
Sabah For. Rec. 5 (1964) 65, pl. 2 (habit), f. 8.

Very tall buttressed tree with dark brown square-
section fissured bark. Twig, panicle, petiole, bud,
stipule and nervation beneath + persistently scabrid
rust pubescent. Twig c. 1.5 mm @g apically, slender,
much branched, terete, becoming smooth. Bud to 1.2
by | mm, globose to ellipsoid, small. Stipule c. 6 by 3.5
mm, oblong-falcate, base obtuse or subcordate; cadu-
cous, more persistent in young trees. Leaves 7-10 by
3-4 cm, ovate, sometimes cream to brown lepidote
beneath; base obtuse; acumen c. 7 mm long; short,
acute; margin revolute; nerves 9-12 pairs, prominent
beneath, at c. 40°—-50°; tertiary nerves dense, scalari-
form, distinct; midrib prominent beneath, depressed
above; petiole 1-1.5 cm long, slender. Panicle to 8 cm
long, terminal or axillary, terete, singly branched;
branchlets to 2 cm long, bearing to 6 secund flowers;
bracteoles to 2 by 2 mm, broadly oblong, obtuse,
puberulent. Flower bud to 5 by 3 mm, ellipsoid,
subacute. Calyx densely pubescent outside, glabrous
within; lobes broadly ovate, subequal, obtuse; inner 2
lobes thinner towards margin, narrower at base, than
outer 3. Petals shortly pubescent on parts exposed in
bud, imbricate at base after opening forming a cup,
spreading and contorted distally. Stamens 15, of 2
lengths the longest 5 exceeding the ovary; filaments
broad, compressed, tapering, somewhat gibbous;
anthers broadly oblong; appendage to connective
longer than anthers, slender, puberulent towards
apex. Ovary and stylopodium narrowly ovoid, with a
band of short hairs towards the apex; style as long as
ovary and stylopodium, slender, glabrous. Fruit sub-
sessile, with prominent obconical receptacle; calyx
puberulent or glabrescent; 3 longer lobes to 6 by 1.3
cm, spatulate, obtuse, to 3 mm broad above the to 4 by
4mm somewhat tuberculate thickened base; 2 shorter
lobes to 4.5 by 0.8 cm, otherwise similar. Nut to 2 by
0.7 cm, narrowly ovoid, densely pale rufous pubes-
cent.

Distr. Malesia: N.E. Borneo (from Baram valley
N.E. to Sabah).

Ecol. Local, Mixed Dipterocarp forest on sandy
clay soils on hills below 500 m, usually near the coast.

Vern. Séraya kuning runching (Sabah), damar
hitam runching (Brun.), barun runching (Sar.).

66. Shorea gibbosa BRANDIS, J. Linn. Soc. Bot. 31
(1895) 99; Rip. J. Str. Br. R. As. Soc. 54 (1909) 23;
Burk. ibid. 76 (1917) 165, fig.; RipL. Fl. Mal. Pen. 1
(1922) 289; Foxw. Mal. For. Rec. 10 (1932) 208; Sym.
Gard. Bull. S.S. 7 (1933) 143, pl. 42; Mal. For. Rec. 16
(1943) 51. f. 29; BRowNneE, For. Trees Sarawak &
Brunei (1955) 162; MEER & Woop, Sabah For. Rec.
5 (1964) 71, pl. 4a, f. 9; ASHTON, Man. Dipt. Brun.
Suppl. (1968) 83, f. 10. — Hopea grisea BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 63; MerR. En. Born. (1921)
404.

FLORA MALESIANA

[ser. I, vol. 92

Very tall, buttressed tree. Twig, bud, petiole and
stipule persistently shortly evenly pale rufous pubes-
cent, leaf nervation beneath sparsely so, glabrescent.
Twig c. 1 mm @ apically, slender, much branched, at
first frequently rugulose. Stipules to 3 by | mm,
narrowly elliptic, acute, fugaceous. Leaves pink when
opening, 5-13 by 2-6 cm, ovate, chartaceous, undu-
late; base broadly cuneate to obtuse; acumen to 1.5cm
long; nerves 7-9 pairs, slender but prominent beneath,
at 50°—65°; tertiary nerves subreticulate; midrib some-
what depressed above, prominent beneath; petiole
8-12(-16) mm long, short. Panicle to 10 cm long,
terminal or axillary, terete, densely persistently buff to
pale rufous pubescent; singly branched, branchlets to
2 cm long, bearing to 6 + distichous flowers;
bracteoles to 2 by 1 mm, oblong, obtuse, shortly
pubescent, fugaceous. Bud to 5 by 3 mm, lanceolate.
Sepals ovate, shortly densely pubescent on parts
exposed in bud; outer 3 acute, inner 2 shortly
acuminate, relatively broader, shorter, thinner at the
margin. Petals cream with a pink base, narrowly
oblong, strongly contorted, sericeous on parts
exposed in bud. Stamens 15, in 3 unequal verticils, the
longest reaching the style apex; filaments compressed
and gibbous at base, tapering and filiform below the
small broadly ellipsoid anthers; appendage to connec-
tive somewhat longer than anther, slender, glabrous.
Ovary ovoid, densely pubescent; style columnar,
pubescent in the basal 5, otherwise glabrous. Fruit
pedicel to 2 mm long, slender. Calyx shortly sparsely
pubescent; 3 longer lobes to 9 by 2 cm, spatulate,
obtuse, c. 4mm broad above the to 10 by 6 mm ovate
saccate thickened tuberculate base; 2 shorter lobes to
6 by 0.5 cm, narrowly spatulate, acute, similar at base.
Nut to 1.8 by 1.2 cm, narrowly ellipsoid, densely buff
pubescent, acute.

Distr. Malesia: Malaya (Johore), Singapore, S.E.
Sumatra (Palembang, Lampong), Borneo.

Ecol. Locally common on deep fertile clay-rich
soils, on undulating land and low hills below 650 m.

Vern. Damar hitam gajah (Mal.), lun gajah (Sar.),
séraya kuning gajah (Sab.), madilan, mérakunyit,
damar ténkuyung, d. kétuyung, awang pakit, kelepeh,
kénuar, méranti kuning, dahu mentola, bangkirai,
lampong kuning (Indon. Borneo), damar buah, d. b.
kuning (Sumatra).

Note. A variable species distinguished by the
sparsely pubescent leaf nervation and leaves crinkling
on drying. Collections from East Borneo typically
bear lanceolate leaves drying pale grey-green beneath,
and a denser fulyous tomentum on perianth and
panicle.

67. Shorea hopeifolia (HEIM) Sym. Gard. Bull. S. S. 8
(1933) 150, pl. 46; ibid. 8 (1934) 36; Mal. For. Rec. 16
(1943) 52, f. 29; BROwNE, For. Trees Sarawak &
Brunei (1955) 163; MEIER & Woop, Sabah For. Rec.
5 (1964) 72; ASHTON, Man. Dipt. Brun. Suppl. (1968)
84, f. 10; Gard. Bull. Sing. 31 (1978) 43. — Cotylelo-
bium hopeifolium HEM, Bull. Mens. Soc. Linn. Paris 1
(1891) 971; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 95;

1982]

BRUHL & KING, Ann. R. Bot. Gard. Calc. 5, 2 (1896)
153. — S. ridleyana KiNG, J. R. As. Soc. Beng. Sc. 62,
2 (1893) 115, p.p.; Rip. Fl. Mal. Pen. 1 (1922) 226,
p.p.; Foxw. Mal. For. Rec. 2 (1927) 353; ibid. 10
(1932) 209, p.p.; SLooT. ex Merr. Pl. Elm. Born.
(1929) 204. — Hopea heimiana BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 63; MerR. En. Born. (1921) 402. — Ho-
pea albescens Riot. J. Str. Br. R. As. Soc. 73 (1916)
142; Fl. Mal. Pen. 1 (1922) 236. — S. kalunti MERR.
Philip. J. Sc. 26 (1925) 475; Descu, Mal. For. Rec. 12
(1936) 35, 39; Foxw. Philip. J. Sc. 67 (1938)
307. — Hopea hopeifolia SLoot. Bull. Jard. Bot. Btzg
IIT, 10 (1929) 396.

Very tall buttressed tree. Vegetative parts glabrous.
Twigs c. 1 mm @ apically, terete, slender, much
branched, minutely rugulose; stipule scars short,
horizontal, obscure. Bud c. 1 by 1 mm, small, ovoid,
acute, glabrous. Stipule to 5 by 2 mm, lanceolate,
acute, fugaceous. Leaves 3.5—8 by 24cm, small, ovate,
somewhat chartaceous and undulate; base broadly
cuneate, usually with prominent pore-like glabrous
domatia on either side of the base of the midrib;
acumen to | cm long, prominent; nerves 9-11 pairs,
slender, hardly raised beneath, arched, at 55°-65°,
with distinct secondary nerves; tertiary nerves reticu-
late; midrib applanate above, prominent beneath,
drying dark red or black; petiole 8-10 mm long,
slender, geniculate. Panicle to 5 cm long, terminal or
axillary, terete, densely persistently pale buff pubes-
cent; singly branched, branchlets bearing to 9 disti-
chous flowers; bracteoles to 2 by 2 mn, elliptic,
obtuse, puberulent outside, glabrous within, cadu-
cous. Bud to 5 by 2 mm, small, lanceolate. Sepals
pubescent on parts exposed in bud, ovate, acute; the
inner 2 relatively broader and thinner at the margins
than the outer 3. Petals pale yellow, narrowly lanceo-
late, densely pubescent on parts exposed in bud,
strongly contorted. Stamens 15, in 3 unequal verticils;
filaments compressed at base, tapering and filiform
below the ellipsoid anthers; appendage to connective
slightly shorter than anther, glabrous. Ovary ovoid,
pubescent, surmounted by a columnar tapering style
equal in length and pubescent in the basal 5. Fruit
pedicel and base of calyx sparsely buff pubescent,
calyx elsewhere glabrescent. Pedicel c. 3 mm long. 3
longer calyx lobes to 7 by 1.5 cm, spatulate, obtuse, c.
4mm broad above the to 5 by 5 mm deltoid thickened
saccate tuberculate base; 2 shorter lobes to 4 by 0.7
cm, spatulate, acute, similar at base. Nut to 2.7 by 1.2
cm, ellipsoid, shortly apiculate, densely shortly evenly
buff pubescent.

Distr. Malesia: Malaya (seasonal area excepted),
Sumatra (Palembang, Lampong, and E. Coast Res.:
Tapanuli to Pariaman in west), Borneo, Philippines
(Mindanao).

Ecol. Scattered on fertile clay rich soil on undulat-
ing land and hills below 600 m, often in moist places.

Vern. Damar siput jantan, séraya, s. labu (Mal.),
suranti limau manis, aruing, méranti kuning, damar
buah (Sumatra), séraya kuning jantan (Sabah), Jun
siput jantan (Sar.), tali makas (Dus.), sélangan kacha,

DIPTEROCARPACEAE (Ashton)

483

merakunyit, bérémbuku, lampong bémbéring, damar
b., utuhatup (Indon. Borneo).

Note. Some specimens from S.E. Sabah and N.E.
Kalimantan suggest that hybridization with S. gib-
bosa may sometimes occur there.

68. Shorea kudatensis Woop ex MElWER, Act. Bot.
Neerl. 12 (1963) 346, pl. 12; Sabah For. Rec. 5 (1964)
74, pl. 5 (bark).

Medium-sized buttressed tree. Panicle, parts of
petals exposed in bud and ovary persistently cream
puberulent; calyx at first densely so, becoming sparse;
twig caducously so. Twigs c. 3 mm @, rather stout,
terete, + rugulose. Bud small, ovoid, obtuse; stipules
fugaceous, not seen. Leaves 8-15 by 5—9 cm, ovate,
coriaceous, lustrous above; base + obtuse, subequal,
shortly decurrent; apex shortly broadly acuminate;
nerves 7—9 pairs, very slender but distinctly elevated
on both surfaces as also the midrib, strongly arched, at
45°-55°; tertiary nerves subreticulate, distinctly ele-
vated beneath, evident above; petiole 17-22 mm long,
rather stout. Panicle to 14 cm long, terminal or
axillary, many-flowered; doubly branched, branchlets
to 3 cm long. Flower buds c. 3 by 2 mm, small,
ellipsoid. Sepals broadly ovoid, the outer 3 subacu-
minate, the inner 2 acute. Petals cream. Stamens 15,
unequal; filaments broadly compressed at base, taper-
ing and filiform beneath the oblong anthers; appen-
dages slender but short, + equal in length to anthers,
scarious towards apices. Ovary ovoid, tapering into
the short glabrous style. Fruit subsessile; 2 longer
calyx lobes to 6 by | cm, spatulate, obtuse, c. 3 mm
broad above the c. 4 by 3 mm narrowly ovate
thickened saccate tuberculate base; 2 shorter lobes to
43 by 5mm, subacute, otherwise similar. Nut to 20 by
8 mm, narrowly ovoid.

Distr. Malesia: N.E. Borneo (Kudat to Kilias and
Sandakan).

Ecol. Locally common on dry hills near coast.

Vern. Séraya kuning kudat.

69. Shorea cuspidata ASHTON, Gard. Bull. Sing. 22
(1967) 290, pl. 35, 350 (phot. habit); Man. Dipt. Brun.
Suppl. (1968) 82, f. 10, pl. 14 (stem-base).
Medium-sized tree. Leaf bud and stipule shortly
buff pubescent, young twigs caducously so; leaf
glabrous. Twig c. | mm g@ towards apex, slender,
terete, minutely striated. Bud c. 1 mm long, small,
ovoid. Leaves 5—9 by 2-6 cm, small, broadly ovate,
subcoriaceous, with broadly cuneate base; acumen to
1.5 cm long, subcaudate; nerves 5—7 pairs, slender,
hardly raised beneath, at 40°-S0°; tertiary nerves
reticulate, obscure; midrib slender, applanate or
slightly raised above, prominent beneath; petiole 7-11
mm long, slender. Panicle to 9 cm long, terminal or
axillary, slender, terete, densely evenly shortly buff
pubescent; singly branched, branchlets to 1.5 cm long.
Flowers secund; buds to 3 by 2 mm, small, ellipsoid.
Calyx pubescent on parts exposed in bud; sepals
broadly ovate, small, subequal, acute, the inner 2
broader and thinner at the margin than the outer 3.

484

Petals pale lime-yellow, lanceolate, densely pubescent
on parts exposed in bud, connate at base, strongly
contorted. Stamens 15, in 3 unequal verticils; fila-
ments compressed at base, tapering and filiform below
the ellipsoid anther; appendage to connective c. 15
times length of anther, slender, pubescent in the distal
4, almost reaching to style apex at anthesis. Ovary and
stylopodium pyriform, sericeous, crowned with a short
columnar glabrous style. Fruit pedicel and calyx
shortly sparsely buff pubescent. Pedicel c. 1 mm long,
short. 3 longer calyx lobes to 5 by 1.5 cm, broadly
spatulate, obtuse, c. 3 mm broad above the to 5 mm
broad saccate thickened base; 2 shorter lobes to 4 by 1
cm, otherwise as in longer lobes. Nut to 2.5 by 1.5 cm,
obovoid, mucronate, densely shortly evenly buff
pubescent.

Distr. Malesia: Borneo (Sarawak w. of the Lupar).

Ecol. Locally common on undulating land and low
hills in Mixed Dipterocarp forest, to 500 m.

Vern. Lun runching padi.

70. Shorea mujongensis ASHTON, Gard. Bull. Sing. 22
(1967) 292, pl. 38, 351 (phot. habit); Man. Dipt. Brun.
Suppl. (1968) 87, f. 11, pl. 17 (stem-base).

Tall buttressed tree. Young twig, bud and petiole
shortly buff pubescent, glabrescent. Twig c. 2mm @
apically, terete, rugulose; stipule unknown. Leaves
6-14 by 2.5-5.5 cm, ovate or elliptic, margin +
revolute, base obtuse to cuneate; acumen to 8 mm
long; nerves 8-13 pairs, slender but prominent
beneath, curved, at c. 50°; tertiary nerves scalariform,
unraised, narrowly channelled, beneath; midrib
applanate above, prominently terete beneath; petiole
10-16 mm long, terete. Flowers unknown. Panicle to 6
cm long, singly branched, terete, rugulose, shortly
evenly buff puberulent. Flower bud to 6 by 4 mm,
fusiform. Perianth puberulent on parts exposed in
bud; sepals ovate, subacute. Stamens 15, in 3 unequal
verticils; filaments broad at base, tapering; anthers
ellipsoid; appendages same length as anthers, slender,
glabrous. Ovary and stylopodium pyriform, puberu-
lent except at apex, surmounted by a short glabrous
style. Fruit pedicel to 2 mm long, puberulent. Fruit
calyx sparsely puberulent to glabrescent; 3 longer
lobes to 7 by 1.5 cm, spatulate, obtuse, 4 mm wide
above the 6 by 5 mm tuberculate incrassate base; 2
shorter lobes to 4.5 by 0.7 cm, otherwise similar. Nut
to 2.2 by 0.7 cm, narrowly ovoid, densely shortly pale
buff pubescent.

Distr. Malesia: Borneo (Central Sarawak, E. Sabah).

Ecol. Local, on fertile soils overlying or influenced
by basic volcanic rocks, to 1100 m.

Note. Close to S. gibbosa but the minutely chan-
nelled, hardly or unraised tertiary nerves beneath
(sapling excluded), and the glabrous subcoriaceous
applanate leaves serve to distinguish it. Collections
from Sarawak have larger more prominently revolute
ovate leaves.

71. Shorea faguetiana Hem, Bull. Mens. Soc. Linn.
Paris 2 (1891) 975; BrRANpbiIs, J. Linn. Soc. Bot. 31

FLORA MALESIANA

[ser. I, vol. 9?

(1895) 95, p.p.; MERR. En. Born. (1921) 414; Sym.
Gard Bull. S.S. 7 (1933) 148, pl. 45; ibid. 10 (1939) 381;
Mal. For. Rec. 16 (1943) 50, f. 29, 31; BROWNE, For.
Trees Sarawak & Brunei (1955) 162; ASHTON, Man.
Dipt. Brun. (1964) 152, f. 14; ibid. Suppl. (1968) 83;
Meer & Woop, Sabah For. Rec. 5 (1964) 69, pl. 3,
4b. — S. ridleyana KING, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 115, p.p.; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
99, p.p.; BRUHL & KING, Ann. R. Bot. Gard, Calc. 5, 2
(1896) 153, t. 185, p.p.; RiDL. Fl. Mal. Pen. 1 (1922)
226, p.p.; Foxw. Mal. For. Rec. 3 (1927) 38, p.p.; ibid.
10 (1932) 209, p.p.; BURK. Dict. (1935) 2021. —S.
dryobalanoides DYER ex BRANDIS, J. Linn. Soc. Bot. 31
(1895) 95, nomen in syn.

Medium-sized or large buttressed tree. Bud,
stipule outside, panicle and young twig pale greyish
pubescent; fresh young leaves sparsely fugaceous
pubescent beneath. Twig slender, dotted with large
pale lenticels, with a minutely wrinkled striated
papery flaked surface. Bud c. 2 by 1.5 mm, small,
ovoid to conical. Stipule to 4 by 1.5 cm, hastate,
pubescent. Leaves 7-12 by 3-5 cm, elliptic to oblong-
lanceolate or ovate, coriaceous; base obtuse or
cuneate, symmetrical or subequal; acumen to 1 cm
long; margin generally slightly revolute; nerves 9-12
pairs, slender but distinctly elevated beneath, at c.
30°-50°; tertiary nerves scalariform, rather dense,
oblique to the nerves; petiole 1-1.5 cm long, fairly
stout. Panicle to 20 cm long, terminal or axillary,
terete or angular, lax; regularly doubly or trebly
branched, branchlets to 1 cm long, bearing to 8
flowers; bracteoles small, fugaceous. Flower bud to 3
mm long, small, elliptic to subglobose. Calyx shortly
pubescent outside, glabrous within; lobes ovate, sub-
equal, subacute, 2 inner lobes thinner towards margin,
more constricted basally. Petals cream, linear, the
basal half imbricate, forming a cup on opening,
spreading and strongly twisted distally. Stamens 15, in
3 unequal verticils; filaments broad at base, tapering
rapidly and filiform distally; anthers subglobose;
appendage to connective as long as anthers and
reaching almost to the style apex, sparsely ciliate
towards the apex. Ovary subglobose, shortly pubes-
cent; stylopodium as long as ovary, pubescent; style
short, glabrous. Fruit sparsely pubescent; 3 longer
calyx lobes to 6 by 1.2 cm, thinly coriaceous, spatulate,
obtuse, 2.5 mm broad above the c. 3.5 mm broad
thickened tuberculate base; 2 shorter lobes c. 4.5 cm
long, narrower, with a narrower base. Nut to 15 by 5
mm, narrowly ellipsoid to subobovoid, shortly buff
pubescent; style remnant short, acute.

Distr. Thailand (Pattani) and in Malesia: Malaya
(non-seasonal parts), Borneo (Sarawak, Sabah, S.E.
Borneo to Muara Tewe).

Ecol. Well drained clay soils on low hills and
particularly ridge tops at 150-1000 m (mainly to 700m).

Vern. Damar siput, kala, k. jantan, séraya, s. kitan,
bam, rinchong, (Mal.), lun siput (Sar.), séraya kuning
siput (Sabah), katukan (Kinababangan), selangan
kacha, sérangan kacha, pangkang puteh (Tidung,
Berau).

1982] DIPTEROCARPACEAE (Ashton) 485

5. Section Anthoshorea

Heim, Rech. Dipt. (1892) 41; BRANDis, J. Linn. Soc. Bot. (1895) 84, p.p.; SYM.
Mal. For. Rec. 16 (1943) 27, f. 17 (map); AsHTon, Gard. Bull. Sing. 20 (1963) 268;
Man. Dipt. Brun. (1964) 116. — Shorea sect. Hopeoides HE, Rech. Dipt. (1892)
43. — Anthoshorea PIERRE ex HEIM, l.c. in syn.— Parahopea Heim, L.c. 66.
— Shorea Meranti Pa‘ang group Sy. Mal. For. Rec. 16 (1943) 27. — Shoreasubg.
Anthoshorea (HEIM) MEER, Act. Bot. Neerl. 12 (1963) 322. — Fig. 91B-C, 92.

Flowers usually large. Petals white, sometimes with a median pink suffusion,
broadly elliptic or ovate-lanceolate, contorted imbricate at base at anthesis
forming a rather large broad goblet enclosing the anthers, loosely connate on
falling. Stamens 15-30 in 3 verticils, or ©; filaments broad at base, gradually
tapering; anthers with 4 pollen sacs, narrowly oblong to linear; appendage to
connective unreflexed, prominent, usually at least half as long as anther, stout or
slender, scabrous or glabrous. Ovary pubescent or glabrous, without distinct
stylopodium; style longer than ovary, + distinctly trifid distally. Stipules
caducous, often large; bracts and bracteoles frequently large, somewhat persis-
tent. Leaf with scalariform tertiary nerves; midrib obscure, depressed, above.
Medium sized or more usually large stoutly buttressed trees. Bark surface with
irregular section fissures, frequently short and anastomosing; inner edge of outer
bark ill-defined; outer surface rotting off, rarely flaking regularly; periderms
undulate, incomplete or absent; inner bark simply laminate. Wood cream or
yellow, rarely red (S. montigena) with ray cells bearing silica deposits; vessels
arranged in a reticulum as seen in transverse section, solitary or occasionally in

pairs.

Distr. Ceylon and Peninsular India to Indochina and through Malesia to the Moluccas.

Ecol. Semi-evergreen and evergreen lowland forests. Several species (S. roxburghii, S. hypochra, S. polita, S.
assamica, S. retinodes) can occur in gregarious stands, especially in seasonal forests.

Vern. White méranti, méranti pa’ang (Mal.), raruk (Iban), mélapi (Sabah).

Note. S. bracteolata appears to be pollinated by thrips (APPANAH) whereas S. roxburghii has been found to be
visited by bees in Thailand (SMITINAND, pers. comm.). Most species are rather uniform and well defined, the
notable exception being the widespread S. assamica and its siblings S. agamii of Borneo, which has two
subspecies, one of which can be triploid, and S. resinosa in which abundant apoximis through adventive
polyembryony is well known (FOXWworRTHY, KAUwR).

72. Shorea dealbata Foxw. Mal. For. Rec. 10 (1932)
192, pl. 14; BurRK. Dict. (1935) 2010; Sym. Mal. For.
Rec. 16 (1943) 35, f. 19; ASHTON, Man. Dipt. Brun.
Suppl. (1968) 93, f. 12.—S. aff. hypochra (non
HANCE) BROWNE, For. Trees Sarawak & Brunei
(1955) 519.

Medium-sized, hardly buttressed tree. Young
twigs, buds, stipules outside and petiole densely
shortly evenly dark fulvous pubescent, nervation
beneath fugaceously so; leaf beneath pale pink-brown
lepidote. Twig. c. 4 by 2 mm @ towards apex,
compressed, smooth to finely rugulose; stipule scars c.
2 mm long on young twigs, subhorizontal, prominent
with a prominent rib descending the twig from their
ends. Bud to 6 by 4 mm, ovoid, conical, subacute.
Stipule to 20 by 5 mm, elliptic-oblong, subacute.
Leaves alternate, 8-16 by 4~7.5 cm, ovate to elliptic,

coriaceous; base obtuse, rarely cuneate; acumen to 1
cm long, narrow; nerves (11—)20—24 pairs, obscure
and slightly depressed above, slender and hardly
elevated beneath, at up to 75° towards the base,
40°-60° towards the apex; tertiary nerves dense,
parallel, slender, obscure; midrib obscure, depressed
above, prominent, subacute beneath; petiole 1.5—2.5
cm long, stout, rugose especially distally when dry.
Panicle to 8 cm long, terete or angular, densely pale
fulvous hirsute, axillary or terminal; unbranched or
singly branched, branchlets to 1.5 cm long; bracteoles
fugaceous, unknown. Flowers distichous; bud to 12 by
5 mm, lanceolate. Calyx densely pale fulvous hirsute
outside, glabrous within; 3 outer lobes narrowly
deltoid, subacute, 2 inner lobes shorter, ovate, cuspi-
date. Petals oblong-lanceolate, white tinged with pink
at base, densely pubescent on parts exposed in bud.

486 FLORA MALESIANA [ser. I, vol. 9?

B3

Fig. 91. Flower details in Shorea sect. Anthoshorea HEIM (B and C) and sect. Brachypterae HEM subsect.

Brachypterae (D). Sepals drawn from inside. — S. roxburghii G. Don. B. Bud, B/. outer sepal, B2. inner sepal,

B3. stamens from inside, B4. pistil, all x 10. — S. henryana Pierre. C. Stamens and pistil, C/. outer sepal, C2.

inner sepal, C3. one stamen in lateral view and pistil, all x 6. — S. balangeran (KorTH.) BuRCK. D. Bud, D1.

outer sepal, D2. inner sepal, D3. stamens from outside, D4. pistil, all x 10 (B KERR 8392, C Kerr 17713, D Cult.
Hort. Bog. VIII-D-6).

1982]

Stamens 17, unequal; filaments lorate, somewhat
tapering, + as long as anthers; anthers narrowly
oblong; appendage to connective filiform, stout at
base, rapidly tapering to a very slender apex, 3 times
length of anther, the inner 5 reaching the style apex,
sericeous in the distal 5. Ovary ovoid, sparsely seri-
ceous; style filiform, trifurcate apically, sericeous in
the basal 4. Fruit pedicel to 5 mm long, slender. 3
longer calyx lobes to 9 by 1.3 cm, narrowly spatulate,
subacute, 3.7 mm broad above the c. 8 by 8 mm
subauriculate thickened base; 2 shorter lobes to 6.5 by
0.4m, lorate, acute, similar but narrower at base. Nur
to 18 by 8 mm, narrowly ovoid, densely pale fulvous
pubescent; style remnant to 3 mm long.

Distr. Malesia: Malaya (Pahang), Sumatra (P.
Lingga; sterile coll.), Borneo (W. Sarawak).

Ecol. Locally frequent on yellow sandy soil on
coastal hills and flat sometimes swampy land.

Vern. Méranti bunbong (Mal.).

73. Shorea farinosa C.E.C. FISCHER, Kew Bull. (1926)
461; ibid. (1927) 81; Sym. Mal. For. Rec. 16 (1943) 36,
f. 19; SMITINAND, Thai For. Bull. (Bot.) 1 (1954) 23.

Large semi-deciduous tree. Leaf bud and parts of
petals exposed in bud persistently cream pubescent,
calyx and ovary thus at first, becoming sparsely
pubescent; parts otherwise sparsely fugaceous puber-
ulent, glabrescent. Twig 2-3 mm @ apically, smooth,
pale brown; stipule scars short, pale, slightly down-
curved. Leaves 7.5-15 by (2-)2.5-6 cm, elliptic-
oblong, thinly coriaceous; base obtuse; apex shortly
acuminate or retuse; nerves 13-20 pairs, very slender,
elevated on both surfaces, arched, at 60°—70’; tertiary
nerves scalariform, evident on both surfaces; midrib
slender but prominent and terete beneath, obscure
and depressed above; petiole (13—)20-30 mm long,
slender. Panicles to 10 cm long, lax, slender, terminal
or axillary; singly branched, branchlets to 8 cm long,
bearing to 6 distant secund flowers; flower buds to 6 by
4 mm, broadly ellipsoid; 3 outer sepals ovate-elliptic,
acute; 3 inner smaller, ovate, acute; stamens 25,
filaments broad and compressed at base, gradually
tapering; appendages filiform, villous in the distal 4, c.
15 times length of the narrowly oblong tapering
anthers; ovary narrowly ovoid, tapering into the
equally long columnar deeply trifid style. Fruit pedicel
to 4 mm long, expanding into the base of the nut; 3
longer calyx lobes to 14 by 2 cm, spatulate, obtuse, c. 8
mm wide above the 18 by 14 mm elliptic saccate
thickened base; 2 shorter lobes to 10 by 0.7 cm,
slender. Nut to 3 by 1.5 cm, ovoid, glabrous, tapering
to the prominent style remnant.

Distr. S. Tenasserim, Peninsular Thailand; doubt-
fully from Malesia: Malaya (fallen leaves from N.
Perlis and Kelantan).

74. Shorea polita VIDAL, Sinopsis (1883) 15, t. 15D;
Rev. Pl. Vasc. Filip. (1886) 61; BRANDIs, J. Linn. Soc.
Bot. 31 (1895) 88; Foxw. Philip. J. Sc. 13 (1918) Bot.
190; Merr. En. Philip. 3 (1923) 98; Descu, Mal. For.
Rec. 12 (1936) 39; Foxw. Philip. J. Sc. 67 (1938)

DIPTEROCARPACEAE (Ashton)

487

304. — S. mindanensis Foxw. Philip. J. Sc. 13 (1918)
Bot. 192; Merr. En. Philip. 3 (1923) 97; Reyes, Philip.
J. Sc. 22 (1923) 330. — Fig. 92.

Large tree. Parts of petals exposed in bud densely
pubescent, young parts + caducously tawny puberu-
lent, more persistent in young trees. Twigs c.2 mm @
apically, terete, smooth. Leaf buds small, ovoid.
Stipules caducous, not seen. Leaves (3.5—)6.5—9(—14)
by (1.5—)4-5(-11.5) em, + elliptic, thinly coriaceous,
occasionally cream lepidote beneath; margin nar-
rowly subrevolute; base cuneate to obscure (young
trees); apex shortly broadly acuminate to retuse;
nerves 11-13 pairs, slender but distinctly elevated
beneath, + applanate above, at 55°-80°; tertiary
nerves remotely scalariform, hardly elevated beneath,
+ obscure; midrib prominent beneath, obscurely
depressed above; petiole 1-2 cm long, slender, genicu-
late. Panicle to 14 cm long, slender, lax, terminal or
axillary; singly branched, branchlets to 8 cm long,
with to 8 secund flowers; bracteoles to 7 by 2 mm,
lanceolate, acute, caducous. Flower bud to 10 by 3
mm, fusiform; sepals ovate, the outer 3 subacute, the
inner 2 acuminate; stamens 21-25, becoming +
shorter than style at anthesis; anthers linear-oblong,
slender; appendages aristate, scabrous distally, +
twice length of anthers; ovary ovoid, puberulent; style
columnar, slender, + twice length of ovary, scarious
in the basal4, with + deeply trifid stigma. Fruit pedicel
short, stout, broadening into fruit. 3 longer calyx lobes
to 13 by 2 cm, lorate, obtuse, c. 6 mm wide above the
to 15 by 12 mm elliptic saccate thickened base; 2
shorter lobes to 7.5 by 0.5 cm, linear-lorate, acute,
similarly saccate at base; nut to 15 by 14 mm, ovoid,
tapering to a long slender filiform style remnant.

Distr. Malesia: Philippines.

Ecol. Widespread but scattered in Semi-evergreen
and Evergreen Lowland Dipterocarp forests.

Vern. Lauan, malaanonan.

Note. Closely allied to S. gratissima and S. monti-
gena, forming a group of three allopatric species
characteristic of seasonally or periodically dry forests
from Thailand to the Moluccas.

75. Shorea gratissima (WALL. ex KURZ) Dyer, FI. Br.
Ind. | (1874) 307; KiNG, J. R. As. Soc. Beng. Sc. 62, 2
(1893) 115; BRANDis, J. Linn. Soc. Bot. 31 (1895) 88,
89; Ript. Agr. Bull. Str. & F.M.S. 1 (1901) 59;
BRANDIS, Indian Trees (1906) 70; BurK. J. Str. Br. R.
As. Soc. 16 (1917) 167; ibid. 81 (1920) 51, 68; RIDL. FI.
Mal. Pen. 1 (1922) 226; Crars, Fl. Siam. Enum. 1
(1925) 143; Foxw. Mal. For. Rec. 10 (1932) 189, p.p.;
Burk. Dict. (1935) 2011; Sym. Mal. For. Rec. 16
(1943) 36, f. 19; SMITINAND, Thai For. Bull. 1 (1954)
22; MEUER & Woop, Sabah For. Rec. 5 (1964) 54, f.
5. — Hopea gratissima WALL. [Cat. (1828) n. 960,
nomen; DC. Prod. 16, 2 (1868) 635, nomen] ex KURZ,
J. R. As. Soc. Beng. Sc. 42, 2 (1873) 61.

Large tree. Young parts entirely shortly evenly
densely pale fulvous pubescent, glabrescent except on
stipules Twig c. 1.5 by 1 mm g apically, at first
somewhat compressed, becoming terete, smooth,

488 FLORA MALESIANA [ser. I, vol. 9?

Fig. 92. Flower details in Shorea sect. Anthoshorea HEM. — S. polita ViDAL. A. Bud, A/. outer sepal, A2. inner
sepal, both from inside, A3. stamens from inside, A4. pistil, all x 5 (CANICOSA 9734).

1982]

DIPTEROCARPACEAE (Ashton)

489

dark brown; stipule scars short, horizontal, obscure.
Buds to 2 by 1 mm, ovoid, subacute. Stipules to 10 by 3
mm, lanceolate, acute, caducous. Leaves 4-10 by
1.54.5 cm, ovate to elliptic, coriaceous, with undulate
margin; base cuneate; apex acute or with to | cm long
slender acumen; nerves c. 12-14 pairs, slender, hardly
raised beneath, arched, at 55°—65°; secondary nerves
scalariform to subreticulate, remote, obscure; midrib
obscure and depressed above, slender and acute
beneath; petiole 8-15 mm long, slender. Panicles to 10
cm long, terete or ribbed, terminal or axillary, singly
(if axillary) or doubly branched; branchlets to 4 cm
long, ascending, bearing to 8 flowers; bracteoles to 2
mm long, linear, fugaceous. Flower bud to 5 by 3 mm,
lanceolate. Sepals narrowly deltoid, 3 outer acute, 2
inner acuminate. Stamens 25; filaments compressed,
tapering, c. 14 times length of anthers; anther oblong,
tapering; appendages very slender, c. 3 times length of
anthers, villous in the distal 5. Ovary narrowly ovoid,
glabrescent, tapering into the style; style c. 15 times
length of ovary, with prominently trifid stigma. Fruit
glabrescent. Pedicel to 1.5 mm long and @; 3 longer
calyx lobes to 7 by 1.3cm, lorate-spatulate, obtuse, c. 6
mm wide above the to 8 by 5 mm saccate somewhat
thickened base; 2 shorter lobes to 5.5 by 0.6 cm,
otherwise similar. Nut to 15 by 8 mm, ovoid, apicu-
late.

Distr. Tenasserim, Peninsular Thailand, and in
Malesia: Malaya (Kelantan, Selangor, Singapore),
W. Sumatra (Bangkinang), Borneo (N.E. Sabah from
Kudat to W. Kutei).

Ecol. Locally abundant on low hills near the coast.

Vern. Meranti laut (Malaya), penggiran (Sabah).

76. Shorea henryana PIERRE in Lanessan, Pl. Util.
Colon. Fr. (1886) 302; For. Fl. Coch. 3 (1889) t. 229;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 89; Crarp, FI.
Siam. Enum. | (1925) 143; AsHToN, Gard. Bull. Sing.
31 (1978) 41.—S. sericeiflora FISCHER & HUTCH.
Kew Bull. (1926) 433; Sym. ex Descu, Mal. For. Rec.
12 (1936) 27; PARKINSON, Ind. For. Rec. (Bot.) 1
(1937) 43; Sym. J. Mal. Br. R. As. Soc. 19 (1941) 160;
Mal. For. Rec. 16 (1943) 41, f. 19, 25; SMITINAND, Thai
For. Bull. 1 (1954) 22. — S. gratissima (non (WALL. ex
Kurz) Dyer) Foxw. Mal. For. Rec. 10 (1932)
189. — S. longestipulata TarpiEU, Not. Syst. 10
(1942) 132. — Fig. 91 C—C3.

Large buttressed tree. Young twig and petiole
densely evenly pale rufous caducous pubescent; leaf
bud, panicles, sepals and parts of petals exposed in
bud persistently so; leaves, stipules and ovary sparsely
fugaceous pubescent; leaves pale pink-brown lepidote
beneath. Twigs 1-2 mm @g apically, smooth, terete,
becoming chocolate-brown; stipule scars slender,
short, horizontal. Buds to 2 by 1 mm, small, ellipsoid.
Stipules to 20 by 3 mm, narrowly elliptic, obtuse,
not at first caducous. Leaves (3—-)4-8(-12) by
(1-)2.5-4.5(-5) cm, broadly lanceolate to elliptic,
coriaceous; base obtuse or rarely broadly cuneate;
apex acute or with to | cm long broad acumen; nerves
17-20 pairs, slender, hardly raised beneath, at 80° at

the base, down to 45° towards the apex; secondary
nerves densely scalariform, slender, obscure; midrib
obscure and depressed above, slender but prominent
and terete beneath; petiole 10-18 mm long, slender.
Panicles to 11 cm long, slender, lax, terete or ribbed,
terminal or more rarely axillary; doubly (if terminal)
or singly branched; branchlets to 5 mm long, bearing
to 6 flowers; bracteoles unknown. Flower buds to 10 by
5 mm, ellipsoid. Sepals lanceolate, the inner 2 some-
what shorter, more narrowly acute, than the outer 3.
Petals white. Stamens 25(—30); filaments slender,
tapering, c. 15 times length of the narrowly oblong
anthers; appendages very slender, c. 3 times length of
anther, villous towards apex. Ovary ovoid; style
columnar, tapering, + twice length of ovary, villous
in the basal 4, prominently trifid at apex. Fruit pedicel
to 3 mm long, to 2mm g; 3 longer fruit calyx lobes to
9.5 by 1.6cm, lorate, obtuse, to 8 mm wide above the 8
by 8 mm suborbicular somewhat thickened saccate
base; 2 shorter lobes to 5.5 by 0.4 cm, otherwise
similar. Nut to 22 by 10 mm, narrowly ovoid,
apiculate, glabrous.

Distr. Lower Burma, S.E. and Peninsular Thai-
land, and in Malesia: N.W. Malaya (Kedah, Perlis,
Langkawi).

Ecol. Locally common in Semi-evergreen Diptero-
carp forest on well drained red soils.

77. Shorea montigena SLOOT. Reinwardtia 2 (1952) 57,
f. 18, 19. —S. balangeran var. binnendijkii BOERL.
Cat. Hort. Bog. 2 (1901) 108.

Huge buttressed tree. Young parts fugaceous
puberulent, ovary persistently so, panicle and parts of
perianth exposed in bud persistently greyish ocherous
pubescent; parts otherwise glabrous. Twigs c. 2 by |
mm apically, distinctly compressed, smooth, dark
brown. Buds minute, falcate-lanceolate. Stipules to 20
by 3 mm, linear-lanceolate, acute. Leaves 5-12 by
2-5.5 cm, ovate-lanceolate, thinly coriaceous, undu-
late; base cuneate; apex with to 1.5 cm long slender
tapering acumen; nerves 7—9(—13 in young trees) pairs,
very slender and hardly elevated on either surface,
ascending at 50°-65°, sometimes with a few scattered
shorter secondary nerves; tertiary nerves densely
scalariform, + obscure; midrib slender but prominent
and acute beneath, + obscure and depressed above;
petiole 22-25 mm long, slender, geniculate. Panicles to
17cm long, axillary or terminal, compressed, singly or
doubly branched; branchlets to 12 cm long; bracts
fugaceous. Flower bud to 12 by 4 mm, large; sepals
broadly ovate, acute, subequal; stamens (55—)65—72,
unequal; filaments compressed, tapering; appendages
slender, 15-2 times length of the narrowly oblong
anthers; ovary ovoid, tapering into somewhat longer
columnar style. Fruit pedicel short; 3 longer calyx
lobes to 11 by 1.5 cm, spatulate, obtuse, 1.5 mm wide
above the 1 by 1 cm ovate saccate thickened base; 2
shorter lobes to 35 by 3 mm, linear, similar at base.
Nut to 2 by 1.5 cm, ovoid, apiculate.

Distr. Malesia: Moluccas (Buru, Ceram), N.E.
Celebes? (leaves only).

490 FLORA MALESIANA ser. 1, voleg?

Fig. 93. Shorea assamica Dyer ssp. globifera (RIDL.) SYM. a. Habit, b. seedling, c. leaf, showing hairs in the left
half, veins in the right, d. fruit, e. nut, all x 4 (a YATES 1675, b bb. 19473, c bb. 20776, d-e KEP 87603).

1982]

Ecol. Common, frequently gregarious, on hills in
S. selanica forest.

Vern. Bahut, kayu bapa, gawa, babat (Buru), umale
(Ceram).

Notes. Differing from the Philippine species S.
polita principally in the greater number of stamens
and related, through it, to the widespread but disjunct
S. gratissima of drought prone evergreen forest.
GOTTWALD & PARAMESWARAN (Bot. Jahrb. 85, 1966,
457, 458) placed this species in sect. Shorea on the
basis of xylem anatomy, but noted its unusually low
density for that section, the presence of oxalate
crystals and red colour.

78. Shorea assamica Dyer, FI. Br. Ind. | (1874) 307;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 85; Indian Trees
(1906) 70; Foxw. Philip. J. Sc. 4 (1909) Bot. 516;
Troup, Ind. Woods and Uses (1909) 241; Silv. Ind.
Trees (1921) 133; GAMBLE, Man. Ind. Timb. (1922) 83;
PEARSON & BROWN, Commerc. Timb. Ind. | (1932)
119; KANJILAL & Das, Fl. Assam (1934) 136; PARKIN-
SON, Ind. For. Rec. (Bot.) 1 (1937) 40; Sym. Gard.
Bull. S. S. 9 (1938) 331; ASHTON, Gard. Bull. Sing. 31
(1978) 43. — Fig. 93.

KEY TO THE SUBSPECIES

1. Branchlets of panicle usually simple
ssp. assamica
1. Branchlets of panicle bifurcate or fascicled.
2. Appendage to connective as long as anther
a. ssp. globifera
2. Appendage to connective twice length of anthers.
3. Leaves epilose beneath b. ssp. koordersii
3. Leaves usually sparsely velutinate beneath
c. ssp. philippinensis

a. ssp. globifera (RIDL.) Sym. Gard. Bull. S. S. 9 (1938)
331, pl. 20; Mal. For. Rec. 16 (1943) 31, f. 19, 20;
ASHTON, Gard. Bull. Sing. 31 (1978) 44. — S. globi-
fera Riwv. Fl. Mal. Pen. 1 (1922) 232; Foxw. Mal.
For. Rec. 3 (1927) 35; ibid. 10 (1932) 191; BurK. Dict.
(1935) 2010; Sym. ex Descu, Mal. For. Rec. 12 (1936)
27. — S. sororia SLooT. Bull. Bot. Gard. Btzg III, 18
(1949) 247, f. 9, 10. — Fig. 93.

Large buttressed tree. Twigs, panicles, parts of
perianth exposed in buds, stipules outside (subgla-
brous to sparsely puberulent within), petioles, midribs
above and leaf undersurface persistently + densely
shortly evenly pale ocherous buff to rufous pubescent;
fruit sparsely persistently puberulent; leaf undersur-
face densely pale rufous lepidote (mature trees only).
Twigs to 2 by 1.5 mm @ apically, + compressed,
smooth, becoming dark brown; stipule scars short,
pale, transverse, with a prominent rib proceeding
down the internode from the base. Buds to 3 by 2mm,
ovoid, subacute, compressed. Stipules to 15 by 4(—7 in
young trees) mm, broadly or narrowly ovate, acute,
subauriculate, not at first caducous. Leaves
(4-)5-9(-10) by 2-4(-6) cm, ovate, elliptic or rarely
obovate, + coriaceous; base obtuse to subcordate;
acumen to 8 mm long, narrow; nerves 13-18 pairs,

DIPTEROCARPACEAE (Ashton)

49]

slender, obscure above, at 50°-65°, arched near the
margin; tertiary nerves densely scalariform, slender;
midrib obscurely depressed above, terete and promi-
nent beneath; petiole 5-7 mm long, short, slender.
Panicles to 6 cm long, short, ribbed, terminal or
axillary; branchlets usually bifurcate or fascicled, lax.
Flower bud to 8 by 4 mm, ellipsoid. Sepals narrowly
deltoid, subacute, the 3 outer somewhat narrower,
longer than the inner 2. Petals white, pink at base
within, broadly elliptic, obtuse. Stamens 15, shorter
than the style at anthesis; filaments slender, tapering,
as long as the slender lorate anthers; appendages to
connectives very slender, }-1 times length of anthers.
Ovary ovoid, glabrous; style columnar, c. 3 times
length of ovary, distinctly trifid at apex. Fruit pedicel
to 2 by 2 mm. 3 longer calyx lobes to 11 by 2 cm,
sublorate, obtuse, c. 5mm broad above the to 12 by 10
mm saccate thickened base; 2 shorter lobes to 9 by 0.8
cm, linear, subacute, similar at base. Nut to 15 by 15
mm, ovoid, tapering to an up to 6 mm long slender
style remnant.

Distr. Peninsular Thailand, and in Malesia:
Malaya (Perak and Central Pahang northwards),
Sumatra (west of the Barisan Divide from S. Atjeh to
Bencoolen), ?S.E. Borneo (sterile coll.).

Ecol. Evergreen forests on fertile clay soil; on well
drained flat land, especially by streams, and slopes to
1000 m; in areas liable to periodic drought. Locally
common, especially in northern Malaya.

Vern. Meéranti pipit, m. lampong, m. pasir, lémsa
kulat, 1. néram (Mal.), sogar baringin nabotar, s. b.
narara, bayang ayér, kélikung, ngérawan, mérkunyit
(Sum.).

Note. Indistinguishable from ssp. philippinensis
when sterile.

b. ssp. koordersii (BRANDIS) SYM. Gard. Bull. S. S. 9
(1938) 331. — S. koordersii BRANDIS ex KoorD. Med.
Lands Pl. Tuin 19 (1898) 355; Ic. Bog. 1 (1901) t. 80;
Fl. N.O. Celebes Suppl. 2 (1922) 8; ibid. Suppl. 3
(1922) 44, t. 91; Koorp.-ScHum. Syst. Verz. 3 (1914)
88: HeyNngE, Nutt. Pl. ed. 1, 3 (1917) 303; ibid. ed. 2
(1927) 1107, 1120; ENDERT, Tectona 28 (1935) 258;
SLooT. Reinwardtia 2 (1952) 42, f. 14, 15. — Aporosa
minahassae Koorp. Med. Lands Pl. Tuin 19 (1898)
625. — Vatica celebica KoorD. ex SYM. Gard. Bull. S.
S. 9 (1938) 331, nomen in syn.

Defining characters: Leaves peltate lepidote but
epilose beneath. Branchlets of panicle usually bifur-
cate or fascicled. Appendage to connective + twice
length of anther. Stigma + unlobed. 3 longer fruit
calyx lobes to 7 cm long.

Distr. Malesia: Celebes (widespread), Moluccas
(Sula, Batjan & Obi Is., 7Ambon), Philippines (Min-
danao, Luzon).

Ecol. Common, often gregarious, on fertile soil in
primary semi-evergreen forest on hills in lowlands.

Uses. The major timber source where it grows; the
clear crystalline resin, ‘damar tenang’, was once much
exploited.

Vern. Malueh, haro, waro, rama wuring, induk,

492

FLORA MALESIANA

[ser. I, vol. 92

tambija, damar lari, d. larieh, maru waru, karmungku
(Celebes), honi, pini, p. boti pien, ténang, t. puteh, t.
merah, t. babudo (Moluccas).

c. ssp. philippinensis (BRANDIS) SyM. Gard. Bull. S.S.9
(1938) 331. — S. philippinensis BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 88; Foxw. Philip. J. Sc. 6 (1911) Bot.
272: ibid. 13 (1918) Bot. 190; ibid. 67 (1938) 302;
Merr. En. Philip. 3 (1923) 98; Descu, Mal. For. Rec.
12 (1936) 27. — S. aff. harmandii (non PIERRE) Foxw.
Philip. J. Sc. 6 (1911) Bot. 272. — S. pallida Foxw.
Philip. J. Sc. 13 (1918) Bot. 190.

Defining characters as ssp. koordersii but leaves
sparsely velutinate beneath.

Distr. Malesia: Philippines (widespread), S.E. Bor-
neo.

Ecol. Scattered in semi-evergreen and evergreen
forest both in seasonal and relatively non-seasonal
regions.

79. Shorea ochracea Sym. Gard. Bull. S. S. 8 (1935)
285, pl. 27; BRowneE, For. Trees Sarawak & Brunei
(1955) 158; ASHTON, Man. Dipt. Brun. (1964) 165, f.
15; ibid. Suppl. (1968) 94, pl. 19 (bark); MEER &
Woop, Sabah For. Rec. 5 (1964) 57.—‘Majaw
DuRANT, Rep. For. Brunei (1933) 43.

Medium-sized to large tree with dark brown bark.
Twig, leaf bud, stipule and petiole densely persistently
rufous-brown powdery pubescent; partially cadu-
cous, sparse, on leaf beneath. Twig 4-S mm @ apically,
stout, terete, becoming smooth, with 2 mm long,
short, cuneate, horizontal or slightly ascending stipule
scars. Bud 3-6 by 3-5 mm, globose to ovoid, obtuse.
Stipule to 2 by 1.5 cm, elliptic, obtuse, caducous.
Leaves 12-18 by 7-10 cm, broadly elliptic-oblong,
coriaceous, bright yellow lepidote beneath; base cor-
date; acumen c. 5 mm broad; nerves 25—30 pairs,
prominent beneath, curved, at c. 110° at base, c. 20
towards the apex, with slender indistinct scalariform
tertiary nerves; petiole 1.3-2 cm long, short, stout.
Panicle to 10 cm long, terminal or axillary, terete, lax,
densely greenish yellow puberulent; regularly singly
or doubly branched, branchlets bearing to 8 flowers;
bracts and bracteoles to 2 by 1| cm, large, elliptic,
obtuse, subpersistent, puberulent. Flower bud to 8 by 3
mm, lanceolate, subacute. Calyx densely yellow-
brown tomentose outside, glabrous within; lobes
deltoid, slightly revolute, acuminate; 3 outer lobes
slightly longer, broader, obtuse; 2 inner lobes acute,
narrow at base. Petals lanceolate, subacute, shortly
pubescent on parts exposed in bud, puberulent else-
where. Stamens 15, subequal; filaments slightly longer
than anther, rather narrow, tapering gradually;
anther oblong, tapering; appendage to connective
somewhat longer than anther, reaching below style
apex, stout, acute. Ovary ovoid, densely pubescent;
style stoutly filiform, pubescent in the basal half,
otherwise glabrous, obscurely trifurcate apically.
Fruit calyx lobes shortly pubescent, densely so at base;
3 longer lobes to 10 by 2 cm, oblong, obtuse, to 1 cm
broad above the slightly broadened shortly auriculate

centrally thickened and saccate base; 2 shorter lobes
unequal, to 4.5 by 0.5 cm, obtuse, similar at base. Nut
c. 1.5 by 1 cm, ovoid, shortly densely tomentose; style
remnant to 2.5 mm long.

Distr. Malesia: Borneo (Lower Kapuas, Sarawak
to S.W. Sabah, Kinabatangan, Tidung to Puruktjau).

Ecol. Scattered on undulating land and hills to 750
m, in Mixed Dipterocarp forest.

Vern. Raruk (Iban), mélapi daun bésar (Sabah),
maro, m. hitam, kodahang (S.E. Borneo), kontoi
témbaga (W. Borneo).

80. Shorea virescens Paris in Fedde, Rep. 33 (1933)
244: Bijdr. Kennis Oost-Ind. Damarhars (1933) 120;
SLooT. Bull. Bot. Gard. Btzg III, 18 (1949) 240, f. 5, 6;
Rojo, Kalikasan 5 (1976) 99, f. 1: ASHTON, Gard. Bull.
Sing. 31 (1978) 44. — S. lamellata (non Foxw.) ASH-
TON, Man. Dipt. Brun. (1964) 167, f. 15, p.p.; ibid.
Suppl. (1968) 94, p.p.

Twig, panicle, bud, stipule, petiole, midrib (both
surfaces) and nervation beneath shortly densely per-
sistently grey-buff tomentose. Twig 2.5-4 by 1.5-3
mm, compressed, becoming finely cracked, stipule
scars c. 2.5mm long, glabrous, horizontal. Bud 2-3 by
4mm, globose to stoutly ovoid, obtuse. Stipule to 25
by 3-4 mm, linear, acute. Leaves alternate, 8-15 by
4-8 cm, obovate; base subcordate; acumen to 7.5mm
long, nerves 20-26 pairs, at 90° at base, c. 40°—-5S°
along the midrib; tertiary nerves very slender, scalari-
form, dense at 90° to the nerves; petiole 1.5—2 cm long.
Panicle to 10 cm long, terminal or axillary + com-
pressed, slender, straight; unbranched or singly
branched: bracteoles to 8 mm long, narrowly lanceo-
late, densely pubescent outside, puberulent within,
caducous. Flower bud to 6 by 3 mm, elllipsoid, obtuse.
Calyx densely puberulent outside, glabrescent within;
3 outer lobes ovate, subacute; 2 inner lobes shorter,
smaller, thinner, ovate-acuminate. Petals narrowly
lanceolate, densely pubescent on parts exposed in
bud. Stamens 15, of 2 lengths; filaments broad at base,
tapering and filiform distally; anthers narrowly
oblong; appendage to connective about twice length
of anther, exceeding style apex. Ovary ovoid, minutely
puberulent; style filiform, as long as ovary, distinctly
trifid. Fruit calyx shortly puberulent or glabrous when
mature; 3 longer lobes to 8 by 1.3 cm, spatulate, +
obtuse, to 6 mm broad above the to 8 by 7 mm elliptic
somewhat thickened saccate base; 2 shorter lobes to
5.5 by 0.5 cm, linear, similar at base. Nut to 1.3 by 1
cm, glabrescent; style remnant to 3 mm long, tapering.

Distr. Malesia: Borneo, Philippines (Mindanao,
Samar).

Ecol. Widespread but very local, flat and undulat-
ing land and low hills, to 500 m.

Vern. Kébang, baung raja, pakit, kontoi raba, k.
tjongil, madja kéruing, m. lilin (W. Borneo), mahum-
bong (S. Borneo), tégélam, bélobunio, pélépak batu
(S.E. Borneo), manggasinorong-lakihan (Mindanao).

81. Shorea javanica K. & V. Bull. Inst. Bot. Btzg 2
(1899) 3; Bijdr. (1900) 121; MoLLt & JANssONIUS,

1982]

Mikrogr. Holz (1906) 361; BACKER, Schoolfl. (1911)
110; Koorp. Exk. Fl. Java 2 (1912) 622; Koorp.-
ScuHuM. Syst. Verz. | (1913) Dipt. 4; ENDERT, Tectona
28 (1935) 288, 488: SLoor. Bull. Bot. Gard. Btzg III,
18 (1949) 230, f. 1; BACKER & BaAKH. f/f. FI. Java 1
(1963) 331. — S. vandekoppelii Paris in Fedde, Rep.
33 (1933) 244; Bijdr. Kennis Oost-Ind. Damarhars
(1933) 112, incl. var. grandifolia Paris, lL.c.
118. — Fig. 16.

Large tree. Twig, leaf buds, stipule outside, pani-
cles, calyx, parts of petals exposed in bud, ovary and
nut persistently evenly tawny brown pubescent;
becoming sparse, scabrous, on fruit calyx, stipule
within, petiole and leaf nervation beneath. Twig 2-3
mm @ apically, terete, becoming smooth. Leaf bud to
7 by 4 mm, ovoid-falcate, acute. Leaves (6.5—)10-15
by (3.5—)4-8 cm, elliptic-oblong to ovate, occasionally
obovate, + thinly coriaceous; base obtuse to shal-
lowly caudate; acumen to 7 mm long, short, + abrupt;
nerves 19-25 pairs, very slender but + prominent
beneath, applanate above, arched, at 65° —70’; tertiary
nerves densely scalariform, very slender but evident
and slightly elevated beneath; midrib slender but
prominent beneath, obscure and depressed above;
petiole 16-22 mm long, slender. Panicle to 14cm long,
slender, terminal or axillary, lax; singly branched,
branchlets to 4 cm long, bearing to 3 secund flowers.
Flower buds to 10 by 5 mm, ellipsoid; sepals narrowly
ovoid, acuminate, somewhat unequal; petals white;
stamens 15, shorter than style at anthesis; filaments
very long and slender, with scabrous apices,
23-3 x length of anthers. Ovary small, ovoid, puberu-
lent, tapering imperceptibly into a tapering puberu-
lent stylopodium twice its length and long filiform
glabrous style thrice its length. Fruit pedicel c. 2 mm
long and @, base of fruit obtuse. 3 longer calyx lobes to
18 by 1.5 cm, spatulate, obtuse, c. 7 mm broad above
the 11 by 10 mm elliptic saccate thickened base; 2
shorter lobes to 7 by 0.5 cm, lorate, subacute, similar
at base. Nut to 14 by 10 mm, ovoid, prominently
apiculate.

Distr. Malesia: Sumatra (W. coast from southern
Atjeh southwards; east from Palembang southwards),
Central Java (Subah in Pekalongan Res.; rare).

Ecol. Scattered in north, becoming gregarious in
many areas in south Sumatra, in primary and second-
ary lowland forests.

Uses. A once valuable producer of clear crystalline
resin as well as timber; grown in plantations in S.
Sumatra.

Vern. Damar puteh (Atjeh), d. sibosa, sibosa
(Tapanuli), d. saga (W. coast), d. kacha, d. mata
kuching (Palembang), méségar, méntégar, kacha (Ben-
coolen), d. ata, d. dacha, d. mata kuching (Lampong),
pélalar, p. léngo (Java).

82. Shorea lamellata Foxw. Mal. For. Rec. 10 (1932)
278; BuRK. Dict. (1935) 2014; Descu, Mal. For. Rec.
12 (1936) 27, 28; ibid. 14 (1941) 16, 17; Sym. Mal. For.
Rec. 16 (1943) 39, f. 19, 23; SLoor. Bull. Bot. Gard.
Btzg III, 18 (1949) 236, f. 3; ASHTON, Man. Dipt.

DIPTEROCARPACEAE (Ashton)

493

Brun. (1964) 164, f. 15, pl. 40 (stem-base), p.p.; ibid.
Suppl. (1968) 94, p.p.; MEER & Woop, Sabah For.
Rec. 5 (1964) 56, pl. la (bark).

Differing from S. javanica only as follows: Tomen-
tum uneven, scabrid. Anthers twice as long as broad,
broadly oblong.

Distr. Malesia: Malaya (Perak), Sumatra (Lingga,
Singkep), throughout Borneo but for N.E. Sarawak
and N. and W. Sabah.

Ecol. Local, undulating land and low hills; rarely
to 650 m in Malaya.

Vern. Méranti lapis (Mal.), tenam (W. Borneo),
buniau (W. Kutei), pakit (S. Borneo).

Note. The distribution of this species and S.

javanica is entirely allopatric; both are widespread,

and the many collections of each confirm that their
differences, though apparently slight, are consistent
and serve always to allow indisputable determina-
tion.

83. Shorea roxburghii G. Don, Gen. Hist. (1831) 813;
KASHYAPA, J. Bomb. Nat. Hist. Soc. 58 (1961)
543. — Hopea floribunda WALL. Cat. (1828) 964,
nomen; DC. Prod. 16, 2 (1868) 635, nomen. — S.
talura RoxB. [Hort. Beng. (1814) 93, nomen] FI. Ind.
ed. Carey 2 (1832) 618; Dyer, FI. Br. Ind. 1 (1874) 304;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 84; BRANDIS &
GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 265, fig.;
BRANDIS, Indian Trees (1906) 70; TALBoT, For. FI.
Bombay | (1909) 110; Troup, Silv. Ind. Trees (1921)
133; GiLG in E. & P. Pfl. Fam. ed. 2, 21 (1925) 261;
PARKINSON, Ind. For. Rec. (Bot.) | (1937) 38; Sym. J.
Mal. Br. R. As. Soc. 19 (1941) 160; Mal. For. Rec. 16
(1943) 42, f. 19, 26; SMITINAND, Thai For. Bull. 1
(1954) 5, 6, 23. — S. laccifera HEYNE ex WALL. [Cat.
(1828) 967, nomen] in DC. Prod. 16, 2 (1868) 630;
BepD. FI. Sylv. (1869) t. 6. — Vatica laccifera (WALL.
ex DC.) W. & A. Prod. 1 (1834) 84; WiGur, Ic. Pl. Ind.
Or. | (1839) t. 164. — Saul iallarea RoxB. ex W. & A.
Prod. 1 (1834) 84, nomen. — S. laurifolia WALL. Cat.
(1828) 967, nomen; ex STEUD. Nom. Bot. ed. 2, 2
(1841) 575, nomen; DC. Prod. 16, 2 (1868) 632,
nomen. — Vatica laurifolia (WALL.) STEUD. Nom.
Bot. ed. 2, 2 (1841) 745, nomen. — S. floribunda
(WALL.) Kurz ex Dyer, Fl. Br. Ind. | (1874) 304;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 85, t. 2, f. 19, 20;
Indian Trees (1906) 70; Sym. J. Mal. Br. R. As. Soc. 19
(1941) 160. —.S. harmandii PieERRE ex LANESSAN, Pl.
Util. Colon. Fr. (1886) 302; PrERRE, For. Fl. Coch. 3
(1889) t. 231; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 85;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895)
266. — S. cochinchinensis PIERRE, For. Fl. Coch. 3
(1889) t. 230; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 84;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895)
266; GUERIN, FI. Gén. I.-C. 1 (1910) 381, fig., incl. var.
saigonensis GUERIN; CRAIB, Fl. Siam. Enum. | (1925)
142; LecomTE, Bois Indochine (1926) 112; Foxw.
Mal. For. Rec. 10 (1932) 182; Burk. Dict. (1935)
2008; CORNER, Wayside Trees 1 (1940) 213.—S.
saigonensis PIERRE, For. Fl. Coch. 3 (1889) 257. — S.
attopoensis PIERRE, /.c. 257, t. 232. — S. obtusa WALL.

494

var. subevenis BOERL. Cat. Hort. Bog. 2 (1900)
106. — Fig. 91 B—B4.

Small or large, hardly buttressed, deciduous or
evergreen tree. Leaf bud persistently shortly evenly
pale rust pubescent, leaf undersurface, petiole, sti-
pules and young twigs usually at first + sparsely
pubescent, caducous (in Malesia) or sometimes per-
sistent. Twigs c.2 mm @ apically, + terete, becoming
smooth, pale to dark brown, sometimes minutely pale
lenticellate; stipule scars short, horizontal, the inter-
node frequently ribbed beneath their apices. Buds to 4
by 2 mm, ellipsoid. Stipules to 10 by 3 mm, linear
obtuse. Leaves 7.5-19 by 2.5—7 cm, elliptic-oblong,
thin; margin undulate; base broadly cuneate to sub-
cordate; apex acute or with to 6 mm long, short, broad
acumen; nerves | 2—20 pairs, slender, rather prominent
beneath, tending to bifurcate near the margin, rather
straight, at c. 60°-70°; secondary nerves slender,
remote, subscalariform; midrib evident towards base,
applanate to somewhat depressed, above, prominent
and terete beneath; petiole |1.4—4.5 cm long, frequently
somewhat swollen in the distal half and geniculate.
Panicle to 8 cm long, axillary to ramiflorous, rarely
terminal, slender, terete, glabrescent, lax; singly
branched, branchlets to 4 cm long, ascending, bearing
to 3 flowers; bracteoles fugaceous. Flower buds to 8 by
4 mm, lanceolate. Sepals glabrescent, fimbriate, del-
toid, acute, the 2 inner somewhat the shorter, subacu-
minate. Petals white, tinged red at base within.
Stamens 15; filaments slender, compressed, tapering,
equal to the lorate anthers as also the slender filiform
appendages. Ovary ovoid, glabrous; style slender
filiform, c. 4 times length of ovary, obscurely trifid.
Fruit glabrous. Pedicel obscure, expanding into the
receptacle; 3 longer calyx lobes to 9 by 1.2 cm,
spatulate, obtuse, c.4mm @ above the to 10 by 7mm
ovate saccate thickened lustrous base; 2 shorter lobes
to 4.5 by 0.6cm, lorate, otherwise similar. Nut to 15 by
10 mm, narrowly ovoid, tapering to an up to 5 mm
slender filiform style remnant.

Distr. Peninsular India, Burma, Thailand, Indo-
china and in Malesia: N.W. Malaya (Kedah, Perlis,
Langkawi).

Ecol. Common, locally gregarious in Schima-bam-
boo forests and on limestone; also occurring in
Semi-evergreen Dipterocarp forests (outside Malesia
in teak and other deciduous forests and in semi-ever-
green forests).

Vern. Témak (Mal.).

84. Shorea bentongensis Foxw. Mal. For. Rec. 10
(1932) 169, pl. 12; Burk. Dict. (1935) 2007; Sym.
Gard. Bull. S.S. 8 (1935) 280, pl. 24; Mal. For. Rec. 16
(1943) 33, f. 19. — S. pahangensis Foxw. Mal. For.
Rec. 10 (1939) 193, pl. 15; Burk. Dict. (1935) 2019.

Large buttressed tree. Young twigs, leaf buds,
stipules outside (glabrous within) petiole and raceme
+ persistently evenly shortly buff to pale rufous
pubescent; leaf nervation beneath and midrib above
sparsely puberulent or glabrescent; ovary densely
caducous puberulent. Twigs c.2 mm @ apically, terete

FLORA MALESIANA

[ser. I, vole9?

to slightly compressed, becoming smooth, dark
brown; stipule scars short, pale, transverse. Buds to 3
by 2 mm, ellipsoid, obtuse. Stipules to 12 by 7 mm,
broadly elliptic, obtuse, caducous. Leaves 7-15 by 4-8
cm, broadly ovate, rarely obovate, coriaceous; margin
subrevolute; base subcordate to broadly cuneate;
acumen to 1.5 cm long, prominent, slender; nerves
13-16 pairs, prominent beneath, curved, set at
50°—-65°; secondary nerves scalariform, slender; mid-
rib obscure, depressed above, terete and prominent
beneath; petiole 8-13 mm long, short, stout. Panicles
to 6 cm long, terminal or axillary, terete, singly
branched; bracteoles unknown. Mature flowers un-
known: Sepals narrowly deltoid, the 3 outer acute, the
2 inner prominently acuminate. Petals pale yellow.
Stamens 15; filaments equal to anthers, lorate, taper-
ing; anthers oblong-lorate; appendages very slender,
c. 3 times length of anthers. Ovary ovoid, surmounted
by an equally long puberulent cylindrical tapering
stylopodium and equally long tapering style. Fruit
glabrous. Pedicel to 3 by 3 mm, stout, becoming
impressed. Calyx lobes vestigial; 3 longer lobes to 7.5
by 0.8 cm, lorate, obtuse, tapering slightly above the
to 20 by 23 mm large prominently saccate thickened
base; 2 shorter lobes with only to 12 by 2 mm linear
projection above the similar base. Nut to 20 by 25 mm,
subglobose, resinous, hidden in the sepals, crowned
by an up to 4 mm long linear style remnant.

Distr. Malesia: Malaya (Selangor,
Johore).

Ecol. Local, low lying land in deep valleys.

Vern. Méranti méngkai, m. séga, bok.

Pahang,

85. Shorea hypochra HANCcE, J. Bot. 14 (1876) 242;
PiERRE, For. Fl. Coch. 3 (1889) t. 228; BRANDis, J.
Linn. Soc. Bot. 31 (1895) 89; BRANDIS & GILG in E. &
P. Pfl. Fam. ed. 1, 3, 6 (1895) 266; GUERIN, FI. Gen.
I.-C. 1 (1910) 383; Cratp, Fl. Siam. Enum. | (1925)
143; Foxw. Mal. For. Rec. 10 (1932) 187; Burk. Dict.
(1935) 2012; Sym. Mal. For. Rec. 16 (1943) 37, f. 19,
22.— S. maritima PIERRE in Lanessan, Fl. Util.
Colon. Fr. (1886) 302; For. Fl. Coch. 3 (1889) t. 229;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 89.—S.
cambodiana PiERRE, For. Fl. Coch. 3 (1889) t. 229;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 89.—S.
crassifolia RIDL. Fl. Mal. Pen. | (1922) 231.

Very large tree. Young parts evenly densely pale
golden pubescent, fugaceous except on stipule, pani-
cle, sepals and parts of petals exposed in bud; leaf
undersurface and petiole persistently pinkish to cream
lepidote (young tree excepted). Twig c. 4 by 3 mm
towards apex, stout, somewhat compressed, becom-
ing smooth to slightly rugulose, terete, pale brown;
stipule scars short, horizontal, obscure. Buds to 4 by 3
mm, broadly ovoid, acute. Stipules to 4 by 3 mm,
ovate, obtuse, caducous. Leaves 7-18 by 4.5-8 cm,
ovate to elliptic, thickly coriaceous; base broadly
cuneate to obtuse; apex obtuse, acute or with to 1 cm
long stout acumen; nerves |5—20 pairs, slender, rather
straight, at to 90° at the base, to 50° near the apex;
secondary nerves slender, subreticulate; midrib ob-

1982]

DIPTEROCARPACEAE (Ashton)

495

scure and depressed above, prominent and terete
beneath; petiole 2-4 cm long, terete, stout. Panicles to
20 cm long, terminal or axillary, somewhat com-
pressed, singly branched; branchlets to 7 cm long, lax,
bearing to 4 flowers; bracteoles unknown. Flower bud
to 10 by 5 mm, lanceolate. Sepals lanceolate, acute,
subequal, the outer 3 acute, the inner 2 acuminate.
Petals pale yellow. Stamens 15; filaments slender,
tapering, equal in length to the narrowly oblong
anthers; appendages slender, c. 14 x length of anthers.
Ovary ovoid, deeply trifid at apex. Fruit pedicel to 6
mm long, c. 4mm @, stout, expanding into the base of
the fruit. 3 Jonger calyx lobes to 17 by 2.6 cm, lorate,
obtuse or subacute, c. 8 mm wide above the to 2 by 1.5
cm elliptic saccate thickened base; 2 shorter lobes to
12 by 0.9 cm, slender, otherwise similar. Nut to 4.5 by
2.5 cm, ovoid, glabrous, tapering to the 4 mm long
tapering style remnant.

Distr. Cochinchina, Cambodia, S.E. and Peninsu-
lar Thailand, and in Malesia: Malaya (Selangor and
Pahang northwards) and N.E. Sumatra (Riouw,
Lingga).

Ecol. Locally common on flat land and undulating
hills near the coast, and on inland hills in more
seasonal zone, in Semi-evergreen Dipterocarp forest
and Evergreen Dipterocarp forest prone to periodic
drought.

Vern. Méranti témak, témak, t. bunga, t. nasi, t.
kacha, térbak, t. paya, m. térbak.

86. Shorea symingtonii Woop, Gard. Bull. Sing. 17
(1960) 493; Meer & Woop, Sabah For. Rec. 5 (1964)
57, pl. 1b (phot.), f. 6.

Very large buttressed tree. Parts of petals exposed in
bud densely persistently pubescent; twig apices, leaf
buds and stipules + persistently densely tawny puber-
ulent; petiole, nervation beneath, calyx and panicle
sparsely caducously so. Twig c. 4 by 2 mm, stout, +
compressed, becoming dark brown, terete. Buds c. 4
by 3 mm, ovoid-falcate, acute. Stipules to 12 by 5mm,
lanceolate, acute, fugaceous. Leaves (9-)10-18 by
(4—)5-8 cm, oblong to narrowly obovate, coriaceous;
base cordate or obtuse; apex tapering abruptly to | cm
long acumen; nerves | 8—22 pairs, prominent beneath,
typically shallowly depressed above, at 45°—70° the
base excepted; tertiary nerves densely scalariform,
elevated beneath, evident above; midrib stout
beneath, evident and + depressed above; petiole 1-2
cm long, stout. Panicle to 17 cm long, lax, pendant,
terminal or axillary; singly branched, branchlets to 11
cm long, bearing to 5 flowers. Flower bud to 10 by 4
mm, large, fusiform. Sepals lanceolate, slender, sub-
acuminate, slightly unequal. Stamens 15, shorter than
style; filaments rather short, broad and compressed at
base, tapering; anthers narrowly oblong; appendages
slender, scarious near apices, c. 25 x length of anthers;
ovary ovoid, puberulent; style stoutly columnar, ob-
scurely trifid at apex, sparsely puberulent in the basal
3. Fruit pedicel c. 5 mm g, very stout and prominent,
expanding into the receptacle. 3 longer calyx lobes to
18 by 2.5 cm, lorate-lanceolate, obtuse to subacute,

not tapering above the concave incrassate base; 2
shorter lobes to 13 by 1.5 cm, narrower, acute but
otherwise as in longer lobes. Nut to 2.5 by 1 cm,
narrowly ellipsoid-ovoid, prominently apiculate.

Distr. Malesia: Borneo (E. Sabah).

Ecol. Scattered on well drained undulating land in
Mixed Dipterocarp forest below 250 m.

Vern. Meélapi bunga.

87. Shorea retinodes SLoorT. Bull. Bot. Gard. Btzg III,
18 (1949) 243, f. 7-8.

Large buttressed tree. Young twigs, leaf buds,
outside of stipules and parts of petals exposed in bud
+ persistently grey-brown puberulent; panicle, calyx,
leaf nervation beneath and petiole fugaceously so
(densely pubescent in young trees). Twig c. 1 mm @
apically, terete, slender, much branched. Leaf bud
minute. Stipules to 8 mm long, linear, fugaceous.
Leaves 5.5-12 by 1.8-4.8 cm, narrowly elliptic to
lanceolate, thinly coriaceous, dull densely lepidote
and drying coppery brown beneath; base broadly
cuneate to obtuse; acumen to | cm long, tapering;
nerves 15-18 pairs, slender but distinctly elevated
beneath, evident above, ascending at 50°—65°; tertiary
nerves densely scalariform, + unraised, + obscure;
midrib slender but prominent beneath, obscurely
depressed above; petiole 12-20 mm long, slender,
geniculate. Panicle to 12 cm long, slender, terminal or
axillary, + doubly branched; flowers secund; brac-
teoles fugaceous. Flower buds to 6 by 3 mm, fusiform.
Sepals ovate, the outer 3 subacuminate, the inner 2
prominently subcaudate. Stamens 15, subequal,
shorter than style at anthesis; filaments slender,
tapering; anthers narrowly oblong; appendages
slender, distinctly scarious, c. 3 x length of anther.
Ovary ovoid, glabrous; style slender, filiform, ob-
scurely trifid. Fruit pedicel to 2mm long, slender, base
of fruit obtuse; 3 longer calyx lobes to 7.5 by 1 cm,
spatulate, obtuse, tapering to c. 5mm broad above the
c. 7 by 5 mm elliptic saccate thickened base; 2 shorter
lobes to 4 by 0.3 cm, linear, similar at base. Nut to 8 by
5 mm, ovoid, apiculate.

Distr. Malesia: Sumatra (Barisan Range from
Toba to Musi Ulu, and down West coast; also
Kuantan Distr. in east; Pulau Musala, Pulau Sitam-
barat).

Ecol. Scattered, rarely common, in lowland coastal
and hill forests to 1000 m.

Vern. Damar meérilem, mérilem, marilem
(Tapanuli), d. mansarai, mansarai, bélamsarai
(Tapanuli, Padang), serga, s. gunong (Panobasan),
meéranti saga, saga, damar saga, banio sapek, b. rawan,
d. puteh (W. coast).

88. Shorea cordata ASHTON, Gard. Bull. Sing. 22
(1967) 285, pl. 31; Man. Dipt. Brun. Suppl. (1968) 93.
its, ID.

Large tree. Young twig and petiole densely cadu-
cous puberulent; leaf nervation beneath sparsely so,
bud persistently so, stipule outside sparsely persis-
tently so. Twig c. 3 by 2 mm @ apically, somewhat

496

compressed, becoming terete, smooth to rugulose;
stipule scars horizontal, broad, pale, prominent. Bud
to 7 by 4mm, ellipsoid, compressed, acute. Stipule to
12 by 6 mm, elliptic, obtuse. Leaves 8-15 by 5.5—10.5
cm, oblong to obovate; base typically cordate, some-
times obtuse; apex obtuse, retuse or abruptly to 8 mm
long acuminate; nerves 15-18 pairs, prominent
beneath, at 30°—40°; tertiary nerves slender, densely
scalariform, set diagonally to nerves; midrib
depressed above, prominent beneath; petiole 12-25
mm long, terete, frequently rugulose. Panicle to 12 cm
long, terminal or axillary, lax, compressed or ribbed;
singly branched, branchlets to 5 cm long, bearing to 5
+ secund flowers; bracts to 10 by 3 mm, lanceolate,
acute, puberulent, fugaceous; bracteoles to 10 by 5
mm, elliptic, obtuse, not at first caducous. Flower bud
to 10 by 5 mm, fusiform. Calyx sericeous outside,
glabrous within; lobes narrowly deltoid-lanceolate,
narrowly obtuse, the 3 outer somewhat longer than
the 2 inner. Petals oblong-lanceolate, sericeous out-
side, glabrous within. Stamens 15, in 3 unequal
verticils or with the outer vestigal and devoid of
anthers; filaments longer than anther, narrow, com-
pressed, tapering; anthers elliptic-oblong; appendage
to connective to 6 times as long as anther, filiform,
tapering somewhat shorter than style apex. Ovary
ovoid, glabrous; style columnar, c. 2xlength of
ovary, tapering, sericeous in the basal 3, obscurely
trifurcate apically. Fruit glabrous. Pedicel obscure,
tapering into the base of the fruit. 3 longer calyx lobes
to 13 by 2.5 cm, lorate, obtuse, c. 8 mm broad above
the c. 12 by 12 mm orbicular saccate thickened base; 2
shorter lobes to 6.5 by 0.8 cm, narowly lanceolate,
acute, similar at base. Nut to 14 by 10 mm, globose,
closely and completely enveloped by the calyx; style
remnant c. 4 mm long.

Distr. Malesia: Borneo (West Borneo to Central
Sarawak).

Ecol. Rare, fertile soils on igneous rocks.

89. Shorea bracteolata Dyer, FI. Br. Ind. | (1874) 305;
KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 117;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895)
264; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 85; BuRK. J.
Str. Br. R. As. Soc. 76 (1917) 164, fig.; ibid. 81 (1920)
73, fig.; Gard. Bull. S. S. 3 (1923) 36; Rip. Fl. Mal.
Pen. | (1922) 229; Baker f. J. Bot. 62, Suppl. (1924)
10; GitG in E. & P. Pfl. Fam. ed. 2, 21 (1925) 260;
HEYNE, Nutt. Pl. ed. 2 (1927) 1115; Foxw. Mal. For.
Rec. 1 (1921) 80; ibid. 3 (1927) 28, 3 pl.; ibid. 10 (1932)
183, pl. 1 (root syst. seedling); SLoor. in Merr. Pl. Elm.
Born. (1929) 202; Epbwarps, Mal. For. Rec. 9 (1931)
141; Burk. Dict. (1935) 2007; Descu, Mal. For. Rec.
12 (1936) 27; ibid. 14 (1941) 16; Sym. Mal. For. Rec. 16
(1943) 34, f. 18, 19,21; SLoor. Bull. Jard. Bot. Btzg III,
18 (1949) 259, f. 13: BRowne, For. Trees Sarawak &
Brunei (1955) 158; ASHTON, Man. Dipt. Brun. (1964)
163, f. 15; ibid. Suppl. (1968) 93; Meer & Woop,
Sabah For. Rec. 5 (1964) 53, f. la. — S. foveolata
ScorT. ex Foxw. Mal. For. Rec. 10 (1932) 183, nomen
in syn.

FLORA MALESIANA

[ser. I, vol. 9?

Medium-sized to large tree. All parts at first
sparsely pale brown pubescent, surfaces covered with
a scurfy waxy deposit; becoming glabrous except for
the pubescent bud and panicle. Twig to 2 mm g@
apically, somewhat compressed, becoming smooth,
glabrous; stipule scars short, inconspicuous. Bud 3—5 by
2 mm, shortly falcate, compressed, subacute. Stipule
c. | cm long, linear, fugaceous. Leaf 9-14 by 4-6 cm,
oblong-ovate to elliptic, + thinly coriaceous; base
obtuse; acumen 8-15 mm long; nerves 12-15 pairs,
slender, curved, at c. 60°—70°; tertiary nerves distant,
scalariform to subreticulate; petiole 1-2 cm long,
slender. Panicle to 10 cm long, terminal or axillary,
slender, terete, straight, usually singly branched;
bracteoles to 12 by S mm, lanceolate, sparsely pubes-
cent. Flower bud to 10 by 5 mm, ellipsoid, obtuse.
Calyx densely pubescent outside, sparsely so within; 3
outer lobes broadly ovate-hastate, obtuse; 2 inner
lobes narrowly ovate, acuminate, slightly shorter,
thinner. Petals pale yellow tinged pink at base within,
large, oblong-lanceolate, obtuse, shortly pubescent on
parts exposed in bud. Stamens 15, the inner 5
somewhat longer than the outer 10; filaments broad,
compressed at base, tapering gradually and filiform
distally; anthers narrowly oblong; appendage to con-
nective 3-4xlength of anther, slender, scabrous.
Ovary ovoid, sparsely pubescent towards apex; style c.
2xlength of ovary, distinctly trifid towards apex.
Fruit calyx puberulent, + glabrescent; 3 longer lobes
to 10 by 1.7 cm, spatulate, obtuse, to 4-5 mm broad
above the c. 10 by 7 mm thickened saccate base; 2
shorter lobes 5S—8 by 0.7 cm, unequal and sometimes
almost as long as the larger lobes, spatulate, obtuse,
similar at base. Nut c. 2 by | cm, narrowly ovoid; style
remnant to 4 mm long, acute, short.

Distr. Malesia: Malaya, Singapore, Sumatra
(excluding Atjeh), Borneo.

Ecol. Coastal hills and undulating land, in valleys
occasionally to 600 m; on deep well drained soils.

Vern. Méranti pa’ang, m. kétapak, m. ségor, méluit,
mémbantai, sama rupa meéranti, sépa pétoi, témak
(Mal.), kédontang, k. abang, méranti kasih, m. kunyit,
m. képala, manisan (Sumatra), lampong bahei, 1. mit, 1.
meérahan, bényau (S.E. Borneo).

Note. A widespread and variable species.

90. Shorea resinosa Foxw. Mal. For. Rec. 10 (1932)
234, pl. 19; Burk. Dict. (1935) 2021; Sym. Mal. For.
Rec. 16 (1943) 40, f. 19, 24; ASHTon, Man. Dipt. Brun.
Suppl. (1968) 94, f. 12.

Large tree. Young twigs, buds, stipules outside and
petioles shortly buff fugaceous puberulent. Twig c. 2
mm @ apically, terete, smooth; stipule scars short,
horizontal, obscure. Bud to 3 by 2 mm, ellipsoid,
obtuse. Stipule to 12 by 8 mm, elliptic obtuse. Leaves
6.5-13 by 3.5—7 cm, lanceolate to elliptic, undulate;
base narrowly to broadly cuneate; acumen to 12 mm
long, slender; nerves 10-13 pairs, slender but promi-
nent beneath, at 40°—50°; tertiary nerves slender,
scalariform, at c. 90° to the nerves; midrib depressed
above, prominent beneath; petiole 9-17 mm long,

1982]

drying rugose. Panicle to 8 cm long, terminal or
axillary, terete, caducous puberulent, — singly
branched. Flower bud to 12 by 4mm, fusiform. Calyx
glabrous; 3 outer lobes narrowly lanceolate, acute; 2
inner somewhat shorter, broader. Petals pale yellow,
lorate, obtuse, puberulent on parts exposed in bud.
Stamens 15, in 2 unequal verticils; filaments com-
pressed at base, tapering; anthers narrowly oblong;
appendage to connective c. 2} length of anthers,
filiform, slender, glabrous. Ovary ovoid, glabrous,
surmounted by a long slender glabrous style c.
3x length of ovary; style trifid towards apex. Fruit
glabrous. Pedice/ to 4mm long, c. 3 mm @, terminat-
ing abruptly at the obtuse base of the fruit. 3 longer
calyx lobes to 9.5 by 2.5 cm, spatulate, obtuse, c. 7 mm
broad above the to 14 by 14 mm subglobose saccate
thickened base; 2 shorter lobes to 7 by 0.6 cm, linear,
similar at base. Nut to 1.5 by 1.5 cm, ovoid; style
remnant to 6 mm long, persistent, frequently lustrous
with a film of resin.

Distr. Malesia: Malaya, Central Sumatra (Indra-
giri, sterile coll.), Borneo (W. Sarawak).

Ecol. Scattered on undulating land or hills to 500
m.

Vern. Méranti bélang, lemesa, sama rupa méranti

(Mal.).

91. Shorea agamii ASHTON, Gard. Bull. Sing. 19 (1962)
270, pl. 9.—S. assamica ssp. philippinensis (non
SyM.) BROWNE, For. Trees Sarawak & Brunei (1955)
TS

a. ssp. agamii ASHTON, Man. Dipt. Brun. (1964) 161, f.
15, pl. 41 (stem-base, bark); ibid. Suppl. (1968) 93;
Gard. Bull. Sing. 22 (1967) 285; MEIJER & Woop,
Sabah For. Rec. 5 (1964) 51, f. 4.

Large tree. Young twig, panicle, bud, stipule,
petiole and midrib beneath caducous pale brown
pubescent. Twig 2-3 mm @ at the apex, becoming
smooth, terete; stipule scars short, inconspicuous. Bud
c. 4 by 2.5 mm, ovoid to falcate, slightly pubescent or
glabrous, obtuse. Stipule c. 10 by 3 mm, oblong,
obtuse, shortly caducous pubescent outside, glabrous
within. Leaves 10-15 by 6-10 cm, broadly ovate to
oblong, coriaceous; base obtuse or subcordate;
acumen ().5S—1 cm long, broad; nerves 9-13 pairs, well
spaced, curved, at c. 90° at the leaf base, c. 40° at the
apex; tertiary nerves densely scalariform, at 90° to
nerves; petiole 1-1.5 cm long. Panicle 6-10 cm long,
terminal or axillary, terete to angular apically; singly
or doubly irregularly branched, branchlets short,
bearing to 5 flowers; bracts fugaceous; bracteoles to 3
mm long, narrowly elliptic, subacute, shortly pale
buff-tawny puberulent, caducous. Flower bud to 5 by
2.5 mm, small, narrowly ellipsoid, subacute. Calyx
densely shortly pale yellowish buff pubescent; 3 outer
lobes narrowly ovate, subacuminate; 2 inner lobes 7 as
long, narrow, prominently caudate. Petals small,
ovate-elliptic, acute, densely pubescent on parts
exposed in bud, puberulent elsewhere on outer sur-
face. Stamens 15 in 3 subequal verticils; filaments +

DIPTEROCARPACEAE (Ashton)

497

same length as anthers, rather narrow, tapering;
anthers oblong, somewhat tapering; appendage to
connective c. 3 times length of anther, slender,
extending to style apex. Ovary ovoid, densely pubes-
cent; style slightly longer than the ovary, sparsely
pubescent in the basal half, glabrous distally, stoutly
filiform, tapering, obscurely trifurcate. Fruit calyx
shortly fugaceous pubescent; 3 longer lobes 10-12 by
1.5—2.2 cm, spatulate, obtuse, tapering to 5mm broad
above the 1—1.5 cm broad strongly saccate thickened
base; 2 shorter lobes to 6 by 0.4 cm, linear, unequal,
similar at base. Nut to 2 by 1.5 cm, ovoid, glabrescent;
style remnant 3-4 mm long, acute; base of fruit
impressed at the short pedicel.

Distr. Malesia: Borneo (Bintulu north-eastwards
to Sabah and Tidung).

Ecol. Leached soils on undulating land and shale
hills below 700 m.

Vern. Méranti puteh timbul (Brun.), mélapi agama
(Sabah).

b. ssp. diminuta ASHTON, Gard. Bull. Sing. 22 (1967)
285, pl. 30; Man. Dipt. Brun. Suppl. (1968) 93.

Leaf 4.5—10 by 2.5—4 cm, narrowly ovate.

Distr. Malesia: Borneo (W. Sarawak, Kapuas and
Rejang hinterlands, Muara Tewe).

Ecol. As ssp. agamii.

92. Shorea confusa ASHTON, Gard. Bull. Sing. 31
(1978) 44. — S. virescens (non PARIS) ASHTON, Man.
Dipt. Brun. (1964) 167; ibid. Suppl. (1968) 95; MEWER
& Woop, Sabah For. Rec. 5 (1964) 60, f. 6.

Large tree. Leaf bud, panicle, stipule outside,
petiole, and very young twig caducous pubescent.
Twig 2-3.5 mm @ apically, frequently rugulose;
stipule scar short, obscure. Bud 3-4 by 2 mm, conical,
acute. Stipule c. 8 by 3mm, linear to deltoid, subacute.
Leaves 6-12 by 3.5—3 cm, elliptic to slightly obovate;
base obtuse; acumen broad, 0.5—1.0 cm long; nerves
(10—)13-18 pairs, curved, at 90° at the base, c. 40° at
the apex; tertiary nerves densely scalariform, at 90° to
nerves; petiole 1-1.5 cm long, c. | mm Qg, rather
slender. Panicle to 22 cm long, terminal or axillary,
terete, lax; singly or doubly branched, branchlets to 10
cm long, bearing to 6 flowers; bracts and bracteoles
unknown. Flower bud to 9 by 5 mm, narrowly ovoid,
obtuse. Calyx puberulent outside, glabrous within;
lobes equal, narrowly deltoid, obtuse. Petals large,
narrowly ovate, acute, puberulent on parts exposed in
bud. Stamens 15, in 3 subequal verticils; filaments c.
15xlength of anther, slender, tapering gradually:
anthers oblong, tapering; appendage to connective c.
3 x length of anther, reaching 4 length of style. Ovary
small, ovoid, puberulent; style stoutly filiform, c.
3 x length of ovary, glabrous in the apical 4, otherwise
puberulent, shallowly trifurcate. Fruit pedicel stout.
Calyx glabrous; 3 longer lobes to 12 by 1.5 cm,
narrowly spatulate, obtuse, hardly tapering, slightly
broadening at the thickened saccate base; 2 shorter
lobes to 6 by 0.5 cm, unequal, linear, similar at base;
base of calyx obconical, tapering into the pedicel. Nut

498 FLORA MALESIANA [ser. I, vol. 9?

Qi At
Fig. 94. Flower details in Shorea sect. Rubella ASHTON. All x 10. Sepals drawn from inside. — S. albida Sym. A.
Outer sepal, A/. inner sepal, 42. stamens from inside, 43. stamens and pistil. — S. e/liptica BuRCK. B. Bud, B1/.

outer sepal, B2. inner sepal, B3. stamens from inside, B4. pistil (A TAHIR 12250, B Rosii & ASAH 3373).

to 2 cm long, ovoid, glabrescent; style remnant c. 6 Ecol. Scattered, undulating land and hills below

mm long, slender. 650 m, on leached clay soils in Mixed Dipterocarp
Distr. Malesia: N.E. Borneo (N.E. Sarawak, forest.

Sabah S.E. to Sangkulirang). Vern. Meranti sulang saling.

6. Section Rubella

ASHTON, Gard. Bull. Sing. 20 (1963) 267; Man. Dipt. Brun. (1964) 117. — Fig. 94,
95.

Flower buds medium size, fusiform. Petals generally cream suffused with pink,
lanceolate, contorted and imbricate at base forming a goblet enclosing the

1982} DIPTEROCARPACEAE (Ashton) 499

anthers, connate on falling. Stamens 15-50; filaments lorate, long, tapering
gradually or abruptly below the anthers; anthers with 4 glabrous linear-oblong
pollen sacs; appendage to connective short or to } length of the pollen sacs,
becoming + reflexed at anthesis, glabrous. Ovary with or without distinct
stylopodium. Leaf white lepidote beneath, with scalariform tertiary nerves;
midrib above evident or obscure (S. a/bida). Large stoutly buttressed trees. Bark
surface prominently V-section fissured. Wood red, without silica deposits; vessels

solitary or in small clusters.
Distr. Malesia: Philippines, northern Borneo.

Ecol. Lowland evergreen forests, scattered or gregarious (S. negrosensis, S. albida).

Note. The androecium and gynoecium characters in most respects resemble those of sect. Anthoshorea.
WHITMORE (Gard. Bull. Sing. 19, 1962, 2) has shown that the bark anatomy of S. albida also in most respects
conforms with that section. The wood anatomy conforms by contrast entirely with SYMINGTON’s (Mal. For. Rec.
16, 1943, 58) and Descn’s (ibid. 14, 1941, 34) Red Meranti Group.

93. Shorea albida Sym. [ex THomas, Mal. For. 3 (1934)
147, nomen] Gard. Bull. S. S. 8 (1935) 283, pl. 26;
BROWNE, For. Trees Sarawak & Brunei (1955) 137;
ANDERSON, Gard. Bull. Sing. 20 (1963) 158; ASHTON,
Man. Dipt. Brun. (1964) 175, f. 19, pl. 42 (stem), 43
(stand), 44 (stand in swamp); ibid. Suppl. (1968) 104;
BRUNIG, Commonw. For. Rev. 52 (1973) 260. — Fig.
8, 94 A-A3, 96, 97.

Small, medium-sized or vast buttressed tree. Twig,
leaf bud, stipule, petiole and leaf nervation beneath
shortly densely persistently red-brown puberulent;
leaf beneath cream lepidote. Twig to 5 by 2 mm @
apically, compressed, becoming smooth; stipule scars
short, straight, thin. Bud to 10 by 6 mm, ovoid to
hastate, compressed, acute. Stipules to 20 by 8 mm,
oblong, obtuse. Leaves 7.5—15 by 4.5—6.5 cm, oblong-
elliptic, coriaceous; base obtuse; acumen to 6 mm
long, broad; nerves 16-20 pairs, slender, hardly raised
beneath, at 55°—70°, with shorter intermediates; ter-
tiary nerves slender, densely scalariform, diagonal to
nerves; midrib prominent beneath, obscure and
depressed above; petiole 2—-3.5 cm long. Panicle to 18
cm long, terminal or axillary, compressed, shortly
densely cream puberulent; doubly branched, branch-
lets bearing to 3 flowers; bracteoles to 8 by 3 mm,
lanceolate, acute, densely pubescent outside, glabrous
within. Bud to 9 by 5 mm, broadly ellipsoid, acute.
Calyx densely puberulent outside, glabrous within; 3
outer lobes deltoid, obtuse; 2 inner shorter, deltoid,
acuminate. Petals cream, ovate, obtuse, pubescent on
parts exposed in bud. Stamens 20-25, of variable
length, the inner 5 somewhat longer than the others;
filament broad at base, tapering and filiform distally;
anther narrowly oblong, longer than the filament;
appendage to connective short, slender, erect, less
than 5 length of anther. Ovary ovoid, shortly pubes-
cent; style twice length of ovary, densely pubescent in
basal half. Fruit calyx persistently sparsely pubescent
at base, glabrescent elsewhere; 3 longer lobes to 8 by
1.4 cm, narrowly spatulate, obtuse, to 4 mm broad
above the to 7 by 7 mm suborbicular thickened saccate
base. Nut to 12 by 9 mm, ovoid, densely greyish buff
pubescent; style remnant to 2 mm long.

Distr. Malesia: N.W. Borneo (Kapuas swamps
through Sarawak to the Limbang Valley).

Ecol. Local on podsols on terraces and plateaux in
Heath forest to 1200 m. Gregarious and dominant on
oligotrophic peat swamps except at the margin and
sometimes centre.

The peat swamp forests where S. albida occurs can
be classified into a succession of concentric ‘phasic’
communities according to the performance of this and
other dominant species (ANDERSON, 1963): 1. A mixed
species forest at the periphery lacking S. albida. 2.
With S. a/bida dominant and sometimes reaching 6.5
m tall, but forming an incomplete heterogeneous
canopy and rarely successfully regenerating. 3. With
S. albida forming a complete even canopy, regenerat-
ing patchily and becoming smaller towards the centre.
4. With Litsea palustris a dominant or codominant,
forming an even canopy but not exceeding 40 m tall.
Here regeneration is abundant though largely
through coppicing.

S. albida does not occur in the innermost communi-
ties. Lightning and wind damage form conspicuous
gaps in the even canopy of phasic community
(BRUNiG, 1973), but mortality over much greater
areas, in large sharply defined patches totalling thou-
sands of hectares in all, is attributed to an unidentified
moth larva, belonging to the Himantidae (cf. Imp.
For. Rev. 40, 1961, 19).

Uses. The largest trees, towards the margins of the
swamps, are all hollow and are very hard wooded;
those towards the centre are too small and the wood is
soft; trees in the intermediate zone are an important
source of red meranti timber.

Vern. Alan (Sar.), séringawan (Brun.).

Note. The vegetative parts are strikingly similar to
those of S. balangeran in sect. Brachypterae, which
shows an apparently identical ecological range and
replaces it in Indonesian Borneo. The stamens at once
distinguish these species, but the presence of abnor-
mal stamens of the S. ba/angeran type in the flowers
examined by SYMINGTON suggests that the affinity is
genuine. The obscure midrib above and slender nerves
distinguish S. albida from other species in this section

500

eu Tan

Gi

Fig. 95. Flower details in Shorea sect. Rubella ASH-

TON. All x 10. Sepals drawn from inside. — S. dispar

ASHTON. C. Bud, C/. outer sepal, C2. inner sepal, C3.

stamens from inside, C4. pistil. — S. rubella ASHTON.

D. Stamens, from outside, D/. pistil (C S 29208, D
BRUN 3078).

FLORA MALESIANA

[ser. I, vol. 9?

and evoke S. coriacea and its allies in sect. Brachy-
pterae, as well as sect. Anthoshorea generally.

94. Shorea rubella ASHTON, Gard. Bull. Sing. 19 (1962)
307, pl. 27; Man. Dipt. Brun. (1964) 216, f. 19, pl. 57
(habit & bark); ibid. Suppl. (1968) 118; MEER &
Woop, Sabah For. Rec. 5 (1964) 143. — Fig. 1, 95
D-D1.

Medium-sized to large tree. Young parts evenly
shortly pale cream to pink-buff pubescent, persistent
only on panicle, bud and stipule. Twig 2-3 mm @ at
apices, ridged and compressed apically on drying,
becoming terete, smooth, stout; stipule scars c. 1.5
mm longat first, + horizontal, narrowly cuneate. Bud
6-9 by 2-3 mm, narrowly ovoid or slightly falcate.
Stipule to 18 by 6mm, hastate to falcate, acute. Leaves
9.5-14 by 6-8 cm, broadly ovate, thickly coriaceous,
cream lepidote beneath; base obtuse; acumen to 1 cm
long; margin sometimes slightly revolute; nerves 5—7
pairs, prominent beneath, curved, well spaced, at c.
45°-60°, with or without minute axillary domatia;
tertiary nerves at c. 90°, slender, densely scalariform;
midrib applanate on upper surface or slightly
depressed. Petiole 2.3-3.5 cm long. Panicles to 15 cm
long, terminal or axillary, ribbed on drying; singly or
doubly branched, branchlets to 5 cm long, lax, bearing
to 5 distichous flowers; bracts and bracteoles un-
known. Flower bud to 7 by 3 mm, narrowly ovoid,
acute. Calyx shortly densely pale pink-grey pubescent
outside, glabrous within; lobes ovate, subequal, the
inner 2 slightly broader and shorter than the outer 3.
Petals pale pink, darkening towards base, oblong,
obtuse, only slightly contorted, hardly adhering on
falling, shortly pubescent on parts exposed in bud.
Stamens 15, closely congested round the ovary and
style, subequal; filaments reaching the ovary apex,
lorate, abruptly tapering apically; anther as long as
filament, narrowly oblong; appendage to connective
short, stout, becoming slightly recurved. Ovary small,
globose, glabrous; stylopodium indistinct; style 3
times as long as ovary, glabrous, filiform. Fruit calyx
subglabrous; 3 longer lobes to 8 by 1.5 cm, spatulate,
obtuse, with to 10 by 7 mm ovate saccate thickened
base: 2 shorter lobes to 5 by 0.6 cm, linear, similar at
base. Nut to 18 by 12 mm, ovoid, densely buff
pubescent, acute.

Distr. Malesia: Borneo (Rejang valley to S.W.
Sabah).

Ecol. Local on deep yellow sandy soils on coastal
hills below 400 m.

95. Shorea elliptica BURCK, Ann. Jard, Bot. Btzg 6
(1887) 215; BRANDis J. Linn. Soc. Bot. 31 (1895) 99;
Merr. En. Born. (1921) 404; BRowneE, For. Trees
Sarawak & Brunei (1955) 138; ASHTON, Man. Dipt.
Brun. Suppl. (1968) 151, f. 13, pl. 20 (bark). — Fig. 94
B-B4.

Medium-sized tree. Twig, bud, stipule outside,
nerves and midrib beneath densely scabrid ocherous
to cream pubescent; midrib above evenly so; nerves
above and stipule within sparsely evenly so; leaf

1982]

DIPTEROCARPACEAE (Ashton)

SO]

Fig. 96. Shorea albida SyM., alan-forest, centre of peat swamp. Brunei (Photogr. ASHTON).

undersurface densely cream lepidote. Twig c. 2mm @
apically, much branched, terete to subcompressed,
smooth or rugulose; stipule scars short, horizontal.
Bud to 8 by 6 mm, ovoid, obtuse, compressed. Stipule
to 10 by 6 mm, narrowly ovate, acute, caducous.
Leaves 7-10 by 4-9 cm, elliptic to oblong, thickly
coriaceous; base broadly cuneate to subcordate; apex
obtuse or shortly broadly acuminate; nerves 11-16
pairs, prominent beneath, at 75° at base, 45°—SS5°
towards the apex; tertiary nerves slender, densely
scalariform, at c. 90° to the midrib; midrib shallowly
depressed above, prominent beneath; petiole 1.5—3.3
cm long, terete, rugose distally. Panicle to 12 cm long,
terminal or axillary, + compressed at first, becoming
terete, densely persistently ocherous scabrid pubes-
cent; singly or doubly branched, branchlets to 2 cm
long, bearing to 3 + distichous flowers; bracteoles to 2

by | mm, small, elliptic, obtuse, pubescent, caducous.
Flower bud to 4 by 3 mm, narrowly ovoid. Sepals
pubescent on parts exposed in bud, ovate, acute, the
inner 2 relatively shorter, narrower than the outer 3.
Petals pale yellow, pubescent on parts exposed in bud,
narrowly oblong. Stamens 19-20; filaments com-
pressed, slender, somewhat tapering; anthers oblong,
almost as long as filaments, crowned by a short
relatively stout glabrous appendage somewhat
shorter than the anthers. Ovary ovoid, glabrous; style
filiform, c. 2 times as long as ovary, trifid at apex,
glabrous. Fruit pedicel to 3 mm long, swelling into the
base of the fruit. Calyx sparsely shortly pubescent; 3
longer lobes to 8 by 1.5 cm, narrowly spatulate to
sublorate, obtuse, c. 4mm broad above the to 10 by 6
mm elliptic saccate + thickened base. Nut to 15 by 12
mm, ovoid, acute, densely evenly buff pubescent.

FLORA

MALESIANA

[ser. I, vol. 97

Fig. 97. Trunk of Shorea albida Sym. Brunei (Photogr.
ASHTON).

Distr. Malesia: Borneo (N.W. Kalimantan, W.
Sarawak).

Ecol. Local, leached soils in Mixed Dipterocarp
forest below 500 m.

Vern. Meéranti lang (Sar.).

96. Shorea dispar ASHTON, Gard. Bull. Sing. 31 (1978)
45. — Fig. 95 C-C4.

Large buttressed tree. Twigs, petioles, bud, pani-
cles, perianth outside and ovary densely + persis-
tently pale tawny pubescent, leaf nervation beneath
sparsely so. Twigs c. 2mm g apically, much branched,
terete, at first rugulose and + ribbed, becoming
smooth, dark brown dappled; stipule scars short,
dark, horizontal. Buds to 3 by 2mm, ellipsoid, obtuse.
Stipules unknown. Leaves 4-7 by 2-3.5 cm, elliptic,
coriaceous, + distinctly but sparsely cream lepidote
beneath; margin subrevolute; base cuneate; acumen
short, broad; nerves 9-11 pairs, ascending, prominent
beneath, obscure and narrowly depressed above as
also the midrib; petiole 12-16 mm long. Panicles to 8
cm long, terminal or axillary, rigid, ascending; singly

or doubly branched, branchlets to 3 cm long. Flower
buds to 6 by 4mm, long, ovoid. Sepals broadly ovate,
subequal, shortly subacuminate. Stamens c. 25; fila-
ments long, lorate, somewhat tapering to the oblong
anthers; appendages c. 3 length of anthers, slender,
tapering, glabrous, becoming + reflexed at anthesis;
ovary small, ovoid, glabrous, surmounted by a slender
filiform style c. twice its height. Fruit unknown.

Distr. Malesia: Borneo (Central Sarawak; once
collected).

Ecol. Mixed Dipterocarp forest on inland hills.

Note. Superficially resembling S. parvifolia DYER
ssp. velutinata ASHTON; the flowers betray the close
relationship of this rare tree with red lauan, S.
negrosensis FOxw., the celebrated timber tree of the
Philippines, and assign both to sect. Rubella ASHTON.

97. Shorea negrosensis Foxw. Philip. J. Sc. 6 (1911)
Bot. 274, pl. 44; ibid. 13 (1918) Bot. 192; ibid. 67 (1938)
315; Merr. En. Philip. 3 (1923) 97; Reyes, Philip. J.
Sc. 22 (1923) 327.

Large buttressed tree. Twigs, buds, stipules, midrib
above, panicles, parts of petals exposed in bud and
ovary densely persistently yellow-brown pubescent;
petiole, leaf beneath and calyx sparsely so; bracteoles
puberulent. Twig c. 2mm g apically, terete, becoming
smooth; stipule scars short, obscure. Buds to 2 by 1
mm, conical; stipules to 15 by 4mm, lanceolate, acute.
Leaves 6.5-17 by 3-7.5 cm, ovate to elliptic, thinly
coriaceous; base cuneate to obtuse, frequently un-
equal; acumen to 2 cm long, slender, prominent,
tapering; nerves (8—)11—15 pairs, slender but promi-
nent beneath, arched, at 60°—-70°, frequently with
small tomentose axillary domatia; tertiary nerves
scalariform, elevated beneath; midrib prominent
beneath, evident but shallowly depressed above;
petiole 18-25 mm long, slender but geniculate. Panicle
to 14 cm long, slender; singly branched, branchlets to
15mm long, rather short, bearing to 4 secund flowers;
bracteoles to 4 by 3 mm, ovate, obtuse, caducous.
Flower bud to 7 by 4 mm, broadly ellipsoid; sepals
ovate, the outer 3 acute, the inner 2 acuminate;
stamens c. 48, subequal, exceeding style at anthesis;
filaments very long and slender, lorate, slightly taper-
ing; anthers linear-oblong; appendages to 1/8 length
of anther but usually much shorter, slender, glabrous;
ovary ovoid, puberulent; style shorter than ovary,
columnar, glabrous. Mature fruit unknown. Fruit
subsessile; 3 longer calyx lobes c. 7 by 1.3 cm,
spatulate, obtuse, c. 9 mm broad above the 10 by 10
mm suborbicular thickened saccate base; 2 shorter
lobes to 30 by 4 mm, linear-lorate, acute, similar at
base; nut ovoid, apiculate.

Distr. Malesia: Philippines.

Ecol. Widespread, common and often gregarious,
in Evergreen and Semi-evergreen + seasonal Diptero-
carp forests in lowlands.

Vern. Red lauan, mangachapuy.

Note. Some sterile collections, apparently of this
species, bear significantly smaller leaves; flowering
material is needed to establish their identity.

1982] DIPTEROCARPACEAE (Ashton) 503

7. Section Brachypterae

HeEIM, Rech. Dipt. (1892) 46; BRANDis, J. Linn. Soc. Bot. 31 (1895) 76, p.p.;
ASHTON, Gard. Bull. Sing. 20 (1963) 270; Man. Dipt. Brun. (1964) 117. — Shorea,
Red Meranti group, S. pauciflora subgroup Sym. Mal. For. Rec. 16 (1943)
58. — Fig. 91 D, 98-100.

Flower buds + ovoid. Corolla as in sect. Rubella. Filament broad and
compressed at base, tapering + abruptly medially and filiform below anthers;
anthers 4-celled, subglobose or broadly oblong; appendage to connective aristate,
14-34 times length of anthers. Ovary with distinct stylopodium: ovary and
stylopodium + pyriform; or without distinct stylopodium but with style
frequently pubescent towards base. Panicle lax, branchlets long. Stipules and
bracteoles sometimes somewhat persistent. Leaf with scalariform tertiary nerves;
midrib evident, elevated, or obscurely depressed above. Vessels solitary or in
small groups; ray cells without silica. Medium-sized or large stoutly prominently

buttressed trees.

Distr. Non-seasonal western Malesia including the Philippines; one species in the Moluccas.

Ecol. Scattered in lowland forests and on mountain ridges (S. venulosa, S. monticola) to 1800 m; a few species
(S. balangeran, S. selanica) are gregarious, even forming pure stands.

Vern. Red méranti, méranti merah (Mal., Sum.), mérawan (Iban), red lauan (Philippines).

Note. The floral morphology is closely similar to the pattern of sect. Anthoshorea, making assignment
difficult without anatomical investigation. With the notable exception of S. parvistipulata and the clearly closely
related short-sepalled S. fallax, species in this section are well defined; though several, e.g., S. pauciflora, S.
Johorensis, S. almon, and S. kunstleri do exhibit some local variation.

7a. Subsection Smithiana

ASHTON, Gard. Bull. Sing. 20 (1963) 270; Man. Dipt. Brun. (1964) 118. — Fig. 98.

Stamens 22-26. Style very short, ovary with prominent columnar stylopodium.
Bark surface with deep V-section fissures, inflaked, as in sect. Mutica but with
sheet-like rhytidome layers as in sect. subsect. Brachypterae and other flaky-

barked groups.

98. Shorea smithiana Sym. Gard. Bull. S. S. 9 (1938)
345, pl. 26; ASHTON, Man. Dipt. Brun. (1964) 223, f.
17; ibid. Suppl. (1968) 119; Meer & Woop, Sabah
For. Rec. 5 (1964) 148, pl. 10 (habit), f. 16. — Shorea
sp. (ELMER 21606) SLoot. ex Merr. Pl. Elm. Born.
(1929) 205. — Fig. 98.

Large buttressed tree. Young twig, leaf bud, stipule
outside (puberulent within), petiole, leaf beneath,
midrib above and panicle persistently grey-brown to
rust scabrid tomentose, leaf above caducous puberu-
lent. Twig 3-4 mm @g apically, stout, ribbed, smooth,
glabrous; stipule scars to 2 mm long at first, broad,
falcate, descending. Bud 5-8 by 3-4.5 mm, ovoid,
slightly compressed. Stipule to 20 by 6 mm, broadly
hastate, subacute. Leaves 12-20 by 6.5-11.5 cm,
broadly obovate to oblong or ovate, grey lepidote
beneath, coriaceous; base obtuse or subcordate;
acumen to 1.5 cm long, broad; nerves 14-17 pairs,
prominent beneath, curved, at 50°—60° distally, more
spreading at base; tertiary nerves scalariform, well

spaced, sinuate; midrib prominent, stout, rounded,
beneath, rather narrow and slightly depressed above;
petiole 2.2-2.8 cm long, stout. Panicle to 25 cm long,
terminal or axillary, terete, regularly singly branched,
the branchlets bearing to 7 secund flowers; bracteoles
to 5 by 4 mm, ovate, subacute, shortly pubescent on
both surfaces, falling before the corolla opens. Flower
bud to 10 by 3 mm, narrowly ovoid, subacute. Calyx
densely pubescent outside, glabrescent within; lobes
broadly deltoid to ovate, acute, subequal, the 2 inner
the narrower. Petals pink, narrowly oblong, short,
densely yellowish brown pubescent on parts exposed
in bud. Stamens 22-26, of 3 sizes; filaments basally
expanded, abruptly tapering and filiform distally;
anthers broadly oblong; appendage to connective
almost twice as long as anthers, reaching to } length of
ovary and style, not reflexed. Ovary and stylopodium
narrowly pyriform, puberulent; style cylindrical, gla-
brous. Fruit calyx glabrescent; 3 longer lobes to 20 by
2.8 cm, narrowly spatulate, coriaceous, narrowly

504

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 98. Flower details in Shorea sect. Brachypterae HEIM subsect. Smithiana ASHTON. — S. smithiana Sy. A.
Bud, B/. outer sepal, B2. inner sepal, C/. stamens from outside, C2. stamens from inside, D. pistil, all x 10 (bb.
34816).

obtuse, c. 1.3 cm broad and often slightly revolute
above the to 1.5 by 1.8 cm broadly ovate shallowly
saccate thickened base; 2 shorter lobes to 13 by 0.9 cm,
linear, similarly expanded at base. Nut to 2.7 by 1.8
cm, ovoid; style remnant to 3.5 mm long, taper-
ing, densely evenly shortly pale cream-buff pubes-
cent.

Distr. Malesia: Eastern Borneo (N.E. Sarawak,
Sabah, S.E. Borneo to Sampit).

Ecol. Frequent on deep sandy clay soils on undu-
lating land and to 400 m.

Uses. One of the chief sources of light red meranti
timber in N.E. Borneo.

Vern. Chémpaga (Nunukan), lempong mérémbong,
l. tembaga, awang (Kutei), mahambung (Sampit),
séraya timbau (Sabah), méranti rambai, méraka belong
(Brunei), éngkabang (mangkabang), rambai (Iban),
bérat (Murut).

7b. Subsection Brachypterae

ASHTON, Gard. Bull. Sing. 20 (1963) 271. — Shorea sect. Pachychlamys DYER ex
KING, J. As. Soc. Beng. Sc. 62, 2 (1893) 109. — Pachychlamys (DYER ex KING)
DYER ex RIDL. FI. Mal. Pen. 1 (1922) 233. — Fig. 91D, 99-100.

Stamens 15. Style generally as long as or longer than ovary. Bark surface
square-section fissured, generally appearing flaky rather than fissured; phloem
matrix proliferation tissue with pale stone cells in conspicuous simple laminae;

phelloderm thin, inconspicuous.

1982]

99. Shorea inaequilateralis Sym. Gard. Bull. S. S. 8
(1935) 281, pl. 25; BRowng, For. Trees Sarawak &
Brunei (1955) 153; ANDERSON, Gard. Bull. Sing. 20
(1963) 158; ASHTON, Man. Dipt. Brun. (1964) 191, f.
17; ibid. Suppl. (1968) 109. —*Semayor’ DuRANT,
Rep. For. Brunei (1933) 37, 42.

Large, buttressed, fissure-barked tree with pendent
branches. Young parts shortly densely evenly pale
tanny tomentose, persistent on young twig, panicle,
leaf bud, outside of stipule, petiole and midrib above;
persistently puberulent, appearing glabrous, on inside
of stipule, lamina on both surfaces, and midrib
beneath. Twig c. 2.5 by 1.5 mm @ apically, slightly
compressed at first, becoming terete with a decurrent
rib below the petiole insertion each side, becoming
glabrous, smooth; stipule scars to 1.5 mm long,
narrowly falcate, descending, obscured by tomentum.
Bud 3-6 by 1.5—2.5 mm, ovoid, slightly compressed,
acute. Stipule to 20 by 5 mm, subpersistent, narrowly
hastate, acute. Leaves 9-14 by 4.5—7.5 cm, ovate, thin;
base subcordate, distinctly unequal; acumen to 2 cm
long, prominent, caudate; nerves | 1—13 pairs, slender,
raised beneath, at more than 90° at base, ascending
towards apex; tertiary nerves slender, scalariform,
well spaced, at c. 90° to nerves; midrib narrow,
prominent, rounded beneath, evident but applanate
above; petiole 5-8 mm long, short, stout. Panicle to 7
cm long, terminal or axillary, terete, pendent; singly
branched, branchlets to 1.5 cm long, bearing to 4
flowers; bracteoles to 10 by 3 mm, narrowly deltoid,
pubescent outside, glabrescent within. Flower bud to
12 by 3 mm, linear, acute. Calyx densely shortly
pubescent on both surfaces; 3 outer lobes deltoid,
subacute; 2 inner lobes shorter, prominently acu-
minate. Petals narrowly hastate, subacute, sparsely
pubescent on parts exposed in bud. Stamens 15, in 3
subequal verticils; filaments broad, compressed at
base, tapering abruptly medially and filiform distally;
anther oblong; appendage to connective c. 3 times
length of anther, reaching 4 length of style, slender,
erect, scabrous towards apex. Ovary ovoid, tapering,
sparsely pubescent, without distinct stylopodium;
style about twice length of ovary, filiform, sparsely
pubescent in the basal half, otherwise glabrous. Fruit
calyx glabrescent but for the sparsely tawny pubes-
cent base; 3 longer lobes to 14 by 2mm, elliptic saccate
thickened base; 2 shorter lobes to 12 by 0.9 cm, long,
narrowly spatulate, acute, to 4mm broad above the to
13 by 9 cm, unequal, narrowly spatulate, similar at
base. Nut to 1.8 by 1.4 cm, ovoid, densely buff
pubescent; style remnant to 6 mm long, slender,
subpersistent, filiform.

Distr. Malesia: N.W. Borneo (Sarawak, Brunei).

Ecol. Locally abundant in Mixed Peat Swamp
forest.

Uses. A valuable heavy construction timber.

Vern. Sémayur (Brun.).

Note. An isolated species apparently related to the
following.

100. Shorea selanica BL. Mus. Bot. Lugd.-Bat. 2

DIPTEROCARPACEAE (Ashton)

505

(1852) 33, incl. var. latifolia BL.; WALP. Ann. 4 (1857)
338; DC. Prod. 16, 2 (1868) 629; HaNceE, J. Bot. 14
(1876) 242; BurcK, Ann. Jard. Bot. Btzg 6 (1887) 216,
incl. var. obtusa BURCK; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 86; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1,
3, 6 (1895) 266; Merr. Int. Rumph. (1917) 375;
HEYNE, Nutt. Pl. ed. 1, 3 (1917) 306; ibid. ed. 2 (1927)
1124; Stoor. Reinwardtia 2 (1952) 50, f. 16,
17. — Dammara selanica RumpH. [Herb. Amb. 2
(1743) 168, t. 56] ex LAMK, Encycl. 2 (1786) 259,
inval. — Unona selanica DC. Prod. | (1824) 92. — En-
gelhardtia selanica BL. Fl. Jav. 1 (1828) 8. — Hopea
selanica W. & A. Prod. (1834) 85; Hassk. Cat. Hort.
Bog. (1844) 209; WaALP. Rep. 5 (1845) 128; Mia. FI.
Ind. Bat. 1, 2 (1959) 504; DC. Prod. 16, 2 (1868) 635.

Vast buttressed tree. Young parts, twigs, buds,
stipules, leaf undersurface, petiole, panicle, ovary and
parts of petals exposed in bud densely persistently pale
cream-brown pubescent; densely so at first on calyx,
becoming sparsely so; young trees sparsely darker
pubescent. Twigs 2-3 mm @ apically, ribbed and
sometimes compressed at first, becoming terete,
smooth; stipule scars short, pale, horizontal. Bud c. 2
by 1 mm, ovoid, obtuse; stipules to 10 by 4 mm,
lanceolate, acute, + fugaceous. Leaves (7—-)9-18 by
3-7 cm (larger in immature trees), oblong-ovate,
thinly coriaceous, cream below (mature trees); base
obtuse to subcordate, + unequal, apex shortly acu-
minate; nerves 19-23 pairs, slender but prominent
beneath, + applanate above, at c. 65°; tertiary nerves
densely scalariform, hardly elevated beneath; midrib
prominent beneath obscurely depressed above; petiole
8-12 mm long, slender. Panicles to 15 cm long,
slender, somewhat lax, terminal or axillary; singly
branched, branchlets to 3 cm long, bearing to 7
flowers. Flower buds to 6 by 3 mm, fusiform; sepals
broadly ovate, the outer 3 acute, the inner 2 acu-
minate; stamens 15, unequal; filaments broadly com-
pressed at base, tapering medially and filiform
beneath the broadly ellipsoid anthers; appendages
very slender, scarious near the apices, twice length of
anthers; ovary ovoid, puberulent; style filiform,
slender, twice length of ovary, glabrous. Fruit pedicel
c. 1mm long, slender; 3 longer calyx lobes to 10 by 1.5
cm, narrowly spatulate, obtuse, c. 5 mm wide above
the 7 to 8 mm ovate saccate thickened base; 2 shorter
lobes to 4.5 by 0.6 cm, linear, similar at base. Nut to 15
by 8 mm, ovoid, apiculate.

Distr. Malesia: Moluccas (Buru; Sula Is.: Man-
gole, Sanana; Obi Is.; Ambon).

Ecol. Gregarious and dominant in the Moluccan
semi-evergreen lowland forests on well drained land
with fertile soils, sometime overlying limestone.

Uses. The most valuable construction timber of the
Moluccas.

Vern. Kayo bapa (Buru, Ambon, Sanana, Mangole),
sehu, boba (Samuja), luma, bahut, biahgawa (Buru).

Note. Two sterile collections (bb. 22808, 31349
from Buru) differ in their larger leaves drying distinct
purplish beneath, with relatively longer petioles (cf.
VAN SLOOTEN, /.c. 60: Shorea ? spec. nov.).

506 FLORA MALESIANA [ser. I, vole

Fig. 99. Flower details in Shorea sect. Brachypterae HE subsect. Brachypterae. All x 10. Sepals drawn from

inside. — S. scaberrima BuRcK. A. Bud, A/. outer sepal, A2. inner sepal, 43. stamens from outside, A4.

pistil. — S. venulosa Woop ex Meurer. B. Bud, B/. outer sepal, B2. inner sepal, B3. stamens from outside, B4.
pistil (A Cult. Hort. Bog. VIII-D-54, B Saini 9486).

1982]

The only red meranti east of Wallace’s line, this
striking species enigmatically shares its characteristic
leaf shape, as well as its pendent inflorescence, flower
structure and very hard durable wood with that
equally distinctive denizen of the N.W. Borneo peat
swamps S. inaequilateralis. If only their evolution
could be traced!

101. Shorea flemmichii Sym. Gard. Bull. S.S. 10 (1939)
378, pl. 26; BROwNE, For. Trees Sarawak & Brunei
(1955) 147; AsHton, Man. Dipt. Brun. (1964) 190, f.
17, pl. 48 (bark); ibid. Suppl. (1968) 108.

Large tree with dark deeply fissured flaky bark and
stout buttresses. Young twig, panicle, leaf bud, stipule
outside (glabrescent within), petiole, midrib beneath
and at base above and nervation beneath densely
persistently scabrid golden-tawny tufted tomentose;
leaf above scabrid puberulent. Twig stout though
tapering to 2 mm @ apically, much branched, terete,
verruculose; stipule scars short, horizontal, obscured
by the tomentum. Bud 3-6 by 2-4 mm, ovoid, slightly
compressed, subacute. Stipule to 8 by 2.5 mm,
narrowly hastate, acute, fugaceous. Leaves 5-9 by
34.5 cm, broadly elliptic or ovate (young trees),
cream lepidote beneath; base obtuse; acumen to 5mm
long, narrow; margin revolute; nerves 14-17 pairs,
dense, prominent beneath, at 45°65"; tertiary nerves
scalariform, sinuate, diagonal to nerves; midrib nar-
row and slightly depressed above, prominent beneath;
petiole 7-9 mm long, short. Panicle to 8 cm long,
terminal or axillary, terete or slightly compressed;
singly or doubly irregularly branched, branchlets to 5
cm long, lax, rather zig-zag, bearing to 8 distichous
flowers; bracteoles to 6 by 5mm, suborbicular, obtuse,
shortly puberulent outside, glabrous within. Flower
bud to 4.5 by 3.5 mm, subglobose, obtuse. Calyx
densely tawny pubescent outside, glabrous within; 3
outer lobes somewhat longer, narrowly ovate, sub-
acute; 2inner lobes narrowly deltoid, subacute. Corolla
dark wine-red, strongly contorted, apically recurved,
the imbricate bases forming a deep cup; petals
oblong-lanceolate, obtuse, twisted, densely tomentose
on parts exposed in bud. Stamens 15, the 5 inner
slightly longer than the others; filaments broad at
base, tapering abruptly and filiform distally; anthers
subglobose; appendage to connective c. 3 times as
long as anther, reaching almost to style apex. Ovary
and stylopodium ovoid-conical, puberulent towards
the apex, tapering; style glabrous, filiform, as long as
ovary. Fruit subsessile. Calyx puberulent; 3 longer
lobes to 6.5 by 1.5 cm, spatulate, narrowly obtuse, to
2.5 mm broad above the to 7 by 6 mm ovate-elliptic
shallowly saccate slightly thickened base; 2 shorter
lobes to 20 by 1.5 mm, linear, similar at base. Nut
subacute, ovoid, to 1.5 by | cm, shortly densely pale
buff pubescent.

Distr. Malesia: N.W. Borneo (Sarawak & Brunei).

Ecol. Very local, deep yellow sandy soils in Mixed
Dipterocarp forest below 400 m, on present or
Pleistocene coastal hills.

Vern. Méranti raya (Brun.).

DIPTEROCARPACEAE (Ashton)

507

102. Shorea almon Foxw. Philip. J. Sc. 67 (1938) 313,
pl. 7; BROWNE, For. Trees Sarawak & Brunei (1955)
150; MEUER & Woop, Sabah For. Rec. 5 (1964) 93, f.
1; ASHTON, Man. Dipt. Brun. Suppl. (1968) 104, f.
13.— S. furfuracea (non Mig.) ROLFE, J. Bot. 23
(1885) 110; VipaL, Rev. Pl. Vasc. Filip. (1886) 62;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 98; Foxw.
Philip. J. Sc. 4 (1909) Bot. 517; WHITFORD, Philip. J.
Sc. 4 (1910) Bot. 712; Bull. Bur. For. Philip. 10 (1911)
63. — S. eximia (non (Mi1Q.) SCHEFF.) Foxw. Philip. J.
Sc. 6 (1911) Bot. 276; ibid. 13 (1918) Bot. 191; MERR.
En. Philip. 3 (1923) 96; Reyes, Philip. J. Sc. 22 (1923)
331. — Fig. 7.

Large buttressed tree. Twig, bud, stipule outside,
petiole and leaf nervation beneath densely persistently
pink-brown scabrid tomentose; stipule inside and
nervation above sparsely, midrib above densely
evenly so. Twig c. 2 mm g apically, terete, smooth;
stipule scars short, horizontal, obscure. Bud to 4 by 3
mm, ovate, compressed, acute. Stipule to 10 by 5mm,
lanceolate, acute, caducous. Leaves 9-16 by 3.5—7 cm,
chartaceous, undulate, the lower surface markedly
concave; base broadly cuneate to obtuse; acumen to 8
mm long, short, broad; nerves 1 7—20 pairs, slender but
prominent beneath, at 40°—60° (to 100° at the base);
tertiary nerves densely scalariform, at 90° to midrib,
applanate above, prominent beneath; petiole 10-18
mm long, slender. Panicle 20 cm long, terminal or
axillary, terete, slender, ascending, lax, densely
shortly unevenly persistently pale pink-brown pubes-
cent; singly branched, branchlets to 2.5 cm long,
bearing to 10 distichous flowers; bracteoles to 5 by 3
mm, elliptic, obtuse, yellow-brown puberulent, per-
sistent until the formation of young fruit. Flower bud
to 7 by 3 mm, lanceolate, small. Sepals pubescent on
parts exposed in bud; 3 outer deltoid, acute, 2 inner
ovate, acuminate, smaller. Petals lanceolate, hirsute
on parts exposed in bud. Stamens 15, in 3 unequal
verticils; filaments compressed at base, abruptly
tapering medially and filiform below the subglobose
anthers; appendage to connective slender, glabrous, c.
3xlength of anther. Ovary ovoid, pubescent; style
filiform, to 15 x length of ovary, pubescent in the basal
4. Fruit pedicel to 3 mm long, prominent. Calyx pale
yellow-brown pubescent at base, glabrescent distally;
3 longer lobes to 14 by 2.5 cm, broadly spatulate,
obtuse, c. 8 mm broad above the to 13 by 12 mm ovate
thickened saccate base; 2 shorter lobes to 8 by | cm,
lorate, obtuse, similar at base. Nut to 2 by 1.5 cm,
ovoid, very shortly mucronate at the otherwise obtuse
apex, shortly evenly pale buff pubescent.

Distr. Malesia: N.E. Borneo (N.E. Sarawak,
rare; E. Sabah, Tidung), Philippines (non-seasonal
areas).

Ecol. Undulating hills in Mixed Dipterocarp for-
est, clay soil.

Vern. Séraya kérukup (Sabah), almon (Philip-
pines).

Note. Particularly variable in Mindanao, where
the leaves are very small, and the tomentum more
sparse and scabrid in some trees.

508

FLORA MALESIANA

[ser. I, vol. 9?

D1

Fig. 100. Flower details in Shorea sect. Brachypterae HEIM subsect. Brachypterae. All x 10. Sepals drawn from
inside. — S. flaviflora Woop ex ASHTON. C. Bud, C/. stamens from outside, C2. pistil. — S. johorensis Foxw. D.
Outer sepal, D/. inner sepal, D2. stamens from outside, D3. pistil (C ASHTON 5664, D SAN 31246).

103. Shorea parvistipulata Heim, Bull. Mens. Soc.
Linn. Paris 2 (1891) 974; ASHTON, Gard. Bull. Sing. 31
(1978) 46.

KEY TO THE SUBSPECIES

i Leaf not lepidote beneath.
. Nerves 13-21 pairs. Longer fruit calyx lobes to 20
cm long a. ssp. parvistipulata
2. Nerves at most 15 pairs. Longer fruit calyx lobes
to 9 cm long . . b. ssp. nebulosa
. Leaf silvery lepidote beneath. Nerves 13-21 pairs
c. ssp. albifolia

a. ssp. parvistipulata. — BRANDIS, J. Linn. Soc. Bot. 31
(1895) 95; Merr. En. Born. (1921) 406 (‘parvistipula’);
MEER & Woop, Sabah For. Rec. 5 (1964) 132;
ASHTON, Man. Dipt. Brun. Suppl. (1968) 114, f.

14. — S. squamata (non BENTH. & HooK. f:) BRANDIS,
J. Linn. Soc. Bot. 31 (1895) 92; MerrR. En. Born.
(1921) 407, p.p. quoad sp. Born. — S. dyeri (non THW.
ex TRIM.) HEIM, Bull. Mens. Soc. Linn. Paris 2 (1891)
957. — S. cristata BRANDIS, J. Linn. Soc. Bot. 31
(1895) 97; MerR. En. Born. (1921) 404; ASHTON, Man.
Dipt. Brun. (1964) 106, f. 17; ibid. Suppl. (1968) 106;
MEWER & Woop, Sabah For. Rec. 5 (1964) 102.
Very large thinly scaly-barked buttressed tree. Leaf
bud, stipule outside, twig, petiole and leaf nervation
beneath shortly persistently pale yellow to grey-brown
scabrid tomentose, stipule inside and midrib above
evenly, not scabridly, so. Twig c. 2 mm @, terete,
slender, becoming smooth; stipule scars short, pale,
prominent, horizontal or descending. Bud to 6 by 4
mm, ovoid, acute. Stipule to 14 by 7 mm, ovate,
subacute. Leaves 6-20 by 3-9 cm, very variable in size
and shape, + oblong, chartaceous; base typically

1982]

DIPTEROCARPACEAE (Ashton)

509

cordate, sometimes broadly cuneate or obtuse;
acumen to | cm long; nerves 13-21 pairs, usually
slender, at 50°—80° except at the base, without second-
ary nerves; tertiary nerves slender, densely scalari-
form; midrib evident but applanate to depressed
above, prominent beneath; petiole 12-15 mm long,
short, terete. Panicle to 16 cm long, terminal or
axillary, terete, lax, straight, ascending, densely
shortly persistently pale brown scabrid pubescent:
singly branched, branchlets to 4cm long, bearing to 11
distichous flowers; bracteoles to 5 by 3 mm, deltoid,
acute, puberulent, not at first caducous. Flower bud to
7 by 3 mm, lanceolate. Sepals pubescent on parts
exposed in bud; 3 outer narrowly deltoid, acute, 2
inner ovate, narrowly acuminate, shorter and thinner
at margins than outer 3. Petals cream suffused with
pink at base, lanceolate, pubescent on parts exposed
in bud. Stamens 15, in 3 unequal verticils; filaments
broad and compressed at base, tapering abruptly and
filiform beneath the anthers; appendage to connective
slender, glabrous, about twice length of anthers.
Ovary ovoid, densely pubescent; style filiform, about
as long as ovary, glabrous but for a pubescent ring at
base. Fruit pedicel to 2 mm long, to 3 mm @. Calyx
puberulent towards base, glabrescent distally; 3
longer lobes to 20 by 2.5 cm, lorate-spatulate, obtuse,
c. 11 mm broad above the to 14 by 18 mm broadly
ovate saccate thickened base; 2 shorter lobes to 8 by
0.4 cm, linear, similar at base. Nut to 2.5 by 2 cm,
ovoid, frequently crowned by a persistent filiform
style remnant, densely evenly buff pubescent.

Distr. Malesia: Borneo (excepting S.W.).

Ecol. Widespread on clay-rich soils on alluvium
and especially hillsides and low ndges to 1300 m.

Vern. Séraya lupa (Sabah), éngkabang pinang, e. p.
bérsisek (Sarawak), kawang pinang (Brunei), ténger-
awan kuning (Nunukan), kénuar kepas (S.E. Borneo).

Note. I no longer consider that S. cristata can be
maintained as a separate species. S. palosapis (BLCO)
Merk. is undoubtedly closely allied but differs in its
amplexicaul stipule scars, large, deltoid, subpersistent
stipules, and its oblong irregular crown, branching
from low on the bole.

b. ssp. nebulosa (MEIJER) ASHTON, Gard. Bull. Sing. 31
(1978) 46. — S. nebulosa MEUER, Act. Bot. Neerl. 12
(1963) 337; MEIER & Woop, Sabah For. Rec. 5 (1964)
123:

Leaves at most 13 by 6 cm, with at most 15 pairs of
nerves. Fruit calyx lobes at most 9 by 1.8 cm.

Distr. Malesia: N.E. Borneo (Crocker Range and
Mt Kinabalu region).

Ecol. Hill forests between 800—1300 m.

c. ssp. albifolia ASHTON, Gard. Bull. Sing. 31 (1978) 46.
Differing in the silvery pink lamina undersurface.
Distr. Malesia: Borneo: N.E. Sarawak (Niah),

Brunei and S.E. Sabah southwards to Balikpapan.
Ecol. Fertile soils on undulating land and periodi-

cally inundated alluvium.

Note. A collection of this subspecies from E.

Kalimantan with short fruit sepals emphasises the
doubtful distinctness of S. fallax from this species.

104. Shorea balangeran (KoRTH.) BuRCK, Ann. Jard.
Bot. Btzg 6 (1887) 214; BRANDIs & GiLG in E. & P. Pf.
Fam. ed. 1, 3, 6 (1895) 297; BRANpbis, J. Linn. Soc. Bot.
31 (1895) 86; BoERL. Cat. Hort. Bog. 2(1901) 108, incl.
var. angustifolia BOERL.; HEYNE, Nutt. Pl. ed. 1, 3
(1917) 297; ibid. ed. 2 (1927) 1115; Merr. En. Born.
(1921) 404; SLoor. Bull. Bot. Gard. Btzg III, 18 (1949)
256, f. 12.— Hopea balangeran Kortu. Kruidk.
(1841) 74, t. 7 f. 1-14; BL. Mus. Bot. Lugd.-Bat. 2
(1852) 34; Wacp. Rep. 5 (1845) 128; Ann. 4(1857) 339:
Mia. FI. Ind. Bat. 1, 2 (1859) 503; Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 84, 85; DC. Prod. 16, 2 (1868) 634:
HEYNE, Nutt. Pl. ed. 2 (1927) 1115.— Parahopea
balangeran Heim, Rech. Dipt. (1892) 66.— Para-
shorea balangeran Mer. En. Born. (1921) 404 — Fig.
91 D-D4.

Large buttressed tree. Twig, leaf bud, stipule.
petiole, midrib and nerves beneath, panicle and calyx
outside shortly densely persistently red-brown puber-
ulent; leaves beneath cream lepidote; nut apex spar-
sely puberulent. Twigs c. 2 mm @ apically, slender,
terete, smooth. Buds 5 by 3 mm, ovoid, acute; stipule
to 7 by 4 mm, ovate, acute. Leaves 7-18 by 3-8 cm.
oblong-elliptic to lanceolate, coriaceous. conspi-
cuously cream beneath: base broadly cuneate to
obtuse; acumen to 1.5 cm long, slender, tapering:
nerves 13-18 pairs, slender, barely elevated beneath,
evident above, at 45°—55°; tertiary nerves densely
scalariform, obscure; midrib slender but prominent
beneath, obscurely depressed above; petioles 2-3 cm
long, slender. Panicle to 8 cm long, slender, terminal
or axillary; doubly branched, branchlets to 3 cm long.
with to 7 secund flowers. Flower buds to 6 by 3 mm,
fusiform; 3 outer sepals narrowly lanceolate, sub-
acute; 2 inner short, ovoid, prominently acuminate.
Stamens 15, shorter than style at anthesis; filaments
compressed at base, gradually tapering and filiform
below the broadly ovoid anthers; appendages very
slender, glabrous, twice length of anthers; ovary
ovoid, puberulent; style columnar, c. 14 x length of
ovary, puberulent towards base, not trifid. Fruit
pedicel short, slender. 3 longer calyx lobes to 50 by 7
mm, spatulate, obtuse, c. 2 mm broad above the to 4
by 3 mm ovate saccate thickened base; 2 shorter lobes
to 15 by 2 mm, linear, similar at base. Nut to 6 by 4
mm, ovoid, apiculate.

Distr. Malesia: Borneo (throughout Indonesian
part), Sumatra (Banka, Billiton).

Ecol. Common, often gregarious, in peat swamp
forests.

Vern. Balangeran, kawi, kahoi, tomi, kelansauw
(Borneo), malangsir (Banka).

Uses. A major timber produced in the swamps of
Indonesian Borneo.

Note. In many ways resembling S. albida (q.v.).

105. Shorea coriacea BuRcCK, Ann. Jard. Bot. Btzg 6
(1887) 214; BRaANpis, J. Linn. Soc. Bot. 31 (1895) 100;

510

Mere. En. Born. (1921) 404; BRowngE, For. Trees
Sarawak & Brunei (1955) 146; ASHTON, Man. Dipt.
Brun. (1964) 182, f. 17; ibid. Suppl. (1968) 106; MEER
& Woop, Sabah For. Rec. 5 (1964) 102.

Large buttressed tree with dark brown, fissured,
flaky bark. Young parts densely clothed in minute
adpressed pale grey hairs, caducous on all but panicle,
stipule outside and bud. Twig 2-3 mm @ at apex,
stout, terete or slightly compressed, smooth or rugu-
lose; stipule scars at first c. 1 mm wide, c. 1.5 mm long,
cuneate, pale brown, horizontal or slightly descend-
ing. Bud to 6 by 3 mm, ovoid. Stipule to 20 by 5 mm,
narrowly oblong, acute. Leaves 10-15 by 5-8 cm,
ovate, coriaceous, lustrous; base obtuse, subpeltate;
acumen c. | cm long; margin revolute; nerves 16—20
pairs, slender, hardly raised beneath, at 60°—70°,
curved towards margin; tertiary nerves slender, den-
sely scalariform; midrib prominent beneath, obscure
and slightly depressed above; petiole 2.5—4 cm long,
geniculate, glabrous. Panicle to 14 cm long, terminal
or axillary, + terete; singly (doubly if terminal)
branched, branchlets regular, short, bearing to 10
secund flowers; bracteoles to 4 by 2.5 mm, ovate,
subacute, sparsely pubescent outside, glabrescent
within. Flower bud to 6 by 1.2 mm, narrowly lanceo-
late, acute. Calyx densely pale grey pubescent outside,
glabrous within; 3 outer lobes narrowly deltoid,
subacute, slightly revolute; 2 inner lobes smaller,
acute, not revolute. Petals pink, linear, densely pubes-
cent on parts exposed in bud. Stamens 15, of 2 lengths,
the inner 5 an anther’s length longer; anther subglo-
bose; appendage to connective 2—3 times as long as
anther, slender, not reflexed, reaching almost to style
apex. Ovary ovoid, densely shortly pubescent except
at the base; style filiform, as long as ovary, glabrous
but for the setose base; stylopodium indistinct. Fruit
calyx shortly sparsely pubescent or glabrescent except
towards base; 3 longer lobes to 9 by 1.5 cm, spatulate;
chartaceous, narrowly obtuse, 3-4 mm broad above
the c. 8 by 5 mm deltoid thickened saccate base; 2
shorter lobes to 3.5 cm long, subequal, linear, similar
at base. Nut c. 18 by 9 mm, ovoid, shortly grey
pubescent distally; style remnant c. 3 mm long,
slender, acute.

Distr. Malesia: Borneo (Lower Kapuas, Sarawak,
Brunei, S.W. Sabah, Sandakan Distr., W. Kutei, and
Muara Teweh).

Ecol. Heath forest on podsols, terraces and pla-
teaux to 1000 m; on ultrabasics in E. Borneo.

Vern. Samar benua, méranti jurai (W. Borneo),
lampong méngkabang (S.E. Borneo), méranti tangkai
panjang (Sarawak), séraya t.p. (Sabah).

106. Shorea venulosa Woop ex MeverR, Act. Bot.
Neerl. 12 (1963) 342, p. 10; Sabah For. Rec. 5 (1964)
153; ASHTON, Man. Dipt. Brun. (1964) 226, f. 17; ibid.
Suppl. (1968) 122. — Fig. 99 B-B4.

Large buttressed tree with dark brown, fissured,
flaky bark. Young parts at first densely dotted with
minute adpressed hair tufts, fugaceous on all but
stipule outside and leaf bud; stipule persistently

FLORA MALESIANA

[ser. I, vol. 9?

shortly pubescent within. Twig c. 1 mm g apically,
slender, much branched, terete, rugulose or smooth,
glabrous; stipule scars narrow, slightly paler at first,
almost or completely amplexicaul, + ascending. Bud
to 7 by 2 mm, fulvous pale grey, lanceolate, com-
pressed, acute. Stipule to 20 by 3.5 mm, hastate, acute,
fugaceous. Leaves 6-10 by 3-5 cm (rarely much
smaller in exposed places), ovate, coriaceous; base
obtuse; acumen to 6 mm long; margin sometimes
slightly revolute; nerves 15-18 pairs, slender, hardly
or not elevated beneath, dense, at 35°—-40°, curved
towards the margin, sometimes with prominent axil-
lary pore-like domatia; tertiary nerves very slender,
densely scalariform, diagonal to nerves; midrib raised
beneath, obscure and depressed above; petiole 1.7-3
cm long, slender, geniculate. Panicle to 11 cm long,
terminal or axillary, slender, lax, + terete, shortly
densely persistently cream to pale grey pubescent;
singly or doubly branched, the branchlets short,
bearing to 10 secund flowers; bracteoles to 3.5 by 2
mm, ovate, obtuse, sparsely pubescent outside, gla-
brous within. Bud to 5 by 2.5 mm, narrowly ovoid,
subacute. Calyx densely pale grey pubescent outside,
glabrous within; 3 outer lobes narrowly deltoid,
subacute, slightly revolute; 2 inner lobes smaller,
acute, not revolute. Petals pink, oblong-lanceolate,
pubescent on parts exposed in bud. Stamens 15, the
inner 5 slightly longer than the others; filaments broad
at base, rather abruptly tapering and filiform distally;
anther oblong; appendage to connective over twice as
long as anther, reaching almost to style apex. Ovary
narrowly ovoid, shortly pubescent, longer pubescent
towards apex; stylopodium indistinct; style filiform,
somewhat shorter than ovary, glabrous but for
shortly pubescent base. Fruit calyx sparsely puberu-
lent, more densely so at base; 3 longer lobes to 6 by 1.3
cm, chartaceous, spatulate, narrowly obtuse, c. 3.5
mm broad above the c. 7 by 5 mm deltoid saccate
thickened base; shorter lobes to 3 by 0.3 cm, subequal,
linear, similar at base. Nut c. 20 by 8 mm, narrowly
ovoid, shortly pubescent towards the apex, otherwise
glabrous; style remnant acute.

Distr. Malesia: Northern Borneo (Sarawak,
Brunei, Sabah).

Ecol. On podsols in Heath forest, on terraces,
plateaux and ridges to 1600 m; locally common; on
ultrabasics in E. Sabah.

Vern. Méranti tangkai panjang padi (Brunei), séraya
kérangas (Sabah).

Note. Closely resembling S. coriacea, within whose
range it is confined, and occupying similar habitats.
No record yet exists though of intermediate forms.

107. Shorea waltoni Woop ex MEvWeR, Act. Bot. Neerl.
12 (1963) 344, pl. 11; Sabah For. Rec. 5 (1964) 155.
Tall buttressed tree with deeply fissured bark.
Twigs, leaf buds, stipules, petioles, nervation beneath
and panicle + densely cream lepidote; calyx at first
densely so, becoming sparse, + caducous in fruit;
midrib above and sometimes nerves, ovary and parts
of petals exposed in bud densely persistently cream-

1982]

DIPTEROCARPACEAE (Ashton)

S11

brown pubescent. Twigs c. 3 mm @ apically, stout,
terete, becoming smooth, dark brown; stipule scars +
amplexicaul, slender. Buds small, conical; stipules to
20 by 8mm, elliptic-lanceolate, acute. Leaves 10-25 by
5.5—12 cm, oblong to ovate or obovate, coriaceous,
cream lepidote beneath; base obtuse or shallowly
cordate; apex abruptly to | cm long acuminate; nerves
18-22 pairs, slender but prominent beneath, drying
dark, + applanate above; tertiary nerves scalariform,
elevated beneath; midrib prominent beneath, obscure
and depressed above, petiole 2.5—4.5 cm long, stout.
Panicle to 12 cm long, rather stout, terminal or
axillary; singly or doubly branched, branchlets to S5cm
long; bracts to 10 mm long, amplexicaul. Flower bud to
8 by 3 mm, fusiform; sepals ovate, the outer 3 longer,
acute, the inner 2 broader, acuminate. Stamens 15,
unequal; filaments broadly compressed at base,
abruptly tapering and filiform distally; anthers
broadly oblong; appendages slender, scarious api-
cally, c. 2x length of anthers. Ovary broadly ovoid;
style equal in length to ovary, rather short, puberulent
in the basal 4. Fruit pedicel c. 3 mm long, stout; 3
longer calyx lobes to 14 by 2.3 cm, spatulate, obtuse,
coriaceous, c. 11 mm wide above the c. 8 by 15 mm
transversely elliptic saccate thickened base; 2 shorter
lobes to 10 by 0.8 cm, lorate, spatulate, narrowly
obtuse. Nut to 25 by 18 mm, ovoid, apiculate.

Distr. Malesia: N. E. Borneo (Sandakan Distr.).

Ecol. Scattered on well or moderately drained land
in lowlands.

Vern. Séraya kélabu.

108. Shorea pachyphylla RIpL. [DURANT, Rep. For.
Brunei (1933) 41, nomen] ex Sym. J. Mal. Br. R. As.
Soc. 19 (1941) 163, pl. 7; BRowng, For. Trees Sarawak
& Brunei (1955) 148; ANDERSON, Gard. Bull. Sing. 20
(1963) 159; ASHTON, Man. Dipt. Brun. (1964) 205, f.
17, pl. 54 (habit, bark); ibid. Suppl. (1968) 112. — Fig.
101.

Large buttressed tree with pale flaky and promi-
nently fissured bark. Young parts very shortly densely
evenly tawny pubescent, caducous on all but leaf bud,
panicle and stipule (puberulent on inner surface).
Twig c. 4 mm @, stout, terete, slightly ridged and
compressed apically, becoming smooth or rugulose;
stipule scars c. 0.5 mm broad, horizontal, normally
amplexicaul. Bud 5-8 by 2-4 mm, compressed or
terete, narrowly ovoid to broadly falcate. Stipule to 20
by 7 mm, lanceolate, acute, fugaceous. Leaves 10-20
by 9-16 cm, broadly ovate to suborbicular, thickly
coriaceous; base obtuse or broadly cuneate; apex
obtuse or with short, to 5 mm long, acumen; margin
slightly but distinctly revolute; nerves 7—9 pairs, well
spaced, prominent beneath, arched at margin, at c.
40°-55°; midrib applanate and broad above, terete
and prominent beneath; tertiary nerves slender, den-
sely scalariform, at 90° to midrib; petiole 4-6 cm long,
stout. Panicle to 11 cm long, terminal or axillary,
compressed, ridged when dry; doubly branched, the
branchlets bearing to 5 distichous flowers; bracteoles
to 3 by 2 mm, ovate, acute, shortly pubescent. Flower

bud to 6 by 4 mm, narrowly ovoid, acute. Calyx
densely pubescent outside; lobes ovate, deltoid, acu-
minate, the 3 outer somewhat longer, more obtuse,
than the 2 inner lobes. Petals broadly lanceolate,
obtuse, hardly contorted, densely pubescent on parts
exposed in bud. Stamens 15, in 3 lengths, the inner 5 an
anther’s length longer than the outer 5; filaments
broad at base, tapering and stoutly filiform distally;
anthers subglobose; appendage to connective
3-4 xlength of anther, pubescent towards apex,
otherwise glabrous; stylopodium indistinct, sparsely
pubescent, tapering into style; style and stylopodium
longer than ovary. Fruit calyx entirely glabrous; 3
longer lobes to 16 by 3.5 cm, spatulate, obtuse, c. 7
mm broad above the to 2 by 1.5 cm thickened saccate
base; 2 shorter lobes to 8 by 0.6 cm, narrowly
spatulate, similarly saccate at base. Nut to 3.3 by 1.8
cm, ovoid, pruinose, buff pubescent towards the apex;
style remnant c. 1.5 mm long, acute.

Distr. Malesia: N.W. Borneo (Sarawak, Brunei
and Kalimantan: Lower Kapuas and Barito).

Ecol. Locally gregarious, mixed peat swamp for-
ests, especially on and beside white sand raised
beaches.

Vern. Kukup, kérukup, méranti kérukup (Brunei).

109. Shorea pauciflora KiNG, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 116; BRANDIS, J. Linn. Soc. Bot. 31 (1895)
98; BurK. J. Str. Br. R. As. Soc. 81 (1920) 51, 69, fig.;
RIDL. Fl. Mal. Pen. 1 (1922) 228; Foxw. Mal. For.
Rec. 3 (1927) 51, p.p.; ibid. 10 (1932) 218; SLooT. in
Merr. Pl. Elm. Born. (1929) 203; BuRK. Dict. (1935)
2020; Sym. Mal. For. Rec. 16 (1943) 87, f. 37A, 38, 52;
BROWNE, For. Trees Sarawak & Brunei (1955) 149;
ASHTON, Man. Dipt. Brun. (1964) 207, f. 17, pl. 46
(bark); ibid. Suppl. (1968) 114; MeuerR & Woop,
Sabah For. Rec. 5 (1964) 133, f. 15; ASHTON, Gard.
Bull. Sing. 31 (1978) 47.

Large buttressed tree. Leaf bud, stipule and panicle
persistently shortly pale gold-brown pubescent; ner-
vation, petiole and young twig sparsely dotted with
minute caducous hair tufts. Twig 1.5—2.5mm @, terete
at first smooth, becoming flaky; stipule scars pale,
short, narrow, horizontal. Bud 4-8 by 2-3.5 mm,
ovoid, acute. Stipule to 13 by 4 mm, hastate, acute.
Leaves 9-15 by 4—5.5 cm, ovate, + thinly coriaceous,
frequently subequal; base obtuse to broadly cuneate;
acumen ().6—1.2 cm long, narrow; nerves 8-9 pairs,
slender but prominent beneath, at c. 40°—S0°, straight,
slightly curved at the margin, occasionally with a few
glabrous domatia; tertiary nerves very slender, den-
sely scalariform, diagonal to the nerves; midrib nar-
rowly depressed above, prominent beneath; petiole
1.3-1.8 cm long. Panicle to 15 cm long, terminal or
axillary, ribbed or slightly compressed, lax; doubly or
trebly branched, the branchlets bearing to 19 secund
flowers; bracteoles to 4 by 2.5 mm, oblong, subacute,
shortly densely pale yellow-brown to pubescent fuga-
ceous. Flower bud c. 7 by 3.5 mm, ovoid, subacute.
Calyx densely pale yellowish buff pubescent outside,
glabrous within; lobes subequal, deltoid, obtuse, the 2

Dl FLORA MALESIANA [ser. I, vol. 97

Fig. 101. Shorea pachyphylla Riv. ex Sym. a. Habit, b. twig apex with leaf, c. fruit, d. nut, all x 4 (a bb. 31414,
b-d § 11245).

1982]

inner slightly narrower. Petals pale yellow, oblong,
obtuse, densely pubescent on parts exposed in bud.
Stamens 15, the inner 5 slightly longer than the other
10; filaments expanded at base, abruptly narrowing
distally; anthers broadly oblong; appendage to con-
nective c. 2xlength of anther, setose towards the
apex, the inner 5 reaching the style apex. Ovary and
stylopodium ovoid to conical, shortly densely pubes-
cent; style glabrous, slightly shorter than ovary and
stylopodium. Fruit subsessile. Calyx sparsely pubes-
cent, more densely so at base; 3 longer lobes to 9 by 1.5
cm, spatulate, narrowly obtuse, c. 4mm broad above
the c. 8 by 11 mm rounded thickened saccate base; 2
shorter lobes to 5 by 0.5 cm, linear, similar at base. Nut
c. 1.4 by 1.2 cm, broadly ovoid, obtuse or with a short
acute style remnant, densely pale buff tomentose.

Distr. Malesia: Malaya, Sumatra (extreme north
and the south excepted, incl. P. Siberut), Borneo.

Ecol. Scattered, rarely common, on deep soils on
undulating land and hills below 700 m.

Uses. A valuable heavy red meranti.

Vern. Némesu, lamsa, lemesa, lemesa samak, mésu,
méranti benia, m. samak, kala samak, méndamak,
pinang baik, séraya, s. batu, témpayan mas (Malaya).
katuko, k. andilan, k. tikau, méranti udang (Sumatra),
ubar suluk (Sabah), méranti samak, éngkabang cheriak
(Sar.), pérawan samak (Iban), lampong merantas, !.
tahan, pelépak, kélapak, k. undang, putang lanan,
awang kulat, a. laying, mérkabong (S.E. Borneo),
téngkawang tijang, t. rambai, kontoi, k. lemak (W.
Borneo), etc.

Note. Variable in leaf size and colour: small-leaved
forms tend to have leaves drying rust-red, large-leaved
chocolate-brown with the nerves and petiole distinctly
paler.

110. Shorea johorensis Foxw. Mal. For. Rec. 10
(1932) 236, pl. 21; BurK. Dict. (1935) 2012; Sym. Mal.
For. Rec. 16 (1943) 72, f. 38; ASHTON. Gard. Bull.
Sing. 22 (1967) 294. — S. polysperma (non MERR.)
KeiTH, North Born. For. Rec. 2 (1938) 238.— S.
leptoclados Sym. Gard. Bull. S. S. 10 (1939) 376, pl. 25;
Mal. For. Rec. 16 (1943) 77: BRowne, For. Trees
Sarawak & Brunei (1955) 140; AsHTON, Man. Dipt.
Brun. (1964) 195, f. 17; ibid. Suppl. (1968) 110; MEER
& Woop, Sabah For. Rec. 5 (1964) 112, f. 13, pl. 6
(habit). — Fig. 100 D—D3.

Large buttressed tree. Twig, panicle, leaf bud,
stipule, petiole, nervation beneath and midrib above
shortly evenly persistently grey-buff pubescent; young
leaf caducously so. Twig 2-3 mm g apically, much
branched, slightly ribbed and compressed when
young, becoming terete, drying glabrous, smooth;
stipule scars 1.5—2 mm long, pale, falcate, descending.
Bud 6-8 by 2-3 mm, ovoid, compressed. Stipule to 3.5
by 0.7mm, + caducous, lanceolate, acute, constricted
at base. Leaves 9-14 by 4.2-7.5 cm, chartaceous to
thinly coriaceous, ovate; base obtuse to subcordate:
acumen to 7 mm long; nerves 10-12 slender but
prominent pairs, at 90° at base, c. 40°—-50° towards
apex; first 3-6 pairs usually with prominent scale-like

DIPTEROCARPACEAE (Ashton)

+53

domatia, the basal two pairs of which fused along the
midrib on each side; tertiary nerves very slender,
densely scalariform, sinuate, diagonal to nerves;
petiole slender, 1.5—2 cm long. Panicle to 15 cm long,
terminal or axillary, terete or slightly compressed,
slender; regularly singly or doubly branched, branch-
lets long, lax, bearing to 18 secund flowers; bracts to 7
mm long, narrowly ovate, obtuse, pubescent on both
surfaces, fugaceous. Flower bud to 8 by 3.5 mm, ovoid
to lanceolate, acute. Calyx densely pale grey pubes-
cent outside, glabrous within; lobes subequal. acute,
slightly revolute towards the apices. Petals pale
yellow, narrowly lanceolate, obtuse, densely pubescent
on parts exposed in bud. Stamens 15, the 5 innermost
only slightly longest; filaments expanded at base,
abruptly narrowing and filiform distally; anthers
oblong, somewhat tapering: appendage to connective
c. 3xlength of anther, reaching 3 length of style.
glabrous. Ovary globose, shortly densely pubescent in
apical half, stylopodium indistinct; style over twice as
long as ovary, filiform, glabrous but for base. Fruit
calyx shortly pubescent towards base, glabrescent: 3
longer lobes to 12 by 2.3 cm, subequal, spatulate,
narrowly obtuse, c. 6 mm broad above the to 1.4 by
1.2 cm deltoid saccate thickened base; 2 shorter lobes
to 6.5 by 0.6 cm, unequal, linear but similar at base.
Nut to 2 by 1.4cm, broadly ovoid, densely shortly pale
buff tomentose: style remnant short.

Distr. Malesia: Malaya (E. Johore), Sumatra (East
coast: Simelungun, Djambi, Palembang), Borneo
(Sarawak, Sabah, S.E. Borneo to Muara Tewe and
Pleihari).

Ecol. Common in Palembang and E. Borneo, local
or rare elsewhere, on fertile soils on hillsides, well-
drained alluvium and undulating land below 600 m
alt.

Vern. Méranti pepijat (Mal.), mérukuyong (Sum.),
kénuar, kénbéwar, langko, pélépak (S.E. Borneo),
majau (Sabah), sélangan pélandok (Brunei), mérampu
(Iban).

Note. Collections from West Sarawak and the
Malay Peninsula lack the basal domatia that so
facilitate distinction of sterile material from S. palem-
banica.

111. Shorea palembanica Mig. Sum. (1861) 487: DC.
Prod. 16, 2 (1868) 632; WaLp. Ann. 7 (1868) 379:
Burck, Ann. Jard. Bot. Btzg 6 (1887) 219; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 103; Sym. Gard. Bull. S. S.7
(1933) 141, pl. 41; Mal. For. Rec. 16 (1943) 83, f. 38:
BROWNE, For. Trees Sarawak & Brunei (1955) 141:
ASHTON, Man. Dipt. Brun. Suppl. (1968) 112, f.
13. — S. aptera BurcK, Ann. Jard. Bot. Btzg 6 (1887)
210, t.22; BRANDIS & GitGin E. & P. Pfl. Fam. ed. 1, 3,
6 (1895) 264; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 78:
HEyYneE, Nutt. Pl. ed. 1, 3 (1917) 296; MeRR. En. Born.
(1921) 403; Gita in E. & P. Pfi. Fam. ed. 2, 21 (1925)
260. — S. brachyptera Hem, Bull. Mens. Soc. Linn.
Paris 2 (1891) 571; Merr. En. Born. (1921)
404. — Pachychlamys beccarianus, P. brachypterus
Dyer ex BRANDIS, J. Linn. Soc. Bot. 31 (1895) 77,

514

nomina pro syn. — S. lepidota (non BL.) Foxw. Mal.
For. Rec. 10 (1932) 166, p.p.

Small or medium-sized, rarely large tree. Young
twig, leaf bud, and stipule outside densely evenly pale
buff pubescent, stipule inside, petiole and leaf ner-
vation sparsely so, sometimes glabrescent. Twig c. 2
mm @, compressed apically, becoming smooth, terete;
stipule scar short, horizontal. Bud to 13 by 6 mm,
lanceolate, falcate, acute. Stipule to 15 by 3 mm,
lanceolate, acute, caducous. Leaves 8-25 by 4-10 cm,
ovate to oblong, chartaceous, undulating; base
broadly cuneate to subcordate; acumen to | cm long,
broad; nerves 12-16 pairs, slender but prominent
beneath, at 45°-65°; tertiary nerves slender, dense;
midrib applanate to depressed above, prominent
beneath: petiole 14-25 mm long, drying ribbed.
Panicle to 16 cm long, terminal or axillary, terete,
shortly evenly persistently pale buff pubescent; doubly
branched, branchlets to 5 cm long, bearing to 10
secund flowers; bracteoles to 4 by 2 mm, ovate, acute,
puberulent, caducous. Flower bud to 7 by 3 mm,
fusiform. Sepals ovate, densely pubescent on parts
exposed in bud; 3 outer acute, 2 inner smaller, thinner
towards the fimbriate margins, acuminate. Petals pale
yellow, lanceolate, pubescent on parts exposed in bud.
Stamens 15, in 3 unequal verticils; filaments com-
pressed at base, tapering medially and filiform below
the broadly ellipsoid anthers; appendage to connec-
tive slender, c. 2xlength of anthers. Ovary ovoid,
tapering, pubescent; style filiform, c. 2xlength of
ovary, pubescent in the basal }. Fruit pedicel to 2 mm
long. Calyx puberulent, glabrescent, short, 3 longer
lobes to 5 by 0.7 cm, lorate, obtuse, with to 2.5 by 2.5
cm ovate thickened prominently saccate base; 2
shorter lobes to 3 cm long, otherwise similar. Nut to
3.5 by 2.5 cm, ovoid, apiculate, densely evenly shortly
persistently pale yellow-brown pubescent.

Distr. Malesia: Malaya (Perak and E. coast),
Sumatra (Palembang), Borneo (excluding Sabah,
Brunei and Sarawak west of the Lupar).

Ecol. River banks, fresh water swamps and rarely
low moist hillsides, locally abundant.

Vern. Méranti tengkawang ayér, singkawang, téng-
kawang, mérpak (Malaya), mérkuyung, mélebékan
(Sumatra), éngkabang asu (Sarawak), kélépak, majau,
ténkawang, t. rambut, t. ringgit, méngkabang (W.
Borneo).

112. Shorea andulensis ASHTON, Gard. Bull. Sing. 19
(1962) 275, pl. 11; Man. Dipt. Brun. (1964) 178, f. 17;
ibid. Suppl. (1968) 105; MEER & Woop, Sabah For.
Rec. 5 (1964) 97.

Medium-sized to large tree. Twig, panicle, leaf bud,
midrib above. Stipule (both surfaces) and petiole
shortly densely persistently yellowish buff puberulent,
more sparsely so on nervation beneath and midrib
above. Twig 1.5-2 mm @ apically, slender, com-
pressed when young, becoming terete, smooth, gla-
brous; stipule scars short, inconspicuous. Bud 3-6 by
1.5—3 mm, ovoid, compressed, subacute. Stipule to 14
by 5 mm, hastate, subacute. Leaves 6-9 by 3-5 cm,

FLORA MALESIANA

[ser. I, vol. 9?

elliptic-ovate, lepidote beneath in mature tree; base
obtuse; acumen to 8 mm long; nerves 10-13 pairs,
straight, abruptly curved near margin, slender but
prominent beneath, at 40°—50° except at base; tertiary
nerves slender, scalariform, sinuate, diagonal to
nerves; petiole 1-1.2 cm long, 1-1.5 mm @, shortly
cream puberulent and rugose on drying. Panicle to 12
cm long, terminal or axillary, terete or slightly
compressed; singly or doubly branched, the branch-
lets bearing to 10 distichous flowers; bracts and
bracteoles to 3 by 1.2 mm, ovate, subacute, shortly
grey puberulent outside, glabrescent within. Flower
bud to 8 by 2.5 mm, narrowly ovoid, subacute, the
calyx spreading. Calyx densely shortly pubescent
outside, puberulent within; lobes subequal, ovate,
acute to subacuminate, the inner 2 more attenuate
than the outer 3. Petals bright lilac within, paler
outside, narrowly lanceolate, acute, twisted, shortly
pubescent on parts exposed in bud, saccate at base.
Stamens 15, the inner 5 only slightly longer than the
others; filaments basally expanded, abruptly tapering;
anther broadly oblong; appendage to connective c.
3xlength of anther, straight, unreflexed, reaching
almost to the style apex on the 5 inner stamens. Ovary
small, globose, glabrous; stylopodium indistinct; style
c. 2x length of ovary, filiform, sparsely pubescent in
the basal half. Fruit calyx shortly pubescent, more
densely so at base; 3 longer lobes to 5 by 1.2 cm,
broadly spatulate, narrowly obtuse, c. 2.5 mm broad
above the c. 6 by 4 mm narrowly ovate thickened
saccate base; 2 shorter lobes to 2.5 by 0.2 cm, linear,
subequal, similar at base. Nut to 14 by 7 mm, ovoid,
abruptly shortly apiculate, shortly densely grey-buff
pubescent.

Distr. Malesia: Northern Borneo (Ulu Kapuas,
Sarawak, Brunei, E. Sabah).

Ecol. Rare, on sandy soils in Mixed Dipterocarp
forest; on ultrabasics in E. Sabah.

Vern. Méranti daun puteh (Brun.).

113. Shorea polysperma (BLCO) MerRR. Publ. Govt.
Lab. Philip. 27 (1905) 22; ibid. 29 (1905) 29; Philip. J.
Sc. 1 (1906) Suppl. 98; EveReTT & WHITFORD, Bull.
Bur. For. Philip. 5 (1906) 26; Foxw. Philip. J. Sc. 2
(1907) Bot. 356, 357, 394; MERRITT, Bull. Bur. For.
Philip. 8 (1908) 16, 48; Foxw. Philip. J. Sc. 4 (1909)
Bot. 423, 510, 518; WHITFORD, ibid. 4 (1910) Bot. 703;
Bull. Bur. For. Philip. 10 (1911) 66; Foxw. Philip. J.
Sc. 6 (1911) Bot. 277; ibid. 13 (1918) Bot. 191; MErRR.
Sp. Blanc. (1918) 269; En. Philip. 3 (1923) 99; REYEs,
Philip. J. Sc. 22 (1923) 328; Foxw. ibid. 67 (1938)
309. — Mocanera polysperma Buco, FI. Filip. 1 (1837)
448. — Dipterocarpus polyspermus BLco, F1. Filip. ed.
2 (1845) 312; ibid. ed. 3, 2 (1878) 213; DC. Prod. 16, 2
(1868) 614; Dyer, J. Bot. 12 (1874) 108. — Hopea
tangili BL. Mus. Bot. Lugd.-Bat. 2 (1852) 35, nom.
illeg.; WALP. Ann. 4(1857) 339; DC. Prod. 16, 2 (1868)
635.— S. talura (non Roxs.) F.-VILL. Nov. App.
(1880) 21. — S. warburgii GiLG, Bot. Jahrb. 18, Beibl.
45 (1894) 38; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 98;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895)

1982]

264; Foxw. Philip. J. Sc. 6 (1911) Bot. 278; ibid. 13
(1918) Bot. 191; Gi_G in E. & P. Pfl. Fam. ed. 2, 21
(1925) 260.— S. teysmanniana (non DYER) Foxw.
Philip. J. Sc. 6 (1911) Bot. 278; ibid. 13 (1918) Bot. 192;
WHITFORD, Bull. Bur. For. Philip. 10 (1911) 68; MerRR.
En. Philip. 3 (1923) 99; Reyes, Philip. J. Sc. 22 (1923)
326. — Fig. 7.

Large buttressed tree. Twigs, buds, stipules, pani-
cles, bracts and parts of petals exposed in bud +
densely persistently cream puberulent, petioles, ner-
vation beneath, calyx and ovary so at first becoming
glabrous on all but ovary. Twig c. 2-3 mm @ apically,
compressed at first, becoming terete, smooth, black-
ish; stipule scars short, horizontal. Buds c. 2 by 1 mm,
small, ellipsoid, stipules to 15 by 8 mm, lanceolate,
acute, caducous. Leaves 7.5—15 by 2.5—6.5 cm, elliptic,
thinly coriaceous, occasionally cream _ lepidote
beneath; base cuneate; acumen to 2 cm long, tapering;
nerves 9-12 pairs, slender but prominent beneath,
slightly elevated above, arched, at 55°—75°, frequently
with glabrous pore-like axillary domatia; tertiary
nerves densely scalariform, sinuate, very slender,
barely elevated beneath, obscure above; midrib
slender but prominent beneath, obscurely depressed
above; petiole 16-22 mm long, slender geniculate.
Panicle to 16 cm long, terminal or axillary, doubly
branching, spreading, many-flowered; branchlets to 7
cm long; bracts to 5 by 3 mm, ovate-deltoid, acute.
Buds to 4 by 2 mm, small, fusiform; sepals broadly
ovate, the outer 3 acute, the inner 3 smaller, shortly
acuminate; stamens 15, shorter than style at anthesis;
filaments broadly compressed at base, tapering
abruptly medially and filiform beneath the subglobose
anthers; appendage slender, + glabrous, c. 25 x
length of anthers; ovary subglobose, abruptly
crowned by a slender glabrous style c. 2 x its length.
Fruit pedicel to 3 mm long, stout, broadening into the
receptacle; 3 longer calyx lobes to 9 by 1.8 cm,
spatulate, obtuse, coriaceous, c. 6 mm broad above
the c. 10 by 10 mm broadly ovate saccate thickened
base; 2 shorter lobes to 5 by 0.4 cm, lorate, obtuse,
similar at base; nut to 18 by 14 mm, ovoid, apiculate.

Distr. Malesia: Philippines.

Ecol. Widespread and often common, evergreen
Mixed Dipterocarp forest on hills.

Vern. Tangile (Tag.), abuhungan (Al.), adamini
(Bis.), araka (Ilk.), balakbakan (Neg.), belagayan
(Mangyan), damilang (Ibn.), maligmal (Tang.),
manaog (C. Bis.), mayapis (Tayabas), pata (Pang.).

114. Shorea platyclados SLooT. ex [ENDERT, M. O.
Born. Exp. 1925 (1926) 261, 266, 267, 272, nomen]
Foxw. Mal. For. Rec. 10 (1932) 214; ENDERT, Tec-
tona 26 (1933) 410; DE Vooap, ibid. 26 (1933) 703;
Boon, ibid. 32 (1939) 839; BurK. Dict. (1935) 2020;
Descu, Mal. For. Rec. 12 (1936) 2, 43, 63, 65; Sym. J.
Mal. Br. R. As. Soc. 14 (1936) 336, 339, 340, 348;
Gard. Bull. S. S. 10 (1939) 377; Mal. For. Rec. 16
(1943) 89, f. 38, 54; DEscH & THoMas, Mal. For. Rec.
13 (1940) 4; SLoor. Bull. Bot. Gard. Btzg III, 17 (1941)
110, f. 16; ASHTON, Man. Dipt. Brun. (1964) 212, f. 17,

DIPTEROCARPACEAE (Ashton)

515

pl. 47 (stem-base); ibid. Suppl. (1968) 115; MEER &
Woop, Sabah For. Rec. 5 (1964) 139; NG, Mal. For.
39 (1976) 91, f. 1-9.

Large tree with dark, flaky and fissured bark.
Young parts grey puberulent, fugaceous except on
stipule and panicle. Twig c. 3 by 1.5 mm @ apically,
compressed, glabrous, smooth; stipule scars c. 2 mm
long, linear, pale, horizontal. Leaf bud to 4 by 1.5mm
@ apically, compressed, acute. Stipule to 13 by 3.5
mm, hastate, acute. Leaves alternate, 6—9 by 2—3 cm,
lanceolate; base cuneate and with revolute margin;
acumen c. | cm long; margin undulate; nerves 15-18
pairs, slender, hardly elevated beneath, + obscure,
with short secondary nerves running } way to margin;
tertiary nerves very slender, densely scalariform,
diagonal to nerves; midrib narrow, obscure and
depressed above, slender but sharply prominent
beneath; petiole 1—1.5 cm long, slender, compressed
laterally. Panicle to 7 cm long, terminal or axillary,
compressed; singly branched, branchlets to 1.8 cm
long, bearing to 7 secund flowers; bracteoles un-
known. Flower bud to 8 by 3 mm, ovoid, ellipsoid.
Calyx densely shortly puberulent outside, glabrous
within; 3 outer lobes ovate, acuminate, 2 inner lobes
smaller, more narrowly acuminate. Petals pale yellow,
lanceolate, slender, acute, densely pubescent on parts
exposed in bud. Stamens 15, of 2 lengths; filament
broad at base, tapering + abruptly and filiform
distally; anther subglobose; appendages to connective
c. 4 x length of anther, slender, glabrous. Ovary and
stylopodium pyriform, puberulent towards apex; style
glabrous, as long as ovary and stylopodium. Fruit
calyx glabrescent; 3 longer lobes to 10 by 1 cm,
narrowly spatulate, obtuse, to 4 mm broad above the
to 7 by 7 mm broadly ovate saccate thickened base; 2
shorter lobes to 5 by 0.4 cm, linear, acute, similar at
base. Nut to 1.5 by 1 cm, glabrescent; style remnant
acute.

Distr. Malesia: Sumatra, Borneo.

Ecol. Widespread in mountainous districts, usually
between 700-1300 m, but occasionally down to 200 m
in valley bottoms near mountains.

Vern. Méranti bukit, jalak (Malaya).

Note. NG /.c. found that if the orthotropic leader
shoot is damaged it is replaced by new orthotropic
leaders from dormant accessory buds, not by lateral
plagiotropic shoots.

115. Shorea scaberrima BuRCK, Med. Lands PI. Tuin
3 (1886) 22; Ann. Jard. Bot. Btzg 6 (1887) 208;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 78; MERR. En.
Born. (1921) 406; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 306;
ibid. ed. 2 (1927) 1124; BRowNE, For. Trees Sarawak
& Brunei (1955) 143; AsHton, Man. Dipt. Brun.
(1964) 219, f. 17, pl. 49 (bark). — Fig. 99 A—A4.
Medium-sized tree. Young twigs, panicle, leaf bud,
stipule outside (puberulent within), petiole, leaf
beneath and midrib above densely persistent-scabrid
tawny tomentose; leaf above fugaceous pubescent.
Twig 2-3 mm @ apically, terete, becoming rugulose,
verrucose, much branched; stipule scars to 1.5 mm

516

FLORA MALESIANA

[ser. I, vol. 9?

long at first, 0.5 mm thick, shortly cuneate, obscured
by tomentum. Bud 4-6 by 4-5 mm, broadly ovoid,
slightly compressed, subacute. Stipule to 18 by 8 mm,
broadly hastate, subacute, constricted at base, cadu-
cous. Leaves 7-20 by 4-9 cm, oblong-obovate to
oblong-ovate, thinly coriaceous; base obtuse, rarely
subcordate; acumen to 8 mm long, short, deltoid:
nerves 14-17 pairs, rather slender, curved, at 40°—50°,
more widely at base; tertiary nerves distinct, rather
densely scalariform; midrib prominently terete
beneath, slightly depressed or applanate above:
petiole 1.8—2.5 cm long. Panicle to 8 cm long, short,
lax, terminal or axillary, terete or ribbed; singly
branched, the branchlets bearing to 3 + secund
flowers; bracteoles to 4.5 by 3 mm, narrowly deltoid,
acute, scabrid pale fulvous pubescent outside, puberu-
lent within, fugaceous. Flower bud to 8 by 5 mm,
ovoid, obtuse. Calyx densely yellow-brown pubes-
cent; 3 outer lobes deltoid to ovate, frequently
acuminate, acute; 2 inner lobes similar but smaller.
Petals pink, narrowly lanceolate, densely setose on
parts exposed in bud. Stamens 15, the 5 innermost
somewhat longer than the other; filaments basally
expanded, abruptly contracted and filiform distally;
anther + oblong; appendage to connective reaching
the style apex on 5 inner stamens, more than twice as
long as anther cells, becoming curved but not reflexed.
Ovary subglobose, small, shortly densely pubescent;
style and stylopodium 3 times as long as ovary,
filiform with glabrous apex, swelling slightly below it
and densely shortly pubescent; frequently further
swollen in the basal half. Fruit calyx shortly evenly
fulvous pubescent; 3 longer lobes to 4.5 by 1 cm or
shorter, unequal, variable, shortly spatulate, sub-
acute, to 7mm broad above the to 2 by 3.5 cm broadly
ovate shallowly saccate thickened base, closely
adpressed to the base of nut; 2 shorter lobes to 3 by 0.3
cm, linear, similarly expanded at base. Nut to 5 by 2.5
cm large, obovoid, tapering to an acute apical style
remnant, densely evenly fulvous pubescent, becoming
the same length or slightly longer than the calyx.

Distr. Malesia: N.W. Borneo (through Sarawak to
S.W. Sabah); S.E. Borneo (Puruktjau, sterile collec-
tion?).

Ecol. Frequent, sandy clay soils on low hills,
alluvium and locally on ridges and volcanic plateaux
to 850 m.

Vern. Engkabang bintang, kantoi, k. lilin, k. tém-
baga, téntung, tengkawang kijang (W. Borneo), mér-
anti paya, m. paya bérsisek (Brunei), éngkabang pinang
(Iban), séraya mémpelas (Sabah).

116. Shorea fallax MEER, Act. Bot. Neerl. 12 (1963)
335, pl. 7; MEVER & Woop, Sabah For. Rec. 5 (1964)
105; ASHTON, Man. Dipt. Brun. (1964) 186, f. 17; ibid.
Suppl. (1968) 107.— S. oleosa Meurer, Act. Bot.
Neerl. 12 (1963) 338, pl. 8; MEER & Woop, Sabah
For. Rec. 5 (1964) 124. — S. squamata (non BENTH. &
Hook. f.) BRowne, For. Trees Sarawak & Brunei
(1955) 144.

Medium-sized to large buttressed tree. Young twig,

panicle, leaf bud, stipule outside (glabrescent or
shortly pubescent within), petiole beneath, leaf
beneath and midrib above sparsely persistently pale
brown scabrid tomentose. Twig 2—2.5 mm @ apically,
terete, ribbed below the stipule scars, becoming
smooth; stipule scar c. 2 mm long, c. 0.5 mm thick,
pale, cuneate, slightly falcate, horizontal or slightly
ascending. Leaf bud 5-8 by 4-6 mm, ovoid, subacute.
Stipule to 15 by 5mm, hastate, acute. Leaves 12-24 by
5.5—11 cm, broadly oblong to ovate, + thinly coria-
ceous; base obtuse; acumen to 1.2 cm long; nerves
15-19 pairs, prominent, at 60°—70°, curved distally;
short secondary nerves frequently present; tertiary
nerves prominent, well spaced, at 90° to nerves;
midrib prominent, terete beneath, applanate but
evident above; petiole 1-1.5 cm long, short. Panicle to
22 cm long, terminal or axillary, terete or ribbed, lax,
regularly alternately singly or doubly branched, the
branchlets bearing to 7 secund flowers; bracts and
bracteoles to 7 by 3 mm, ovate, acute, pubescent
outside, glabrous within, not at first caducous. Flower
bud to 8 mm long, narrowly ovoid, acute. Calyx
shortly densely pale grey-brown pubescent outside,
glabrous within; 3 outer lobes longer, + deltoid to
ovate, subacute; 2 inner lobes shorter, narrowly
deltoid to ovate, acute, acuminate. Petals pale cream-
yellow, narrowly oblong, densely pubescent on parts
exposed in bud. Stamens 15, the 5 inner an anther’s
length longer than the others; filaments basally
expanded, abruptly tapering and filiform distally;
anthers oblong; appendage to connective reaching to
style apex on 5 inner stamens, more than twice as long
as anthers, curved but not reflexed. Ovary and
stylopodium pyriform, densely pubescent in the distal
half, shortly densely pubescent basally; style as long as
ovary and stylopodium, filiform, glabrous. Fruit
subsessile. Calyx glabrescent; 3 longer lobes to 5 by
0.8 cm, spatulate, subequal, to 3.5 mm broad above
the to 1.5 by 1.2 cm ovate glabrous shallowly saccate
thickened base; 2 shorter lobes to 25 by 1.5 mm, linear,
similar at base. Nut to 2.7 by 1.0 cm, ovoid, shortly
buff pubescent, apiculate, shorter than longer calyx
lobes.

Distr. Malesia: Borneo (N.E. Sarawak and Sabah
to Berau).

Ecol. Widespread on clay soils in Mixed Diptero-
carp forest, on well drained alluvium, undulating land
and hillsides to 600 m.

Vern. Engkabang layar, e. pinang (Sar.), méntalun,
meéranti sépit undang (Brun.), séraya minyak (Sabah),
tuntong séluing, kontoi (S.E. Borneo).

Note. Doubtfully distinct from S. parvistipulata
(see sub ssp. albifolia).

117. Shorea pubistyla AsHTON, Gard. Bull. Sing. 22
(1967) 297, pl. 43; Man. Dipt. Brun. Suppl. (1968) 116,
f. 14, pl. 21 (bark). — Fig. 102.

Large tree. Twig, leaf bud, petiole and midrib
beneath densely dark fulvous scabrid tomentose;
more shortly, sparsely so on nervation beneath,
midrib above and stipule outside; stipule within

1982]

DIPTEROCARPACEAE (Ashton)

517

evenly densely pale fulvous caducous pubescent. Twig
c. 7 mm @ towards the apex, stout, prominently
ribbed at first, becoming terete; stipule scars c. 2mm
long, horizontal, obscure. Bud to 1.3 mm long and @,
ovoid, compressed. Stipule to 2.5 by 0.7 cm, lanceo-
late, subcaudate, caducous. Leaves 11-19 by 7-12 cm,
broadly oblong to obovate, thickly coriaceous; base
obtuse to cordate; apex obtuse, subretuse or shortly
acuminate; nerves 14-18 pairs, obscurely depressed
above, prominent beneath, at 110° at the base, 45° at
the apex; tertiary nerves remotely scalariform, promi-
nent beneath; petiole 2.54 cm long, stout. Panicle to
20 cm long, ribbed, densely shortly fulvous scabrid
tomentose, lax, axillary; singly branched, branchlets
to 6 cm long, bearing to 12 secund flowers; bracteoles
to 8 by 5 mm, oblong-ovate, subacute, shortly evenly
caducous pubescent. Flower bud to 7 by 3 mm,
lanceolate. Sepals densely pubescent; 3 outer ovate,
acute, 2 inner ovate, thinner at the fimbriate margins,
narrower at base, than the outer 3. Petals pink
outside, crimson within, elliptic-lanceolate, pubescent
on parts exposed in bud. Stamens 15, the inner 5
slightly longer than the outer 10 and reaching the style
apex at anthesis; filaments broad and compressed at
base, tapering abruptly medially and filiform below
the broadly ellipsoid anthers; appendage to connec-
tive slender. 2-3 times length of anther, glabrous.
Ovary ovoid, densely pubescent, crowned by a some-
what longer stoutly columnar densely pubescent
stylopodium and short glabrous style. Fruit pedicel to
3 mm long, to 3 mm @g, stout. Fruit calyx at first
sparsely pubescent towards base, becoming entirely
glabrous; 3 longer lobes to 14 by 2.5 cm, spatulate,
obtuse, c. 8 mm broad above the to 1.3 by 1.4 cm
broadly ovate saccate thickened base; 2 shorter lobes
to 12 by 1.2 cm, lorate, obtuse, otherwise as in longer
lobes. Nut to 3 by 1.8 cm, narrowly ovoid, densely
shortly evenly pale cream-brown pubescent; stylopo-
dium to 4 mm long, apiculate.

Distr. Malesia: N.W. Borneo (W. and Central
Sarawak).

Ecol. Scattered on leached clay soils on low hills in
Mixed Dipterocarp forest.

Vern. Méranti bulu merah.

118. Shorea palosapis (BLCO) Merr. Sp. Blanc. (1918)
271; En. Philip. 3 (1923) 98; Reyes, Philip. J. Sc. 22
(1923) 325.— Dipterocarpus palosapis Btico, FI.
Filip. ed. 2 (1845) 312; ibid. ed. 3, 2 (1878) 214; DC.
Prod. 16, 2 (1868) 614; Dyer. J. Bot. 12 (1874)
108. — Hopea squamata Turcz. Bull. Soc. Nat.
Mosc. 31 (1858) 239: DC. Prod. 16, 2 (1868) 635:
Wacp. Ann. 7 (1868) 379. — S. squamata BENTH. &
Hook. f. [Gen. Pl. 1 (1862) 193] ex DC. Prod. 16, 2
(1868) 632; VipaL, Phan. Cuming. (1885) 97; Rev. Pl.
Vasc. Filip. (1886) 62; BRANbIs, J. Linn. Soc. Bot. 31
(1895) 92; BRANDIS & Gitc in E. & P. Pfl. Fam. ed. 1,
3, 6 (1895) 267: MerR. Philip. J. Sc. 2 (1907) Bot. 285;
Foxw. Philip. J. Sc. 2 (1907) Bot. 386; MERRITT, Bull.
Bur. For. Philip. 8 (1908) 16, 48; Foxw. Philip. J. Sc. 4
(1909) Bot. 519; WuitForD, ibid. 4 (1910) Bot. 715:

Bull. Bur. For. Philip. 10 (1911) 66; Foxw. Philip. J.
Sc. 6 (1911) Bot. 275; in Elmer, Leafi. Philip. Bot. 6
(1913) 1957; Philip. J. Sc. 13 (1918) Bot. 191; MERR.
En. Born. (1921) 407; Foxw. Philip. J. Sc. 67 (1938)
310. — S. floribunda (non KuRZ) F.-VILL. Nov. App.
(1880) 21. — S. rugosa (non HEM) Foxw. Philip. J. Sc.
13 (1918) Bot. 191.

Large buttressed tree. Midrib and nerves above,
branchlets of panicle, bracteoles, ovary and parts of
petals exposed in bud + densely persistently cream
puberulent; calyx at first so, becoming sparse, glabres-
cent; twigs, buds, bracts, stipules, petioles and ner-
vation beneath + sparsely persistently scabrid pubes-
cent. Twig 2-3 cm @ apically, rather stout, ribbed,
becoming terete, smooth; stipule scars pale, distinct,
amplexicaul. Bud to 15 by 10 mm, ovate, acute,
compressed. Stipules to 1.5 by 11 mm, ovate-auricu-
late, prominent, not at first caducous. Leaves 12-24 by
8-11 cm, large, oblong, thinly coriaceous; base +
shallowly cordate or sometimes obtuse; apex obtuse
or to 2 cm long slender acuminate (young trees):
nerves 14-19 pairs, prominent beneath, + depressed
above as also the midrib and tertiary nerves, set at c.
45° towards the apex, at c. 90° at the base, usually with
short less prominent secondary nerves; tertiary nerves
remotely scalariform; petiole 1.5 to 2.5 cm long, rather
stout. Panicles to 25 cm long, terminal or subterminal
axillary, stout, spreading, many-flowered, singly (if
axillary) or doubly branched; branchlets to 3 cm long.
bearing to 8 flowers; bracts as stipules but somewhat
smaller, bracteoles to 6 by 3 mm, elliptic, subacute,
neither at first caducous. Flower bud to 6 by 3 mm,
fusiform. Sepals narrowly ovate; outer 3 acute, inner 2
slightly shorter, prominently acuminate. Stamens 15,
unequal, slightly shorter than style at anthesis; fila-
ments broadly compressed at base, abruptly tapering
and filiform distally; appendages slender, glabrous,
34 x length of the subglobose anthers. Ovary and
stylopodium pyriform, densely pubescent; style some-
what shorter, filiform, glabrous. Fruit pedicel to 2mm
long, stout; 3 longer calyx lobes to 12(—17) by 1.5 cm,
spatulate, coriaceous, obtuse, c. 6 mm broad above
the to 12 by 10 mm ovate saccate thickened base; 2
shorter lobes to 3.5 by 0.3 cm, linear, similar at base;
nut to 23 by 14 mm, ovoid, acute.

Distr. Malesia: Philippines.

Ecol. Widespread, often abundant, in lowland
evergreen Mixed Dipterocarp forest on fertile well-
drained soils in non-seasonal areas below 300 m.

Vern. Mayapis, tabak (Tag.), pura (Bik.), kalian
(Lan.), alam (Tag., Mang.).

119. Shorea bullata AsHTON, Gard. Bull. Sing. 19
(1962) 283, pl. 15; Man. Dipt. Brun. (1964) 181, f. 17;
ibid. Suppl. (1968) 105.

Medium-sized tree with golden brown cracked and
patchily flaked bark. Outside of stipule, twig, panicle,
petiole and nervation beneath persistently scabrid
fulvous pubescent; more densely tomentose on leaf
buds and midrib above; caducous hispid on lamina
above; shortly pubescent on stipule within. Twig c. 2

FLORA MALESIANA

[ser. I, volP9?

DPBS + 5 Se ‘f : <
Fig. 102. Trunk of Shorea pubistyla ASHTON. Sar-
awak, Semengoh Arboretum (Photogr. ASHTON).

mm @ apically, becoming thinly papery flaked, terete,
much branched; stipule scars short, horizontal, ob-
scured by tomentum. Bud 3-4 by 2-3 mm, broadly
ovoid. Stipule to 7 by 2 mm, narrowly hastate, acute.
Leaves 6.5-10 by 3-4.5 cm, chartaceous, frequently
cupped, elliptic to oblong-ovate; base narrowly
obtuse or broadly cuneate; apex obtuse or with to 5
mm long short acumen; margin usually revolute;
nerves 10—12 pairs; tertiary nerves well spaced, scalari-
form, at 90° to nerves; nervation depressed above;
petiole 0.8—1 cm long, short. Panicle to 11 cm long,
terminal or axillary, terete, lax, regularly singly (or
doubly if terminal) branched; branchlets to 3.5 cm
long, bearing to 8 + secund flowers; bracts unknown;
bracteoles to 6 by 3.5 mm, broadly elliptic, obtuse,

pale grey puberulent. Flower bud to 15 by 3 mm,
fusiform, obtuse. Calyx small, densely pale greyish
pubescent outside, glabrous within; lobes subequal,
broadly ovate to suborbicular. Petals pale cream pink
at base, linear, obtuse, densely pubescent on parts
exposed in bud, loosely adhering on falling. Stamens
15, congested, the 5 inner longer by an anthers length,
the outer 10 subequal; filaments connate at base,
broad, tapering abruptly and filiform distally; anthers
oblong; appendages to connective to 4 times length of
anther, exceeding style apex, very slender, twisted.
Ovary and stylopodium ovoid, densely shortly pubes-
cent; style almost twice their length, the basal half
thickened and setose. Fruit calyx persistently sparsely
pale buff pubescent at base, glabrescent towards apex;
2 longer lobes to 9 by 1.3 cm, spatulate, obtuse, to 5
mm broad above the to 6 by 6 mm prominently
saccate thickened base; 2 shorter lobes to 5 by 3 mm,
linear, obtuse, similar at base. Nut to 10 by 7 mm,
ovoid, shortly densely cream pubescent, acute.

Distr. Malesia: Northern Borneo (N.E. Sarawak,
Brunei).

Ecol. Rare, yellow sandy soils in Mixed Diptero-
carp forest below 800 m.

Vern. Méranti mélechur.

120. Shorea flaviflora Woop ex ASHTON, Gard. Bull.
Sing. 19 (1962) 289, pl. 18; Man. Dipt. Brun. (1964)
188, f. 17; ibid. Suppl. (1968) 108; MEER & Woop,
Sabah For. Rec. 5 (1964) 107. — Fig. 100 C—C2.
Medium-sized tree, often misshapen. Young parts
shortly pale grey pubescent with minute adpressed
hair tufts, caducous except on panicle, leaf bud and
stipule. Twig 2-3 mm @ apically, terete, glabrous,
smooth, pale brown; stipule scars 1.5—2 mm long at
first, narrow, ascending, half encircling twigs. Bud
6-12 by 1.5-3 mm, acute, falcate. Stipule to 24 by 7
mm, narrowly hastate, acute, fugaceous. Leaves
12-18 by 5.5—7 cm, thinly coriaceous, narrowly ovate;
base obtuse or broadly cuneate; acumen to 1.5 cm
long, slender; nerves 6—7 pairs, curved, well spaced,
slender, prominent beneath, at c. 40°—50°, with small
axillary domatia; midrib applanate above, prominent
beneath; tertiary nerves very slender, diagonal to
nerves, densely scalariform; petiole 2-3 cm long.
Panicle to 15 cm long, terminal or axillary, pendent,
terete, rugose on drying; doubly branched at to 3 cm
intervals, the branchlets bearing to 12 distichous
flowers; bracteoles to 5 by 2.5 mm, elliptic, obtuse,
densely shortly pale grey pubescent outside, sparsely
pubescent within. Flower bud to 14 by 4.5 mm, large,
fusiform, obtuse. Ca/yx glabrous but for the setose
margin: lobes equal, broadly ovate, thickened, closely
adpressed at the corolla. Petals rich yellow, narrowly
lanceolate, acute, firmly connate at base, strongly
contorted and twisted, imbricate, the basal half
forming a broad cup when open, shortly pubescent on
parts exposed in bud. Stamens 15, of 3 distinct lengths;
filaments expanded at base, abruptly tapering and
filiform distally; anthers large, reniform, cells indis-
tinct; appendage to connective c. 3 x length of anther,

1982]

DIPTEROCARPACEAE (Ashton)

519

the longest reaching half length of style. Ovary and
stylopodium pyriform, shortly densely pubescent; style
as long as ovary and stylopodium, filiform, glabrous.
Fruit calyx glabrous; 3 longer lobes to 13 by 3 cm,
broadly spatulate, obtuse, to 11 cm broad above the to
1.0 by 1.2 cm deeply saccate thickened base; 2 shorter
lobes to 8 by 1.3 cm, subequal, similarly saccate. Nut
to 2.5 by 1.3 cm, ovoid, shortly densely buff pubes-
cent; stylopodium to | cm long, tapering.

Distr. Malesia: N. Borneo (Rejang hinterland,
Crocker Range and Kinabalu).

Ecol. High hill ridges (150—)400—1300 m; locally
common.

Vern. Sélangan merah bukit (Brunei), séraya daun
bésar (Sabah).

121. Shorea monticola ASHTON, Gard. Bull. Sing. 19
(1962) 297, pl. 22; Man. Dipt. Brun. (1964) 200, f. 17:
ibid. Suppl. (1968) 111; MEUER & Woop, Sabah For.
Rec. 5 (1964) 121.

Medium-sized or relatively large tree. Young twig,
leaf bud, petiole and nervation at first shortly yellow-
ish buff pubescent, caducous on all but bud. Twig to 3
mm @ apically, stout, terete, becoming smooth but for
the minute pale elliptic warty lenticels; stipule scars
subamplexicaul or amplexicaul, pale, prominent. Bud
5-10 by 24 mm, narrowly ovate, acute, falcate.
Stipule unknown. Leaves 8-13 by 5-8 cm, thickly
coriaceous, elliptic; undersurface golden lepidote at
first, becoming mauve-grey; base obtuse or broadly
cuneate; acumen to 5 mm long, short, broad; nerves
13-16 pairs, prominent beneath, curved towards
margin, at 45°—55°; tertiary nerves slender, densely
scalariform, diagonal to nerves; midrib stout, promi-
nent, sharply ridged beneath, obscurely depressed
above; petiole 2-2.5 cm long. Panicle to 12 cm long,
terminal or axillary, terete, densely pale tawny pubes-
cent; singly branched, branchlets bearing to 12 disti-
chous flowers. Bracts and bracteoles unknown. Flower
bud to 7 by 3 mm, lanceolate, subacute. Calyx shortly
pubescent outside, glabrous within; 3 outer lobes
narrowly ovate, obtuse; 2 inner lobes c. 3 as long,
broader, ovate, acute, constricted at base. Petals
oblong, obtuse, shortly pubescent on parts exposed in
bud. Stamens 15, in 3 verticils, the inner 5 somewhat
longer than the rest; filaments exposed in bud, broad
at base, tapering abruptly, filiform distally; anthers
broadly oblong; appendage to connective 2-3 times
length of anther, slender. Ovary and stylopodium
pyriform, shortly pubescent; style as long as ovary,
filiform, glabrous. Fruit calyx puberulent at base,
otherwise glabrous; 3 longer lobes to 7 by 1.5 cm,
spatulate, obtuse, only slightly tapering above the to
1.5 by 1.5 cm shallowly saccate thickened base; 2
shorter lobes to 3.5 by 0.6 cm, narrowly oblong,
similar at base. Nut to 1.6 by 1 cm, ovoid, shortly
acute, densely shortly pubescent.

Distr. Malesia: N. Borneo (West Borneo to Kina-
balu and Trusmadi).

Ecol. Mountains, between 1000-1500 m:; locally
common.

Vern. Méranti gunong (Brunei), séraya gunong

(Sabah).

122. Shorea kunstleri KiNG, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 116; BRUHL & KING, Ann. R. Bot. Gard.
Calc. 5, 2 (1896) 154, t. 186; BRANDis, J. Linn. Soc.
Bot. 31 (1895) 93; Ripv. Fl. Mal. Pen. 1 (1922) 228;
Foxw. Mal. For. Rec. 3 (1927) 51; ibid. 10 (1932) 204;
Burk. Dict. (1935) 2012; Sym. Mal. For. Rec. 16
(1943) 72, f. 38, 44; BROWNE, For. Trees Sarawak &
Brunei (1955) 154; ASHTON, Man. Dipt. Brun. (1964)
192. f. 17, pl. 50 (habit, bark); ibid. Suppl. (1968) 109;
MEER & Woop, Sabah For. Rec. 5 (1964) 109.

Large buttressed tree. Young parts shortly pale buff
puberulent, + caducous on twig, petiole and leaf,
more dense and persistent on panicle, leaf bud and
stipule outside (sparse within). Twig c. 2mm @ at
apices, much branched, ribbed at first, becoming
terete, smooth or rugulose; stipule scars + horizontal,
c. 2mm long, cuneate, cream-grey. Bud 5—7 by 2-2.5
mm, falcate. Stipule to 1.2 by 0.4 cm, narrowly
lanceolate, acute, fugaceous. Leaves 8-12 by 4.5—7 cm,
broadly ovate, coriaceous, frequently twisted to one
side, normally glabrous (rarely pubescent); base
obtuse or broadly cuneate; acumen to | cm long;
nerves 6-8 pairs, slender, hardly raised beneath,
curved, at c. 50°-60°, with or without minute axillary
domatia; midrib applanate; tertiary nerves slender,
scalariform, at 90° to midrib; petiole 2-3.5 cm long.
Panicle to 15 cm long, terminal (rarely axillary),
slender, slightly compressed on drying; regularly
singly or doubly branched, the branchlets lax, bearing
to 9 secund flowers; bracteoles to 5 by 3.5 mm, elliptic,
obtuse, shortly densely pale greyish pubescent. Flower
bud to 9 by 3 mm, narrowly ovoid-lanceolate, acute. 3
outer sepals longer, deltoid, acuminate; 2 inner ovate,
acuminate. Corolla pale yellow, pink at base, shortly
pubescent on parts exposed in bud; petals narrowly
lanceolate, obtuse, twisted, slightly saccate. Stamens
15, the inner 5 only slightly longest; filaments
expanded at base, narrowing abruptly distally;
anthers broadly oblong; appendage to connective
more than twice length of anther, the inner 5 reaching
almost to style apex. Ovary and stylopodium pyriform,
shortly pubescent; style filiform, glabrous but for the
sparsely pubescent base, somewhat shorter than
ovary. Fruit calyx entirely glabrous; 3 longer lobes to
8.5 by 1.8 cm, spatulate, narrowly obtuse, c. 5 mm
broad above the c. 1.2 cm long and broad strongly
saccate thickened base; 2 shorter lobes to 4 by 0.4 cm,
unequal, linear, similarly saccate at base. Nut to 2 by
1.5 cm, ovoid, densely shortly buff pubescent; style
remnant c. 3 mm long, conical.

Distr. Malesia: Malaya (Perak and E. coast), N.
Sumatra (Langsa, Atjeh), Borneo (Lower Kapuas,
West Borneo; Sarawak; E. Sabah and S.E. Borneo to
Sampit).

Ecol. Local on infertile leached sandy clay soils on
low hills and ridges, rarely to 800 m; on ultrabasics in
Sabah.

Vern. Damar laut mérah, méranti pahang, sélimbar,

520 FLORA MALESIANA [ser. I, vol. 92

A B2

Fig. 103. Flower details in Shorea sect. Pachycarpae HEM. All x 10. — S. pinanga ScuerF. A. Bud, A/. stamens
from inside, A2. pistil. — S. rotundifolia ASHTON. B. Bud, B/. stamens from inside, B2. pistil (A Jacoss 5330,
BS 29226).

1982]

DIPTEROCARPACEAE (Ashton) 521

téngkawang bukit, t. batu, térbak, séraya, s. kitan,
balau mérah (Malaya).
Note. Collections from the Kapuas valley differ in

having the lamina somewhat bullate between the
nerves, which are slightly more numerous and more
ascending than in the type.

8. Section Pachycarpae

Hem, Rech. Dipt. (1892) 44; AsHton, Gard. Bull. Sing. 20 (1963) 269
(‘Pachycarpa’); Man. Dipt. Brun. (1964) 118. — Shorea sect. Pinanga BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 90. — Fig. 103, 104.

Flower buds ovoid or fusiform; corolla as in sect. Rubella; stamens 15, in 3
subequal verticils; filaments lorate, adnate along their margins thus forming a
tube round the ovary, tapering + abruptly below the anthers; anthers subglobose
or broadly oblong; appendage to connective filiform, slender, glabrous, erect,
2—co times length of anther; ovary small, glabrescent or glabrous; style filiform,
stylopodium indistinct, or both spindle-shaped, tapering distally and basally. 3
outer fruit calyx lobes lorate, broad at base. Stipules, bracts and bracteoles
persistent, large. Leaf with scalariform tertiary nerves; midrib above evident, +
depressed. Small or large stoutly buttressed trees. Bark surface remaining smooth
and hoop-marked longer than in other sections, later becoming + flaky,
sometimes scroll marked. Wood as in sect. Brachypterae.

Distr. Malesia: Borneo.

Ecol. In lowland forest below 1200 m.

Vern. Langgai. éngkabang (Iban, Sarawak), téngkawang, abang.

Note. Flowers of S. splendida and S. stenoptera are visited by small Hymenoptera as well as thrips (S.
APPANAH) and are thought to be pollinated by them. The only uniform and constantly distinguishable
widespread species is S. mecistopteryx; all others exhibit much geographical variability, with high uniformity
within populations nevertheless, and apparent hybridization with one or more species in some part of their
range. Embryogenesis appears to be normal, self-incompatibility high (CHAN H. T.) and the group appears

therefore to be undergoing active speciation.

123. Shorea pilosa ASHTON, Gard. Bull. Sing. 19
(1962) 304, pl. 25; Man. Dipt. Brun. (1964) 209, f. 18,
pl. 55 (slash, bark); ibid. Suppl. (1968) 115; MEUER &
Woop, Sabah For. Rec. 5 (1964) 136.

Medium-sized to large tree. Twig, leaf bud, petiole,
both surfaces of midrib, leaf beneath and panicle
densely persistently scabrid gold tomentose; leaf
above puberulent or glabrescent. Twig 2-2.5 mm @
apically, slightly compressed, becoming terete, much
branched; stipule scars narrow, almost horizontal.
amplexicaul. Leaf bud 3-7 by 1.5—3 mm, ovoid, obtuse
or subacute. Stipule to 3.5 by 1.3 cm, hastate, acute;
base slightly constricted; relatively persistent. Leaves
10-17 by 47.5 cm, ovate or elliptic, somewhat
coriaceous: base obtuse; acumen to 1.3 cm, long,
narrow; margin frequently narrowly revolute: nerves
12-15 close slightly curved pairs, at 45°—-50°, with
hispid axillary domatia; tertiary nerves densely scalar-
iform, at c. 90° to the nerves; petiole 1—-1.3 cm long,
short. Panicle to 14cm long, singly branched, terminal
or axillary, terete. Flower bud to 8 by 3 mm, lanceolate.
Sepals narrowly deltoid, subequal, densely pubescent
on parts exposed in bud. Petals cream tinted with

pink, lanceolate, densely pubescent on parts exposed
in bud. Stamens 15, the inner 5 slightly the longest:
filaments compressed, connate, abruptly tapering
below the broadly ellipsoid anthers; appendage to
connective aristate, c. 4 x length of anthers. Ovary
ovoid, glabrous; style and stylopodium spindle-
shaped, glabrous. Fruit calyx glabrescent: 3 longer
lobes by 17 by 2.5 cm, subequal, oblong-spatulate,
obtuse, c. 1.3 cm broad above the c. 1.3 by 1.5 cm
thickened deeply saccate base, adpressed to and
hiding the nut; 2 shorter lobes to 12 by 1 cm, spatulate,
similar at base. Nut to 2 by 1.8 cm, ovoid, abruptly
constricted below the to 1 cm long narrow-conical apex:
style remnant acute, densely shortly buff pubescent.

Distr. Malesia: N.W. Borneo (Kapuas hinterland,
W. Borneo, to W. Sabah).

Ecol. Local, on sandy clay soils on undulating land
and low hills.

Vern. Kawang bulu (Brun.), langgai (Iban).

Note. Collections from Sabah and Brunei differ in
their distinctly buff tomentum and prominent tomen-
tose leaf domatia. Those from W. Borneo could be
interpreted as a hybrid with S. amplexicaulis.

FLORA

CG

Fig. 104. Flower details in Shorea sect. Pachycarpae

Hem. All x 10.— S. beccariana BuRCK. C. Outer

sepal, C/. inner sepal, both from inside, C2. stamens

from inside, C3. pistil. — S. macrophylla (DE VRIESE)

ASHTON. D. Stamens from inside, D/. pistil (C JAcoBs
5563, D bb. 29722).

MALESIANA

[ser. I, vol. 9?

124. Shorea splendida (DE VRIESE) ASHTON, Gard.
Bull. Sing. 20 (1963) 279; Man. Dipt. Brun. Suppl.
(1968) 119, f. 15.— Hopea splendida DE VRIESE,
Minjak Tengkawang (1861) 28.—S. martiniana
SCHEFF. Nat. Tijd. N. I. 32 (1873) 408; BuRCK, Ann.
Jard. Bot. Btzg 6 (1887) 208, t. 29, f. 2; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 78; HEYNE, Nutt. Pl. ed. 1, 3
(1917) 306; MerrR. En. Born. (1921) 405; HEYNE, Nutt.
Pl. ed. 2 (1927) 1123; BACKER & BAKH. f. Fl. Java |
(1963) 331; BROwNE, For. Trees Sarawak & Brunei
(1955) 140.

Small to medium-sized tree. Vegetative parts gla-
brous. Twig c. 3 by 1 mm @ apically, smooth,
compressed, ribbed; stipule scars to 1 mm thick,
prominent, amplexicaul, pale. Bud c. 8 by 6 mm, a
loose aggregation of young leaves and stipules. Stipule
to 2.5 by 1.5 cm, prominent, subpersistent, ovate,
acute or subacute, with cordate subequal base. Leaves
8.5—23 by 4.2-11 cm, oblong, chartaceous, undulate,
+ lustrous; base typically cordate, sometimes obtuse
or broadly cuneate; acumen to | cm long; nerves 9-12
pairs, slender but prominent beneath, at 45°-65° to
the midrib; tertiary nerves sinuous, remotely scalari-
form, vertical to the nerves; midrib prominent
beneath, applanate above; petiole 11-22 mm long,
drying rugose. Panicle to 20 cm long, terminal or
axillary, glabrous, terete, ascending; singly branched,
branchlets to 7 cm long, bearing to 10 flowers; bracts
to 18 by 8mm, as stipules, caducous; bracteoles to 6 by
4 mm, elliptic, obtuse, subpersistent, prominent.
Flower bud to 10 by 3 mm, lanceolate. Sepals narrowly
deltoid, acute, subequal, sparsely pubescent on parts
exposed in bud. Petals lanceolate, sparsely pubescent
on parts exposed in bud. Stamens 15, the inner 5
longer than the outer 10; filaments compressed and
connate along ? of their length, abruptly tapering
below the broadly ellipsoid anthers; appendage to
connective 4-5 times length of anthers, almost reach-
ing style apex, slender, sericeous in the distal 4. Ovary
ovoid, glabrous; style and stylopodium spindle-
shaped, glabrous. Fruit pedicel to 7 mm long, to 4mm
@, large. Calyx glabrous; 3 major lobes to 7.5 by 2.3
cm, narrowly oblong to broadly spatulate, c. 12 mm
broad above the to 22 by 18 mm ovate saccate
thickened base; 2 inner lobes to 6.5 by 1.2 cm, as long
as or somewhat exceeding the nut, oblong, similar at
base. Nut to 5.5 by 3 cm, ovoid, acute, shortly densely
evenly pale buff pubescent.

Distr. Malesia: Borneo (Kapuas valley, Sarawak
west of the Lupar; Muara Tewe, planted?).

Ecol. Frequent, locally abundant, on clay-rich
periodically flooded alluvium, sometimes planted.

Uses. An important producer of Borneo illipe fat
(buah téngkawang).

Vern. Téngkawang, t. rambai (W. Borneo), mélin-
dang, t. lémying (Muara Tewe), éngkabang rambai
(Sarawak).

Notes. Though variable, the leaf shape and broad,
cordate, subpersistent stipules make this species
generally easy to recognize; possibly occasionally
hybridizing with S. pinanga.

1982]

Some populations in Kapuas appear to hybridize
with S. stenoptera.

125. Shorea stenoptera BurcK, Med. Lands PI. Tuin
3 (1886) 11; Ann. Jard. Bot. Btzg 6 (1887) 209, t. 21;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 78; BRANDIS &
GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895) 264; HEYNE,
Nutt. Pl. ed. 1, 3 (1917) 308; Merr. En. Born. (1921)
407; GiLG in E. & P. Pfl. Fam. ed. 2, 21 (1925) 260;
HeEYNE, Nutt. Pl. ed. 2 (1927) 1113, 1125; BROWNE,
For. Trees Sarawak & Brunei (1955) 144; ASHTON,
Man. Dipt. Brun. Suppl. (1968) 120, f. 15.

Small tree. Twig apex, petiole and midrib above
sometimes densely ocherous pubescent (immature
tree?), more commonly glabrous; stipule outside
occasionally sparsely pubescent. Twig 3-4 mm @
apically, somewhat compressed to terete, becoming
smooth; stipule scars pale, prominent, descending,
amplexicaul. Bud 4 by 3mm, ellipsoid, obtuse, usually
obscured by stipules. Stipule to 2 by | cm, ovate,
deltoid, obtuse, subauriculate at base, subpersistent.
Leaves 18-40 by 8-22 cm, large, oblong, thickly
coriaceous; base broadly cuneate to cordate; acumen
to 2cm long, + prominent; nerves 10-14 pairs, stout,
prominent, beneath, at 45°-60° along the midrib, to
110° near its base; tertiary nerves remotely scalari-
form; midrib evident but applanate above, prominent
and terete beneath; petiole 2.3-4.5 cm long, stout.
Panicle to 35 cm long, terminal or axillary, glabrous or
(rarely) sparsely pubescent towards the base; borne
behind the twig apices in the axils of fugaceous
rudimentary leaves, a chain of short internodes being
concealed in a dense profusion of the straight terete
inflorescences with ascending branchlets; branchlets
to 10 cm long; bracts and bracteoles to 6 by 4 mm,
identical, ovate, acute, glabrous, not at first caducous.
Flower bud to 7 by 3 mm, lanceolate. Sepals deltoid,
acute, densely pubescent outside, sparsely so within;
inner 2 smaller, relatively narrower than outer 3.
Petals deep pink, lanceolate, sparsely pubescent on
parts exposed in bud. Stamens 15, the inner 5
somewhat longer than the others and exceeding the
style apex; filaments compressed, lorate, connate at
margins along 2 of their length, tapering abruptly
below the ellipsoid anthers; appendage to connective
slender, 3-4 x length of anther, sericeous towards the
apex. Ovary narrowly ovoid, glabrous; style and
stylopodium spindle-shaped, slender, glabrous. Fruit
pedicel to 4 mm long and @, prominent. Calyx
glabrous; 3 longer lobes to 7.5 by 2 cm, spatulate,
obtuse, to 1 cm broad above the to 2.5 by 2 cm ovate
thickened saccate base; 2 shorter lobes to 5.5 by 0.8
cm, lorate-spatulate, obtuse, similar at base. Nut to 5
by 3 cm, ovoid, apiculate, large, densely shortly evenly
buff pubescent.

Distr. Malesia: Borneo (Kapuas valley, Sarawak
west of the Saribas; Muara Tewe, planted?).

Ecol. Locally common on + poorly drained sandy
soils on alluvium and plateaux at low altitudes.

Vern. Téngkawang layar, t. tungkul, t. rambai, t.
tajan, t. télur (W. Borneo), éngkabang rusa (Sarawak).

DIPTEROCARPACEAE (Ashton)

323

Notes. A remarkably variable species. Sometimes a
big tree, when it may differ from S. macrophylla only
in the glabrous midrib and could be a hybrid. Usually
small, but even then differing greatly between pro-
venances; in West Sarawak the inflorescences are
always axillary, born in groups between short inter-
nodes and subtended by modified or no leaves.

Some cultivated plants at Kepong flower almost
annually, while one of the provenances planted at
Haurbentes in West Java flowers almost con-
tinuously; it is said that the wild parents of the latter
also did so.

126. Shorea macrophylla (DE VRIESE) ASHTON, Gard.
Bull. Sing. 20 (1963) 278; Man. Dipt. Brun. (1964)
196, f. 18, 20; ibid. Suppl. (1968) 110, pl. 22 (illipe
nuts); MEVER & Woop, Sabah For. Rec. 5 (1964)
156. — Hopea macrophylla DE VRIESE, Minjak Teng-
kawang (1861) 28.— S. gysbertsiana BURCK, Med.
Lands Pl. Tuin 3 (1886) 15; Ann. Jard. Bot. Btzg 6
(1887) 211, t. 23, 30, f. 2-3, incl. var. scabra BURCK,
l.c., p.p.; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 93;
HEyYNngE, Nutt. Pl. ed. 1, 3 (1917) 301; Merr. En. Born.
(1921) 405; HeyNg, Nutt. Pl. ed. 2 (1927) 1113, 1114,
1119; FiscHER, Kew Bull. (1932) 177; BROWNE, For.
Trees Sarawak & Brunei (1955) 139; MEWER & Woop,
Sabah For. Rec. 5 (1964) 45, 108, 156, f. 1d. — S.
bakeriana Hem, Bull. Mens. Soc. Linn. Paris 2 (1891)
974; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 96; MERR.
En. Born. (1921) 404. — Pachychlamys gysbertsiana
Ript. Fl. Mal. Pen. 1 (1922) 233 (‘ghysber-
tiana’). — Fig. 104 D-D1.

Medium-sized to large tree. Young twig, leaf bud,
stipules, petiole, leaf beneath and midrib above
persistently densely evenly caducous pale brown
pubescent. Twig 2.5—4 by 2-3 mm @ apically, com-
pressed, becoming smooth, glabrous; stipule scars to |
mm thick, amplexicaul, pale cream, + horizontal,
smooth, glabrous. Bud 12-18 by 4-6 mm, compressed,
hastate, narrowly obtuse. Stipule to 5 by 1.3 cm,
broadly hastate, obtuse, constricted at base. Leaves
17-35 by 10-14 cm, elliptic-oblong, + coriaceous;
base obtuse or subcordate; acumen to 1.5 cm long;
nerves | 1—20 pairs, prominent beneath, well spaced, at
55°-65°; tertiary nerves distinct, well spaced, scalari-
form, at 90° to nerves; midrib prominent, rounded,
beneath, broad and applanate above; petiole 1.5—3 cm
long. Panicle to 17cm long, terminal or axillary, ribbed
and compressed when dry, shortly evenly persistently
buff pubescent, singly branched; bracteoles to 1.2 by
0.5 cm, oblong, subacute. Flower bud to 8 by 5 mm,
broadly or narrowly ellipsoid. Calyx lobes densely
pale brown pubescent outside, glabrous within; lobes
subequal, broadly elliptic to deltoid, obtuse to acu-
minate. Petals pale pink, broadly ovate, obtuse,
densely pubescent on parts exposed in bud. Stamens
15, of 2 lengths, the inner 5 an anther’s length longer
than the outer 10; filaments broad, compressed,
united in a tube round the ovary, tapering distally and
filiform below the oblong anthers; appendage to
connective c. 2 x length of anther, slender, glabrous.

524

Ovary ovoid, densely pubescent in the distal half;
stylopodium and style glabrous, spindle shaped, as
long as ovary. Fruit calyx glabrous; 3 longer lobes to
11 by 3 cm, coriaceous, oblong, obtuse, to 1.5 cm
broad above the to 1.8 by 2.3 cm horizontally elliptic
deeply saccate thickened base; 2 shorter lobes to 8 by
1.5 cm, similar at base, base obscured by larger lobes.
Nut to 6 by 4cm, obovoid, persistently evenly shortly
pale buff pubescent; style remnant, small, short, acute.

Distr. Malesia: Borneo (widespread, but especially
W. and Central Sarawak, Kapuas valley, Tidung and
Berau).

Ecol. Locally abundant on clay-rich periodically
flooded alluvium and river banks, uncommon on
hillsides, below 600 m.

Uses. The principal source of the Bornean IIlippe
nut; sometimes planted.

Vern. Téngkawang, t. jantong, t. guntjong, tukung
(West Borneo), éngkabang jantong (Sarawak, Iban),
kawang, k. jantong (Brun.), abang (Mur., Dusun, S.
Borneo), kékawang buah (Berau), méngkalang
(Tidung), orai tolloi (Muara Tewe).

Note. The form occurring in E. Sabah shares
certain characters (stipule scars, length of petiole,
smaller nut) of S. pinanga and suggests hybridisation.

127. Shorea praestans ASHTON, Gard. Bull. Sing. 22
(1967) 297, pl. 42; Man. Dipt. Brun. Suppl. (1968) 115,
f. 14.

Small tree. All parts apparently glabrous. Twigs c. 5
by 3 mm towards the apices, stout, compressed,
smooth; stipule scars slender but clearly evident,
amplexicaul. Bud to 25 by 7 mm, falcate-lanceolate,
compressed, acute. Stipules to 11 by S cm, very large,
elliptic, subacute, not at first caducous. Leaves 24-35
by 11-18 cm, large, + broadly oblong, thickly
coriaceous; base cordate; apex obtuse or shortly
broadly acuminate; nerves 11-13 pairs, obscure
above, prominent beneath, at 45°—70°, remotely sub-
scalariform, slender but evident beneath; midrib
applanate above, prominent beneath; petiole 4—-6.5 cm
long, stout, drying rugose. Flowers and inflorescences
unknown. Fruit entirely glabrous. Pedicel to 5 mm
long and g, stout. 3 longer calyx lobes to 15 by 3 cm,
spatulate, obtuse, glabrous, tapering to c. 1.5 cm
broad above the to 1.5 by 1.8 cm broadly elliptic
saccate thickened base; 2 shorter lobes to 10.5 by 1.5
cm, spatulate, subacute, similar at base. Nut to 2 by 2
cm, ovoid, glabrous; style remnant to 4 mm long,
filiform.

Distr. Malesia: Borneo (Central Sarawak).

Ecol. Rare, deep yellow sandy soils in Mixed
Dipterocarp forest on coastal hills.

Note. Vicarious with S. stenoptera, of which it
appears to be a segregate; specimens from the Saribas
valley in particular approach an intermediate leaf-
shape.

128. Shorea rotundifolia ASHTON, Gard. Bull. Sing. 22
(1967) 299, pl. 44; Man. Dipt. Brun. Suppl. (1968) 117,
f. 14. — Fig. 103 B-B2.

FLORA MALESIANA

[ser. I, vol. 92

Medium-sized tree. Young twig, petiole and stipule
pruinose, otherwise glabrous. Twig c. 3 by 2mm @
apically, terete or slightly compressed, smooth; stipule
scars amplexicaul, prominent. Bud to 20 by 8 mm,
large, lanceolate, compressed. Stipule to 6 by 2 cm,
lanceolate-falcate, obtuse, subpersistent. Leaves alter-
nate, 9-21 by 8-14 cm, broadly ovate to suborbicular,
coriaceous, lustrous; base obtuse to cordate; acumen
to 1 cm long, cuspidate; nerves 9-11 pairs, curved,
prominent beneath, at to 115° towards the base, 45°
towards the apex; tertiary nerves remotely scalari-
form; midrib applanate above, prominent and terete
beneath. Petiole 3-4 cm long, very long. Panicle to 15
cm long, terminal or axillary, lax, glabrous, com-
pressed; regularly singly branched, branchlets to 2cm
long, bearing remote distichous flowers; bracts to 10
by 6 mm, elliptic-lanceolate, acute, glabrous, fuga-
ceous; bracteoles to 8 by 4mm, similar. Flower bud to
10 by 4mm, fusiform. Calyx glabrous; lobes narrowly
deltoid, subequal, subacute. Petals lanceolate, shortly
pubescent on parts exposed in bud. Stamens 15, the 5
innermost slightly longest; filaments broad, connate,
tapering abruptly beneath the ellipsoid anthers;
appendages c. 3 x length of anther, aristate. Ovary
ovoid, glabrous, surmounted by a spindle-shaped
style and stylopodium. Fruit glabrous. Pedicelc.6mm
long, 3 longer fruit calyx lobes c. 13 by 2.5 cm,
spatulate, obtuse, c. 6mm broad above the c. 15 by 18
mm somewhat saccate thickened base; shorter lobes
to 8 by 0.7 cm, narrowly lanceolate, subacute, similar
at base. Nut c. 2.5 by 1.08 cm, ovoid, glabrous; style
remnant to 7 mm long.

Distr. Malesia: Borneo (Central Sarawak).

Ecol. Local on inland ridges at 300-500 m.

Notes. A local species apparently derived from S.
amplexicaulis with which it occurs.

Occurring as small groups or scattered individuals,
with some local differentiation even within its small
range.

129. Shorea amplexicaulis ASHTON, Gard. Bull. Sing.
19 (1962) 273, pl. 10; Man. Dipt. Brun. (1964) 177, f.
18, 20, pl. 53 (bark); ibid. Suppl. (1968) 104; MEVER &
Woop, Sabah For. Rec. 5 (1964) 95, f. 11. — Fig. 105.

Medium-sized to large tree. All parts, including
stipule (both surfaces), caducous or persistently pale
buff to golden pubescent. Twig to 3 by 2 mm @
apically, slightly compressed when young, slender,
becoming terete, glabrous, smooth; stipule scars c. 0.5
mm thick, horizontal, + amplexicaul. Bud 10-15 by
3-5 mm, hastate, obtuse. Stipule to 25 by 8 mm,
hastate, subacute, subpersistent. Leaves 11-21 by 5-8
cm, elliptic, coriaceous; base cuneate; acumen to | cm
long; nerves 9-12 pairs, curved, at 50’—70°, prominent
beneath; tertiary nerves distinct, remotely scalari-
form; midrib prominent and terete beneath, +
applanate above; petiole 15—2.5 cm long. Panicle to 24
cm long, terminal or axillary, lax, slender, com-
pressed; regularly singly branched, branchlets to 10
cm long, bearing to 11 distichous flowers; bracts to 13
by 8 mm, lanceolate, acute, glabrous, fugaceous;

1982]

v.Os

Vl
f/ =
_——
(ee

Fig. 105. Shorea amplexicaulis ASHTON. a. Habit, b.
fruit, c. nut, with part of indumentum, all x 5(aSAN
22690, b-c S 6502).

bracteoles to 11 by 5mm, oblong, obtuse, glabrescent
or persistently puberulent outside, glabrous within,
subpersistent. Flower bud to 10 by 3.5 mm, lanceolate,
obtuse. Calyx densely shortly pale grey pubescent
outside, glabrous within; lobes subequal, narrowly
deltoid, obtuse, slightly expanded at base. Petals pale
yellow, narrowly lanceolate, subacute, shortly pubes-
cent on parts exposed in bud. Stamens 15, the 5 inner
slightly longest; filaments broad, united in a ring,

DIPTEROCARPACEAE (Ashton)

525

tapering abruptly beneath the narrowly oblong
anthers; appendage to connective over twice as long as
anther, not reflexed. Ovary ovoid, somewhat longer
than ovary; no distinct stylopodium. Fruit impressed
at base. Calyx puberulent or glabrescent; 3 longer
lobes to 18 by 3 cm, oblong-spatulate, obtuse, to 2cm
broad above the to 1.6 by 2.2 cm broadly ovate saccate
thickened base; 2 shorter lobes to 13 by 0.8 cm, similar
at base. Nut to 3.7 by 2.5 cm, broadly ovoid,
persistently evenly densely gold-buff pubescent; style
remnant short, acute.

Distr. Malesia: Borneo (S.W. excepted).

Ecol. Widespread, often common, on clay soils,
especially ridges, from sea level to 700 m.

Vern. Kawang pinang, méranti kawang pinang lichin
(Brun.), /anggai (Iban), abang (Dus.), kawang bukit
(Sabah), orai lanyung, awang rambut (S.E. Borneo),
éngkabang pipit, tengkawang megeh télur (W. Bor-
neo).

Notes. Sometimes difficult to distinguish from S.
beccariana, especially when young; the amplexicaul
stipule scars and crimson young parts are typical of
the present species, and the golden-yellow stellate-
hairy young parts and leaf undersurface of S. beccar-
iana typically distinguish them.

In S.E. Borneo apparently hybridizing with S.
pinanga, in W. Borneo with S. pilosa (q.v.).

130. Shorea mecistopteryx RipL. Kew Bull. (1925)
280; SLoot. in Merr. Pl. Elm. Born. (1929) 203; Sym.
Gard. Bull. S. S. 9 (1938) 348; AsHTon, Man. Dipt.
Brun. (1964) 198, f. 18, pl. 52 (bark), ibid. Suppl.
(1968) 111; MEER & Woop, Sabah For. Rec. 5 (1964)
119, f. 2a, pl. 7 (habit). — S. chrysophylla RiwL. Kew
Bull. (1926) 470; SLooT. in Merr. Pl. Elm. Born. (1929)
202.

Large tree. Young twig, panicle, leaf bud, stipule
outside, petiole and leaf beneath shortly evenly +
persistently gold-tomentose. Twig to 6 by 2.5mm @
apically, compressed, becoming terete, smooth; sti-
pule scars initially to 2.5 mm long, cuneate, horizon-
tal. Leaf bud 5-8 by 2.5-5 mm, ovoid, compressed,
obtuse. Stipule to 25 by 7 mm, hastate, acute. Leaves
13-20(-30) by 6-10(—12) cm, oblong, chartaceous;
base cordate; acumen to 8 mm long, broad; nerves
16-20 pairs, curved, at 45°-65° along midrib and to
120° at base; tertiary nerves densely scalariform, at
90° to nerves; midrib prominent and terete beneath,
rather narrow, + applanate above; petiole 2.5—3.5 cm
long. Panicle to 12 cm long, terete or compressed,
terminal or axillary, singly branched; bracteoles to 7
by 2 mm, oblong, subacute, densely pubescent out-
side, sparsely so within. Flower bud to 12 by 4 mm,
lanceolate. Calyx densely shortly greyish pubescent
outside, glabrous within; lobes narrowly deltoid,
subequal, + patent, the 2 inner slightly shorter,
thinner, slightly expanded at base. Corolla shortly
pubescent on parts exposed in bud; petals linear,
subacute. Stamens 15, the 5 inner somewhat longer
than the others; filaments united in the basal half,
tapering to the anthers in the distal half; anthers

526

oblong; appendage to connective over twice as long as
anthers, not reflexed. Ovary narrowly ovoid-conical,
glabrous; stylopodium somewhat longer than ovary,
thickened in the distal half; style stout. Fruit pedicel to
6 mm long, stout. Fruit calyx glabrescent or persis-
tently puberulent at base; 3 longer lobes to 23 by 3.3
cm, spatulate, narrowly obtuse, to 1.8 cm broad above
the to 2 by 2.5 cm broadly ovate saccate thickened
base; 2 shorter lobes to 15 by 1.2 cm, linear, similar at
base, base enveloped by larger lobes. Nut to 4.2 by 2.5
cm, ovoid, persistently shortly evenly yellowish buff
pubescent; style remnant acute.

Distr. Malesia: Borneo.

Ecol. Local: undulating land and low hills on sandy
clay soils in Mixed Dipterocarp forest below 400 m.

Vern. Téngkawang layar (Kapuas), téntang pakar
(M. Tewe), éngkabang larai (Sarawak), rébah (Iban),
kawang tikus, kawang burong (Sabah).

Note. The most uniform and distinct species in the
section.

131. Shorea beccariana Burck, Ann. Jard. Bot. Btzg
6 (1887) 213; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 87;
Mere. En. Born. (1921) 404; BRowne, For. Trees
Sarawak & Brunei (1955) 138; ASHTON, Gard. Bull.
Sing. 20 (1963) 280; Man. Dipt. Brun. (1964) 180, f.
18, 20: ibid. Suppl. (1968) 105; MEER & Woop, Sabah
For. Rec. 5 (1964) 100. — S. franchetiana HEM, Bull.
Mens. Soc. Linn. Paris 2 (1891) 956; Mem. Ass. Frang.
Besancon 1892 (1893) 459. — S. beccarii DYER ex
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 87, nomen in
syn. — Fig. 104 C—C3.

Medium-sized or large tree. Young twig, panicle,
leaf bud, stipule (both surfaces), and petiole shortly
evenly densely deep rufous-brown puberulent,
becoming pale mauve-grey, sparsely so on nervation
beneath and midrib above. Twig 3-6 by 1-2.5 mm @
apically, compressed, becoming smooth, glabrous;
stipule scars to 2 mm long, shortly cuneate or falcate,
+ ascending, sometimes horizontal, prominent. Bud
7-11 by 3.5-4.5 mm, hastate, compressed, acute.
Stipule to 14 by 5 mm, small, oblong, obtuse,
fugaceous. Leaves 11—20 by 5.5—7 cm, elliptic to ovate;
coriaceous, gold lepidote beneath, becoming mauve-
grey in mature trees; base obtuse or broadly cuneate;
acumen to 8 mm long, broad; nerves 11-14 pairs,
curved, not strongly raised beneath, at 50°—60°, to 90°
at base; tertiary nerves remotely scalariform, slightly
diagonal to nerves; midrib prominent beneath, +
depressed above; petiole 2-4 cm long. Panicle to 20cm
long, terminal or axillary, lax, + compressed; singly
branched, branchlets to 2.5 cm long, short, bearing to
8 distichous flowers; bracts to 15 by 9 mm, oblong,
obtuse, shortly pubescent outside, puberulent within,
fugaceous; bracteoles to 13 by 8 mm, otherwise as with
bracts, caducous. Flower bud to 9 by 3 mm, lanceolate,
obtuse. Calyx outside shortly pubescent or glabres-
cent, glabrous within; lobes subequal, deltoid, the
inner 2 slightly shorter, thinner, somewhat expanded
at base. Petals pink, paler along margins, lanceolate,
obtuse, sparsely pubescent on parts exposed in bud.

FLORA MALESIANA

[ser. I, voleg:

Stamens 15, the inner 5 slightly longer; filaments
broad, tapering in the distal half, united ina ring at the
base; anthers oblong; appendage to connective 2-3 x
length of anther, reaching almost to style apex. Ovary
ovoid, frequently sparsely pubescent apically; style
and stylopodium cylindrical, 15-2 x length of ovary,
rather short, glabrous. Fruit base frequently im-
pressed. Calyx glabrescent, sparsely dotted with
minute hair tufts; 3 longer lobes to 19 by 2.7 cm,
spatulate, obtuse, to 1.2 cm broad above the to 1.2 by
1.8 cm broadly ovate deeply saccate thickened base; 2
shorter lobes to 10.5 by 0.9 cm, subequal, linear,
similar at base. Nut to 4 by 2.8 cm, broadly ovoid,
shortly persistently evenly pale buff tomentose; style
remnant to 4 mm long, conical.

Distr. Malesia: Northern Borneo (Ulu Kapuas,
Sarawak to Sabah and Nunukan).

Ecol. Common on deep leached soils in lowlands
and along dry sandstone and shale ridges to 1350 m.

Vern. Méranti langgai, kawang (Brun.), langgai
(Iban), abang (Dus.), séraya langgai (Sabah), téng-
kawang téngkal, t. raraing, t. bagok, t. layar, t. bénua,
t. tangga, éngkabang maha (W. Borneo).

Note. Closely allied to S. amplexicaulis.

132. Shorea pinanga Scuerr. Nat. Tijd. N. I. 31 (1870)
350; BurcK, Ann. Jard. Bot. Btzg 6 (1887) 211, t. 24,
30, f. 1; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 94, t. 2, f.
17-18; BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6
(1895) 267; Becc. For. Born. (1902) 570; Bull. Ort.
Bot. Napoli 2 (1904) 92; Merr. En. Born. (1921) 406;
SLOoT. in Merr. Pl. Elm. Born. (1929) 203; BACKER &
BAKH. f. Fl. Java 1 (1963) 331; ASHTON, Gard. Bull.
Sing. 20 (1963) 281; Man. Dipt. Brun. (1964) 210, f.
18, 20; ibid. Suppl. (1968) 115; MEWER & Woop, Sabah
For. Rec. 5 (1964) 137, pl. 9. — S. gysbertsiana BURCK
var. scabra BURCK, Med. Lands PI. Tuin 3 (1886) 17,

p.p. — S. compressa BuRcK, Med. Lands PI. Tuin 3
(1886) 26; Ann. Jard. Bot. Btzg 6 (1887) 212; BRANDIS,
J. Linn. Soc. Bot. 31 (1895) 87; MerrR. En. Born.
(1921) 404. — Fig. 103 A—A2.

Medium-sized or large tree. Young twig, panicle,
leaf bud, stipule and petiole shortly densely fuga-
ceously to persistently gold-brown pubescent; spar-
sely so on leaf beneath. Twig 4-6 by 2-3 mm @
apically, compressed, becoming smooth; stipule scars
to 3mm long, to 1.5mm thick at base, strongly falcate,
descending. Bud to 10 by 4 mm, hastate, subacute,
usually hidden by stipules. Stipule to 6 by 1.5 cm,
glabrescent, hastate, subacute, somewhat persistent.
Leaves 11-24 by 4-9 cm, elliptic to narrowly ovate, +
thinly coriaceous, with broadly cuneate to subcordate
base and to 1.5 cm long deltoid acumen; nerves 10—20
pairs, slender, curved, at 50°—70° but to 90° at base;
tertiary nerves densely scalariform, diagonal to
nerves; midrib prominent and terete beneath, +
applanate and narrow above; petiole 1.5—2.3 cm long,
slender. Panicle to 24 cm long, terminal or axillary,
lax, compressed; singly branched, the branchlets long,
bearing to 15 distichous flowers; bracts to 15 by 7mm,
lanceolate, acute, glabrous to pubescent, fugaceous;

1982] DIPTEROCARPACEAE (Ashton) 527

Fig. 106. Flower details in Shorea sect. Mutica BRANDIS. All x 10. Sepals drawn from inside. — S. slootenii

Woop ex AsHTon. A. Outer sepal, A/. inner sepal, A2. stamens from outside, A3. pistil. — S. platycarpa HEIM.

B. Outer sepal, B/. inner sepal, B2. stamens from outside, B3. pistil.— S. macrantha BRANDIS. C. Young

stamens from outside, C/. older stamens from outside, C2. pistil. — S. macroptera DyER. D. Outer sepal, D/.

inner sepal, D2. stamens from outside, D3. pistil (A Rost & AsaH 3365, B Neth. Ind. For. Serv. -E-2P-694. G
HAVILAND 2119, D SAN 36703 = NT 592).

528

Bt

FLORA MALESIANA

[ser. I;, volMg?

Fig. 107. Flower details in Shorea sect. Mutica BRANDIS subsect. Auriculatae ASHTON. — S. myrionerva SYM. ex
ASHTON. A. Bud, B/. outer sepal, B2. inner sepal, C. petal, D/. stamens from a young bud, D2. stamens from
mature bud, E. pistil, all x 10 (JAcoBs 5371).

bracteoles to 10 by 8 mm, broadly ovate, acute,
glabrous to pubescent, subpersistent. Flower bud to 10
by 3 mm, linear, subacute. Calyx glabrous or pubes-
cent outside; lobes subequal, linear, subacute. Petals
deep pink, hardly contorted, long, linear-lanceolate,
sparsely pubescent on parts exposed in bud. Stamens
15, the innermost 5 slightly the longest; filaments
broad, united in a ring round the ovary, tapering
abruptly distally; appendage to connective many
times length of anther, reaching to style apex. Ovary
subglobose, surmounted by a long narrow style and
stylopodium about twice its length, the latter 2
distended in the distal half; entirely glabrous. Fruit
calyx glabrescent; 3 longer lobes to 28 by 3.5 cm,
coriaceous, narrowly spatulate, narrowly obtuse, to
2.3 cm broad above the to 1.5 by 2.5 cm broadly ovate
saccate thickened base; 2 shorter lobes to 17 by 1.2 cm,
subequal, linear, similar at base. Nut to 2.3 cm long
and broad, broadly ovoid, shortly evenly pale yellow-

ish buff pubescent; style remnant to 2 mm long,
slender, acute.

Distr. Malesia: Borneo.

Ecol. Clay-rich soils especially on ridges below 700
m; locally common.

Vern. Langgai (Iban), méranti langgai bukit
(Brunei), kKawang pinang (Sabah), awang lanying, a.
kélalai, a. boi, a. labuan, abang burong, kakan merah (S.E.
Borneo), éngkabong bukit, tengkawang amung, t. télur,
t. kérayung, t. gunong, t. sambai, t. bunga, t. minggi, t.
réput, t. lumut, t. umba, t. télaga, etc. (W. Borneo).

Notes. This, and to a lesser extent the two previous
species, varies greatly and almost continuously in
tomentum density, and also in leaf shape, number of
nerves and relative length of petiole; the twig shape,
and stipule scars are the most reliable characters to
distinguish them.

The species may hybridize with S. macrophylla and
S. amplexicaulis in restricted parts of its range.

1982] DIPTEROCARPACEAE (Ashton) 529

9. Section Mutica

BRANDIS, J. Linn. Soc. Bot. 31 (1895) 100; ASHTON, Gard. Bull. Sing. 20 (1963)
268; Man. Dipt. Brun. (1964) 117. — Shorea, Red Meranti group, S. parvifolia
subgroup, SyM. Mal. For. Rec. 16 (1943) 59, f. 36 (map). — Fig. 106, 107.

Flower buds + ovoid. Corolla as in sect. Rubella. Stamens 15, in 3 verticils;
filaments broad at base, tapering gradually to anthers; anthers with 4 pollen sacs,
broadly oblong to subglobose; appendage to connective aristate, rather short,
becoming reflexed at least on outer anthers. Ovary with distinct stylopodium,
both + densely tomentose; style shorter than ovary or very slightly longer.
Branchlets of raceme short; flowers dense. Stipules, bracts and bracteoles usually
caducous, -rarely subpersistent (S. quadrinervis, S. acuminata). Leaf with
scalariform tertiary nerves; midrib + evident above. Bark surface usually
becoming V-section fissured. Wood as in sect. Brachypterae.

Distr. Peninsular Thailand and Malesia: Malaya, Sumatra, Banka, Borneo.
Ecol. Scattered in lowland evergreen forest below 1600 m.
Vern. Red méranti, méranti mérah (Mal., Sum.), pérawan, lop (Iban).

9a. Subsection Auriculatae

ASHTON, Gard. Bull. Sing. 22 (1967) 300. — Fig. 107.
Fruit calyx lobes auriculate at base. Bark surface remaining smooth or, after
passing through an ephemeral shallowly V-section fissured stage, becoming flaky.

Distr. As section.

Note. This subsection consists of two very distinct species, S. slootenii and S. myrionerva, and those others
which cluster round the geographically very variable S. macroptera. S. macroptera is known in Malaya to be
thrip pollinated, and variably apomictic through adventive polyembryony (CHAN H. T.), all but S. macroptera
are endemic to Borneo: S. macroptera itself manifests great morphological uniformity except in that island

where it is highly variable.

133. Shorea slootenii Woop ex ASHTON, Gard. Bull.
Sing. 19 (1962) 312, pl. 29; Man. Dipt. Brun. (1964)
222, f. 16; ibid. Suppl. (1968) 119; MEER & Woop,
Sabah For. Rec. 5 (1964) 147. — Fig. 106 A—A3.

Twig, panicle, leaf bud, stipule outside (pubescent
within), petiole and leaf beneath densely persistently
scabrid pale fulvous tufted tomentose, the tufts short,
hard and knob-like; midrib above shortly pubescent.
Twig to 5 by 3 @ apically, stout, compressed and
ribbed at first, later terete, verrucose owing to the
persistent bases of the hair-tufts; stipule scars to 3 mm
long, 0.5 mm thick at base at first, long cuneate,
slightly ascending. Bud 6-10 by 5-8 mm, broadly
ovoid, slightly compressed, obtuse. Stipule to 25 by 8
mm, oblong, obtuse. Leaves 11—22 by 4—7 cm, broadly
oblong, coriaceous, deeply cupped; base obtuse;
acumen to 1.5 cm long; nerves (25—)27—34 pairs,
dense, depressed above, prominent beneath, arched at
margin at 50°-60°; tertiary nerves scalariform, promi-
nent beneath, slightly diagonal to nerves or at 90°;
midrib prominently terete beneath, depressed above;
petiole \.7—2.3 cm long, stout. Panicle to 18 cm long,

terminal or axillary, stout, brittle, + compressed;
singly or doubly regularly branched, branchlets short,
bearing to 4 flowers; bracteoles to 3 by 2 mm, ovate to
deltoid, obtuse, densely yellowish brown pubescent
outside, glabrous within, caducous. Flower bud to 12
by 7 mm, ovoid to ellipsoid, obtuse. Calyx shortly
densely tawny pubescent outside, glabrous within; 3
outer lobes ovate acuminate, longer than the 2 acute
inner lobes. Petals pale yellow, lanceolate, subacute,
densely shortly tawny pubescent on parts exposed in
bud. Stamens 15, the 5 inner reaching the base of the
style, about twice as long as the other 10, the latter of
two different heights; filaments tapering gradually,
slightly twisted in bud; anthers globose; appendage to
connective shorter than anther, reflexed on 5 inner
stamens; twice as long as anthers, hardly reflexed, on
10 outer stamens. Ovary and stylopodium narrowly
ovoid to lanceolate, sparsely setose in distal half,
surmounted by a broadly filiform style almost as long
as the ovary, glabrous. Fruit calyx lustrous, puberu-
lent, more densely so towards the base; 3 longer lobes
to 17 by 1.8 cm, oblong, obtuse, hardly tapering
towards base; base with an ovoid to 2 by 1.2 cm

530

U.S

FLORA MALESIANA

[ser. I, vol. 9?

Fig. 108. Shorea myrionerva SYM. ex ASHTON. a. Habit, b. fruit, c. seed, all x 4 (aS 31986, with terminal bud
from bb. 14614, b—c S 15591).

saccate thickened central area adpressed to nut, the
lobes continuing laterally as to 1 cm broad auricles,
tapering abruptly at pedicel; 2 shorter lobes 5.5 by 0.7
cm, unequal, linear, similarly expanded and auricu-
late at base. Nut to 3.5 by 1.3 cm, ovoid, densely
shortly pale brown to buff pubescent; style remnant to
4 mm long, tapering, acute.

Distr. Malesia: N.W. Borneo (Lower Kapuas,
Sarawak to S.W. Sabah).

Ecol. Local, sandy clay soils on hills near the
present or Pleistocene coastline, to 400 m.

Vern. Méranti képong, m.k. kasar (Sar., Brun.),
kawang raung (Dusun).

134. Shorea myrionerva Sym. ex ASHTON, Gard. Bull.
Sing. 19 (1962) 299, pl. 23; Man. Dipt. Brun. (1964)
201, f. 16; ibid. Suppl. (1968) 111; MevER & Woop,
Sabah For. Rec. 5 (1964) 122. — Shorea sp. BROWNE,
For. Trees Sarawak & Brunei (1955) 145. — Fig.
107-109.

Medium-sized, occasionally large, tree. Twig and
petiole persistently dotted with scattered pale fulvous-

brown long hair tufts; nervation beneath and stipule
more shortly so; fully expanded leaf glabrescent. Twig
2.5-3.5mm g apically, stout, straight, little branched,
terete; stipule scars to 2 mm long, to | mm wide, pale,
cuneate. Bud to 10 by 7 mm, a loose group of
bud-scales, compressed. Stipule to 17 by 7 mm,
broadly hastate, subauriculate, subacute, subpersis-
tent (more so in young trees and saplings). Leaves
12-22 by 4.5-9 cm, narrowly oblong, chartaceous;
base obtuse; apex tapering abruptly; acumen to 8 mm
long, broad; nerves 24~28 slender close pairs, promi-
nent beneath, curved towards the margin; tertiary
nerves slender, scalariform, at c. 90°; midrib depressed
above; petiole 1.2-2 cm long, short, stout. Panicle to 8
cm long, axillary, rarely terminal, becoming rami-
florous, terete, ribbed on drying, lax, glabrous but for
sparse fulvous tufts towards base; singly branched,
the branches short, bearing to 5 distichous flowers;
bracts to 8 by 1.5 mm, narrowly lanceolate, acute,
puberulent outside, glabrous within; bracteoles to 5 by
3 mm, ovate, obtuse. Flower bud to 5 by 3 mm, elliptic,
obtuse. Calyx shortly densely pubescent outside,

1982]

glabrous within; lobes subequal, ovate, the outer 3
rather narrower and subacuminate. Petals dark red
with a pale margin, narrowly ovate, obtuse, densely
yellowish gold pubescent on parts exposed in bud.
Stamens 15, of 3 lengths, the inner 5 much longer than
the others and almost reaching the style apex; fila-
ments thick, tapering gradually; anthers broadly
oblong, slightly tapering; appendages to connective
short, becoming reflexed. Ovary and stylopodium
narrowly conical to pyriform, shortly pubescent; style
somewhat shorter than ovary and stylopodium,
broadly filiform, glabrous. Fruit calyx glabrous; 3
longer lobes to 17 by 2.5 cm, oblong, obtuse, thinly
chartaceous, not tapering basally; base thickened and
saccate at the centre, with 2 thin, to 1.5 cm broad,
lateral auricles; 2 shorter lobes subequal, to 8 by 0.7
cm, linear, saccate but not auriculate. Nut to 2.5 by 1.3
cm, ellipsoid, pruinose, glabrous; style remnant c. 1.5
mm long, acute.

Distr. Malesia: Borneo (Rejang hinterland to S.W.
Sabah, E. Kutei).

Ecol. On moist clay hillsides, alluvium and river-
banks.

Vern. Abang gunong (Kutei), kawang tikus
(Brunei), séraya urat, banyak (Sabah), pitun (Dus.),
abang (Murut), langgai, sépit undang, éngkabang
(Iban).

135. Shorea sagittata ASHTON, Gard. Bull. Sing. 22
(1967) 299, pl. 45, 352 (phot. habit); Man. Dipt. Brun.
Suppl. (1968) 118, f. 15, pl. 23 (stem-base).

Large tree. Leaf above evenly caducous cream-buff
pubescent, other vegetative parts densely shortly
persistently pink-brown scabrid tomentose. Twig c. 2
mm @g at first, terete, ribbed, becoming red-brown
verruculose: stipule scars short, ascending. Bud to 7 by
7 mm, broadly ovoid. Stipule to 10 by 8 mm, ovate,
subacute, saccate, caducous. Leaves 7-15 by 3-6 cm,
narrowly oblong to lanceolate; base obtuse; acumen
to 1.5 cm long, slender; nerves 19-25 pairs, slender but
prominent beneath, at 70°—80° near the base, 50° near
the apex; tertiary nerves slender, densely scalariform;
midrib depressed above, prominent beneath; petiole
9-17 mm long. Panicle imperfectly known, at least 6
cm long, singly branched, densely persistently pink-
brown scabrid tomentose. Fruit calyx sparsely puber-
ulent; 3 longer lobes to 11 by 2.5 cm, lorate-spatulate,
obtuse, tapering only slightly above the prominently
sagittate-auriculate base, base in all to 18 mm broad,
with to 6 by 4 mm elliptic thickened saccate central
disc; 2 shorter lobes to 2 cm long, c. 3 mm wide, linear,
acute, not auriculate at base. Nut 15 by 8 mm,
ellipsoid, shortly apiculate, puberulent towards the
apex.

Distr. Malesia: N.W. Borneo (Central and W.
Sarawak, Ulu Kapuas in W. Borneo; ?Tidung, ?Pur-
aktjau, sterile collections).

Ecol. Locally frequent on sandy clay soils on low
hills and ridges to 1000 m, in Mixed Dipterocarp
forest.

Vern. Méranti luang (Sarawak).

DIPTEROCARPACEAE (Ashton)

531

—

Ze

/
vA

Le Z 3

LE
S : . \

=

Fig. 109. Shorea myrionerva SYM. ex ASHTON. Part of
twig and leaf of 3 m high sapling, x +(BRUN 5200).

532

FLORA MALESIANA

[ser. I, vol. 9?

a —__ EE

136. Shorea macroptera Dyer, FI. Br. Ind. 1 (1874)
308: BRANDIS, J. Linn. Soc. Bot. 31 (1895) 90; BRANDIS
&Gwe in’ Bs Pe. Pi-peam. ed: 1 3576 (11895)
267. — Fig. 106 D-D3.

Note. Beside the subspecies described at least two
forms (one in Melawi, one in S. Central Kalimantan)
exist in Indonesian Borneo, for which more complete
material is required. See also under the next species, S.
ferruginea.

KEY TO THE SUBSPECIES

1. Panicles singly branched or doubly branched at
base.
2. Twigs, petioles and panicles densely evenly
shortly buff pubescent a. ssp. Macroptera
2. Twigs, petioles and panicles sparsely tufted
tomentose or glabrescent b. ssp. baillonii
1. Panicles doubly branched; tomentum dense, even,
persistent.
3. Leaves (9-)18-23 by (4-)6.5-9.5 cm; nerves
13=I5-pairs) = 2 4. c. ssp. sandakanensis
3. Leaves 8-16 by 4-6 cm; nerves 10-14 pairs
d. ssp. macropterifolia

a. ssp. macroptera. — KING, J. R. As. Soc. Beng. Sc.
62, 2 (1893) 113; Rpt. Agr. Bull. Str. & F.M.S. 1
(1901) 57; Fl. Mal. Pen. 1 (1922) 225; Burk. J. Str. Br.
R. As. Soc. 76 (1917) 164, fig.; ibid. 81 (1920) 51, 70,
75, fig.; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 305; ibid. ed. 2
(1927) 1123; Foxw. Mal. For. Rec. 3 (1927) 30; ibid. 8
(1930) 21; ibid. 10 (1932) 195; Epwarps, Mal. For.
Rec. 9 (1932) 145; Burk. Dict. (1935) 2016; Sym. Mal.
For. Rec. 16 (1943) 78, f. 38, 48; ASHTON, Gard. Bull.
Sing. 20 (1963) 276. — S. auriculata ScorRT. ex Foxw.
Mal. For. Rec. 10 (1932) 195, nom. in syn.

Large tree. Young twig, panicle, leaf bud, stipule
outside (puberulent within), petiole, midrib above at
base and nervation beneath densely evenly shortly
pale brown pubescent. Twig to 2 mm @ apically,
frequently slightly compressed, becoming terete, gla-
brous; stipule scars short, narrowly cuneate, +
horizontal. Bud 4-6 by 2.5-4 mm, ovoid, subacute.
Stipules to 8 by 3 mm, oblong, subacute, fugaceous.
Leaves 10-15 by 3-5 cm, elliptic or oblong, coria-
ceous; base broadly cuneate; acumen to | cm long;
nerves 12-15(—18) pairs, prominent beneath, curved,
at 40°-50°; tertiary nerves slender, densely scalari-
form, sinuate, diagonal to nerves; midrib prominent
and terete beneath, slightly depressed above; petiole
1.5 cm long. Panicle to 10 cm long, terminal or
axillary, slightly compressed or terete, lax; singly
branched or doubly branched near base, the branch-
lets bearing to 7 distichous flowers; bracteoles to 3.5
by 2.5 mm, elliptic, obtuse, shortly yellowish brown
pubescent outside, glabrous within, fugaceous. Flower
bud to 5 by 3mm, ovoid. Calyx outside densely shortly
pale yellowish brown pubescent, glabrescent within;
lobes subequal, ovate, the 3 outer acuminate, the 2
inner acute. Petals cream to pink at base, linear, acute,
twisted, yellow pubescent on parts exposed in bud.

Stamens 15, of 3 lengths, the longest reaching just
above the ovary; filaments tapering gradually; anthers
broadly oblong; appendage to connective short,
becoming reflexed. Ovary and stylopodium + conical,
densely pale grey pubescent, glabrous near base; style
c. 1/2 length of ovary, slender, glabrous. Fruit calyx
shortly sparsely puberulent, glabrescent; 3 longer
lobes to 12 by 2.3 cm, spatulate, obtuse, to 1 cm broad
above base; base with to 7 by 5 mm ovoid saccate
thickened centre and 2 lateral narrow auricles, the
whole to 1.2 cm broad, abruptly tapering at pedicel; 2
shorter lobes to 6 by 0.6 cm, narrowly oblong, similar
at base. Nut to 1.8 by 1.2 cm, ovoid, densely evenly
shortly pale buff pubescent; style remnant 2 mm long,
acute.

Distr. Peninsular Thailand, and Malesia: Malaya
(excepting seasonal areas), Singapore, Singkep,
Lingga, E. Sumatra (Djambi, Indragiri, Kuantan
Distr., Langkat).

Ecol. Well drained Mixed Dipterocarp forest,
especially in hills, to 900 m, common.

Vern. Mélanta, méranti mélantai, m. bélantai, m.
kunyit, m. kétapah, képong labu, k. ségar, témak
(Malaya), /ukup (Riouw), méranti undang, m. sabut,
m. piring, m. télor, m. mangu, m. tampalu, m. kunyit
(Sumatra).

b. ssp. baillonii (HEIM) ASHTON, Gard. Bull. Sing. 20
(1963) 277, f. 16, pl. 51 (bark); Man. Dipt. Brun.
Suppl. (1968) 111; BRowNE, For. Trees Sarawak &
Brunei (1955) 137. — S. bailloni Heim, Bull. Mens.
Soc. Linn. Paris 2 (1891) 973.

Defining characters: Small tree. Twigs, petioles and
panicle sparsely tufted pubescent or glabrescent,
drying blackish. Leaves 12-19 by 3.5—7 cm, narrowly
elliptic; base narrowly cuneate; apex tapering; nerves
11—14(-15) pairs, prominent but terete beneath. Pani-
cles c. 13 cm long, singly branched or doubly so at
base; branchlets to 2.5 cm long. 3 longer fruit calyx
lobes to 13 cm long.

Distr. Malesia: W. Borneo (Sampit,
Kapuas, Sarawak west of Bintulu).

Ecol. Clay soils on low hills to 600 m, common.

Vern. Méranti mélantai (Sar.), sépit undang
(Iban).

Lower

c. ssp. sandakanensis (SyM.) ASHTON, Gard. Bull. Sing.
20 (1963) 277. — S. macroptera SLooT. in Merr. PI.
Elm. Born. (1929) 203; MEWER & Woop, Sabah For.
Rec. 5 (1964) 116, f. 14, p.p. — S. sandakanensis SYM.
Gard. Bull. S. S. 9 (1938) 343, pl. 25.

Defining characters: Large tree. Leaves (9—)18—23
by (4-)6.5-9.5 cm, narrowly oblong; base obtuse;
apex abruptly attenuate; nerves 13-15 pairs, promi-
nently acute beneath; petiole to 2 cm long. Panicles to
16cm long, doubly branched, branchlets to 8 cm long.
3 longer fruit calyx lobes to 14 cm long.

Distr. Malesia: E. Borneo (Kudat to Balikpapan:
wrongly recorded from Sarawak by BROWNE, For.
Trees Sarawak & Brunei p. 143).

Ecol. As ssp. baillonii.

1982]

Vern. Séraya mélantai (Sabah), tégérangan sibu, t.
batu (Nunukan).

d. ssp. macropterifolia ASHTON, Gard. Bull. Sing. 20
(1963) 277; Man. Dipt. Brun. (1964) 197; ibid. Suppl.
(1968) 111; MeveR & Woop, Sabah For. Rec. 5 (1964)
116.

Defining characters: Large tree. Leaves 8-16 by 4-6
cm, narrowly ovate, coriaceous; base obtuse; apex
tapering; nerves 10-14 pairs, slender but sharply
prominent beneath; petiole to 1.5 cm long. Panicles to
16 cm long, twice branched; branchlets to 8 cm long. 3
longer fruit calyx lobes to 14 cm long.

Distr. Malesia: northern Borneo (Rejang hinter-
land to W. Sabah).

Ecol. Clay soils on undulating land and hillsides to
600 m, local.

Vern. Mérantimélantai(Brun.), sépit undang (Iban).

137. Shorea ferruginea DyeR ex BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 91; Becc. For. Born. (1902) 570;
Merr. En. Born. (1921) 405; BROowNE, For. Trees
Sarawak & Brunei (1955) 139; AsHToN, Gard. Bull.
Sing. 20 (1963) 281; Man. Dipt. Brun. (1964) 187, f.
19; ibid. Suppl. (1968) 108; MEevER & Woop, Sabah
For. Rec. 5 (1964) 106. — S. discolor Heim, Rech.
Dipt. (1892) 67, nomen.

Large tree. Young twig, panicle, leaf bud, stipule
outside (glabrescent or shortly pubescent within),
petiole beneath and leaf above sparsely persistently
pale brown scabrid puberulent. Twig 2-2.5 mm g
apically, terete, ribbed below the stipule scars, becom-
ing smooth; stipule scar c. 2mm long, c. 0.5 mm thick,
pale, cuneate, slightly falcate, horizontal or slightly
ascending. Leaf bud 5-8 by 4-6 mm, ovoid, subacute.
Stipule to 15 by 5mm, hastate, acute. Leaves 12-24 by
5.5—11 cm, broadly oblong to ovate, thinly coriaceous;
base obtuse; acumen to 1.2 cm long; nerves 15-19
pairs, prominent, at 60°—70°, curved apically; short
secondary nerves frequently present; tertiary nerves
elevated, remote, at 90° to nerves; midrib prominent
and terete beneath, depressed above; petiole 1—-1.5 cm
long, short. Panicle to 22 cm long, terminal or axillary,
terete or ribbed, lax, regularly alternately singly or
doubly branched, the branchlets bearing to 7 secund
flowers; bracts and bracteoles to 7 by 3 mm, ovate,
acute. Calyx shortly densely pale grey-brown pubes-
cent, glabrous within; 3 outer lobes longer, + deltoid
to ovate, subacute; 2 inner lobes shorter, narrowly
deltoid to ovate, acute, acuminate. Petals narrowly
oblong, densely pubescent on parts exposed in bud.
Stamens 15, the 5 inner an anther’s length longer than
the others; filaments basally expanded, abruptly
tapering and filiform distally; anthers oblong; appen-
dage to connective reaching to style apex on 5 inner
stamens, more than twice as long as anthers, curved
but not reflexed. Ovary and stylopodium pyriform,
densely pubescent in the distal half, shortly densely
pubescent basally, style as long as ovary and stylopo-
dium, filiform, glabrous. Fruit subsessile. Calyx gla-
brescent; 3 longer lobes to 5 by 0.8 cm, spatulate,

DIPTEROCARPACEAE (Ashton)

353

Fig. 110. Shorea acuta ASHTON. a. Tip of sterile twig,
b. fruit, c. nut, all x 4 (a SAN 17474, b-c BRUN
3291).

subequal, to 3.5 mm broad above the to 1.5 by 1.2 cm
ovate glabrous shallowly saccate thickened base; 2
shorter lobes to 23 by 1.5 mm, linear, similar at base.
Nut to 2.7 by 1.0 cm, ovoid, shortly buff pubescent,
apiculate, shorter than longer calyx lobes.

Distr. Malesia: Borneo.

Ecol. Widespread on skeletal soils along ridges to
1100 m.

Vern. Seraya melantai kechil (Sabah), meranti
menalit (Sar.), tehan betung, t. paru, t. parei, t. lutup,
lampong tahan (S.E. Borneo).

Note. Collections from Melawi which I ascribe to
S. macroptera on account of their prominent ner-
vation possess the indumentum of this species and
underline the close relationship between the two. I
maintain this species separately from S. macroptera
because, throughout its range in East Malaysia I never
saw morphologically intermediate individuals though
the two species commonly grow in mixture.

138. Shorea acuta ASHTON, Gard. Bull. Sing. 19
(1962) 268, pl. 8; Man. Dipt. Brun. (1964) 174, f. 16;
ibid. Suppl. (1968) 104. — Fig. 110.

534

Large tree. Young twig, panicle, leaf bud, stipule
(both surfaces), petiole, nervation beneath and midrib
above evenly persistently shortly pale brown puberu-
lent, glabrescent on nervation and midrib. Twig to 4
mm @, slightly compressed at first, becoming terete,
glabrous, smooth; stipule scars initially to 3 mm long,
to | mm thick, cuneate, horizontal. Bud 6-10 by 4-6
mm, ovoid, subacute. Stipule to 12 by 5 mm, oblong,
obtuse. Leaves 14-26 by 6.5—-12 cm, elliptic, thickly
coriaceous; base obtuse, sometimes broadly cuneate;
acumen to 2 cm long, + narrowly deltoid; nerves
10-13 pairs, prominent beneath, curved, at 45°—55°
along the midrib, to 90° at the base; tertiary nerves
slender, sinuate, densely scalariform, diagonal to
nerves; midrib prominent and terete beneath, nar-
rowly slightly depressed above; petiole 1.5—2 cm long,
stout. Panicle to 8 cm long, terminal or axillary,
compressed; singly or doubly branched at 2 cm
intervals, branchlets to 6 cm long, bearing to 9
distichous flowers; bracteoles to 3.5 by 2.5 mm,
elliptic, obtuse, shortly yellowish brown puberulent
outside, glabrous within, caducous. Flower bud to 9 by
4 mm, narrowly ellipsoid, obtuse. Calyx shortly
densely yellowish buff pubescent outside, glabrous

FLORA MALESIANA

[ser. I, vole@2

within; lobes subequal, ovate, subacute. Corolla dark
crimson, pubescent on parts exposed in bud; petals
linear, subacute. Stamens 15, of 3 lengths, the longest
reaching just above the ovary, filaments tapering
gradually; anthers subglobose; appendage to connec-
tive short, becoming reflexed. Ovary and stylopodium
narrowly conical to ovoid, long tomentose towards
the apex, shortly so at base; style c. 5 length of ovary
and stylopodium filiform, glabrous. Fruit calyx
shortly sparsely glabrescent; 3 longer lobes to 15 by
2.8 cm, spatulate, obtuse, to 1.5 cm broad above the
base; base to 18 mm broad, with to 8 by 7 mm ovate
saccate thickened centre and 2 lateral auricles,
abruptly tapering to the pedicel; 2 shorter lobes to 8 by
0.7 cm, linear, unequal, similar at base. Nut to 3 by 2
cm, ovoid, shortly evenly densely pale fulvous to buff
pubescent; style remnant short, acute.

Distr. Malesia: Borneo (N.E. Sarawak, Brunei).

Ecol. Locally common below 400 m, on deep sandy
soils in Mixed Dipterocarp forest.

Vern. Mélantai, kawang tikus, éngkabang tikus,
meéranti kawang tikus (Brun.), mérabubok, langgai
(Iban).

9b. Subsection Mutica

ASHTON, Gard. Bull. Sing. 22 (1967) 301 (‘Muticae’). — Shorea subg. Rubro-
shorea Meer, Act. Bot. Neerl. 12 (1963) 322. — Fig. 106.

Fruit calyx lobes not auriculate. Bark surface early becoming deeply and
persistently V-section fissured, only rotting off in very large trees.

Distr. As section.

Note. Species of this subsection are, with the exception of S. scabrida and S. retusa easy to identify. Many are
widespread and morphologically remarkably uniform. S. parvifolia, which is usually variable for the subsection,
and consists of at least two ecotypically and partially geographically separate forms is linked with several sibling
species endemic in Borneo (S. rubra, S. foraminifera, S. scabrida, S. retusa, S. revoluta); itis known to be partially
apomictic through adventive polyembryony (MAuRy). Hybridization is known among some species but is
exceptionally rare. All species are thrip pollinated, and sympatric species flower sequentially (CHAN, H. T.).

139. Shorea quadrinervis SLooT. Bull. Bot. Gard.
Btzg III, 17 (1942) 220, f. 21; BROwNg, For. Trees
Sarawak & Brunei (1955) 142; AsHTon, Man. Dipt.
Brun. (1964) 213, f. 16, pl. 56 (habit); ibid. Suppl.
(1968) 116; MEER & Woop, Sabah For. Rec. 5 (1964)
140. — Fig. 111.

Medium-sized, occasionally large, tree. Young
parts shortly densely tawny pubescent, largely cadu-
cous on stipule, leaf bud and leaf beneath; leaf above
glabrous but for midrib. Twig c. 3 mm @ at apex,
terete, stout, strongly ridged below stipule scars,
becoming minutely cracked longitudinally; stipule
scars c. 3 mm long, c. 1 mm broad, pale, falcate or
amplexicaul-descending. Bud to 2 by 1.3 cm, loosely
enclosed by stipular bud scales, compressed, subcor-
date. Stipule to 2.5 by 1.3 cm, ovate to subcordate,
subpersistent, glabrescent, persistently ridged along
the 5 nerves; stipule pairs sometimes united at base.

Leaves 10-18 by 5-8 cm, broadly ovate to elliptic,
strongly cupped; margin usually slightly revolute;
base obtuse; acumen to 1.3 cm long; nerves c. 4 pairs,
strongly curved, at c. 40°—70° with | or more second-
ary nerves of varying length; tertiary nerves slender,
scalariform, at c. 90° to nerves; midrib depressed
above; petiole 8-10 mm long. Panicle to 28 cm long,
terminal or axillary, large, spreading, lax, terete or
angular, densely persistently pale rust pubescent;
doubly, sometimes trebly branched, branchlets 1-8
cm long, straight, bearing to 6 secund flowers; bracts
as the stipules, + caducous; bracteoles to 2 by 1 mm,
ovate to elliptic, obtuse, densely grey pubescent
outside, puberulent within. Flower bud to 4.5 by 5.5
mm, broadly elliptic to subglobose, obtuse. Calyx
shortly pale grey pubescent outside, glabrous within;
lobes + patent, subequal, ovate, the inner 2 slightly
more acute. Petals pink, paler at margins, elliptic,

1982]

Fig. 111. Habit of Shorea quadrinervis SLOOT. with
drooping branches. Brunei (Photogr. ASHTON, 1959).

obtuse, densely pubescent on parts exposed in bud.
Stamens 15, the inner 5 longest, the outer 10 some-
times vestigial; filaments tapering gradually; anther
subglobose; appendage to connective shorter than
anther, becoming reflexed. Ovary and stylopodium
conical, densely pubescent; style short, frequently
expanded at the stigma. Fruit calyx shortly sparsely
pubescent: 3 longer lobes to 8 by 1.3 cm, narrowly
spatulate, obtuse, 2-3 mm broad above the c. 5 by 4
mm thickened base; 2 shorter lobes to 5.5 by 0.7 cm,
unequal, tapering to c. 2 mm broad, base as in longer
lobes. Nut to 1.5 by 0.7 cm, narrowly ovoid, shortly
buff pubescent; style remnant c. 1.5 mm long, slender,
acute.

Distr. Malesia: N.W. Borneo (Ulu Kapuas, Sar-
awak to W. Sabah).

Ecol. Common on sandy clay soil on low hills,
occasionally to 700 m.

Vern. Téngkolong, témpilong, méranti tempilong (W.
Borneo), méranti sudu (Sarawak), séraya sudu (Sabah).

140. Shorea acuminata Dyer, FI. Br. Ind. | (1874)
305; KinG, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 113;

DIPTEROCARPACEAE (Ashton)

535

BRANDIS, J. Linn. Soc. Bot. 31 (1895) 91; RipL. Agr.
Bull. Str. & F.M.S. 1 (1901) 58; Fl. Mal. Pen. 1 (1922)
225; HEYNE, Nutt. Pl. ed. 1, 3 (1917) 296; ibid. ed. 2
(1927) 1115; Foxw. Mal. For. Rec. 3 (1927) 32; ibid. 8
(1930) 23; ibid. 10 (1932) 197; EDwarps, Mal. For.
Rec. 9 (1931) 139; Burk. Dict. (1935) 2006; Sym. Mal.
For. Rec. 16 (1943) 65, f. 38, 39; SLoor. Bull. Bot.
Gard. Btzg III, 18 (1949) 251, f. 11.

Large buttressed tree, to 50 m. Young parts densely
buff pubescent, persistent on twig apices, midrib
above, petioles, panicles, parts of perianth exposed in
bud, ovary and calyx, caducous elsewhere; becoming
sparse on fruit calyx and expanding stipules. Twigs c.
2mm @g apically, terete, smooth, pale brown; stipule
scars prominent, horizontal. Leaf buds small, conical,
acute, generally hidden in the conspicuous, to 30 by 8
mm, oblong cordate obtuse lustrous concave subper-
sistent stipules. Leaves 6-12 by 2.6—-6 cm, ovate-fal-
cate, coriaceous, lustrous beneath: base subequal,
obtuse to broadly cuneate; margin subrevolute:
acumen to 1.8 cm long, slender: nerves 7-11 pairs,
arched, ascending, slender but + prominent beneath,
frequently spaced irregularly along the midrib and
with + prominent short secondary nerves; tertiary
nerves scalariform, obscure; midrib slender but
prominent, terete, beneath, evident above: petiole 7-9
mm long, short. Panicles to 12 cm long, terminal or
axillary, singly (if axillary) or doubly branched:
branchlets to 2 cm long, bearing to 6 congested secund
flowers: bracts as stipules, frequently subpersistent in
terminal inflorescences. Flower bud to 4 by 2 mm,
lanceolate; sepals broadly ovate, the 3 outer somewhat
larger, acute, the 2 inner acuminate: petals dark wine
red, paler towards base: stamens 15, in 3 unequal
verticils: filaments lorate, tapering: anthers small,
subglobose: appendages shorter than anthers, becom-
ing reflexed on inner 2 verticils; ovary ovoid, tapering
into a somewhat longer columnar tapering style. Fruit
pedicel to 2 mm long, slender; 3 longer calyx lobes to 9
by 1.2 cm, narrowly spatulate, obtuse, c. 2 mm broad
above the to 6 by 4 mm ovate thickened saccate base:
2 shorter lobes to 25 by 2 mm, linear, similar at
base. Nut to 1.5 by 6 mm, lanceolate, prominently
apiculate.

Distr. Malesia: Malaya (from Lower Perak and
Trengganu R. southwards), Sumatra (Langsa and P.
Musala south to northern Palembang), Lingga.

Ecol. Common in Mixed Dipterocarp forest, on
undulating land and hills to 400 m.

Vern. Méranti rambai daun, m. hijau (Malaya),
damar téték, m. samarupa, samarupa (Achin), latuko,
méranti bunga, m. gombang (Tapanuli), méranti sar-
ang burung, m. sarang punai, m. béros, m. képala,
sésawoh (E. coast Sumatra), madang kuaun (W. coast),
méranti djuntah, m. képala tupai, m. kérang, bénio, m.
hitam batang, m. samak bénio (Riouw). méranti baw-
ang (Djambi), merakunyit bétina, mélas uwai (Palem-
bang).

Note. Two collections from S.E. Sumatra appear
to represent hybrids between this species and S.
parvifolia.

536

FLORA MALESIANA

[ser. I, voleg2

141. Shorea macrantha BRANDIS, J. Linn. Soc. Bot. 31
(1895) 97; Mere. En. Born. (1921) 405; Sym. Gard.
Bull. S.S. 7 (1933) 131, pl. 34; Mal. For. Rec. 16(1943)
77, f. 38; BROWNE, For. Trees Sarawak & Brunei
(1955) 148; ANDERSON, Gard. Bull. Sing. 20 (1963)
158; ASHTON, Man. Dipt. Brun. Suppl. (1968) 110, f.
13. — S. hemsleyana (non KING) Foxw. Mal. For.
Rec. 10 (1932) 167, p.p. — Fig. 106 C—C2.

Small or medium-sized tree. Twig, bud, stipule
outside, petiole and nervation beneath densely persis-
tently yellow-brown scabrid tomentose; stipule inside
and midrib above densely evenly, nerves above spar-
sely evenly yellow-brown pubescent. Twig 2-4 mm @
apically, terete, becoming smooth; stipule scars short,
dark, horizontal or descending. Bud 6 by 3 mm, ovoid,
subacute. Stipule to 16 by 5mm, lanceolate, subacute,
subpersistent. Leaves 6-17 by 2.5-8 cm, of variable
size, narrowly ovate, coriaceous; base cordate, sub-
equal; acumen to 1.5 cm long, slender; nerves 13-17
pairs, prominent beneath, at 110° towards the base,
35° towards the apex; tertiary nerves scalariform,
prominent beneath, set at c. 90° to the nerves; midrib
beneath prominent; midrib and nervation frequently
somewhat depressed and the lamina bullate above;
petiole S—6 mm long, very short. Panicles to 10 cm
long, terminal or axillary, terete, persistently fulvous
to yellow-brown scabrid pubescent; singly branched,
branchlets short, the flowers congested. Flower bud to
14 by 4 mm, lanceolate. Sepals pubescent on parts
exposed in bud; outer 3 deltoid, acute; inner 2 smaller
than outer 3, narrowly ovate, with thinner margin.
Petals dark red within, paler outside (Borneo) or
white throughout (Malaya), lanceolate, densely
pubescent on parts exposed in bud. Stamens 15, in 3
unequal verticils; filaments compressed, tapering to
the small subglobose anthers; appendage to connec-
tive slender, about as long as anther, becoming
reflexed at anthesis. Ovary ovoid, shortly pubescent;
style filiform, glabrous, about twice as long as ovary.
Fruit subsessile. Calyx sparsely yellow-brown pubes-
cent; lobes to 2.5 by 2 cm, subequal, ovate, acute,
saccate towards the base. Nut to 5.5 by 2.5 cm,
ellipsoid, acute, pale yellow-brown pubescent.

Distr. Malesia: Malaya (Perak, E. coast in Pahang
and Johore), E. Sumatra (Indragiri), N.W. Borneo
(Central and W. Sarawak).

Ecol. Local, in Mixed Peat Swamp forest at their
inland margins, and on white sand terraces (Bor-
neo).

Vern. Kepong, k. hantu, chéngal pasir, méranti pasir
(Malaya), éngkabang bungkus (Sar.), pérawan lam-
pong kijang (Iban).

Note. In Perak the tree has an even, soft tomentum
and applanate leaves with the nervation not depressed
above.

142. Shorea hemsleyana (KING) KING ex Foxw. Mal.
For. Rec. 10 (1932) 167, p.p.; Sym. Gard. Bull. S. S. 7
(1933) 129, pl. 33; Burk. Dict. (1935) 2011; BROWNE,
For. Trees Sarawak & Brunei (1955) 147; ASHTON,
Gard. Bull. Sing. 22 (1967) 293.

a. ssp. hemsleyana. — SyM. Mal. For. Rec. 16 (1943)
70, f. 38, 43. — Balanocarpus hemsleyanus KiNG, J.R.
As. Soc. Beng. Sc. 62, 2 (1893) 134; BRUHL & KING,
Ann. R. Bot. Gard. Calc. 5, 2 (1896) 160; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 109. — Pachychlamys hems-
leyanus RIDL. Fl. Mal. Pen. 1 (1922) 234; Foxw. Mal.
For. Rec. 3 (1927) 38.

Small or medium-sized tree. Twig, bud, stipule
outside, petiole and nervation below densely persis-
tently yellow-brown scabrid tomentose; stipule inside
and midrib above densely evenly, nerves above spar-
sely evenly, yellow-brown pubescent. Twig 2-4mm @
apically, terete, becoming smooth; stipule scars short,
pale, ascending. Bud to 6 by 3 mm, lanceolate,
subacute. Stipule to 16 by 5mm, lanceolate, subacute,
not at first caducous. Leaves 14-35 by 6-15 cm,
oblong, coriaceous; base obtuse; acumen to | cm long,
slender; nerves 14-17 pairs, straight and prominent
beneath, at 45°—S5°; tertiary nerves remote, scalari-
form; midrib prominent beneath; nervation some-
what depressed above; petiole 14-20 mm long, short.
Panicle to 25 cm long, axillary, terete, very shortly
persistently yellow-brown scabrid pubescent; singly
branched, branchlets short, the flowers congested.
Flower bud to 18 by 6 mm, lanceolate, large. Sepals
pubescent on parts exposed in bud; 3 outer ovate,
acuminate, fimbriate along the lateral margins. Petals
dark red within, paler outside, lanceolate, densely
pubescent on parts exposed in bud. Stamens 15, in 3
unequal verticils, the inner 5 exceeding the style apex;
filaments compressed, tapering to the small subglo-
bose anthers; appendage becoming reflexed at
anthesis. Ovary ovoid, pubescent; style filiform, gla-
brous, as long as ovary, surmounting a short pubes-
cent stylopodium. Fruit subsessile; pedice/ to 1 mm
long, obscure and expanding into fruit. Calyx pale
fulvous partially caducous pubescent; 3 outer lobes to
25 by 18 mm, as short as or shorter than the ripe nut,
lanceolate, acute, chartaceous, thickened and some-
what saccate at the base; 2 inner lobes to 20 by 18 mm,
ovate, acute. Nut to 7 by 3 cm, ovate, shortly
apiculate, densely evenly pale fulvous pubescent.

Distr. Peninsular Thailand, in Malesia: N.W.
Malaya (Perak), E. Sumatra (Indragiri).

Ecol. Shallow peat swamps; local.

Vern. Chengal pasir daun bésar, méranti bakau
(Mal.), m. Kunyit (Sumatra).

b. ssp. grandiflora (BRANDIS) ASHTON, Gard. Bull.
Sing. 22 (1967) 293; Man. Dipt. Brun. Suppl. (1968)
109, f. 13. — S. grandiflora BRaNois, J. Linn. Soc.
Bot. 31 (1895) 93; Merr. En. Born. (1921) 405.

Differing as follows: Leaves 10-23 by 4-11 cm;
petiole 6-12 mm; panicle to 4 cm long.

Distr. Malesia: N.W. Borneo (Lower Kapuas,
Central & West Sarawak).

Ecol. Local in Mixed Dipterocarp forest on
leached sandy soils below 400 m.

143. Shorea singkawang (M1Q.) Mia. Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 84; AsHToN, Gard. Bull. Sing. 31
(1978) 47.

1982]

DIPTEROCARPACEAE (Ashton)

Fad

a. ssp. Singkawang. — BurCK, Ann. Jard. Bot. Btzg 6

(1887) 219; BRANbis, J. Linn. Soc. Bot. 31 (1895) 87;
HEYNE, Nutt. Pl. ed. 1, 3 (1917) 307; Foxw. Mal. For.
Rec. 10 (1932) 164; BurK. Dict. (1935) 2022; Sym.
Mal. For. Rec. 16 (1943) 92, f. 38, 56; ASHTON, Gard.
Bull. Sing. 31 (1978) 47. — Hopea singkawang Mia.
Sum. | (1860) 489, 191; DC. Prod. 16, 2 (1868) 635;
Wacp. Ann. 7 (1868) 379; HEYNE, Nutt. Pl. ed. 2
(1927) 1125.—S. thiseltonii KiNG, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 122; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 77, t. 2, f. 13-14; BRANDIS & GILG in E. & P.
Pfi. Fam. ed. 1, 3, 6 (1895) 265; BRUHL & KING, Ann.
R. Bot. Gard. Calc. 5, 2 (1896) 155, t. 188; Burk. J.
Str. Br. R. As. Soc. 86 (1922) 285; HEYNE, Nutt. Pl. ed.
2 (1927) 1125. — S. forbesii KING ex BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 77, nom. in syn. — Pachychlamys
thiseltonii RipL. Fl. Mal. Pen. | (1922) 233; Foxw.
Mal. For. Rec. 3 (1927) 38. — Pachychlamys beccar-
ianus (non DYER ex BRANDIS) RIDL. Fl. Mal. Pen. |
(1922) 233.

Small, occasionally large tree with somewhat scaly
fissured bark. Young twigs, petioles, midrib and fruit
calyx caducous ocherous-buff puberulent; panicles,
parts of perianth exposed in bud and nut persistently
so. Twig 2-S mm @ apically, typically stout, terete,
pale brown, + prominently ribbed; stipule scars
short, horizontal. Leaf bud to 3 by 2 mm, ovoid,
obtuse; stipule to 12 by 6 mm, deltoid-lanceolate,
falcate, acute, fugaceous. Leaves (8—)12-24 by
(2.3-)5.5-9 cm, oblong-lanceolate, coriaceous; base
+ broadly cuneate; acumen to 1.5 cm long, tapering;
nerves 7-12 pairs, ascending, prominent beneath as
also the terete midrib; tertiary nerves densely scalari-
form, sinuate, very slender but evident beneath;
petioles (6-)12—17 mm long, + stout. Panicles to 8cm
long, axillary, erect, with to 1 cm long, short, branch-
lets bearing to 5 flowers. Flower buds to 10 by 4mm,
lanceolate; sepals narrowly ovate, the 3 outer longer,
acute, the 2 inner subacuminate; corolla pale red to
dark purple-red, variable; stamens 15, in 3 unequal
verticils; filaments lorate, tapering; anthers subglo-
bose; appendages shorter than anthers, becoming
reflexed; ovary narrowly ovoid, tapering into the
filiform glabrous somewhat shorter style. Fruit sub-
sessile; 3 longer calyx lobes to 8 by 0.8 cm, short,
lorate, obtuse, expanding into a 14 by 11 mm ovate
thickened saccate base; 2 shorter lobes to 3 cm long,
with similar base; nut to 6 by 2.5 cm, ellipsoid to ovoid
or obovoid, acute.

Distr. Peninsular Thailand, in Malesia: Malaya,
Lingga, E. Sumatra (Indragiri, Palembang, Lampong).

Ecol. Frequent in lowland Mixed Dipterocarp
forest on well drained undulating land, sometimes by
streams, below 400 m.

Vern. Méranti séngkawang, séngkawang (Malaya,
Sumatra), m. séngkawang merah, m. gajah, m. bahru,
m. sékan, siput mélantai (Malaya).

b. ssp. scabrosa ASHTON, Gard. Bull. Sing. 31 (1978)
48. — Shorea sp. C Sym. Mal. For. Rec. 16 (1943) 95.
Differing as follows: Twigs, buds, stipules, petioles,

midrib on both surfaces and nerves beneath, panicles
and calyx outside at first densely scabrid fulvous
pubescent, becoming sparse on nervation beneath,
caducous on calyx but otherwise persistent. Leaves
thickly coriaceous; base usually with obtuse or cor-
date base; apex shortly acuminate or obtuse; nerves
12-17 pairs.

Distr. Malesia: E. Malaya (coastal Pahang and
Trengganu).

Ecol. Forests on low hills, sometimes on soils with
impeded drainage, near the coast.

Vern. Téngkawang lampong.

144. Shorea retusa MEveER, Act. Bot. Neerl. 12 (1963)
340, pl. 9; Sabah For. Rec. 5 (1964) 141; AsHTON,
Man. Dipt. Brun. Suppl. (1968) 117, f. 14.

Small or medium-sized tree. Young parts greyish
sericeous, glabrescent except on buds and stipules
outside. Twig c. 1 mm @ apically, terete, much
branched, smooth; stipule scars pale. short, horizon-
tal. Bud to 3 by 2 mm, ellipsoid, obtuse. Stipule to 7 by
3 mm, elliptic, obtuse, caducous. Leaves 3—9 by 1.5-5
cm, small, elliptic, coriaceous, lustrous; base cuneate;
apex retuse; nerves 7—10 pairs, slender, hardly raised
beneath, at 40°-60°; tertiary nerves slender, obscure,
densely scalariform; midrib prominent beneath,
depressed above: petiole 6-9 mm long, short. Panicle
to 10 cm long, terminal or axillary, terete, caducous
buff sericeous; singly, or doubly if terminal, branched,
branchlets bearing to 7 + distichous flowers; brac-
teoles to 3 by 2mm, elliptic, obtuse, pubescent. Flower
buds to 8 by 3 mm, lanceolate. Sepals densely
pubescent on parts exposed in bud; outer 3 ovate,
acute; inner 2 shorter, narrower, thinner towards
margin than outer 3, fimbriate distally, flanged at
base. Petals pale yellow, lanceolate, fimbriate, pubes-
cent on parts exposed in bud. Stamens 15, in 3 unequal
verticils; filaments compressed, slender, tapering;
anthers subglobose, the inner 5 much longer than the
outer 5; appendage to connective slender, c. 4 x
length of anther on inner stamens, c. 2 x on medium
stamens, about same length on outer stamens, becom-
ing reflexed at anthesis. Ovary and stylopodium nar-
rowly conical, shortly densely buff sericeous, crowned
by a short glabrous style. Fruit pedicel to 1 mm long.
Calyx bud sericeous, caducous except at base; 3 longer
calyx lobes to 10 by 1.8 cm, spatulate, obtuse, c. 4mm
broad above the to 7 by 6 mm ovate saccate thickened
base; 3 shorter lobes to 5.5 by 0.4 cm, lorate, obtuse,
similar at base. Nut to 13 by 2 mm, ovoid; stylopo-
dium to 3 mm long.

Distr. Malesia: Borneo (S.E. and S. Borneo,
Tawau Distr., W. Sarawak).

Ecol. Local on skeletal podsols in Heath forest on
coastal hills.

Vern. Méranti daun tumpul (Sarawak), séraya daun
tumpul (Sabah), /ampong, lanan putéh, awang, a.
péringet, damar lantang (S.E. Borneo).

Note. See note under 154. S. scabrida.

145. Shorea lepidota (KORTH.) BL. Mus. Bot. Lugd.-

538

Bat. 2 (1852) 32; WALP. Ann. 4 (1857) 338; Mia. FI.
Ind. Bat. 1 (1859) 503; Sum. (1860) 191; DC. Prod. 16,
2 (1868) 629; BuRcK, Ann. Jard. Bot. Btzg 6 (1887)
217: BRANDIS, J. Linn. Soc. Bot. 31 (1895) 96; HEYNE,
Nutt. Pl. ed. 1, 3 (1917) 304; ibid. ed. 2 (1927) 1113,
1114, 1121; Foxw. Mal. For. Rec. 10 (1932) 166; SYM.
Gard. Bull. S. S. 7 (1933) 135, pl. 38; Mal. For. Rec. 16
(1943) 73, f. 38, 45; BurK. Dict. (1935) 2014. — Vatica
lepidota KortH. Kruidk. (1841) 73; WALP. Rep. 5
(1845) 127; DC. Prod. 16, 2 (1868) 629. — Vatica
stipulosa MiqQ. Sum. (1860) 486; HEYNE, Nutt. Pl. ed. 2
(1927) 1121. — S. nitens Mig. Sum. (1860) 488, 191;
DC. Prod. 16, 2 (1868) 632; WALP. Ann. 7 (1868) 379;
Burck, Ann. Jard. Bot. Btzg 6 (1887) 219; HEYNE,
Nutt. Pl. ed. 2 (1927) 1121.— S. stipulosa Burck,
Ann. Jard. Bot. Btzg 6 (1887) 220. — S. megistocarpa
Foxw. Mal. For. Rec. 10 (1932) 216, pl. 17; BurK.
Dict. (1935) 2019.

Large, buttressed tree. Panicles, parts of perianth
exposed in bud and nut persistently buff puberulent;
young twigs, midrib above, petioles and fruit calyx
caducously so. Twigs c. 3 by 2 mm @ apically, +
compressed, becoming smooth, dark brown; stipule
scars short, horizontal. Leaf bud small, ovoid, acute;
stipule to 20 by 4mm, + persistent, lanceolate. Leaves
6-14 by 3-6 cm, narrowly obovate to oblong, subcor-
iaceous; base obtuse or broadly cuneate; acumen to 1
cm long, turned down and frequently bent over on
pressing; nerves 14-16 pairs, rather straight and only
arching towards the ends, slender but distinctly
elevated beneath, evident above as also the densely
scalariform tertiary nerves and the midrib; petioles
7-11 mm long, slender. Panicle to 7 cm long, terminal
or axillary, with to 1.5 cm long branchlets bearing to 5
secund flowers; bracteoles to 4 by 3 mm, oblong,
concave, glabrous. Flower bud to 10 by 2 mm,
lanceolate. Sepals ovate, the 3 outer acute, the 2 inner
subacuminate. Petals cream. Stamens 15, in 3 unequal
verticils; filaments lorate, tapering; anthers subglo-
bose, becoming reflexed at least on the inner 2
verticils; appendage as long as anthers; ovary broadly
ovoid, sericeous, crowned by an equally long filiform
style. Fruit pedicel to 3 by 3mm, + impressed into the
receptacle; 3 longer calyx lobes to 11 by 2.5 cm,
broadly lorate-spatulate, subacute, c. 11 mm broad
above the to 10 by 11 mm suborbicular saccate
thickened base; 2 shorter lobes to 7 by 0.7 cm,
linear-lanceolate, similar at base; nut to 16 by 11 mm,
ovoid, apiculate.

Distr. Malesia: Malaya, Sumatra (West coast from
Pariaman and Ophir to Sibolga and Padang; Kuantan
Distr. in east).

Ecol. Frequent or common in Lowland Diptero-
carp forest on undulating land and low hills below 350m.

Vern. Méranti langgong, m. pala, m. sega, m. labu,
damar siput (Malaya), meranti katuko, katuko, m.
taratung, m. sitarah (W. coast Sumatra), m. galur, m.
sabat (Kuantan Distr. Sumatra).

146. Shorea foraminifera ASHTON, Gard. Bull. Sing.
22 (1967) 295, pl. 40; Man. Dipt. Brun. Suppl. (1968)

FLORA MALESIANA

[ser. I, vol. 9?

108. — S. teysmanniana (non DYER ex BRANDIS) ASH-
TON, Man. Dipt. Brun. (1964) 225, p.p.
Medium-sized or large buttressed tree. Young parts
(leaf and stipule excepting) fugaceous puberulent.
Twig 1-2 mm g apically, at first slightly compressed,
becoming terete, smooth; stipule scar short, + hori-
zontal. Bud to 8 by 5 mm, ovoid, compressed,
subacute. Stipule to 10 by 5 mm, oblong to elliptic,
obtuse, caducous. Leaves 6-9 by 4-7 cm, broadly
ovate, coriaceous; base cordate to obtuse; acumen to 5
mm long, somewhat falcate; nerves 8-9 pairs, stout,
prominent beneath, with large pore-like axillary
domatia, arched, at 55°-70° except at the base;
tertiary nerves densely scalariform, obscure; midrib
evident, applanate, above, stout, terete, prominent
beneath. Petiole 10-15 mm long. Flower and inflores-
cence unknown. Fruit pedicel c. 1 mm long, short.
Calyx buff puberulent in the basal half, sparsely so
distally; 3 longer lobes to 7 by 1.2 cm, spatulate,
subacute, c. 4 mm broad above the to 11 by 5 mm
ovate thickened saccate base; 2 shorter lobes to 1.5 by
0.4 cm, lorate, acute, similar at base. Nut to 15 by 10
mm, ovoid, persistently buff sericeous, shortly apicu-
late.
Distr. Malesia: Borneo (Rejang valley to Brunei).
Ecol. Locally common on shallow peat on allu-
vium in small swampy valleys; rare on hillsides.
Vern. Méranti lobang hidong (Brun.).

147. Shorea teysmanniana Dyer ex BRANDIS, J. Linn.
Soc. Bot. 31 (1895) 100; HeyNg, Nutt. Pl. ed. 1, 3
(1917) 309; ibid. ed. 2 (1927) 1126; Sym. Gard. Bull. S.
S. 7 (1933) 134, pl. 37; Mal. For. Rec. 16 (1943) 94, f.
38, 57; BROWNE, For. Trees Sarawak & Brunei (1955)
144; ANDERSON, Gard. Bull. Sing. 20 (1963) 159;
AsHTON, Man. Dipt. Brun. (1964) 225, f. 16, p.p.; ibid.
Suppl. (1968) 121; MEER & Woop, Sabah For. Rec. 5
(1964) 152.— 8S. cochinchinensis var. oligoneura
BOERL. Cat. Hort. Bog. 2 (1901) 107. — S. balanger-
oides BOERL. I.c. — S. paludosa Foxw. Mal. For. Rec.
10 (1932) 277; Burk. Dict. (1935) 2019.
Medium-sized buttressed tree. Young twig and
petiole shortly densely evenly grey-brown caducous
pubescent; outside of stipule and leaf bud persistently
so. Twig c.2mm @ apically, slightly compressed when
young, becoming terete, much branched, smooth;
stipule scars prominent, cuneate, c. 2mm long, | mm
thick, pale, + horizontal. Leaf bud 4-8 by 2-5 mm,
compressed, ovoid, subacute. Stipule to 14 by 5 mm,
ovate to oblong, tapering at base, subacute, fuga-
ceous. Leaves 7.5-11 by 3.5-7 cm, ovate, entirely
glabrous, lustrous, + coriaceous; base obtuse;
acumen to 8 mm long; nerves 8-11 pairs, slightly
raised beneath, strongly curved, at c. 60°—70° at the
base; with or without 1-3 pairs of small basal
domatia; midrib narrow, depressed above, sharply
prominent beneath. Petiole 1.2-1.8 cm long. Panicle
to 8 cm long, terminal or axillary, terete, shortly buff
pubescent; singly branched. Flower bud to 14 by 5mm,
ovoid. Calyx lobes deltoid, subequal, subacute, den-
sely buff pubescent outside, glabrous within. Petals

1982]

linear, sparsely pubescent on parts exposed in bud.
Stamens 15, of 3 lengths; filaments tapering gradually;
anther subglobose; appendage to connective short,
becoming reflexed. Ovary and stylopodium narrowly
conical, the former densely shortly cream pubescent,
the latter more coarsely so; style c. 5 length of ovary
and stylopodium, glabrous. Fruit calyx sparsely
pubescent towards apex, more densely so towards
base; 3 longer lobes to 8 by | cm, spatulate, obtuse, to
8 mm broad above the to 9 by 9 mm expanded saccate
base; 2 shorter lobes to 3 by 0.5 cm, linear, similar at
base. Nut to 12 by 8 mm, ovoid, densely shortly buff
pubescent; style remnant to 2 mm long, short, acute.

Distr. Malesia: Malaya (Selangor), E. Sumatra
(Labuan Batu, Bengkalis, Palembang, Siak), Kari-
mun, Banka, Borneo (Lower Kapuas in West Borneo,
S.E. Borneo to Sampit, Sarawak and Brunei to W.
Sabah).

Ecol. Local, sometimes common, in Mixed Peat
Swamp forest, at sea level (one record at 900 m).

Vern. Méranti bunga (Malaya, Sumatra), m. kait,
m. daun kalus (Sumatra), kélépak (S.E. Borneo),
lintang, bangkirai (Sampit), méranti lilin (Sarawak).

148. Shorea argentifolia Sym. Gard. Bull. Sing. 17
(1960) 489; BRowne, For. Trees Sarawak & Brunei
(1955) 187; ASHTON, Man. Dipt. Brun. (1964) 179, f.
16; ibid. Suppl. (1968) 105; MevER & Woop, Sabah
For. Rec. 5 (1964) 98, f. 12.

Medium-sized or large buttressed tree. Young twig,
panicle, leaf bud, stipule (both surfaces), petiole, leaf
beneath and midrib above shortly evenly densely
persistently pink-gold velutinate. Twig c.2 by 1mm @
apically, compressed when young, becoming terete,
slender, rugulose, glabrous; stipule scars c. 1.5 mm
long, short, thin, horizontal, obscure. Bud c. 4 by 2.5
mm, ovoid, slightly compressed, subacute. Stipule to
20 by 7 mm, not at first caducous, oblong-hastate,
acute, base subcordate. Leaves 6-11 by 2.5—4.5 cm,
oblong-lanceolate, thinly coriaceous; base obtuse;
acumen c. 5 mm long; nerves c. 20-25 pairs, with
prominent short secondary nerves, dense, curved
towards the margin, at c. 66°—70°; tertiary nerves
slender, densely scalariform, at c. 90°; midrib narrow
and slightly depressed above, prominent and terete
beneath; petiole 8-10 mm long, short, rugose. Panicle
to 12 cm long, terminal or axillary, compressed; singly
branched, the branchlets short, compact, bearing to 6
distichous flowers; bracteoles to 8 by 3.5 mm, oblong-
elliptic, obtuse, caducous. Flower bud to 6 by 3 mm,
ovoid, subacute. Calyx + patent, densely pubescent
outside, glabrous within; 3 outer lobes ovate-deltoid,
obtuse; 2 inner lobes c. 4 as long, ovate, subacute.
Petals pink, oblong, obtuse, densely pale golden
pubescent on parts exposed in bud, hardly connate at
base. Stamens 15, the 5 inner almost twice as long as
the others, reaching the base of the style; filaments
tapering gradually; anthers subglobose; appendage to
connective short, becoming reflexed. Ovary and stylo-
podium pyriform, shortly pubescent; style as long as
ovary, glabrous. Fruit calyx shortly pubescent; 3

DIPTEROCARPACEAE (Ashton)

539

longer lobes to 7.5 by 1.2 cm, narrowly spatulate,
narrowly obtuse, 3-4 mm broad above the to 7 by 6
mm ovate thin saccate base; 2 shorter lobes, 10-30 by
1.5mm, unequal, linear, similar at base. Nut to 1.4 by
0.9 cm, ovoid, shortly buff pubescent; style remnant
tapering, c. 2.5 mm long.

Distr. Malesia: N.E. Borneo (Rejang valley north-
eastwards to Sabah and Nunukan).

Ecol. Locally frequent in Mixed Dipterocarp for-
est, especially on clay soils on undulating land and in
valleys, below 600 m.

Vern. Séraya daun mas (Sabah), méranti binatoh,
binatoh (Sar.), sénkajang (Iban), mérangan (Brun.).

149. Shorea uliginosa Foxw. Mal. For. Rec. 10 (1932)
210, 277; BurK. Dict. (1935) 2023; AsHton, Gard.
Bull. Sing. 22 (1967) 294; Man. Dipt. Brun. Suppl.
(1968) 121, f. 15. — Shorea sp. Foxw. Mal. For. Rec.
3 (1927) 36. — S. rugosa (non HEIM) Sym. Gard. Bull.
S.S.7 (1933) 132, p.p. quoad spec. Malay. — S. rugosa
var. uliginosa SYM. Gard. Bull. S. S. 10 (1939) 372;
Mal. For. Rec. 16 (1943) 91, f. 38; ANDERSON, Gard.
Bull. Sing. 20 (1963) 159.

Large buttressed tree with dark fissured bark. Twig,
leaf bud, stipule outside, petiole and leaf beneath
densely shortly pale chocolate-brown scabrid tomen-
tose, stipule within, midrib and nerves above evenly
so; leaf above puberulent or glabrescent. Twig 3-4mm
@ apically, terete, at first prominently ribbed by the
decurrent lateral petiolar bundles, becoming
smooth; stipule scars short, pale, horizontal. Bud to 6
by 4 mm, ovoid, subacute, compressed. Stipule to 14
by 5mm, elliptic, subacute. Leaves 12-22 by 6-12 cm,
medium-sized to large, elliptic-oblong, somewhat
chartaceous, prominently boat-shaped with the lower
surface concave; base broadly cuneate to subcordate;
acumen to | cm long, acute; nerves 16-21 pairs,
prominent beneath, at 50°—70° except at the base,
tertiary nerves scalariform, at 90° to the nerves;
midrib evident but depressed above, prominent
beneath; petiole 2.2-3.2 cm long, stout, terete. Panicle
to 16 cm long, terminal or axillary, ribbed, densely
chocolate-brown scabrid tomentose; doubly
branched, branchlets bearing to 6 + distichous
flowers; bracteoles to 3 by 2 mm, elliptic, subacute,
pubescent outside, glabrous within, fugaceous. Flower
bud to 5 by 3 mm, small, ellipsoid. Sepals densely
pubescent on parts exposed in bud, ovate, acute, the 2
inner smaller than the 3 outer, thinner towards
margin, flanged at base. Petals pale yellow, elliptic-
lanceolate, densely pubescent on parts exposed in
bud. Stamens 15, in 3 unequal verticils; filaments
compressed, slender, tapering; anthers small, subglo-
bose; appendage to connective about as long as
anther, becoming reflexed with anther at anthesis.
Ovary and stylopodium conical, densely pubescent;
style filiform, glabrous, slender, as long as ovary and
stylopodium. Fruit pedicel to 1 mm long, short. Fruit
calyx persistently sericeous towards base, otherwise
glabrescent; 2 longer lobes to 6 by 1.2 cm, spatulate,
obtuse, to 4 mm broad above the to 5 by 4 mm elliptic

540

FLORA MALESIANA

[ser. I, vol. 9?

slightly thickened saccate base; 2 shorter lobes to 2cm
long, linear, similar at base. Nut to 8 by 6 mm, ovoid,
acute, densely buff pubescent.

Distr. Malesia: Malaya (Perak, Selangor), E.
Sumatra (Labuan Batu, Bengkalis, Palembang, In-
dragiri, Asahan), Karimun, Banka, W. Borneo (Sam-
pit, Sukadana, Lower Kapuas and Sarawak west of
the Baram).

Ecol. Mixed Peat Swamp forest, locally abundant.

Uses. Important as a source of dark red meranti
timber.

Vern. Méranti séngkawang, m. s. mérah, séngkaw-
ang, m. gajah, m. bahru, m. sékam, siput mélantai
(Malaya), méranti segar, m. kait kait, m. daun lebar
(Sumatra), séraya (Banka), méranti lang (Borneo),
lanan buaya (Sampit), méranti buaya, m. paya (Sar-
awak), pérawan buaya (Iban).

150. Shorea rugosa Heim, Bull. Mens. Soc. Linn.
Paris 2 (1891) 973; BRaANpis, J. Linn. Soc. Bot. 31
(1895) 96; MeRR. En. Born. (1921) 406; Sym. Gard.
Bull. S. S. 7 (1933) 132, pl. 35; ASHTON, Man. Dipt.
Brun. (1964) 218, f. 16; ibid. Suppl. (1968) 118; MEUER
& Woop, Sabah For. Rec. 5 (1964) 144. — S. verrucu-
losa DYER ex BRANDIS, J. Linn. Soc. Bot. 31 (1895) 96,
nom. in syn. — S. almon (non FOxw.) BROWNE, For.
Trees Sarawak & Brunei (1955) 150.

Medium-sized or large buttressed tree with dark
fissured bark. Young twig, panicle, bud, stipule
outside, petiole and nervation beneath densely persis-
tently purplish brown scabrid tomentose; stipule
inside, midrib above and leaf beneath shortly evenly
so. Twig to 2.5 mm @ apically, ribbed at first,
becoming terete, rugulose; stipule scars 2 mm long at
first, 0.5 mm thick, pale, slightly descending, falcate or
horizontal, obscured by tomentum. Bud 4-6 by 3-5
mm, ovoid, subacute. Stipule to 14 by 5 mm, oblong-
elliptic, subacute. Leaf 9-17 by 4-9 cm, oblong-ovate
to oblong-obovate; base obtuse; acumen to 9 mm
long, broad; margin frequently somewhat revolute;
nerves variable, 14-19 pairs, slightly curved, promi-
nent beneath, at 60°—75"; tertiary nerves scalariform,
slightly diagonal to nerves; midrib terete and promi-
nent beneath, evident but somewhat depressed above;
petiole 1.3-2.3 cm long. Panicle to 12 cm long,
terminal or axillary, lax, terete or slightly compressed;
regularly alternatively singly or doubly branched,
branchlets bearing to 6 distichous flowers; bracteoles
to 4 by 3mm, + elliptic, subacute, pubescent outside,
glabrous within, fugaceous. Flower bud to 7 by 4mm,
broadly ovoid. Calyx densely rust-brown pubescent
on parts exposed in bud; lobes subequal, ovate; 3
outer lobes obtuse, 2 inner acute to subacuminate.
Petals yellow, oblong, obtuse, densely long gold-
brown pubescent on parts exposed in bud. Stamens
15, the 5 inner somewhat longer and reaching the style
apex; filaments much twisted, tapering gradually;
anthers subglobose; appendage to connective very
short, becoming reflexed. Ovary and stylopodium
conical, densely pale grey pubescent in the distal half,
glabrous at base; style short, cylindrical, glabrous.

Fruit calyx shortly sparsely fulvous-brown to buff
pubescent; 3 longer lobes to 10.5 by 2.5 cm, spatulate,
narrowly obtuse, to 4-7 mm broad above the to 1.4 by
1 cm elliptic to ovate saccate thickened base; 2 shorter
lobes subequal, to 5 by 0.4.cm, linear, acute, similar at
base. Nut to 2.3 by 1.5 cm, ovoid, persistently shortly
evenly buff fulvous pubescent; style remnant acute.

Distr. Malesia: Borneo (Puruktjau and Lower
Dayak in S. Borneo, Melawi in W. Borneo, W.
Sarawak to S.W. Sabah).

Ecol. Local on leached yellow and occasionally
white sandy soils in Mixed Dipterocarp forest and its
ecotone with Heath forest below 400 m, subcoastal.

Note. Some collections from South and South-
East Borneo bear larger leaves and closely resemble S.

furfuracea Mia. (q.v.) a Sumatran species that may

prove, when flowers are collected, to be conspecific.

151. Shorea leprosula Mig. Sum. (1860) 487, 191;
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 84; DC. Prod. 16,
2 (1868) 631; WALP. Ann. 7 (1868) 379; SCHEFF. Nat.
Tijd. N. I. 31 (1870) 349; Dyer, FI. Br. Ind. 1 (1874)
305; BurcK, Ann. Jard. Bot. Btzg 6 (1887) 215; KING,
J. R. As. Soc. Beng. Sc. 62, 2 (1893) 110; BRANDIS, J.
Linn. Soc. Bot. 31 (1895) 100; Ripv. Agr. Bull. Str. &
F.M.S. 1 (1901) 55; Becc. For. Born. (1902) 570, 571;
HeEyneE, Nutt. Pl. ed. 1, 3 (1917) 304; Burk. J. Str. Br.
R. As. Soc. 76 (1917) 161, fig.; ibid. 79 (1918) 41; ibid.
81 (1920) 75, fig.; MERR. En. Born. (1921) 405; RIDL.
Fl. Mal. Pen. 1 (1922) 222; Baker f. J. Bot. 68, Suppl.
(1924) 11; THORENAAR, Med. Proefst. Boschw. 16
(1926) 117, t. 20; HEYNE, Nutt. Pl. ed. 2 (1927) 1122;
Foxw. Mal. For. Rec. 3 (1927) 26, 3 pl.; SLooT. in
Merr. Pl. Elm. Born. (1929) 203: E>warps, Mal. For.
Rec. 9 (1931) 144; Foxw. Mal. For. Rec. 10 (1932)
220: DescH, Mal. For. Rec. 12 (1936) 43, pl. 7, f. 1;
ibid. 14 (1941) 36, pl. 15, f. 1; ibid. 15 (1941) 136, pl. 40,
f. 1, pl. 43, f. 3; Sym. Mal. For. Rec. 16 (1943) 75, f. 38,
46, 47: SLooT. Bull. Bot. Gard. Btzg III, 18 (1949) 262,
f. 14; Browne, For. Trees Sarawak & Brunei (1955)
110; BACKER & Baku. f. Fl. Java 1 (1963) 331;
ASHTON, Man. Dipt. Brun. (1964) 193, f. 16; ibid.
Suppl. (1968) 110; MeeR & Woop, Sabah For. Rec. 5
(1964) 110. — Hopea ? maranti Mia. Sum. (1860) 489,
192: DC. Prod. 16, 2 (1868) 635; WALP. Ann. 7 (1868)
379: HEYNE, Nutt. Pl. ed. 2 (1927) 1122. — S. maranti
Burck, Ann. Jard. Bot. Btzg 6 (1887) 217; KinG, J. R.
As. Soc. Beng. Sc. 62, 2 (1893) 120; BRANDis, J. Linn.
Soc. Bot. 31 (1895) 101; HEYNgE, Nutt. Pl. ed. 1, 3
(1917) 306: ibid. ed. 2 (1927) 1122. —S. astrosticta
ScorT. ex Foxw. Mal. For. Rec. 10 (1932) 220, nom.
in syn.

Large buttressed tree. Twig, panicle, leaf bud,
stipule, petiole and nervation beneath persistently
evenly densely shortly pale buff pubescent. Twig c. 1.5
mm @ apically, terete, ridged when young, much
branched, slender, becoming glabrous, smooth; sti-
pule scars short, horizontal, obscure. Bud 3-5 by 2-3
mm, compressed, broadly ovoid, subacute. Stipule to
10 by 3.5 mm, oblong to broadly hastate, obtuse,
fugaceous. Leaves 8-14 by 3.5-5.5 cm, elliptic to

1982]

DIPTEROCARPACEAE (Ashton)

54]

ovate, thinly coriaceous, cream below in mature trees;
base obtuse or broadly cuneate; acumen short, to 8
mm long; nerves 12-15 pairs, slender, curved towards
margin, set at c. 40°—55°; tertiary nerves very slender,
densely scalariform, obscure except in young trees;
midrib narrow and depressed above and prominent
beneath, in young trees beset from the base up more or
less its length with lines of small, pale, scale-like
domatia occasionally extending also on the nerves;
petiole 1.0-1.5 cm long. Panicle to 14 cm long;
terminal or axillary, terete, lax, slender, sparsely or
densely evenly persistently pale brown to cream
pubescent; regularly singly, rarely doubly, branched,
branchlets short, bearing to 12 + secund flowers;
bracteoles to 3 by 2 mm, elliptic, obtuse, shortly
pubescent, fugaceous. Flower bud to 6 by 3 mm,
fusiform, subacute. Calyx densely pale brown pubes-
cent outside, glabrous within; 3 outer lobes narrowly
ovate, obtuse; 2 inner lobes broadly ovate, shorter,
shortly acuminate. Petals pale yellow, narrowly
oblong, densely pale yellowish grey pubescent on
parts exposed in bud. Stamens 15, the inner 5 twice as
long as the others and reaching half the length of the
style; filaments long, tapering gradually; anthers
subglobose; appendage to connective short, becoming
reflexed. Ovary and stylopodium ovoid, glabrous; style
filiform, twice as long as ovary and stylopodium,
glabrous. Fruit calyx glabrescent or persistently
shortly pubescent at base; 3 longer lobes to 10 by 2cm,
spatulate, obtuse, c. 5 mm broad above the to 8 by 6
mm thickened elliptic shallowly saccate base; 2
shorter lobes to 5.5 by 0.3 cm, unequal, similarly
saccate at base. Nut to 2 by 1.3 cm, ovoid, densely pale
buff pubescent; style remnant c. 2 mm long, tapering,
acute.

Distr. Thailand (Pattani) and in Malesia: Malaya
(excluding seasonal areas), Sumatra, Banka, Billiton,
Borneo.

Ecol. Common, often abundant, on deep clay soils
in Mixed Dipterocarp forest below 700 m.

Vern. Meéranti témbaga, m. bétul, m. bunga, m.
lampong, m. témak, m. hijau, m. sabut, m. kait kait
(Malaya, Sumatra), m. sépang (Palembang), /ampong
(Kutei), pérawan lop (Sarawak), séraya témbaga
(Sabah), kontoi (Melawi), /éntang (Sampit) and many
others.

152. Shorea platycarpa Heim, Bull. Mens. Soc. Linn.
Paris 2 (1891) 956; BRANDIS, J. Linn. Soc. Bot. 31
(1895) 94; Becc. For. Born. (1902) 510; Merr. En.
Born. (1921) 406; Sym. Gard. Bull. S. S. 7 (1933) 133,
pl. 36; Mal. For. Rec. 16 (1943) 88, f. 38, 53; BROWNE,
For. Trees Sarawak & Brunei (1955) 142; ANDERSON,
Gard. Bull. Sing. 20 (1963) 159; ASHTON, Man. Dipt.
Brun. (1964) 211; ibid. Suppl. (1968) 115. — S. lepro-
sula (non Mig.) BorRL. Cat. Hort. Bog. 2 (1899)
110. —.S. palustris Riv. Fl. Mal. Pen. 1 (1922) 224;
Foxw. Mal. For. Rec. 3 (1927) 381; ibid. 10 (1932)
212; BurK. Dict. (1935) 2019. — Fig. 106 B—B3.
Large buttressed tree. Young twig, panicle, bud and
petiole densely shortly persistently rust-brown scabrid

pubescent, sparsely so on stipule, midrib above and
leaf beneath, glabrescent on leaf above. Twig c. 3 mm
@ apically, stout, ridged, verrucose; stipule scars
short, narrow, ascending, obscure. Bud 4-6 by 3.5—5
mm, broadly ovoid, obtuse. Stipule to 10 by 6 mm,
ovate, subacute. Leaves 9-17 by 5.5—10 cm, medium-
sized to large, elliptic-oblong or broadly ovate, thinly
coriaceous; base obtuse; acumen to | cm long, short,
broad; nerves 16-20 pairs, close, straight, curved near
margin, with or without scale-like axillary domatia
(without in old trees); tertiary nerves distant, scalari-
form, slender but distinct, slightly diagonal to nerves;
midrib broad, terete beneath, narrow yet hardly
depressed above; petiole 1.5—2 cm long, stout. Panicle
to 9 cm long, terminal or axillary, ribbed, straight,
densely shortly persistently rust-brown scabrid pubes-
cent; singly or doubly branched; bracteoles to 4 by 3
mm, broadly ovate, acute, densely shortly pubescent
outside, glabrous within. Flower bud to 3 by 4 mm,
ellipsoid-ovoid, obtuse. Calyx densely pubescent out-
side, glabrous within; 3 outer lobes ovate, obtuse, c. 2
x length of 2 inner lobes; inner lobes broadly ovate,
prominently caudate. Petals pale yellow, narrowly
lanceolate, densely pubescent on parts exposed in
bud. Stamens 15, in 3 unequal verticils, the longest
almost as long as the ovary; filaments compressed,
tapering gradually; anthers subglobose; appendage to
connective as long as anther, becoming reflexed.
Ovary and stylopodium glabrescent, narrowly ovoid;
style as long as ovary and stylopodium, filiform,
glabrous. Fruit subsessile, calyx shortly persistently
pubescent towards base, otherwise glabrescent; 2
longer lobes to 6.5 by 1.2 cm, narrowly spatulate,
narrowly obtuse, to 4 mm broad above the to 5 by 4
mm elliptic slightly thickened saccate base; 2 shorter
lobes to 2cm long, linear, similar at base. Nut to 8 by 6
mm, ovoid, densely buff pubescent; style remnant
short, acute.

Distr. Malesia: Malaya (both coasts), Sumatra
(Palembang, Siak), Banka, Billiton, Borneo (except E.
Sabah).

Ecol. Locally common, widespread, in Mixed Peat
Swamp forests.

Vern. Méranti paya, m. kait kait, m. ségar
(Malaya).

Note. With close affinities to S. leprosula of the
lowland forests.

153. Shorea curtisii DYER ex KING, J. R. Soc. Beng.
Sc. 62, 2 (1893) 111; BRUHL & KING, Ann. R. Bot.
Gard. Calc. 5, 2 (1896) 152, t. 185; ASHTON, Gard.
Bull. Sing. 31 (1978) 48.

a. ssp. curtisii. — BRANDIS, J. Linn. Soc. Bot. 31 (1895)
101; Rip. Agr. Bull. Str. & F.M.S. 1 (1901) 58; FI.
Mal. Pen. | (1922) 223; HEyNg, Nutt. Pl. ed. 1, 3(1917)
299: ibid. ed. 2 (1927) 1116; Burk. J. Str. Br. R. As.
Soc. 81 (1920) 71, fig.; Foxw. Mal. For. Rec. 1 (1921)
78; ibid. 3 (1927) 39; ibid. 10 (1932) 226; Burk. Dict.
(1935) 2009; CoRNER, Wayside Trees | (1940) 213;
Sym. Mal. For. Rec. 16 (1943) 67, f. 38, 40, 41;

542

FLORA MALESIANA

[ser. 1, voleg?

me nn Spe

Fig. 112. Habit of a big tree of Shorea curtisii DYER ex

KING. Note man at bottom left from tree, one tree

climber halfway up, one in fork of first branch.
Brunei, Labi Road (Photogr. ASHTON).

BROWNE, For. Trees Sarawak & Brunei (1955) 147;
ASHTON, Man. Dipt. Brun. (1964) 185, f. 16, pl. 45
(habit, bark); ibid. Suppl. (1968) 106; MEIJER & Woop,
Sabah For. Rec. 5 (1964) 103; BurRGEss, F.R.I. Res.
Pamphlet 66 (1975). — Fig. 6, 112, 113.

Large buttressed tree. Young twig, panicle, leaf
bud, stipule outside (glabrescent within) petiole, and

midrib and nerves beneath densely shortly evenly
caducous pale pink-brown to grey puberulent. Twig c.
1.5 mm @ apically, terete, much branched, slender,
becoming smooth, glabrous; stipule scars short,
cuneate, horizontal, obscure. Bud 4-9 by 2.5—3.5 mm,
subacute, ovate to slightly falcate, slightly constricted
at base. Stipule to 10 by 4 mm, obtuse, slightly
constricted at base. Leaves 6-9 by 2.5—3.5 cm, ovate-
lanceolate, pale pink to grey lepidote beneath; base
broadly cuneate; acumen to 8 mm long, narrow;
nerves 9-11 pairs, curved, slender, hardly raised
beneath, at 50°-60°; tertiary nerves slender, densely
scalariform, obscure, diagonal to nerves; midrib
slender, prominently terete beneath, narrow
depressed above; petiole 1-1.3 cm long, slender.
Panicle to 6 cm long, terminal or axillary, terete; singly
or doubly branched, branchlets short, bearing to 12
distichous flowers; bracteoles to 2.5 by 2 mm, sub-
acute, shortly pale grey pubescent outside, glabrous
within, fugaceous. Bud to 5 by 3 mm, ovoid, obtuse.
Calyx densely pale grey puberulent outside, glabrous
within; lobes ovate, 3 outer longer, narrow, obtuse; 2
inner broad, acuminate. Petals pale cream yellow
(Malaya) or deep crimson (Borneo), linear, obtuse,
connate at base, densely pubescent on parts exposed
in bud. Stamens 15, the outer 10 sometimes aborting,
the inner 5 longer, reaching the ovary apex; filaments
tapering gradually to the anther, markedly gibbous;
anther subglobose; appendage to connective short on
outer 10 stamens, rudimentary on inner 5, reflexed.
Ovary and stylopodium ovoid, densely pubescent
except at base; style short, broadening at the apex,
glabrous. Fruit pedicel c. | mm long, short. Fruit calyx
puberulent at base, otherwise glabrous; 3 longer lobes
to 7 by 1 cm, narrowly spatulate; apex narrowly
obtuse, tapering to c. 2.5mm broad above the to 6 by 5
mm ovate shallowly saccate thickened base; 2 shorter
lobes to 4 by 0.3 cm, unequal, linear, similarly
expanded at base. Nut to 1.2 by 0.9 cm, ovoid,
tapering to a short | mm long apiculus, shortly
densely pale buff pubescent.

Distr. Peninsular Thailand (Pattani) and in Male-
sia: Malaya, Sumatra (Singkep, Lingga), Borneo
(Rejang valley to S.W. Sabah).

Ecol. Deep dry soils on coastal hills; in Malaya
continuing up ridges and gregarious in Hill Diptero-
carp forests between 300-850 m. See for detailed
ecology BURGESS (1975).

Uses. The major dark red meranti timber in
Malayan hills.

Vern. Séraya, s. sabut, s. bukit, méranti lampong, m.
sutra, m. séraya, penak lampong, bohor, jarang, mén-
tanam (Malaya), séraya (Riouw).

Note. Small groups of hybrid trees between this
species and S. /eprosula exist on Bukit Timah, Singa-
pore and Bukit Lagong, Selangor; this subspecies may
have the same origin. The two species often occur
together at the margin of their respective ecological
ranges, and it is curious that morphologically interme-
diate trees are not more common.

1982]

b. ssp. grandis ASHTON, Gard. Bull. Sing. 31 (1978)
48. — Shorea sp. B SyM. Mal. For. Rec. 16 (1943) 95.

Differing as follows: Twig c. 4 by 2mm @ apically,
compressed. Leaves 11-17 by 5-8 cm, elliptic; nerves
10-13 pairs; petiole 2-2.5 cm long.

Distr. Malesia: Malaya (Perak).

Ecol. As ssp. curtisii.

Vern. Séraya daun besar.

154. Shorea scabrida Sym. Gard. Bull. S. S. 8 (1935)
287, pl. 28; BRowneg, For. Trees Sarawak & Brunei
(1955) 143; ANDERSON, Gard. Bull. Sing. 20 (1963)
159; ASHTON, Man. Dipt. Brun. (1964) 221, f. 16, pl.
58 (habit, stem-base); ibid. Suppl. (1968) 119; MEWER
& Woop, Sabah For. Rec. 5 (1964) 146.

Medium-sized tree. Young twig, panicle, leaf bud,
outside of stipule, midrib on both surfaces and
nervation beneath + persistently densely tawny-
brown scabrid pubescent. Twig c. 2 mm @ apically,
much branched, terete, becoming smooth, glabrous;
stipule scars short, horizontal, obscure. Bud 2.5—5 by
1.5—3 mm, ovoid, compressed, obtuse. Stipule to 6 by
3 mm, oblong, obtuse, fugaceous. Leaves 5—9 by 3-5
cm, small, obovate or elliptic, coriaceous; basv
broadly cuneate, rarely obtuse; apex retuse, obtuse or
with to 5mm long acumen; nerves 8-11 pairs, slightly
curved, at c. 25°-40°, rarely with domatia; tertiary
nerves slender, scalariform, diagonal to nerves; mid-
rib narrow and depressed above, terete and prominent
beneath; petiole 9-11 mm long, short, rugose. Panicle
to 8 cm long, terminal or axillary, short, terete or
slightly compressed; singly, rarely doubly, branched,
branchlets short, rather irregular, bearing to 7 +
distichous flowers; bracteoles to 3.5 by 2.5 mm, ovate,
subacute, shortly buff pubescent outside, glabrescent
within. Flower bud to 8 by 3.5 mm, lanceolate, obtuse.
Calyx densely pale buff pubescent outside, glabrous
within; 3 outer lobes deltoid to ovate, subacute; 2
inner lobes smaller, broadly ovate, subacuminate.
Petals cream, pink at base, lanceolate, acute, densely
pubescent on parts exposed in bud. Stamens 15, in 3
distinct verticils; filaments tapering gradually; anther
subglobose; appendages to connective slightly longer
than anther, the longest reaching to the base of the
style, becoming reflexed. Ovary and stylopodium
narrowly conical, densely pubescent except at the
base; style almost half length of ovary and stylopo-
dium, glabrous. Fruit calyx glabrescent; 3 longer lobes
to 7 by 1.5 cm, spatulate, c. 3 mm wide above the to 7
by 5mm ovate saccate thickened base; 2 shorter lobes
to 3 by 0.3 cm, linear, similar at base. Nur to 10 by 8
mm, ovoid, shortly densely buff pubescent, shortly
apiculate.

Distr. Malesia: E. Sumatra (Lower Langkat,
Langsa, Lingga), Borneo.

Ecol. Local in fresh water swamp forest on shallow
peat overlying sand, and on skeletal sandy soil on
ridges and plateaux, in Heath forest and Mixed
Dipterocarp forest.

Vern. Kulap daun, méranti tembalang, m. pepak
lantai, péngérawan (W. Borneo), lanan lutung, I.

DIPTEROCARPACEAE (Ashton)

543

Fig. 113. Trunk-base of Shorea curtisii DYER ex KING,

with V-shaped fissure bark. Brunei
ASHTON).

(Photogr.

témbaga (S.E. Borneo), bangkirai lutung (S.E. Bor-
neo), séraya lop (Sabah), méranti télur (Kedayan), m.
lop (Sarawak).

Notes. A variable species with several local eco-
types varying principally in leaf shape and persistence
and evenness of tomentum.

Doubtfully distinct from S. retusa; apparently
intermediate forms occur in S.E. Borneo (sterile
collections).

155. Shorea revoluta ASHTON, Gard. Bull. Sing. 19
(1962) 304, pl. 26; Man. Dipt. Brun. (1964) 215, f. 16;
ibid. Suppl. (1968) 117; MEER & Woop, Sabah For.
Rec. 5 (1964) 142.

Medium-sized tree. Twig, panicle, leaf bud, stipule
outside (pubescent within) and petiole densely shortly

544 FLORA MALESIANA [ser. I, vol. 9?

Fig. 114. Hill dipterocarp forest with Shorea ovata DYER ex BRANDIS amidst the common bertam palm,
Eugeissona triste. Malaya (Photogr. WyATT-SMITH, 1948).

pale brown to fulvous somewhat scabrid pubescent;
nervation beneath and midrib above more sparsely so,
glabrescent. Twig c. 2-3 mm g apically, at first slightly
ribbed and compressed, becoming terete, stout,
smooth, glabrous: stipule scars to 2 mm long at first,
horizontal, narrow, obscure. Bud 4-7 by 3-5 mm,
ovoid, compressed, subacute. Stipule to 1.5 by 0.7 cm,
elliptic to oblong, obtuse. Leaves 10-15 by 5.5-10 cm,

ovate, coriaceous, lustrous; base obtuse to subcor-
date; acumen to 1.5 cm long, narrow; margin revolute:
nerves 9-12 pairs, curved, prominent beneath, well
spaced, at to 90° at the base and down to 45° towards
the apex, with small glabrous axillary domatia; ter-
tiary nerves slender, scalariform, diagonal to nerves;
midrib narrow and slightly depressed above, promi-
nent beneath; petiole 1.3—-1.5 cm long. Panicle to 22 cm

1982]

long, singly or doubly branched, terminal or axillary,
straight. Flower bud to 7 by 4 mm. Sepals narrowly
deltoid-ovate, subacute, pubescent on parts exposed
in bud, the inner 2 smaller than the outer 3. Petals
lanceolate, hispid on parts exposed in bud. Stamens
15, in 3 unequal verticils; filaments slender, com-
pressed, tapering to the small subglobose anthers;
appendage to connective slender, short, as long as
anther, anthers and appendages becoming reflexed.
Ovary and stylopodium narrowly conical, densely
pubescent; style slender, filiform, glabrous, c. 5 length
of ovary and stylopodium. Fruit pedicel to 1.5 cm
long. Calyx puberulent; 3 longer lobes to 7.5 by 1.3
cm, narrow, spatulate, obtuse, c. 3 mm broad above
the c. 6 by 5 mm ovate thickened saccate base; 2
shorter lobes to 3.5 by 0.2 cm, subequal, linear, acute,
similar at base. Nut to 1.5 by 0.8 cm, ovoid, pale buff
pubescent; style remnant c. 1.5 mm long, tapering.

Distr. Malesia: Borneo (N.E. Sarawak, Brunei,
S.W. Sabah).

Ecol. Local, Heath forest on terraces and sand-
stone plateaux to 1200 m.

Vern. Méranti kérangas (Sarawak), séraya daun
tajam (Sabah).

156. Shorea ovata DYER ex BRANDIS, J. Linn. Soc.
Bot. 31 (1895) 91; Merr. En. Born. (1921) 406; Sym.
Gard. Bull. S. S. 7 (1933) 140, pl. 40; Mal. For. Rec. 16
(1943) 82, f. 38, 50: BRowneE, For. Trees Sarawak &
Brunei (1955) 148; ASHTON, Man. Dipt. Brun. (1964)
230, f. 16; ibid. Suppl. (1968) 112; MevEeR & Woop,
Sabah For. Rec. 5 (1964) 127, pl. 8A. — S. parvifolia
(non Dyer) KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893)
112, p.p.; Foxw. Mal. For. Rec. 10 (1932) 199. — S.
plagata Foxw. Philip. J. Sc. 13 (1918) Bot. 192; ibid. 67
(1938) 308. — S. agsaboensis W.L. STERN, Brittonia
17 (1965) 36, f. 1-2; Rojo, Pterocarpus 3 (1977) 70,
fig. — Fig. 114.

Small or medium-sized tree. Young twig, panicle,
leaf bud, stipule (both surfaces), petiole, midrib above
and leaf beneath densely persistently evenly ochra-
ceous to rust pubescent, scabrid on raceme and
nervation beneath. Twig c. 1 mm g apically, slender,
much branched, terete, becoming smooth, glabrous;
stipule scars c. 1 mm long, thin, slightly descending.
Bud 3-6 by 1.5—3 mm, ovoid, obtuse. Stipule to 13 by 5
mm, ovate, subacute, fugaceous. Leaves 4-8 by
2.5-4.5 cm, small, coriaceous, broadly ovate; base
subequal to equal, obtuse; acumen to | cm long,
narrow; margin slightly revolute; nerves 8-10 pairs,
curved, prominent beneath, at c. 50°—70°, with (in
Borneo) small pilose axillary domatia; tertiary nerves
slender, densely scalariform, set diagonally to the
nerves; midrib slightly depressed above; petiole 1—-1.3
cm long. Panicle to 11 cm long, terminal or axillary,
terete or slightly compressed: singly or doubly
branched, branchlets to 3 cm long, bearing to 8 +
distichous flowers; bracteoles to 3.5 by 2mm, oblong,
obtuse, sparsely puberulent outside, glabrescent
within. Flower bud to 6 by 2.5 mm, ovoid, subacute.
Calyx densely golden-brown pubescent outside, gla-

DIPTEROCARPACEAE (Ashton)

545

brescent within; 3 outer lobes narrowly ovate, acu-
minate; 2 inner shorter, broader, more acuminate:
petals pale pink with cream margin, lanceolate,
obtuse, pubescent on parts exposed in bud, loosely
connate on falling. Stamens of 3 lengths, the outer 5
often aborting, the inner 5 about twice their length;
filaments tapering gradually; anthers subglobose,
narrower apically; appendage to connective short,
becoming reflexed. Ovary ovoid, with a band of hairs
at the apex, otherwise glabrous, tapering into the
rather short glabrous style. Fruit calyx glabrescent, +
persistently puberulent at base; 3 longer lobes to 5.5
by | cm, spatulate, obtuse, to 4mm broad above the to
7 by 6 mm ovate shallowly saccate thickened base; 2
shorter lobes to 3 by 0.3 cm, subequal, linear, similarly
saccate at base. Nur to 11 by 8 mm, broadly ovoid,
tapering, shortly evenly densely buff pubescent: style
remnant to 2.5 mm long, acute.

Distr. Malesia: Malaya, Sumatra (W. coast:
Tapanuli, Sibolga to Painan, Bangkinang:; P.
Musala), Borneo (Kapuas hinterland, Sarawak to
S.W. Sabah, S.E. Borneo), Philippines (Mindanao).

Ecol. Sandy soils on coastal hills (especially in
Borneo); inland ridges to 1500 m (especially in Malaya
and Sumatra).

Vern. Méranti sarang punai bukit (Malaya), man-
dirawan (Sumatra), séraya punai bukit (Sabah).

Notes. Formerly I pointed out (Gard. Bull. Sing.
31, 1978, 47) that S. plagata and S. agsaboensis (from
Mindanao) represented the same species and referred
these names to S. pauciflora, owing to the leaf shape
and 8 pairs of nerves, pointing out, however, that the
Philippine plants differed in the usually small leaf-size
and the usual but not consistent presence of small
axillary domatia up to the midrib. I also indicated that
flowers would be needed for confirmation. J. P. Rojo
has now collected and described (1977, /.c.) these
flowers (RoJo 292, Agusan del Norte, Mindanao) and
correctly pointed out that they place the species in a
different section, Mutica, S. paucifiora being in sect.
Brachypterae. Roso’s excellent field description
alludes to the fissured bark and bluish-red corolla,
pink at the base. This matches Shorea sect./subsect.
Mutica, while the description of the corolla is distinc-
tive and clearly matches that of S. ovata, in which the
leaves also bear 8 pairs of nerves and pubescent
axillary domatia. Though not closely resembling the
leaves of S. ovata as it occurs in Borneo, the Philippine
specimens so much resemble specimens from Sumatra
and West Malaysia, that I am convinced of their
conspecificy. This conclusion is strengthened by
Rojo’s comment that it is in Mindanao a species of
high ridges, exactly as is S. ovata, except in northern
Borneo where it spreads down onto sandy soils in the
lowlands.

157. Shorea rubra ASHTON, Gard. Bull. Sing. 19
(1962) 309, pl. 28; Man. Dipt. Brun. (1964) 217, f. 16;
ibid. Suppl. (1968) 118; MevER & Woop, Sabah For.
Rec. 5 (1964) 144.

Medium-sized or large buttressed tree with dark

546

FLORA MALESIANA

[ser. I, voleg2

bark. Twig, panicle, bud, stipule (both surfaces,
shorter within), petiole, midrib above and leaf
beneath densely persistently deep rufous-brown sca-
brid pubescent. Twig to 1.5 mm @ apically, terete,
much branched, becoming smooth; stipule scars c. 1
mm long, short, narrow, descending. Bud 4-7 by 3-4.5
mm, compressed, ovoid, broadly acute. Stipule to 16
by 5mm, hastate, subacute. Leaves 8-13 by 4-7.5 cm,
broadly ovate, thinly coriaceous; base obtuse; acumen
to 1.3 cm long; margin narrowly revolute; nerves
11-13 pairs, prominent beneath, curved distally, at
40°—50°; tertiary nerves slender, densely scalariform,
diagonal to nerves; midrib prominently terete
beneath, narrow and shallowly depressed above;
petiole 1-1.4 cm long. Panicle to 14 cm long, terminal
or axillary, terete, rather straight and ridged; singly or
doubly regularly branched, branchlets short, com-
pact, bearing to 5 distichous flowers; bracteoles to 4 by
3.5mm, broadly ovate, subacute, imbricate round the
bud, shortly pale buff pubescent outside, more spar-
sely so within. Flower bud to 7 by 4 mm, ellipsoid to
subglobose, obtuse. Calyx densely pale grey pubes-
cent outside, glabrous within; 3 outer lobes deltoid-
ovate, obtuse; 2 inner lobes smaller, broadly ovate,
acute. Petals pale yellow, narrowly oblong, subacute,
shortly pubescent on parts exposed in bud. Stamens
15, of 3 lengths, the inner 5 being almost twice as long
as the others; filaments of outer 10 tapering gradually
to anther, filaments of inner 5 oblong, broad, tapering
abruptly distally; anthers subglobose; appendage to
connective shorter than anther, becoming slightly
reflexed. Ovary and stylopodium ovoid-conical, den-
sely pubescent except at the base; style shorter than
ovary, glabrous. Fruit calyx sparsely puberulent,
glabrescent; 3 longer lobes to 11 by 1.6 cm, narrowly
spatulate, obtuse, to 5 mm broad above the to 8 by 6
mm deeply saccate thickened base; 3 shorter lobes to
4.5 by 0.4 cm, subequal linear, acute, similarly
expanded at base. Nut to 10 by 9 mm, broadly ovoid,
shortly densely evenly pale buff pubescent. Style
remnant to 2 mm long, tapering.

Distr. Malesia: Northern Borneo (West Borneo;
Rejang valley to West Sabah and Tawau).

Ecol. Uncommon, on sandy clay soils on low hills
and ridges to 1350 m.

Vern. Méranti méerah kesumba (Brun.).

158. Shorea dasyphylla Foxw. Mal. For. Rec. 10
(1932) 224, pl. 18; BurK. Dict. (1935) 2010; Sym. Mal.
For. Rec. 16 (1943) 69, f. 38, 42; BROWNE, For. Trees
Sarawak & Brunei (1955) 138; ASHTON, Man. Dipt.
Brun. Suppl. (1968) 106, f. 13.

Medium-sized or large buttressed tree. Twigs, buds,
stipules, petioles, leaf beneath and midrib above
densely persistently shortly scabrid golden-brown
pubescent, leaf above puberulent. Twig c. 2 mm @
towards apices, terete; stipule scars short, obscure.
Bud to 3 by 2 mm, ovoid, obtuse. Stipules to 6 by 4
mm, broadly ovate, obtuse, caducous. Leaves 7-14 by
3-6 cm, ovate to elliptic, coriaceous, margin some-
what revolute; base obtuse or broadly cuneate;

acumen to | cm long, slender; nerves 11—15 pairs,
prominent beneath, at 50°—65°; tertiary nerves scalari-
form, elevated beneath; midrib + depressed above,
prominently terete, striated, beneath; petiole 12-15
mm long. Panicle to 8 cm long, terminal or axillary,
terete or ribbed, densely persistently shortly golden-
brown scabrid pubescent; singly branched, branchlets
to | cm long, bearing to 4 flowers; bracteoles to 3 by 2
mm, elliptic, obtuse, pubescent, caducous. Flower bud
to 4 by 3 mm, ovoid. Calyx densely pubescent outside,
sparsely so within; 3 outer lobes deltoid-ovate, 2 inner
lobes smaller, broadly ovate. Petals cream-yellow,
oblong, obtuse, densely pubescent on parts exposed in
bud. Stamens 15, of 3 lengths, the inner 5 about twice
the length of the others; filaments tapering gradually
to the subglobose anthers; appendages to connective
short, slender, becoming reflexed. Ovary and stylopo-
dium ovoid to conical, densely pubescent; style shorter
than ovary, glabrous. Fruit pedicel to 2 mm long.
Calyx glabrescent; 3 longer lobes to 9 by 1.3 cm,
narrowly spatulate, subacute, c. 4 mm broad above
the to 8 by 6 mm ovate saccate thickened base; 2
shorter lobes to 40 by 4 mm, linear, similar at base.
Nut to 18 by 9 mm, ovoid, shortly evenly buff
pubescent, shortly apiculate.

Distr. Malesia: Malaya (excluding seasonal area),
Sumatra (Palembang north to Langkat and Labuan
Batu in east), Borneo (Sarawak west of the Lupar).

Ecol. Scattered in Mixed Dipterocarp forest on
well drained flat land and low hills and occasionally to
1000 m.

Vern. Méranti batu, m. tembaga, m. sarang punat,
m. sabut (Mal.), m. gambong, m. sabut, kétuho andilan
(Sumatra).

159. Shorea parvifolia Dyer, Fl. Br. Ind. 1 (1874) 305;
Kina, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 112, p.p.;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 92, p.p.; RIDL.
Agr. Bull. Str. & F. M.S. 1 (1901) 57; Fl. Mal. Pen. 1
(1922) 224: Burk. J. Str. Br. As. Soc. 81 (1921) 517;
HEYNE, Nutt. Pl. ed. 1, 3 (1917) 306; ibid. ed. 2 (1927)
1123; Foxw. Mal. For. Rec. 3 (1927) 31; SLooT. in
Merr. Pl. Elm. Born. (1929) 203; Epwarps, Mal. For.
Rec. 9 (1931) 146; Sym. Gard. Bull. S. S. 7 (1933) 137,
pl. 39; Mal. For. Rec. 16 (1943) 85, f. 37C, 38, 51;
BROWNE, For. Trees Sarawak & Brunei (1955) 141;
AsHToN, Gard. Bull. Sing. 20 (1963) 278; Man. Dipt.
Brun. (1964) 206, f. 16; ibid. Suppl. (1968) 113; MEUER
& Woop, Sabah For. Rec. 5 (1964) 128, f. Ic.

a. ssp. parvifolia. — S. scutulata KiNG, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 110; BRANDIS, J. Linn. Soc. Bot.
31 (1895) 102; BurK. J. Str. Br. R. As. Soc. 81 (1920)
71, fig.; RIDL. Fl. Mal. Pen. 1 (1922) 222; Foxw. Mal.
For. Rec. 10 (1932) 201; Burk. Dict. (1935)
2022. — S. gentilis Parws in Fedde, Rep. 33 (1933)

244.
Large tree. Young twig, panicle, leaf bud, stipule,

petiole and midrib above shortly evenly persistently
pale brown pubescent or glabrous, otherwise gla-
brous. Twig c. 2 mm @ apically, terete, much
branched, becoming glabrous, smooth; stipule scars

1982]

short, inconspicuous. Bud 4-7 by 3-5 mm, ovoid,
compressed, obtuse. Stipule to 12 by 4 mm, oblong to
ovate, obtuse. Leaves S—9 by 2.5—S cm, broadly ovate,
thinly coriaceous, base obtuse or cordate with occa-
sionally a pair of large pale scale-like domatia;
acumen to | cm long, margin slightly revolute; nerves
10-13 pairs, slightly curved, slender, hardly elevated
beneath; tertiary nerves slender, closely scalariform
diagonal to nerves, midrib narrow and depressed
above; petiole 1—1.5 cm long. Panicle to 12 cm long,
terminal or axillary, slender, terete or slightly com-
pressed; regularly singly, rarely doubly, branched,
branchlets short, compact, bearing to 8 + secund
flowers; bracteoles to 6 by 3 mm, elliptic-oblong,
obtuse, shortly sparsely pale buff pubescent. Bud to 7
by 5 mm, broadly ovoid to subglobose, obtuse. Calyx
densely pubescent outside, glabrous within; 3 outer
lobes deltoid-ovate; 2 inner lobes smaller, broadly
ovate, acute, thin at base. Petals cream suffused with
pink at base, oblong, obtuse, shortly pubescent on
parts exposed in bud. Stamens 15, of 3 lengths, the 5
inner about twice length of others; filaments tapering
gradually; anther subglobose; appendage to connec-
tive short, slender, becoming reflexed. Ovary and
stylopodium ovoid to conical, densely pubescent
except at base; style shorter than ovary, glabrous.
Fruit calyx shortly sparsely pale brown pubescent; 3
longer lobes to 9 by 1.5 cm, thin, spatulate, obtuse, c. 4
mm broad above the c. 5.5 by 5 mm elliptic thickened
saccate base; 2 shorter lobes to 3.5 by 0.2 cm, linear,
subequal, similarly saccate at base. Nut to 14 by 7mm,
ovoid, shortly buff pubescent, style remnant c. 2.5mm
long, tapering, acute.

Distr. Thailand (Pattani) and in Malesia: Malaya,
Sumatra, P. Musala, Borneo.

Ecol. Perhaps the commonest dipterocarp in the
region, on clay soils on hills below 800 m.

Uses. A very important timber tree, the main
source of light red meranti.

Vern. Méranti sarang punai, m. bunga, m. samak,
m. daun halus, m. choh, m. lémésa bung (Malaya), m.
sabut, m. kapala tupai, tambong sawa (Sumatra),
sawang puteh, awang belah (Mahakam), ponga pipit, p.
payur, p. bahaya (Melawi), lampung, |. nasi, |. témbaga

DIPTEROCARPACEAE (Ashton)

547

\

Nin \ \\\ \| Hi}

hy \ AW DTA
SAAN PL

\ ’ AN \ \N Wy,
\ WY) ‘ vt // y Wy)
te f n Z,

. IW 4) tf

SSA

Saw!
Qyw 7/
Ny

Fig. 115. Flower details in Shorea sect. Ovalis AsH-

TON. — S. ovalis (KORTH.) BL. A. Bud, B/. outer

sepal, B2. inner sepal, C. stamens and pistil, all x 10
(SAN 19266).

(Kutei), kontoi, k. burong (W. Borneo), merangan, m.
nasi (Nunukan), dangar siak, d. burau (Murut),
pérawan lop (Iban), etc.

b. ssp. velutinata ASHTON, Gard. Bull. Sing. 20 (1963)
278.

Differing as follows: Leaves 6-11 by 3.56 cm, ovate
or elliptic; base obtuse or cuneate; nerves stoutly
prominent evenly sparsely scabrid pubescent,
beneath; margin often narrowly revolute.

Distr. Malesia: Malaya (E. Pahang, E. Johore),
Sumatra, Borneo.

Ecol. As ssp. parvifolia but mainly near the coast.

10. Section Ovalis

ASHTON, Gard. Bull. Sing. 20 (1963) 268; Man. Dipt. Brun. (1964) 117. — Shorea,
Red Meranti group, S. ovalis subgroup, SyM. Mal. For. Rec. 16 (1943) 58. — Fig.

115.

Flower buds broadly ovoid, obtuse; corolla as in sect. Rubella; stamens 50-70;
filaments very long, filiform, folded in bud; anthers subglobose, with 4 pollen
sacs; appendage to connective vestigial; ovary and stylopodium narrowly conical,
densely pubescent; style short. Stipules, bracts and bracteoles subpersistent. Leaf
tertiary nervation scalariform; midrib obscure and depressed above. Bark surface

deeply V-section fissured. Wood as in sect. Brachypterae.
Distr. Malesia: Sumatra, Malaya, Borneo. Monotypic.

548

FLORA MALESIANA

[ser. I, vol/ 9?

Fig. 116. Shorea ovalis (KORTH.) BL. ssp. ovalis. a. Young inflorescence, b. fruit, c. nut, all x $(a bb. 20237, b—c
SAN 16474).

160. Shorea ovalis (KORTH.) BL. Mus. Bot. Lugd.-
Bat. 2 (1852) 33; WALP. Ann. 4 (1857) 338; Mia. FI.
Ind. Bat. 1, 2 (1859) 503; DC. Prod. 16, 2 (1968)
631. — Fig. 115-118.

KEY TO THE SUBSPECIES

1. Leaves oblong-ovate, + bent upwards along the
midrib, base obtuse.
2. Twig and nervation and lamina beneath +
shortly scabrid pink-brown pubescent
a. ssp. ovalis
2. Twig and nervation beneath rufous tufted tomen-
tose, lamina beneath glabrous, lustrous
b. ssp. sarawakensis
1. Leaves oblong-obovate, boat-shaped with the
lower surface concave; base cuneate
c. ssp. sericea

a. ssp. ovalis. — BuRCK, Ann. Jard. Bot. Btzg 6 (1887)
219; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 103; MERR.
En. Born. (1921) 406; Sym. Gard. Bull. S. S. 10 (1939)
370; Mal. For. Rec. 16 (1943) 80, f. 37B, 38, 49;
BROWNE, For. Trees Sarawak & Brunei (1955) 132;
ASHTON, Gard. Bull. Sing. 20 (1963) 274; MEIJER &
Woop, Sabah For. Rec. 5 (1954) 125, f. 1b, pl.
8b. — Dilleniacea ? nervosa WALL. Cat. (1832) 6635,
nomen. — Vatica ovalis KorTH. Kruidk. (1841) 73;
WALP. Rep. 5 (1845) 127; DC. Prod. 16, 2 (1868)
623. — Vatica eximia Mia. Sum. (1861) 486; DC.
Prod. 16, 2 (1868) 623; Warp. Ann. 7 (1868)

378. — Vatica sublacunosa Mia. Sum. (1861) 486, 191;
DC. Prod. 16, 2 (1868) 623; Wap. Ann. 7 (1868)
378. — Hopea aspera DE VRIESE, Minyak Tengkaw-
ang (1861) 28. — S. eximia ScHEFF. Nat. Tijd. N. I. 31
(1870) 349; BurcK, Ann. Jard. Bot. Btzg 6 (1887) 218,
incl. var. angustifolia BURCK; KING, J. R. As. Soc.
Beng. Sc. 62, 2 (1893) 121; HEYNeE, Nutt. Pl. ed. 2
(1927) 1116; Sym. Gard. Bull. S. S. 7 (1933) 143, pl.
43. — S. sublacunosa SCHEFF. Nat. Tijd. N. I. 31
(1870) 350; HEYNE, Nutt. Pl. ed. 2 (1927) 1116. —S.
furfuracea (non MiqQ.) BRANDIS, J. Linn. Soc. Bot. 31
(1895) 98, p.p.; SLOOT. ex HEYNE, Nutt. Pl. ed. 1, 3
(1917), 299; Ripe. El. Mal. Pens 91 4@922) e232
p.p. — Fig. 116, 117.

Large tree. Twig, petiole, nervation beneath, leaf
undersurface, midrib above, panicle and parts of
perianth exposed in bud densely persistently pink-
brown scabrid pubescent. Twig to 3 mm @ apically,
stout, terete, becoming smooth; stipule scars cuneate,
c. 2mm long, slightly ascending. Bud to 8 by 6 mm,
ovoid, obtuse. Stipule to 13 by 7 mm, ovate, acute,
prominently nerved, abruptly constricted at base,
subpersistent. Leaves to 10-18 by 3—7 cm, oblong or
narrowly ovate, attenuate, coriaceous; base obtuse;
acumen to 8 mm long, narrow; margin + revolute;
nerves (20—)22-25 pairs, dense, prominent beneath,
curved, set at 55°—70°; tertiary nerves distinct, scalari-
form, at 90° to nerves; midrib prominently terete
beneath, shallowly depressed above; petiole 7-9 mm
long. Panicle to 18 cm long, terminal or axillary,

1982]

DIPTEROCARPACEAE (Ashton)

549

Fig. 117. Shorea ovalis (KORTH.) BL. ssp. ovalis.

terete; singly or doubly regularly branched, branchlets
short, compact, bearing to 8 flowers: bracteoles to 8 by
6 mm, broadly elliptic, obtuse, tightly cupped round
the buds, pink-brown pubescent outside, glabrous
within, falling at anthesis. Flower bud to 5 mm long.
broadly ovoid to globose. Calyx densely golden-
brown pubescent outside, glabrous within: 3 outer
lobes narrowly ovate, acuminate; 2 inner lobes
shorter, subacute. Petals cream with a pink tinge
towards base, ovate, obtuse, densely pubescent on
parts exposed in bud. Stamens 50-70, of varying
lengths, the innermost almost twice as long as the style
and almost as long as the petals; filaments filiform,
much twisted and folded in bud; anthers elliptic to
oblong, the cells rather narrow; appendage to connec-
tive short, vestigial. Ovary and stylopodium narrowly
conical, densely shortly pubescent in the distal half,
glabrous at base; style short, cylindrical, glabrous.
Fruit calyx puberulent; 3 longer lobes to 11 by 1.4m,
spatulate, narrowly obtuse, to 7 mm broad above the
to 12 by 9 mm elliptic shallowly saccate slightly
thickened base; 2 shorter lobes to 6 by 0.4 cm, linear,

Shoot of 3 m high sapling, x 4 (SAN 19446).

similar at base. Nut to 2.2 by 1.3 cm, shortly evenly
densely rufous pubescent; style remnant to 2.5 mm
long, tapering, acute.

Distr. Malesia: Malaya (Penang, E. coast), Singa-
pore. E. Sumatra (Indragiri), Banka, Billiton, Borneo
(E. Sabah and Indonesian Borneo).

Ecol. Scattered in lowland Mixed Dipterocarp
forest, usually in moist places in valleys and low-lying
ground, to 500 m.

Vern. Méranti kepong, kepong labu, k. segar
(Malaya), méranti méerah, m. sabut (Sumatra), lam-
pong, |. rasa, |. mérambung (E. Borneo), damar
salémsung, d. putang, mésélurang (S.E. Borneo),
sébong gunung puteh (E. Kutei), tahan létup (Lower
Dayak).

b. ssp. sarawakensis ASHTON, Gard. Bull. Sing. 20
(1963) 275; Man. Dipt. Brun. (1964) 202, f. 19; ibid.
Suppl. (1968) 111.

Defining characters: Twigs prominently rufous
tufted tomentose, tufts to 3 mm long; leaf nervation
beneath, petiole and panicle shortly sparsely so:

550

FLORA MALESIANA [ser. Lwolege

ea inal ©

Fig. 118. Shorea ovalis (KORTH.) BL. Forest relict tree in Trengganu (Photogr. CORNER, 1935).

is

St

1982]

lamina beneath glabrous, lustrous. Leaves 12-17 by
2-4.5 cm, narrowly oblong, margin prominently
revolute, base obtuse, nerves at 55°-65°.
Distr. Malesia: Borneo (Sarawak and W. Sabah).
Ecol. As ssp. ovalis.

¢. ssp. sericea (DYER) ASHTON, Gard. Bull. Sing. 20
(1963) 255. — S. sericea Dyer, Fl. Br. Ind. | (1874)
306; KING, J. R. As. Soc. Beng. Sc. 62, 2 (1893) 111;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 101, p.p.;
BRANDIS & GILG in E. & P. Pfl. Fam. ed. 1, 3, 6 (1895)
267; Rip. Agr. Bull. Str. & F.M.S. 1 (1901) 58;
HEYng, Nutt. Pl. ed. 1, 3 (1917) 308; Burk. J. Str. Br.
R. As. Soc. 81 (1920) 73, fig.; Foxw. Mal. For. Rec. 1
(1921) 80; MerR. En. Born. (1921) 406; RiDL. Fl. Mal.
Pen. 1 (1922) 223; HEYNE, Nutt. Pl. ed. 2 (1927) 1116;
Foxw. Mal. For. Rec. 3 (1927) 33; SLooT. in Merr. PI.
Elm. Born. (1929) 204; Foxw. Mal. For. Rec. 8 (1930)
23; EDwWArRpbs, Mal. For. Rec. 9 (1931) 147; Foxw.
ibid. 10 (1932) 229; BurK. Dict. (1935) 2022. — S.
fusca Burck, Ann. Jard. Bot. Btzg 6 (1887) 207;
BRANDIS, J. Linn. Soc. Bot. 31 (1895) 83. — S. rigida
BRANDIS in Hook. f. Ic. Pl. (1895) t. 2402; J. Linn. Soc.
Bot. 31 (1895) 102; RipL. Agr. Bull. Str. & F.M.S. 1
(1901) 58; HeEYNE, Nutt. Pl. ed. 1, 3 (1917) 306; Burk.
J. Str. Br. R. As. Soc. 76 (1917) 164, fig.; ibid. 81 (1920)
73; RwLe. Fl. Mal. Pen. 1 (1922) 223; HEYNE, Nutt. Pl.
ed. 2 (1927) 1124; Foxw. Mal. For. Rec. 3 (1927) 33.

Defining characters: Twigs and leaves beneath +
shortly evenly pink-brown pubescent. Leaves 14-22
by 4-10 cm, broadly oblong or obovate, deeply
boat-shaped with the lower surface concave, base
cuneate, nerves at S0°—S5°.

Distr. Malesia: Malaya, S. Sumatra (Angkola,
Bangkinang and Djambi southwards), Banka, Billi-
ton.

Ecol. As ssp. ovalis.

Vern. As ssp. ovalis; also kujung, kalup, kélukung
daun (Sumatra).

Note. This subspecies is known to be tetraploid
and to reproduce largely apomictically through
adventive polyembryony (KAUR c.s., Nature 271,
1978, 440).

Insufficiently known

Flowers are unknown from the following three species
which can therefore not be placed in a section with
certainty.

161. Shorea carapae ASHTON, Gard. Bull. Sing. 22
(1967) 294, pl. 39; Man. Dipt. Brun. Suppl. (1968) 105,
flee

Medium-sized tree with pale flaky bark. Twig, leaf
bud, stipule and petiole persistently densely evenly
buff sericeous, nervation and midrib above sparsely
so. Twig c. 3 by 2 mm @ apically, compressed, drying
rugose at first, becoming terete, smooth; stipule scar
prominent, almost amplexicaul. Bud c. 7 by 2 mm,
lanceolate, acute. Stipule to 25 by 20 mm, oblong-lan-
ceolate, subacute, caducous. Leaves 14-18 by 7-10

DIPTEROCARPACEAE (Ashton)

55]

cm, broadly ovate, elliptic, coriaceous; base obtuse to
cordate; acumen short, broad; nerves 11-13 pairs,
stout, prominent beneath, at 50°—70°; tertiary nerves
slender, densely scalariform, unraised; midrib
applanate to somewhat depressed above, prominent
beneath; petiole 23-35 mm long, terete. Flowers
unknown. Panicle to 15 cm long, terminal or axillary,
terete, smooth or rugulose, densely shortly evenly pale
buff pubescent, singly branched; bracteoles to 16 by 5
mm, lanceolate, acute, densely pubescent outside,
sparsely so within. Fruit calyx and pedicel sparsely
evenly buff pubescent. Pedice/ short, stout. 3 longer
calyx lobes to 7 by 1.5 cm, spatulate, subacute, c. 5mm
broad above the to 6 by 5 mm ovate saccate thickened
base; 2 shorter lobes to 25 by 3 mm, linear, similar at
base. Nut to 10 by 7 mm, ovoid, glabrous, acute.

Distr. Malesia: Borneo (Central Sarawak, W.
Kutei).

Ecol. Volcanic plateaux at 800-1200 m; locally
abundant.

Vern. Awang jangut (Kutei), abang uloh (Kenyah).

Note. Bark and wood characters suggest that this
species belongs to sect. Brachypterae.

162. Shorea furfuracea Mig. Sum. (1861) 488, 191;
WaALp. Ann. 7 (1868) 379; Burck, Ann. Jard. Bot.
Btzg 6 (1887) 219; BRaANbDis, J. Linn. Soc. Bot. 31
(1895) 98, p.p.; HEYNE, Nutt. Pl.’ed. 1, 3 (1917)
299. — S. purpurea DC. Prod. 16, 2 (1868) 632,
sphalm.

Medium-sized or large tree, up to 50 m. Twigs,
buds, stipules, petioles, leaves beneath and midrib
above densely persistently pale chocolate-brown sca-
brid pubescent. Twigs c. 2 mm @ apically, much
branched, terete, becoming smooth. Buds to 3 by 2
mm, ovoid, acute. Stipules to 6 by 4mm, ovate, acute,
fugaceous. Leaves 6-13 by 2.5—6 cm, ovate-oblong,
thinly coriaceous; base obtuse; apex shortly acu-
minate (more prominently so in young trees); nerves
14-17 pairs, slender but prominent beneath, obscurely
depressed above, at 65°—75°, with short intermediate
secondaries in young trees; tertiary nerves densely
scabriform, distinct and elevated beneath; midrib
prominent beneath, evident but + depressed above;
petiole 9-14 mm long, rather slender. Flower and fruit
unknown.

Distr. Malesia: Sumatra (Atjeh, Ophir, Sibolga; P.
Musala).

Ecol. Mixed primary forest, up to 400 m.

Vern. Méranti térutung, P. Musala, m. udang, m.
bunga, Sibolga, habung banio, suantan hambang, Sum.
W. coast, kétrahan silang, Atjeh.

Note. There are similarities in leaves and buds with
S. dasyphylla and S. rugosa of sect. Mutica, but also
with S. scaberrima of sect. Brachypterae.

163. Shorea pallidifolia ASHTON, Gard. Bull. Sing. 22
(1967) 296, pl. 41; Man. Dipt. Brun. Suppl. (1968) 113,
f. 14.

Small to medium-sized tree with V-section fissured
bark. Twigs, buds, stipule, petiole and nervation

552

beneath persistently pale yellowish brown scabrid
pubescent, lamina beneath very shortly densely persis-
tently ocherous scabrid pubescent. Twig c. 6 by 3 mm
@ towards apex, stout, compressed and ribbed at first,
becoming terete; stipule scars c. 2 mm long, horizon-
tal, prominent. Bud to 10 by 8 mm, broadly ovoid-
conical, compressed. Stipule to 7 by 5 mm, oblong,
obtuse. Leaves 13-22 by 7-13 cm, broadly oblong to
ovate or obovate, thickly coriaceous; base obtuse or
subcordate; acumen short; nerves 12-19 pairs, ob-
scure above, prominent beneath, arched, at 110° at the
base, 45°-70° near the apex; tertiary nerves slender,
hardly elevated, densely scalariform; midrib obscure
and depressed above, prominent beneath; petiole
2-3.5 cm long, stout, drying rugose. Panicle to 15 cm
long, compressed and prominently angled, axillary,
rarely terminal; singly or doubly branched, branchlets
to 3 cm long, bearing to 6 distichous flowers; brac-
teoles to 4 by 3 mm, elliptic, obtuse, shortly evenly
pubescent outside, glabrous within, caducous. Flower
bud to 6 by 3 mm, ellipsoid. Sepals pubescent on parts
exposed in bud; outer 3 deltoid, acute, inner 2 ovate,
smaller and thinner at margin than outer 3. Petals
elliptic, pubescent on parts exposed in bud; outer 3
deltoid, acute, inner 2 ovate, smaller and thinner at
margin than outer 3. Stamens unknown. Ovary ovoid,
glabrous; style filiform, glabrous, somewhat longer
than ovary. Fruit pedicel and calyx shortly sparsely
evenly pubescent. Pedicel/ c. 1 mm long and @, small. 3
longer calyx lobes to 7 by 1.5 cm, broadly spatulate,
obtuse, c. 7 mm broad above the to 10 by 8 mm
broadly ovate saccate somewhat thickened base; 2
shorter lobes to 4 by 0.3 cm, linear, similar but smaller
at base. Nut to 12 by 9 mm, ovoid, shortly apiculate,
densely shortly pale buff pubescent.

Distr. Malesia: N.W. Borneo (W. and Central
Sarawak).

Ecol. Rare, on podsols in Heath forest at low
altitude.

Note. The shape of the ovary, as well as fruit, bark
and wood characters, suggested that this species
belongs to sect./subsect. Mutica.

Dubious

Caryolobis indica GAERTN. Fruct. | (1789) 215, t.
IVE

FLORA MALESIANA

[ser. I, vol. 9?

GAERTNER based this genus on a fruit in the Leyden
Botanical Institute, said to have come from Ceylon,
with the vernacular name berelie. It was mostly
reduced to the genus Doona THw.., possibly in part due
to the fact that acommon name for Doona in Ceylon is
beraliya dun. Unfortunately its type could not be
traced in the Leiden carpologica.

The fruit figured seems not to agree with that of
Shorea sect. Doona, for this is characterized by the
extremely unequal cotyledons, one of which is minute
and acicular, whereas the picture shows two equal
cotyledons. The description is too poor, and in
absence of type material unidentifiable in Dipterocar-
paceae.

Excluded

Cotylelobiopsis beccariana HEIM, Rech. Dipt. (1892)
125; BRANDIS, J. Linn. Soc. Bot. 31 (1895) 116; MERR.
En. Born. (1921) 408; SLoor. Bull. Jard. Bot. Btzg III,
10 (1929) 395.

HeEIM based this monotypic genus on a BECCARI
specimen consisting of sterile material only, the
number of which he did not cite. HEM annotated
many of his types at Kew, but BRANDIS, who conjec-
tured that BeccCARI 467 was the type, was clearly
unsure from which it may be guessed that this
specimen, now lost, was not annotated. The Florence
duplicate of this number fits HEIM’s description, and
represents the fallen leaflets of the Leguminosae
Pseudosindora palustris SyM. Proc. Linn. Soc. 155th
sess. (1944) 285; the specimen, mounted with another
of the same species numbered 3468, is quoted by DE
Wit when he made his new combination Copaifera
palustris (SYM.) DE WiT (Webbia 9, 1954, 462). HEIM’s
detailed description of leaf morphology, and espe-
cially petiole anatomy, conforms with that of this
species and he noted that the latter was atypical of
Dipterocarpaceae. HEIM’s genus and species antedates
that of SYMINGTON. Nevertheless the Paris duplicate
of BECcCARI 467, now also lost, was referred to
Cotylelobium melanoxylon (Hook. f.) PIERRE by the
latter (Fl. For. Coch. 3, 1889, t. 235). There must
therefore remain some question as to the real identity
of the Kew duplicate of BECCARI 467, which anyway
cannot with certainty be accepted as the holotype of
Cotylelobiopsis. Until refound this name must be
regarded as doubtful.

ADDENDA, CORRIGENDA ET EMENDANDA
GyiGwG, J. VAN: STEENTS:-cis:

At times colleagues have asked me whether my effort to collect the Addenda, Corrigenda et Emendanda was
worthwhile.

The main purpose is to keep readers up to date with the plants of Malesia in one work and keep them aware
of additions, name changes, efc.

They are also important as a source for plant-geographical purposes, to correct names of useful plants, e/c.

Another facet of keeping up with the records is that they reflect the degree of completeness of collections
at the time of the original revision, and form a certain check on the degree of exploration.

In an overall review of the ‘Floristic inventory of the Tropics: Where do we stand?’ PRANCE has made use
of the Addenda in comparing the state of exploration in the neotropics with that of Africa and Malesia (Ann.
Mo. Bot. Gard. 64, 1977, 657—685, especially p. 671). He found the number of addenda and novelties much
larger in the neotropics than in Malesia, obviously due to a lower, and especially less varied exploration (collec-
ting density). This comparison tends to support my conviction that the bulk of the Malesian species has
become gradually represented in the herbarium.

It was pleasant to experience that the careful keeping on record of the Addenda serves good purposes and
should therefore be continued.

Printing errors have only been corrected if they might give rise to confusion.

Volume and page number are separated by a colon. Page numbers provided with either a or b denote the
left and right columns of a page respectively.

Alismataceae Araliaceae

5: 319, Caldesia parnassifolia (Bassi ex L.) PARL. 9: 39a Osmoxylon lineare (MERR.) PHILIPSON.

320a In Malesia a very rare plant (see map in FI. Substitute in description after ‘the central
Males. I, 5: 322, fig. 3). In New Guinea it branch 4—5 mm long’: bearing an umbel of
was only known from Cape Vogel Peninsu- c. 12 sterile, globose, bacciform flowers, c.
la, but it is now also collected in West New 8 mm @ when dry.
Guinea: Star Mts, Sibil Valley, 1200—1300 Distr. Add: Negros Occidental (PAN-
m (KALKMAN 4188) and in Papua New Gui- CHO 1845).
nea: Kubor Ra., Nona Minj Divide, 1920 m Ecol. Low bushy shrub along river-
(Vink 16512); Morobe Distr., 15 miles west banks. Flowers orange; fruit dark purple or
of Lae, 150 m (HARTLEY 9778) and near black. Cultivated (recently) as an ornamen-
Mumeng, 950 m (W. Mor 166). In the last- tal for its dark green and shiny foliage.
named collection the inflorescence consists Vern. Miagos
of only one whorl of 3 flowers and ater- 9: 67 Arthrophyllum stonei AH-LAN Lin, Mal.
minal flower. For. 43 (1980) 263, f. 1; STONE, Fed. Mus.
J. n.s. 26 (1) (1981) 71, f. ii.
Anacardiaceae (DinGc Hou) Distr. Malesia: Malaya: Pahang-Selan-
gor (STONE 12358, 13754, 14140, KLU
8: 483 Spondias pinnata (L. f.) Kurz. 27353, 30006, 30007).
KOSTERMANS (Quart. J. Taiwan Mus. 34, Ecol. Montane forest, 1000—1800 m,
1981, 108-111) suggested that what was sometimes dwarfed in elfin forest on sum-
named Spondias pinnata from Malesia mit ridges.
(Malaya to New Guinea) would differ from Notes. Said to be intermediate between
the typical S. pinnata from India and repre- A. montanum and A. alternifolium, dis-
sent an undescribed species, S. malayana tinct from the latter by the purple fruit, de-
KOSTERMANS. gree of branching of the inflorescence, and
When writing the paper he could only re- the narrow, thin-coriaceous leaflets.
ly on his memory of former experience and A. montanum and A. alternifolium were
on only three specimens from Malaya. I ob- already distinguished by PHitipson (FI. Ma-
served that in one of them the leaf differ- les. I, 9: 55) by weak characters and the ad-
ence mentioned in his key does not fit. His dition of a third one, with intermediate leaf
experience can only relate to the size of characters is therefore hardly welcome; the
trees and bears no testimony on the flower differentiating characters are also slight.
details mentioned in the key and their geo- Mr. Lm should have given a clear key for
graphical variability, which is in this genus the three species instead of this haphazard
difficult to judge because of the cultivation, description.
domestication, and running wild of Prof. Pumpson (in litt.) finds ‘A. stonei’
cultivated trees. For the present his represents rather larger than usual speci-
delimitation does not seem convincing, es- mens of A. alternifolium.
pecially as there are also discrepanciesinhis 9: 103 Replace: 16. ELEUTHEROCOCCUS
key and descriptions. Maxim. Mém. Ac. Sc. St. Pétersb. Sav. Etr.

(553)

554

FLORA MALESIANA

7: 797a

7: 798a

7: 802b

91(1859)913823'Ss Ya Huse Arn>s Arb516
(1980) 108. — Panax subg. Acanthopanax
DECNE & PLANCH. Rev. Hort. IV, 3 (1854)
105. — Acanthopanax (DECNE & PLANCH.)
H. Witte, Ann. Hort. Bot. 4 (1861) 89;
Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863)
10; Pumpson, Fl. Males. I, 9 (1979) 103.
Note. Unfortunately Acanthopanax has
to give way. The two Malesian species
should be called:
1. Eleutherococcus trifoliatus (L.) S. Y.
Hu, J. Arn. Arb. 61 (1980) 110. — Acan-
thopanax trifoliatus (L.) Voss. Vilmor.
Blumengartn. 1 (1894) 406; Merr. Philip.
J. Sc. 1 (1906) Suppl. 217; Pumreson, FI.
Males. I, 9 (1979) 103.
2. Eleutherococcus malayanus (M. R. HEN-
DERSON) STONE, Mal. For. 43 (1980) 395. —
Acanthopanax malayanus M. R. HENDER-
son, Gard. Bull. S. S. 7 (1933) 105, pl. 22;
Puripson, Fl. Males. I, 9 (1979) 103.

Balanophoraceae

Balanophora elongata BL.

Add to literature: B. C. SToNE, Mal. Nat.
J. 33 (1979) 129, fig.; Fed. Mus. J. n.s. 26
(1) (1981) 72. — B. papuana (non SCHLTR)
SoEPADMO, Nature Malaysiana 3 (1) (1978)
24, with col. illus. — B. hansenii HAMBALI,
Reinwardtia 9 (1980) 425.

Add to Distr.: Malaya: Selangor/Pahang
border, G. Ulu Kali, in dwarf forest on
ridge at 1500-1700 m, on Pentaphylax
auryoides.

Add to Note: According to Dr. B. HANSEN
(in litt.) the new record from Malaya be-
longs undoubtedly to B. elongata to which
it keys out without difficulty in the key in
Fl. Males. I, 7: 793.

Balanophora abbreviata BL.

Add to Distr.: New Hebrides: Mallicolo I.
(N. HALLE RSNH 6351), parasitic on Ficus.
Cf. Ha.ié, Adansonia 17 (1978) 260.
Add to Note: Mr. G.D. AREKAL & G.R.
SHIVAMURTHY (Phytomorph. 26, 1976,
135-138, 6 fig.) have, for the first time,
been able to examine, in situ — since all for-
mer experimental infections have failed —
the germination of the seed of B. ab-
breviata, a magnificent achievement with
these utterly minute seeds. After the break-
ing of the seed coat the endosperm cells
produce 4—8 narrow tubular extensions
coated with a sticky substance adhering to
the fine rootlets for anchorage. Then the
hardly differentiated embryo produces 1—4
tubular processes to form the primary haus-
torium, penetrating into the rootlet and
making contact with the vascular tissue of
the rootlet of the host; a nodule develops,
throwing off the seed coat, and parasitism

8: 137b

8: 138a

8: 14la

8: 148b

8: 159a,
153b

4: 20,
593a

4: 21

[ser. I, vol. 93

is established. The procedure is established
on uninjured rootlets. Naturally it does not
exclude the possibility that infection can al-
so proceed on injured roots or even stem
bases of host plants.

Bignoniaceae

Deplanchea bancana (SCHEFFER) STEEN.
Add to Distr.: Central Celebes (Baolu, near
Palopo; Usu near Malili; near Matana
lake).

Add to Notes: The Celebes specimens were
poor and originally assigned to D. glabra
(STEEN.) STEEN. With abundant recent ade-
quate material from localities nearby, they
appear to belong to D. bancana.

Change in Fig. 11, map: the localities from
Celebes of Deplanchea glabra belong to D.
bancana.

Deplanchea glabra (STEEN.) STEEN.
Delete Celebes from the distribution.

3. Stereospermum colais (Ham. ex DILLW.)
MABBERLEY, Taxon 27 (1978; publ. 1979)
553. — Bignonia colais HAM. ex DILLw. Re-
view of the references to the Hortus Mala-
baricus etc. (1839) 28, based on padri RHEE-
DE, Hort. Malab. 6, t. 26. — S. personatum
(HassK.) CHATTERJEE; STEEN. Fl. Males. I,
8 (1977) 148.

Nomencl.: On the strength of Hamm-
TON’s opinion that this was different from
Bignonia chelonoides L. f., DILLWUN pro-
posed a new name, which antedates that of
HASSKARL.

Fernandoa macroloba (MiqQ.) STEEN.

The mention of this N. Sumatran endemic
tree (as ‘Heterophragma macrolobium’=
Haplophragma macrolobum (MiaQ.)
STEEN.) by FLENLEY & RICHARDS (eds.),
The Krakatao Centenary Expedition, in
Dept. Geogr., Univ. Hull, Misc. ser. 25
(1982) 16, 27, 48, 166 is derived from a mis-
identification; the specimens belong to the
common Radermachera glandulosa (BL.)
Mia.

These two pages have unfortunately be-
come transposed.

Pandorea pandorana (ANDR.) STEEN.
Add to Distr.: Central Celebes (VAN BAL-
Gooy 3915), and change Fig. 38, map, ac-
cordingly.

Burmanniaceae

Gymnosiphon BL.

Add to Distr.: Lesser Sunda Islands: Flores
(ScHMuTz 4802, prob. G. aphyllus BL.).
The genus is not yet reported from the Mo-
luccas and the Philippines.

Thismia GRIFFITH.

Add the synonym: Geomitra Brcc. Malesia

1982]

4: 22

4: 25

Addenda, corrigenda et emendanda

1 (1877) 250.
Add Note. 15. Thismia clavigera seems to
be better accommodated in Thismia sect.
Sarcosiphon (Bui.) JoNKER, from the three
species of which it is distinct by the free cla-
vate inner tepals.
Insert in the key after the second lead of 1:
la. Inner perianth lobes connate at the tips,
forming an erect mitre with large holes.
Continue to 12.
la. Inner perianth lobes connate at the tips,
forming a mitre with large holes, the
lobes continued above the mitre as 3
free, almost erect, filiform, clavate seg-
15. T. clavigera
Insert after 14. Thismia crocea:
15. Thismia clavigera (Becc.) F.v.M. Vict.
Nat. (1890) 235; Pap. & Proc. R. Soc.
Tasm. for 1890 (1891) 235; Stone, Blumea
26 (1980) 420, pl. — Geomitra clavigera
Becc. Malesia 1 (1877) 250, t. 10, f. 1; Jon-
ker, Fl. Males. I, 4 (1948) 25. — Fig. 1.

Fig. 1. Thismia clavigera (BEcc.) F.v.M., x2 (after
BECCAR)).

5: 232a

53.251

5: 275a

5: 276b

5: 2836

555

Distr. Malesia: Malay Peninsula (?Pe-
rak, G. Hiau above Taiping; Langkawi Is.),
N. Sumatra (Gajo Lands), Borneo (Sara-
wak: Lundu Distr.).

Delete 4. Geomitra etc.

Burseraceae (LEENHOUTS)

Dacryodes rugosa (Bu.) H. J. Lam.
Add to Distr.: Nicobar Is., Katchal I. (var.
rugosa).
Dacryodes longifolia (KING) H. J. Lam.
Add to literature: KOCHUMMEN in Whit-
more, Tree Fl. Malaya 1 (1972) 141.
Add to description: Tree up to 40 m with
buttresses up to 3.5 m (cf. KOCHUMMEN,
1972).
Add to Distr.: Throughout the Malay Pen-
insula (KOCHUMMEN, 1972); var. longifolia
also in Borneo.
Insert: Ecol. Usually on coastal hills or
hills which were coastal in the Pleistocene
(KOCHUMMEN, 1972), up to 300 m. Fr.
April.
Santiria grandiflora KALKMAN.
Add to description: Branchlets up to 1.75
cm thick. Leaflets finally glabrous beneath.
Fruits nearly transversely obovate, 2 by
1.25 by 1.5 cm, stigma more than 90° ex-
centric.
Add to Distr.: N. Borneo (Sipitang Distr.,
Mendalong, Lumaku For. Res.).
Add in the key to the species of Canarium,
after the second lead of 6:
6b. Stipules up to 1.5, rarely 2.5 cm long,
not distinctly exceeding the terminal
bud. Branchlets solid. Stamens 6.
7a. Branchlets 1.5—3 cm g; pith thick and
soft.
7A. Petiole terete with the stipules in-
serted at the base. Fruit glabrous
56. C. batjanense
7A. Petiole sharp-angular with the sti-
pules inserted on the angles up to 5
cm from the base. Fruit prickly-
haiey.- «tektites, Beye 43. C. hirsutum
Canarium grandifolium (RiDt.) H. J. Lam.
Add to Distr.: According to KOCHUMMEN
(in Whitmore, Tree Fl. Malaya 1, 1972,
129) also in Borneo (Brunei).
Canarium decumanum GAERTN.
Add to description: Leaflets of the upper
pairs sometimes on the basiscopic side de-
current till the rachis. Fruits up to 12 cm
long.
Add to Distr.: Admiralty Is. (Manus I.).
Add to Ecol.: sago swamp.
Canarium fusco-calycinum RIDL.
Add to description: Tree up to 21 m high
and 60 cm g. Infructescences c. 15—30 cm
long with few short oblique-erect branches
and with few fruits; calyx funnel-shaped,

556

FLORA MALESIANA

5: 290a

5: 290b,
291a

537296;
6: 928

7: 822a

6: 118

trigonous, c. 1 cm diam. Fruits (immature)
narrowly oblong, tapering to both ends, tri-
gonous in section, 3.5 by 1.25 cm, glabrous;
in section the kernel with flat sides and
slightly rounded angles, the lids with a very
faint median rib, lids c. 1.5 mm thick. Seeds
2, the sterile cell moderately reduced.
Add to Ecol.: On clay, up to 340 m. Fi.
Aug.
Canarium rigidum (Bt.) Mia.
Add to Distr.: Moluccas (Obi I.).
Canarium cestracion LEENH.
Add to description: Twigs up to 11 mm @.
Stipules linear, tapering to an acute point,
4 by 1 mm. Leaves up to 8-jugate; leaflets
from 2.5 cm broad on; acumen sometimes
long and slender; nerves up to 17 pairs.
Fruits up to 3 cm high and 1.75 cm broad.
Add to Distr.: Milne Bay Distr.
Insert after 55. Canarium pimela:
56. Canarium batjanense LEENH. Blumea
27 (1981) 211.

Distr. Malesia: Moluccas: Batjan I. (Mt
Sibela).

Ecol. Canopy or subcanopy tree in pri-
mary forest, 250—1050 m. Fi. fr. Oct.

Note. The present species seems to be
allied with C. acutifolium MERR., C. balsa-
miferum WILLD., and C. oleosum ENGL.,
which is also well in accordance with its
geographical position (cf. LEENHOUTS, Blu-
mea 9, 1959, 317, f. 9). It seems nearest to
C. balsamiferum, which differs by the ab-
sence of stipules, the far stronger reduced
pistil in the male flower, and the fruit cells
that are all three well developed. The only
moderately reduced pistillode is well in ac-
cordance with C. oleosum, different in
both other points mentioned under C. bal-
samiferum, however, and moreover in the
much smaller, nearly ovoid fruits. Vegeta-
tively, C. acutifolium and C. batjanense are
hardly separable but for the longer and
more persistent stipules of the former; C.
acutifolium differs, however, in the strong-
ly reduced pistillode and in the smaller and
more globular fruits.
Dacryodes multijuga LEENH. Add:

Distr. South Sumatra (ForBEs 3073, L),
fruits only. Hitherto only known from one
collection in Malaya.

Campanulaceae

Insert after 4. Wahlenbergia marginata:
5. Wahlenbergia papuana v. Royen, Bot.
J. Linn. Soc. 77 (1978) 121, f. 2.
Branched dwarf herb, prostrate. Twigs,
leaves and calyx hairy. Leaves at end of
twiglets, narrow-oblanceolate, 8-10 mm
long. Filaments glabrous, not widened at
base, thus different from both W. confusa

6: 122,
928

6: 136,
928

4: 175

4: 176

4: 19la

4: 192a

4: 194

[ser. I, vol. 93

and W. marginata.

Distr. Malesia: Papua New Guinea:
along bank leading from Iswan swamp to
Koma Creek (Mt Victoria), c. 2650 m. Fi.
fr. May.

Note. According to the author closely
related to the glabrous New Zealand species
W. albomarginata Hook. f.

P. vAN RoyEN has provided a new key to
the New Guinea species of Lobelia. Cf.
Bot. J. Linn. Soc. 77 (1978) 120.
Add to the species of Lobelia:
Lobelia victoriensis v. ROYEN,
Linn. Soc. 77 (1978) 118, f. 1.

Stemless dwarf, 7-20 mm high, with a
single central flower in a rosette of ovate
leaves. Corolla isomerous, without a dorsal
slit, lobes 5, equal, filiform, long.

Distr. Malesia: Papua New Guinea: Is-
wan swamp on Mt Victoria, 2660 m alt. Fi.
Sr. May.

Bota:

Caprifoliaceae

Add to the family description: Leaves very

rarely scattered or in pseudowhorls (sub-

fam. Alseuosmoideae).

Insert in the key after the second lead of 1:

la. Leaves scattered or in pseudowhorls
5. Periomphale

la. Leaves decussate. Continue to 2.

Sambucus javanica Bu.

Add to literature: STEEN. Blumea 24 (1978)

479.

Add to Distr.: West New Guinea: Arfak

Mts, Mt Lensemoi, 1850 m, BW 12657;

Wissel Lake region, EyMa 4618; Papua

New Guinea: Morobe Distr., Sattelberg,

Timbe R., 1500 m, CLEMENS 7793.

Add to Notes: These records fill the gap of

the generic range in East Malesia, two spe-

cies being recorded from Australia, which

are discussed in VAN STEENIS, /.c.

Insert after 4. Carlemannia:

5. PERIOMPHALE

BaILL. Bull. Mens. Soc. Linn. Paris 1 (1888)

731; STEEN. Blumea 24 (1978) 480. — Pa-

chydiscus GitG & SCHLTR, Bot. Jahrb. 39

(1906) 270. — Memecylanthes Guc &

SCHLTR, /.c. 269.

Shrublets. Leaves scattered or in pseudo-
whorls, entire or with a few faint teeth to-
wards the apex. Flowers actinomorphic,
5-merous, fascicled or solitary axillary.
Ovary inferior, 2-celled, with rather few
(1—4) ovules on the septum. Corolla gamo-
phyllous, somewhat barrel-shaped. Sta-
mens 5, free.

Distr. New Caledonia (4 spp.), Papua
New Guinea (1 sp.).

Notes. This genus belongs to the sub-
family Alseuosmoideae, best known from

1982]

Addenda, corrigenda et emendanda

557

6: 233

6: 243

6: 245

6: 420

the small genus A/seuosmia A. CuUNN. from
New Zealand.

There is no unanimity of opinion about
its taxonomic place. Sometimes it is treated
as a separate family, but mostly it is assign-
ed an isolated place in Caprifoliaceae, to
which I agree. This is another New Caledo-
nian genus now turned up in New Guinea.
A third, yet undescribed monotypic genus
of the subfamily occurs in Queensland (VAN
STEENIS, /.c.).

1. Periomphale papuana STEEN. Blumea 24
(1978) 481.

Glabrous, epiphytic shrublet, c. 75 cm;
stem and branches very slender. Leaves
scattered and in pseudowhorls, lanceola-
te-oblong, 15—25 by 6—8 mm, cuneate at
base, acute at apex, margin entire or mostly
with | or 2 short gland-tipped teeth; nerves
2-3 pairs; very erect; venation impressed
above. Petiole 3—4 mm. Flowers solitary;
pedicels pink, 1—2 mm. Calyx lobes 5,
thickish, blunt-deltoid, 1.25 by 0.5 mm.
Corolla narrowly barrel-shaped, 6 mm
long, pinkish light green; lobes 1 mm, ca-
runculate inside apex. Stamens 5, free, al-
ternipetalous. Ovules 4—5 in each cell, at-
tached to the septum, flattish. Style colum-
nar, as long as the corolla; stigma globular,
rugose.

Distr. Malesia: Papua New Guinea:
West Sepik Distr., Telefomin Subdistr.,
3000 m alt., in Podocarpus-Phyllocladus
woodland with Gahnia tussocks dominat-
ing the undergrowth (LAE 670687).

Celastraceae (DinG Hou)

Celastrus hindsii Btu. Cf. STONE, Mal.
For. 43 (1980) 244.

This species was not treated in the paper,
but its occurrence in Malaya is mentioned
twice in the summary. The specimen in
question (STONE 14039) was misidentified
and belongs to the common C. monosper-
moides LOES.

Xylonymus KALKMAN.

Change in description: Flowers 4—5-mer-
ous.

Xylonymus versteeghii KALKMAN.

Add to Distr.: West New Guinea: Dalman,
Nabire, KANEHTRA & HatTusma 12241;
Darmi Distr., BW 9317. Moluccas: Obi I.,
DE VOGEL 4347.

Salacia kalahiensis KOoRTH.

Hitherto known from the Philippines, Bor-
neo and Java. Now also from the Lesser
Sunda Is.: W. Flores (Paku, 400 m,
ScuMutz 4523; vern.: wase mantur).

4: 104

5: 495

Chenopodiaceae

Arthrocnemum Mog.
After along and careful study of Australian
Salicornieae P. G. WiLson has concluded
that the concept of the genus Arthrocne-
mum is polymorphous, and that the Male-
sian species attributed to it (A. indicum)
should belong to a new genus, Halosarcia,
differing from Arthrocnemum in sclereids
and the adaxial stamen which he finds of
fundamental importance; he suggested that
the two genera are not closely related.

Within Halosarcia indica WiLson distin-
guishes four subspecies, two of which are
endemic to Australia and two others occur
also outside Australia, the type ssp. indica
also in India and Tanzania, and a new sub-
species in South Malesia.

As ssp. indica might also be found in Ma-
lesia, I extract WiILson’s key.

Halosarcia indica (WILLD.) P. G. WiLson,
Nuytsia 3 (1980) 63. — Arthrocnemum in-
dicum (WILtD.) Moa.: BACKER, FI. Males.
I, 4 (1949) 104.

KEY TO THE SUBSPECIES

1. Plant decumbent or prostrate. Articles
of branches corky with age, + trun-
Cate eGnlinees Asse Fe ssp. indica

1. Plant + erect. Cortical tissue of articles
shrivelling with age, often lobed, + ci-
lidlate’s32 22/4. Se see ssp. leiostachya

ssp. leiostachya (BTH.) P. G. WILSON,
Nuytsia 3 (1980) 66. — Arthrocnemum ci-
liolatum BUNGE ex UNG.-STERNB. Versuch
Syst. Salicorneen 69 (1866); UNG.-STERNB.
D. Atti Congr. Int. Bot. Firenze 1874
(1876) 283, based on specimens from Java
and Lesser Sunda Is.; A. J. Scott, Bot. J.
Linn. Soc. 75 (1977) 370. — Salicornia leio-
stachya BTH. Fl. Austr. 5 (1870) 203. — Ar-
throcnemum leiostachya (BTH.) PAULSEN,
Dansk Bot. Ark. 2 (8) (1918) 61.

Distr. Australia, South Malesia: north-
coast of Java, incl. Madura and Kangean
Is., Lesser Sunda Is. (Sumba, Sumbawa,
Timor).

Connaraceae (LEENHOUTS)

Anatomy. W. C. Dicxison (J. Elisha
Mitchell Sc. Soc. 87, 1971, 77—86; ibid. 88,
1972, 120—136; ibid. 89, 1973, 121—138)
concluded that Connaraceae are distinctly
allied with Leguminosae and Rosaceae. His
conclusions regarding relationships within
the family deserve a more critical attitude,
however, as at that time the author was in-

558

FLORA MALESIANA

5: 504a

5: 507b

5: 520a

5: 524b

33526

5: 528

5: 538b

4: 485a

sufficiently aware of the difficulties of in-
terpreting taxonomy, especially regarding
synonymy; he was too much inclined to
take identifications of herbarium speci-
mens for granted.

Palynology. W. C. Dickson, Pollen et
Spores 21 (1979) 31—79. In this extensive
survey of the pollen of the Connaraceae,
the author avoided the mistakes made in his
earlier anatomical papers. This makes his
conclusions of greater importance. The tax-
Onomic opinions brought forward in the
Flora Malesiana revision of this family are
to a high degree confirmed.

Agelaea trinervis (LLANOS) MERR.

Add to Distr.: Hainan (cf. ANon., Fl. Hai-
nan 3, 1974, 113).

Roureopsis asplenifolia SCHELLENB.

Add to Distr.: Borneo.

Rourea prainiana TALBOT.

Add to Distr.: Northern Thailand (cf. V1-
DAL, Fl. Thailand 2, 1972, 124).
Ellipanthus beccarii PIERRE.

Add to Distr.: Sumatra (var. beccarii).
Correct the key to the species of Connarus
as follows:

second lead of 18: ...... endocarp sparse-
ly to densely non-glandular pubescent.
couplet 19: delete ‘Sepals blunt’ versus ‘Se-
pals acute’.

Insert after 2. Connarus euphlebius:

2a. Connarus impressinervis STONE, Mal.
For. 43 (1980) 255, fig.

Differs from C. euphlebius MERR. as fol-
lows: Branches and leaves glabrous. Petio-
lules 0.8 cm. Nerves 5—6(—8) pairs. Petals
(under the fruit) outside appressed short-
hairy. Beak of fruit faint, lateral, slightly
below the apex; pericarp outside glabrous,
inside with a few scattered hairs.

Distr. Malesia: Borneo (Sarawak; S
14731, S 18903, S 24232, S 24715, S 28417).

Note. Already in 1962 I studied the type
material on which this new species is based
and found that it is allied to C. euphlebius,
but differed in several points and seemed to
represent a new species. I pointed out the
differences cited above (in sched.). Because
floral characters are important in the
genus, I refrained from describing it, all
material hitherto known being in fruit.
Connarus monocarpus L. ssp. malayensis
LEENH.

Add to Distr.: SE. Thailand (cf. VAL, FI.
Thailand 2, 1972, 129).

Convolvulaceae

Change 36. Ipomoea illustris etc. as fol-
lows:

36. Ipomoea campanulata LINNE, Sp. PI.
(1753) 160; Moon, Cat. Pl. Ceyl. (1824) 14;

4: 485b

4: 487a

4: 197

fsér.. 1. voleg?

TRIMEN Handb. Fl. Ceyl. 3 (1895) 221;
AUSTIN, POWELL & NICOLSON, Brittonia 30
(1978) 196. — Adamboe RHEEDE, Hort.
Mal. 11 (1692) 115, t. 56 (lectotype). — J.
illustris (CLARKE) PRAIN, Beng. Pl. 2 (1903)
735; OoststR. Fl. Males. I, 4 (1953) 485.

Note. Apparently Adamboe RHEEDE
does not belong to Stictocardia—as VAN
OosTsTROOM hesitantly assumed — and be-
comes the type through the new typifica-
tion. The name change is a nuisance, but
has to be accepted.
Change 37. Ipomoea crassicaulis etc. as fol-
lows:
37. Ipomoea carnea Jacq. ssp. fistulosa
(Mart. ex Cuolsy, in DC.) D. AusTIN,
Taxon 26 (1977) 237. — I. fistulosa MART.
ex CuHotsy in DC. Prod. 9 (Jan. 1845) 349.
— Batatas crassicaulis BTH. Voy. Sulph. 5
(June 1845) 134. — J. crassicaulis (BTH.) B.
L. Ropinson, Proc. Am. Ac. Sc. 51 (1916)
530; Ooststr. Fl. Males. I, 4 (1953) 485.

Note. AustTIN has convincingly shown
that the differences between /. carnea and I.
fistulosa do not merit specific distinction.
As the two forms, which are distinct by mi-
nor vegetative characters, grow in South
America largely geographically isolated (al-
lopatric), a subspecific rank (as a race)
seems the best disposition.
Change 40. Ipomoea tuba etc. as follows:
40. Ipomoea macrantha R. & S. Syst. Veg.
4 (1819) 451; PowELL, NIcoLtson & AUSTIN,
Brittonia 30 (1978) 201. — JI. tuba
(SCHLECHTEND.) G. Don, Gen. Syst. 4
(1838) 271; Ooststr. Fl. Males. I, 4 (1953)
487.

Note. An unfortunate but unavoidable
name change of a pantropical species.

Crassulaceae (H. OuBa, Tokyo)

Insert the key and replace the species de-
scription under the genus Sedum L. by the
following new treatment:

KEY TO THE SPECIES

1. Sepals free.

2. Flowers 4-merous, sepals narrowly ob-
long-spathulate. Styles very short (less
than 0.3 mm). Flowering stems simple.
Cauline leaves spathulate

1. S. erythrospermum ssp. australe

2. Flowers 5-merous, sepals spathulate (to
oblong). Styles long (0.9—1.2 mm).
Flowering stems tri- (rarely bi-)furcate.
Cauline leaves spathulate to broadly
ObOVatelma. ceases 2. S. formosanum

1. Sepals connate for c. 0.5 mm. Flowers
5-merous, sepals linear-lanceolate to
very narrowly oblong. Style long (c. 1.2

1982]

Addenda, corrigenda et emendanda 559

mm). Flowering stems with a short ster-
ile shoot or simple. Cauline leaves oblan-
ceolate to narrowly obovate

3. S. parvisepalum ssp. philippinense

1. Sedum erythrospermum Hayarta, Ic. PI.
Formos. 3 (1913) 110.

ssp. australe MerR. H. Ona, J. Jap. Bot.
52 (1977) 322. — S. australe MERR. Gov.
Lab. Publ. Philip. 29 (1905) 16, non RosE
(1903); Philip. J. Sc. 5 (1910) Bot. 350,
p.p.; En. Philip. 2 (1923) 217, p.p. — S.
ambiflorum R. T. CLAUSEN, Cact. Succ. J.
18 (1946) 58; Backer, FI. Males. I, 4 (1951)
197, versim., p.p. — Fig. 2a—d.

Ae ‘Ade

Fig. 2. Sedum erythrospermum HAYATAssp. australe
H. Oupa. a. Petal with stamen, b. sepal, c. nectar
scale, d. ovaries. — S. parvisepalum YAMAMOTO Ssp.
Dhilippinense H. OuBa.e. Petal with stamen, f. calyx
lobe, g. nectar scale, h. ovaries. All x 5 (a—d ELMER
6568, e—~h Munt 5644).

Succulent, glabrous, perennial herb.
Roots thin. Flowering stems annual, 5—10
cm long, 1.5—2 mm g, simple, erect from
nearly decumbent base, smooth. Leaves al-
ternate (rarely opposite or ternate), remote-
ly arranged, sessile, shortly spurred (the
spur with a round-truncate apex, c. 0.5 mm
long), entire, spathulate, apex round or ob-
tuse, base long attenuate, 7—18(—27) by
2—5(—11) mm, _ thick-herbaceous, flat,
smooth, spreading or ascending. /nflores-
cences terminal, a ternate bracteate cyme.
Bracts leafy, spathulate to obovate.
Flowers mostly 4-merous, erect, sessile.
Sepals free, narrowly oblong-spathulate,
base spurred (the spur with round-truncate
apex, 0.2 mm long), apex round to nearly
truncate, somewhat broadening towards
the base, entire, somewhat unequal in size,
2-4 by 0.8—1 mm, more or less fleshy,
green, smooth, suberect throughout, per-
sistent at fructescence. Petals bright yellow,
connate c. 0.3 mm from the base, lanceo-
late, acuminate, c. 4mm by 1.2 mm, widely
spreading at anthesis. Stamens 8, shorter

than the petals, erect at anthesis; filaments
filiform, 3—4 mm, oppositipetalous ones c.
1 mm from the base connate with the petal.
Nectar-scales flat, narrowly oblong, 0.8 by
0.3 mm. Gynoecium 3—4 mm long, the
ovaries basally for c. 0.5 mm connate, ven-
trally gibbose, dorsally round, suberect, 1.5
mm wide at the middle, style very short, less
than 0.3 mm. Ovules c. 20 in each locule.

Distr. Malesia: Philippines (Luzon,
Benguet Prov.: Mt Santo Tomas, ELMER
6568 = PNH 114365). The ssp. erythrosper-
mum occurs in Formosa.

Ecol. Onledges and gravelly hillside near
the summit of mountain, 2000—2200 m.

Note. Ssp. erythrospermum differs
from ssp. australe in having 5-merous flo-
wers, narrowly oblong-oblanceolate or lin-
ear sepals, subulate or narrowly lanceolate
petals, and usually trifurcate flowering
stems.

2. Sedum formosanum N. E. Brown,
Gard. Chron. n.s. 24 (1885) 134; ForBEs &
HEMSLEY, J. Linn. Soc. 26 (1888) 285; Ha-
YATA, Ic. Pl. Formos. 2 (1912) 12; PRAE-
GER, Not. R. Bot. Gard. Edinb. 13 (1921)
83; J. R. Hort. Soc. 46 (1921) 295; R.-Ha-
MET, Candollea 4 (1929) 32; BERGERin E. &
P. Nat. Pfl. Fam. ed. 2, 18a (1930) 460;
Frop. Act. Hort. Gothob. 6 (1931) app. 97,
f. 778—784, t. 62; OHw1, FI. Jap. Engl. ed.
(1965) 497; Hatus. Mem. Fac. Agr. Kago-
shima Univ. 5 (1966) 31; Fl. Ryukyus (1971)
301; Moranin Walker, Fl. Okinawa (1976)
508; H. Ousa, J. Jap. Bot. 52 (1977) 322.
— §. mariae R.-HAMET in Fedde, Rep. 8
(1910) 143.

Succulent, glabrous, perennial herb, up
to 25 cm high. Flowering stems erect from
a creeping or procumbent base, once or
twice tri- (rarely bi-)furcate, terete, smooth.
Leaves alternate, remotely arranged, ses-
sile, very shortly spurred (the spur less than
1 mm), entire, spathulate to broadly obo-
vate, apex round, base long-attenuate,
1.5—3.5 by 0.6—1.6 cm, fleshy, bright green
above, paler below, smooth. /nflorescences
terminal, a ternate, bracteate, loosely
many-flowered cyme. Bracts leafy. Flowers
5-merous, sessile, 6—9 mm at anthesis. Sep-
als free, very shortly spurred (c. 0.3 mm),
entire, spathulate or rarely oblong, apex
round, somewhat broadening towards the
base, nearly equal in size, 2.8—3.7 by
0.8—1.3 mm, fleshy, ascending at anthesis.
Petals bright yellow, basally connate for c.
0.5 mm, lanceolate to oblong-lanceolate,
acute, 5.2—6.2 by 1.3—1.6 mm, nearly erect
or ascending at anthesis. Stamens 10, shor-
ter than the petal, nearly erect at anthesis;
filaments filiform, c. 4.5 mm, the oppositi-

FLORA MALESIANA

petalous ones for c. 0.8 mm connate with
the petal, anthers oblong, c. 0.4 mm long,
deep yellow before dehiscence. Nectar-
scales broadly oblong-obovate to broadly
oblong, c. 0.5 by c. 0.4 mm, creamy white,
flattish. Gynoecium 5.6—6.2 mm long, the
ovaries for c. 1.5 mm connate, ventrally
gibbose just above the ventral connection,
dorsally round, erect throughout, c. 1.6
mm wide at the middle part, abruptly nar-
rowing near the apex; style 0.9-1.2 mm,
slender, tapering towards the apex; stigma
lowly papillate, yellowish. Ovules 20—24 in
each locule. Follicles 6—7 mm long, whit-
ish, obovate, carpels not spreading. Seeds
oblong, c. 0.7 mm long, testa brown, very
minutely puncticulate throughout.

Distr. Formosa and Japan (Ryukyu and
Kyushu), in Malesia: N. Philippines (Batan
I.: Hatusma & SATO 28624).

Ecol. On rocks at the shore, rarely in-
land. F/. June—August.

3. Sedum parvisepalum YAMAMOTO, Suppl.
Ic. Pl. Formos. 2 (1926) 22, f. 14.

ssp. philippinense H. Ousa, J. Jap. Bot. 52
(1977) 323. — S. australe (non MERR.)
STEEN. Bull. Jard. Bot. Btzg III, 13 (1934)
195. — S. ambiflorum (non R. T. CLAUSEN)
BACKER, FI. Males. I, 4 (1951) 197, p.p.—
Fig. 2e—h.

Succulent, glabrous, perennial herb, up
to 15 cm high. Flowering stems ascending
or erect from creeping, sometimes branch-
ed base, with a short sterile shoot or simple,
terete, smooth. Leaves alternate, remotely
arranged, sessile, very shortly spurred (c.
0.5 mm), entire, narrowly obovate to ob-
lanceolate, apex obtuse, base attenuate,
0.9-1.7 by 4-6 mm, thick-herbaceous,
smooth. Inflorescences a terminal, ternate
cyme, densely 30—50-flowered, sparsely
bracteate. Bracts leafy. Flowers 5-merous,
sessile, c. 10 mm wide at anthesis. Sepals
basally for c. 0.5 mm connate, linear-lan-
ceolate to very narrowly oblong, entire,
slightly unequal in size, 2.5—3.5 by 0.5—0.7
mm, apex round, slightly broadening to-
wards the base, base spurless, fleshy,
ascending or spreading through anthesis.
Petals bright yellow, basally for c. 0.5 mm
connate, lanceolate, apex acute to acumi-
nate-acute, S—6.5 by 0.8—1 mm, spreading
at anthesis. Stamens shorter than the
petals, filaments 3.5—3.7 mm, the oppositi-
petalous ones for c. 1 mm connate with the
petals, anthers ovate, c. 0.6 mm long, red-
dish before dehiscence. Nectar-scales
broadly oblong to square, c. 0.5 by 0.3 mm.
Gynoecium 4—5S mm long, the ovaries for
0.7 mm connate, ventrally gibbose, dorsally
round, c. 1 mm wide, tapering towards the

7: 468

7: 471a

7: 474a

7: 522a

7: 567a

5: 567b

[ser. I, vol. 93

apex from the middle, style c. 1.2 mm, slen-
der, stigma not papillate. Ovules (20—) 24
(—34) in each locule. Follicles brownish,
4—5 byc. 2.5 mm, carpels widely spreading.
Seeds brownish red, oblong-cylindrical, 0.7
mm long, apex round, testa very minutely
puncticulate.

Distr. Malesia: Philippines (Luzon,
Benguet Prov.: BS 4279, 4451, 5373, 5644,
MERRILL 4861, VAN STEENIS 17950, Wi-
LIAMS 117). The ssp. parvisepalum occurs in
Formosa.

Ecol. On rocks or boulders along steep
slopes, c. 300—1700 m.

Note. Ssp. philippinense is distinguish-
ed from the Formosan ssp. parvisepalum
by the narrowly obovate to oblanceolate
leaves, shorter flowering stems, and longer
petals. Ssp. parvisepalum has linear-oblan-
ceolate leaves, flowering stems up to 25 cm
long, and petals attaining 7.5 mm long.
Ssp. philippinense differs from the Philip-
pine S. erythrospermum ssp. australe by the
5-merous flowers, the ovaries with long
style, the broadly connate, linear-lanceo-
late sepals, and the narrowly obovate to
oblanceolate cauline leaves.

Cyperaceae

Add to Fig. 10, map of Mapania: Lesser
Sunda Islands: Flores %

Mapania macrocephala (GaupicH.) K.
SCH.

Add to Distr.: Lesser Sunda Is. (W. Flores:
Paku, Wae Meleng, 700 m, damp stream-
bed in rain-forest, ScHMUTZ 4913).

Add to Notes: The species was known from
the S. Philippines and N.—Central Moluc-
cas eastwards; Flores is an important exten-
sion westwards. The nuts are on the small
side, some 4—4.5 by 3 mm and pearl grey.
Mapania cuspidata (MiqQ.) UITTIEN.

Add to Distr.: Lesser Sunda Is. (W. Flores:
Paku, 500 m, plant 1 m, locally gregarious,
in rain-forest, SCHMUTZ 4816).

Add to Notes: The widest distributed spe-
cies of the genus, of which KERN(FI. Males.
I, 7, 1974, 466) expressly stated that it lack-
ed in the Lesser Sunda Islands, the gap now
being filled.

Lipocarpha chinensis (OsB.) KERN.

Add to Distr.: Lesser Sunda Is. (Flores:
VELDKAMP 7102).

Fimbristylis eragrostis (NEES) HANCE.
Add to literature: VELDKAMP, Reinwardtia
10 (1982) 26.

Add to Distr.: Australia: Northern Terri-
tory (Arnhem Land: P. K. Latz 2836).
Fimbristylis fusca (NEES) CLARKE.

Add to literature: VELDKAMpP, Reinwardtia
10 (1982) 26.

1982]

7: 60565

7: 687a

9: 1494

9: 164a

82239

9: 242

9: 246

9: 246

Addenda, corrigenda et emendanda

Add to Distr.: Australia, Northern Terri-
tory (Katherine Gorge: DuN Lop 3733). A
new record for Australia.

Cyperus esculentus LINNE.

Add to literature: EVERAARTS, Weeds Vege-
tables Java (1981) 79. KERN knew this wide-
ly distributed species only from a single
‘suspected’ collection. This suspicion is
now removed by Mr. A. P. EVERAARTs in
his study on weeds of Java. He collected
this species in fields (with vegetables) near
Lembang and Pengalengan in West Java
and on Mt Tengger (Tosari, Ngadisari) in
East Java at 1300—1400 m.

Cyperus compactus RETZ.

Add to literature: K. L. Wmson, Telopea 1
(1980) 462.

Add to Distr.: Australia: Northern Terri-
tory.

Oreobolus ktikenthalii STEEN.

Add to literature: STEEN. Reinwardtia 10
(1982) 26.

Add to Distr.: NW. Borneo: Sarawak (4th
Div., N. side of Mt Murud), and dot this lo-
cality in Fig. 85.

Add to Ecol.: Forming dense tufts in wet
rock holes, 2100 m, Burtt & MARTIN 5482.
Add Note: Hitherto only known from N.
Sumatra and Malaya; Mt Kinabalu is the
westernmost locality of another species, O.
ambiguus KUK & STEEN.

Carex breviscapa CLARKE.

Add to Distr.: Central Celebes (Mt Lokila-
laki: W. MEUER 9876).

Add to Ecol.: 1700—2200 m alt.

Carex oedorrhampha NELMES.

Add to Distr.: Central Celebes (Mt Lokila-
laki: W. MEDER 9869).

Dipterocarpaceae ( ASHTON)

Line 16 from top: Cotylelobium has 5 spp.,
not 6.

Paragraph 5: replace 2nd and 3rd sentence
by: It is not impossible that they were de-
rived from the Indian subcontinent, as ac-
cording to geophysical theory this rafted
block of land had by the Eocene joined the
Asian plate, while, moreover, Dipterocar-
pus could have migrated through S.W. Asia
before it became arid in the Miocene.
Paragraph 2, lines 10—12: ssp. philippinen-
sis also in S.E. Borneo; the doubtful sterile
specimens belong to ssp. philippinensis.
Line 16 from top: change SAsAH into Sasa-
KI.

Change the last two lines from bottom into:
and its intensity would tend to inhibit vec-
tor numbers from reaching adequate levels
for effective pollination. CHAN (1980)
found most species to have high self-incom-
patibility, but apomictic adventive em-

Ay |

: 267

: 269

S212

AOA Be

: 276

APA bey

: 284

: 290

: 293

: 304a

: 310a

: 339a

: 341

: 342b

: 344

: 35Sa

561

bryony occurs in several species (see p. 263,
270).

Line 13 from top: change SAsaHinto Sasa-
KI.

Paragraph ‘Morphology’, line 2, first
word: change cylindrical into circular.
Paragraph 3, line 4, add after ‘species lev-
el.’: Dipterocarpoideae differ from other
subfamilies in the presence of resin canals
and in their multiseriate rays.

Paragraph 2, last line, add between the
brackets: SomEGo, 1978.

Line 16 from top: change ‘hypochroa’ into
hypochra.

Line 15 from bottom: change DC. into
KURZ).

Line 4 from bottom: change Dyer into
KING.

Line 4 from top: change Mia. into BL.
Line 7 from bottom: omit ‘Balanocarpus’.
Line 5 from bottom: change ‘flora’ into flo-
ral.

Paragraph 2, lines 8 & 9, change sentence
into: The mature fruit of Pakaraimaea is
small, capsular, but the method of germi-
nation is unknown.

Substitute paragraph 5 by: A picture thus
emerges of the Asiatic subfamily originat-
ing in Central Gondwanaland and migrat-
ing eastwards. It appears likely that the
family originated in the seasonal tropics;
subsequent immigration and rapid diversi-
fication in the oceanic climates of S.W.
Ceylon, West Malesia and New Guinea be-
ing accompanied by a reduction in stamen
number and pollen production, and tomen-
tum; and the evolution of a thin-walled and
poorly dispersed fruit.

Line 6 from bottom: change ‘sort’ into
short.

Line 5 from top: replace ‘D. apterus’ by D.
validus.

Paragraph 5, line 3: change ‘but’ into by.
Dipterocarpus baudii KortTH.

Line 11 from top: change ‘D. elongatus
Kortu.’ by D. validus BL.

Dipterocarpus kunstleri KiNG.

Line 16 from top: change ‘D. elongatus
Kortu.’ by D. validus Bt.

In caption Fig. 35 replace ‘place’ by tree,
and add: The collector Musau indicates
scale.

Cotylelobium PIERRE.

Paragraph 2, after ‘Distr.’, change 6 into 5.
Cotylelobium melanoxylon (Hoox. /f.)
PIERRE.

Delete the Note.

In caption Fig. 41: b belongs to Vatica um-
bonata (Hook. f.) BURCK ssp. acrocarpa
(SLoot.) ASHTON, d—e to ssp. umbonata.
Vatica sarawakensis HEM.

Line 11 from top: change ‘V. oblongifolia’

FLORA MALESIANA

562
into V. sarawakensis.

9: 365a_ Vatica mangachapoi BLco ssp. mangacha-
poi.

Under ‘Ecol.’ line 3: change ‘confirmed’ in-
to confined.

9: 404b Hopea latifolia Sym.

In ‘Vern.’ change ‘jongkang’ into jang-
kang.

9: 405b Hopea pierrei HANCE.

Add to Distr.: Sumatra (N.W. coast).

9: 424a Hopea plagata (BLco) VIDAL.

Delete the Note.

9: 429 In Note under Subsection Pierrea, line 1,
change ‘The New Guinea species’ into
Some New Guinea species.

9: 436b Hopea siranda Mia. under ‘Excluded’,
change ‘489’ into 491.

9: 437 Shorea Roxs. ex GAERTN. f.

Under ‘Uses’, line 4, change ‘timber ven-
eer’ into veneer timber.

9: 448b Shorea guiso (Bico) BL., in synonymy:
Lines 1 & 2 from top: change ‘263’ into 45.

9: 459b Shorea superba Sym.

Under ‘Distr.’, line 2, read: Sabah, Tidung,
Berau; Sampit, sterile coll.).

9: 460b Shorea astylosa Foxw.

Delete the third paragraph of the Notes.

9: 472 Line 2 from top in text: change ‘sepalled’
into sepals.

9: 483b Shorea kudatensis Woop ex MEIER.

In Distr. change ‘Kilias’ into Klias.

9: 487a Shorea dealbata Foxw.

Under ‘Vern.’ change ‘bunbong’ into bum-
bong.

9: 491b Shorea assamica DyeER ssp. globifera
(RIDL.) Sym.

Under ‘Distr.’ delete line 4 and read: Ben-
coolen.

9: 499b Shorea albida Sym.

Under ‘Ecol.’ line 10: change ‘6.5’ into 65.
Ditto line 21: add after ‘community’: 3.

9: 503. Under Subsection Smithiana, line 3, delete
‘inflaked’.

9: 504b Shorea smithiana SYM.

In ‘Vern.’ line 3, change ‘belong’ into be-
lang.

9: 515b Shorea platyclados Stoot. ex Foxw.
Delete at the end of the Note: ‘not by lateral
plagiotropic shoots’.

9: 518a Incaption Fig. 102, line 2, read: (Sar)awak,
Semengoh Arboretum, with collector SaL-
LEH standing beside it (Photogr. SMYTHIES).

9: 524a Shorea macrophylla (DE VRIESE) ASHTON.
In ‘Uses’, line 1, read for Illippe: Illipe.

9: 529 In Note under Subsection Auriculatae, end
of line 1, read: three others.

9: 541b Shorea platycarpa HEIM.

Note, line 2, read: lowland, dry land for-
ests.

9: 542b

Shorea curtisii DYER ex KING ssp. curtisil.
Note, first sentence, after ‘Selangor;’, read:
subspecies grandis may have the same

9: 543b

9: 547a

4539
S455;
6: 943

6: 668

6: 483,
493

6: 490b

6: 550

6: 580,
582

6: 568

[ser. I, vol. 93

origin.

In caption Fig. 113, line 2, add after ‘Bru-
nei’: The collector is LADI ANAK BIKAS.
Shorea parvifolia DYER ssp. parvifolia.

In ‘Vern.’ line 2, change ‘bung’ into bunga.

Droseraceae

B. J. ConN (Brunonia 3, 1980, 209—216, 2
fig.) gave a review of Drosera L. in New
Guinea, where all Malesian species occur.
He gave a new key, cited new records, and
added a 7th species, which was hitherto on-
ly known from lowland savannahs in
Northern Australia.

Drosera banksii R.Br. ex DC. Prod. 1
(1824) 319; B. J. Conn, Brunonia 3 (1980)
209, f. 2A.

Similar to D. peltata J. E. Smiru, but dif-
fering by absence of bracts under the flo-
wers and presence of stipules.

Distr. Northern Australia; in Malesia:
Papua New Guinea (W. Div.: near More-
head, JoHNns 2201; Wassi Kussa R., HGF
38747).

Ecol. Under seasonal climatic condi-
tions in open Banksia dentata savannahs,
with Melaleuca, Acacia, and Eucalyptus, at
c. 20 m altitude.

Ericaceae

Add the Rhododendron species 298—301;
see below.

298. Rhododendron capellae P. Korgs,
Blumea 24 (1978) 181, f. 1.

Distr. Malesia: Papua New Guinea, W.
Sepik, Mt Capella (Star Mts), 3800 m (LAE
68056).

Note. Keys out to aff. R. vinkii SLEUM.,
but its affinity is obviously with R. pullea-
num Koorvb.

Rhododendron scortechinii K. & G.

Add to Distr.: Borneo: Sarawak (Mt Mulu,
en route from 4th camp to summit, on west-
ern ridge), 2000—2400 m (Hotta 14907).
299. Rhododendron roseiflorum P. F. STE-
veNns, Adansonia II, 18 (1978) 55, 1 fig.

Distr. Malesia: West New Guinea: Mt
Carstensz, near mining on S._ slope,
2100—2700 m (RAYNAL 17580, 17672).

Note. Keys out near R. ruttenii J.J.S.
300. Rhododendron burttii P. J. B.
Woops, Not. R. Bot. Gard. Edinb. 37
(1978) 157, f. 1 a=d.

Distr. Malesia: Borneo: Sarawak (Mt
Murud), 1500 m.

Note. Belongs to ser. Buxifolia; culti-
vated at Edinburgh. No affinity given but
keys out to R. frey-wysslingii J.J.S. from
N. Sumatra.

301. Rhododendron rubineiflorum Cra-

1982]

Addenda, corrigenda et emendanda

6: 6465

6: 657a

6: 878,
Vit

6: 8925,
895b

6: 895,
892b

6: 914

vEN, Not. R. Bot. Gard. Edinb. 38 (1980)
Ae fale

Distr. Malesia: Papua New Guinea.

Ecol. Alpine shrubberies, 2650—3400 m.

Note. Closely allied to R. anagalliflo-
rum WERNH.

Rhododendron nervulosum SLEUM.
exuberans SLEUM.

Add to literature: P. J. B. Woops, Not. R.
Bot. Gard. Edinb. 37 (1978) 159, f. 1 e—-i.

Note. Additional remarks; attention is
drawn towards similarity with R. steno-
phyllum Hook. f.

Correction as follows:

277. Rhododendron irroratum FRANCH.
ssp. kontumense (SLEUM.) CHAMBERLAIN,
Not. R. Bot. Gard. Edinb. 37 (1978) 117. —
R. atjehense SLEUM. Fl. Males. I, 6 (1966)
657, f. 47 & 48.

Note. R. irroratum is a Chinese and In-

dochinese species.

Add the following species:

242. Vaccinium altiterrae VELDK. Blumea
25 (1979) 479.

Distr. Malesia: Papua New Guinea
(Western Highlands: Mts Burgers and Ke-
gum).

Ecol. Edges of Drimys/Rapanea coppi-
ces, 3400—3675 m.

Note. Allied to V. oranjense J.J.S.
New reduction proposed:

Dimorphanthera amblyornidis (BECC.)
F.v.M. var. steinii STEVENS, J. Arn. Arb.
58 (1977) 439. — D. steinii SLEUM. Fl. Ma-
les. I, 6 (1967) 895.

Reduction proposed to variety:
Dimorphanthera apoana (MERR.) SCHLTR.
var. mindanaensis (MERR.) STEVENS, J.
Arn. Arb. 58 (1977) 440. — D. mindanaensis
MERR.; SLEUM. FI. Males. I, 6 (1967) 892.
Add the following three Dimorphanthera
species:

71. Dimorphanthera napuensis P. F. STE-
vENS, J. Arn. Arb. 58 (1977) 441.

Distr. Malesia: West New Guinea (Ba-
liem; Bele R.; Lake Habbema).

Ecol. Sprawling or scandent shrub in
valley forest, 2000—2350 m.

Note. Specimens were confused with D.
wrightiana (KoorD.) J.J.S. Not assigned to
a section.

72. Dimorphanthera wisselensis P. F. STE-
vENS, J. Arn. Arb. 58 (1977) 442.

Distr. Malesia: West New Guinea (Wis-
sel Lakes); one collection, formerly con-
fused with D. wrightiana (Koorp.) J.J.S.
73. Dimorphanthera albida P.F. STEVENS,
J. Arn. Arb. 58 (1977) 437.

Distr. Malesia: Papua New Guinea,
Morobe Distr., Edie Creek, 2700 m, VAN
DER KLOET 35875, in fern heath; f/. Aug.

Note. Belongs to sect. Pachyanthae,

var.

VTS

1278

7: 280

7: 398

5: 46b

563

probably closest to D. ingens (SLEUM.) STE-
VENS.

Fagaceae

Nothofagus BL.
Unfortunately the typification of the genus
Nothofagus by N. betuloides (Mutrs.)
OkERsT. (Taxon 7, 1958, 145) has been over-
looked, and consequently some names of
infrageneric taxa need correction.
Changes in the key at the bottom of the
page:
Line 1, replace ‘1. Sect. Nothofagus’ by: 1.
Sect. Calucechinus (Hompr. & JAcQ.)
KRASSER, and omit this name from line 2.
Line 6, replace ‘la. Subsect. Antarcticae
STEEN.’ by: la. Subsect. Calucechinus, and
add the former name to its synonymy.
Line 9, replace ‘2. Sect. Calusparassus
(Hompr. & JAcg.) KRASSER’ by: 2. Sect.
Nothofagus, and add the former name to
its synonymy.
Line 13, replace ‘2a. Subsect. Quadriparti-
tae STEEN.’ by: 2a. Subsect. Nothofagus,
and add the former name to its synonymy.
Replace ‘1. Section Calusparassus (HOMBR.
& JAcQ.) KRASSER’ by: 1. Section Nothofa-
gus, and add the former name to its syno-
nymy.
Trigonobalanus FORMAN.
Through the paper by D. H. Mai on the
Tertiary fossils of the genus in the Eocene
in Europe (Jahrb. Geol. 3, 1970, 381—409)
it has become clear that the two living spe-
cies of the genus in West Malesia and Thai-
land are relicts of a former much larger
range. 7. doichangensis, hitherto only
known from Thailand, has now also been
recorded from Yunnan (Hsu, WANG, Wu&
Li, Acta Bot. Yunnan. 3, 1981, 213).
This conclusion is now unexpectedly con-
firmed by the find of a third living species
in the mountains of the National Park of
Colombia (NW. South America). G. Loza-
No-C, J. I. HERNANDES-C & J. E. HENAO-S
published this as T. excelsa nov. sp. (Calde-
sia 12 (n. 60), 1979, recvd June 1980,
517-537, 3 pl.). It grows in mountain forest
at 1550-1800 m, as a large tree, 20—40 m
high. Its leaves are scattered, as in the Thai-
land-Yunnan species. It is clear that the ge-
nus once had an ancient, large i aurasian
range. It should also occur in the fossil state
in the southern U.S.A.

Flacourtiaceae

Ryparosa javanica (Bu.) KuRz.

Add to Distr.: Lesser Sunda Is. (Flores: Pa-
ku, 500 m, ScHMuTz 4818).

Homalium Jacq.

564

FLORA MALESIANA

CRAVEN (Brunonia 2, 1979, 107—124, 9
fig.) has given a new treatment of the Pa-
puasian species by proposing 8 new species,
by which the number of 2 distinguished by
SLEUMER (FI. Males. I, 5, 1954, 51) has in-
creased to 10, apart from the 11th, H. ta-
tambense StEum. from the Solomons which
is included here. This is surprising to me,
the more so as 7 of them are based on a

[ser. I, vol. 93

single specimen. From the key given it ap-
pears that single vegetative differences are
frequently used for their delimitation and
almost all are compared with the widely dis-
tributed, very variable H. foetidum. Future
collections will show whether they will
stand the test of time. CRAVEN’s key and
diagnoses are extracted here:

KEY TO THE PAPUASIAN SPECIES

1. Stamens always solitary before each petal.
2. Flowers sessile or subsessile, in simple spikes or racemes.
3. Leaf-apex long-attenuate, acute; leaf margin entire or obscurely distantly crenate. Papua New Guinea
(MorobetDistr-slroldicoll-anotiseen) ss 4---5... «..- eee ae te nee eee 1. H. acutissimum GILG
3. Leaf-apex shortly acuminate, obtuse; leaf margin distinctly crenate. — Differs from H. foetidum by
sessile flowers and solitary stamens. Papua New Guinea (Western Distr.; 1 coll.)7. H. reductum CRAVEN
2. Flowers distinctly pedicellate, in racemes or panicles.

4. Petiole c. 10 mm long. Leaf margin strongly crenate. — Differs from H. acutissimum by distinctly
crenate leaves and pedicelled flowers in panicles. Papua New Guinea (Madang Distr. and Bagabag I.;
Acoli) ANewnrelandi(acollyAneeeee. 2h. fare Sune Sle ete on be RE 2. H. bismarckense CRAVEN

4. Petiole c. 5—6 mm long. Leaf margin weakly crenate. — Differs from H. acutissimum by elliptic acumi-
nate leaves and distinctly pedicelled flowers. West New Guinea (Vogelkop; 1 coll.)
H. caput-avis CRAVEN
1. Stamens in fascicles of (1) 2 or more before each petal.
5. Stamens constantly in pairs, one behind the other. Papua New Guinea and Bismarcks, many collections
5. H. foetidum (Roxs.) BTH.
5. Stamens generally in fascicles of 3 or more.
6. Leaves broadly ovate to subcordate. Stamens in fascicles of 5—8. — Differs from H. foetidum by broad
subcordate leaves and stamens in fascicles of 5 or more flowers. Solomons (Guadalcanal; 1 coll.)
9. H. subcordatum CRAVEN
6. Leaves elliptic to lanceolate. Stamens in fascicles of not more than 5.
7. Perianth segments sparsely hairy, at least the sepals glandular onthe margins. — Differs from H. foetidum
by glandular perianth. Papua New Guinea (Milne Bay Distr.; 1 coll.) ... 8. H. streimannii CRAVEN
7. Perianth segments densely hairy and non-glandular.
8. Leaf margin weakly crenate to entire. — Possibly allied to H. subcordatum but different in elliptic nar-
rower leaves and stamens usually in fascicles of 3—4. Solomons (Santa Isabel I.; 1 coll.)
10. H. tatambense SLEUM.
8. Leaf margin distinctly crenate.
9. Leaves velutinous. — Differs from H. foetidum by the velutinous leaves and stamens mostly in fasci-
cles of 3—4. Papua New Guinea (Central Distr.; 1 coll.).............. 11. H. velutinum CRAVEN
9. Leaves glabrous.
10. Leaf margin entire in the basal 1/3—1/2. Branchlets glabrescent. — Differs from H. foetidum by:
petals and sepals similar, stamens mostly in fascicles of 3. Normanby I. (1 coll.)
4. H. dentrecasteauxense CRAVEN
10. Leaf margin wholly crenate. Branchlets glabrous. — Differs from H. foetidum by larger flowers
and stamens in fascicles of 3—5. Papua New Guinea (Milne Bay Distr.; 1 coll.)
6. H. maneauense CRAVEN
Flagellariaceae moner form with small, globular fruits clo-
ser set, by spaced, larger, ovate-acute fruit

4: 249b Hanguana malayana (JACK) MERR. up to 1.5—2 by 1.25 cm. SHAw added that

4: 249

Add to Distr.: N. Australia: Northern Ter-
ritory, cf. AIRY SHAW, Kew Bull. 33 (1978)
4.

Note. The specimens belong to var.
anthelminthica (Bu.) BAKH. f.
Hanguana major SHAW, Kew Bull. 35
(1981) 819, 1 fig.

This proposed new species, confined to
the Kinabalu area, differs from the com-

the small-fruited form does not occur in
this area.

The species H. malayana, as conceived
by BACKER (FI. Males. I, 4, 1951, 249) is ad-
mittedly variable, BACKER mentioning the
fruit also to attain 2 cm. Extensive field ob-
servation would be needed to check the
racial and population variability in detail
over the entire range.

1982]

Addenda, corrigenda et emendanda 565

Geraniaceae

Geranium LINNE.

The number of Malesian species, according
to CAROLIN (Fl. Males. I, 6, 1964, 445) 3
and one variety, has unexpectedly been in-
creased by VELDKAMP & MOERMAN (Blumea
24, 1978, 463—477) to 15, including 9 new
species from New Guinea and | from SW.
Celebes. In their introduction the authors
state that the evaluation of taxonomic sta-
tus of these taxa has posed a bit of a prob-

to their satisfaction, but further considera-
tion has induced them ‘to regard the taxa as
species’. These considerations are, amongst
others, that there may be chromosome
taxa, that most taxa are represented by
more than one collection, and that they ap-
pear to be homogeneous, although in sever-
al cases occurring on more than one moun-
tain. In the key the exact description of the
leaves plays an important role. Future re-
search, especially karyologic, must show
whether the specific status of the new spe-

lem, which they have not been able to solve cies can be maintained.
KEY TO THE SPECIES

1. Leaves (sub)compound, middle ‘leaflet’ free for at least 0.9th of its length.
2. ‘Leaflets’ repeatedly 2—3-partite with + linear-lanceolate, overlapping segments; the middle 0.2—0.3 mm

wide at base. Papua New Guinea (Mt Suckling; 4 coll.)................. 9. G. leptodactylon VELDK.
2. ‘Leaflets’ 3-lobed to -fid, segments broader, not overlapping, the middle 0.3—1 mm wide at base. Papua
New Guinea (Star, Saruwaket, Owen Stanley Mts; 12 coll.)........... 13. G. subcompositum VELDK.

1. Leaves palmatifid to -partite, middle segment free for at most 0.9th of its length, usually much less.
3. Lower leaf surface very densely grey to silvery hairy, sometimes brown when dried, the indument obscur-
ing the venation. West New Guinea (Carstensz to Star Mts; 15 coll.)......... 10. G. monticola RiDL.
3. Lower leaf surface variously strigose, the indument not obscuring the venation, often making it more
prominent, instead.
4. Middle leaf segment entire, the laterals sometimes with a lobe, rarely the middle segments of some leaves
2- or 3-lobed, leaves then glabrous on the upper surface, 5-partite, and petals pink.
5. Upper surface of the leaves strigose-setose. Peduncle at anthesis already 18—23 mm long. Papua New
Guinea (ME SaruwaketselycOlls)aetaey sere see elets = -l-'s clayeserscetnel sie raters ote seb eeae 6. G. editum VELDK.
5. Upper surface of the leaves glabrous. Peduncle in fruit 0-11 mm.
6. Leaves 7-fid, middle segment free for 0.5—0.6th of its length, 0.8—2 mm wide at the base of its free

part. Papua New Guinea (Mts Giluwe, Wilhelm, Bangeta; 10 coll.)..... 7. G. hyperacrion VELDK.
6. Leaves 5-partite, middle segment free for 0.8—0.9th of its length, 0.5—0.7 mm wide at the base of its
free part. West New Guinea (Mt Wilhelmina; 3 coll.)................ 15. G. wilhelminae VELDK.

4. Middle leaf segment lobed, in some upper or reduced leaves occasionally entire and rarely also glabrous
on the upper surface, then 5—7-fid or petals purple.
7. Upper leaf surface glabrous or sparsely and patchily long-strigose.
8. Petals 9-16 mm long, purple (? always), 1.7—2.1 times as long as the 5.5—7.6 mm long sepals. West
New. Guinea’ (Garstensz; Star, Piora’ Mts; 9 coll!)2.-0). S222 --- ee ee 12. G. papuanum RDL.
8. Petals 4.5-6 mm long, white to pink, 1.2—1.4 times as long as the 3.5—4.8 mm long sepals.
9. Upper leaf surface patchily long-strigose. West New Guinea (Mt Wilhelmina; 1 coll.)
8. G. lacustre VELDK.
9. Upper leaf surface glabrous or with some long hairs near the margin. Papua New Guinea (Wharton
JES CWO) bs) aa Ai A a ES a ee tS 14. G. whartonianum VELDK.
7. Upper leaf surface evenly strigulose to strigose.
10. Leaf blade fairly large, usually over 15 by 25 mm.
11. Inflorescences strictly 1-flowered. Peduncle 25—41 mm. Sepals 5—6 mm long, in fruit 7.5—9 mm.
Petals 5.5—7.5 mm long. Java (Mt Merbabu eastward to Mt Tengger; 13 coll.)
2. G. ardjunense Z. & M.
11. Inflorescences usually 2-flowered. Pedicels 9-16 mm. Sepals 3.5—5 mm long, in fruit 3.8—7.5 mm.
Petals 3.5—6 mm long. New Zealand, Tasmania, East Australia, Timor, East Java (Mt Tengger); 12
COLL: ; AE a ee es Fs 2 ES PR eee er ere 3. G. homeanum TuRCz.
10. Leaf blade fairly small, usually less than 15 by 25 mm.
12. Middle leaf segment pinnately lobed. Peduncle in fruit 55—150 mm. Ceylon, Nilghiris, Himalayas
(Afghanistan to W. China), N. Sumatra (one coll.)...............+.-.- 1. G. nepalense SWEET
12. Middle leaf segment 3-lobed, its outer lobes rarely with a lateral tooth. Peduncle in fruit 0-45 mm.
13. Leaf blades 7-fid to -partite. Peduncle in fruit 0-3 mm long. Papua New Guinea (Star, Kinkain,
Wilhelm and'Saruwaket Mts: 11 coll)... 225 faces = sen eterna re 5. G. balgooyi VELDK.
13. Leaf blades 5-partite. Peduncle in fruit 6—45 mm.
14. Inflorescences 1- or 2-flowered. Peduncle in fruit 6—20 mm, pedicel then 4—15 mm. Petals red-
purple. SW. Celebes (Mt Bonthain; 2 coll.)..................4.- 4. G. frigidurbis MOERMAN

566

FLORA MALESIANA

[ser. I, vol. 93

14. Inflorescences strictly 1-flowered. Peduncle in fruit 19—45 mm, pedicel then 16—31 mm. Petals
white to pinkish. Papua New Guinea (Sugarloaf, Giluwe, Saruwaket to Dayman Mts; 24 coll.)

Goodeniaceae

5: 339a, 1. Scaevola sericea VAHL, Symb. Bot. 2

6: 95la

5: 396a,
7: 828b

7: 75b

4: 213b

(1791) 37. — S. taccada (GAERTN.) ROXB.
Hort. Beng. (1814) 15. JEFFREY, Kew Bull.
34 (1980) 543.

For extremely formal meticulous-no-
menclatural reasons JEFFREY has advanced
that S. sericea should be the proper combi-
nation.

Hydrocharitaceae

Change 1. Limnobium stoloniferum (laevi-
gata) into:

1. Hydromystria laevigata (H. & B. ex
WILLD.) D1Az-MIRANDA & PHILcox, Bot. J.
Linn. Soc. 83 (1981) 321, 6 fig.

The generic name Hydromystria G.
Meyer, Prim. Fl. Esseq. (1818) 152 ante-
dates the synonym Limnobium RIcH. ex
STEUD. (1841).

Icacinaceae

Gonocaryum Mia.

Add to Distr.: Lesser Sunda Is. (West
Flores), probably G. macrophyllum (BL.)
SLEUM.

Replace the name Nothapodytes foetida by:
1. Nothapodytes nimmoniana (J. GRAH.)
MaABBERLEY in K.S. Manilal (ed.), Bot.
Hist. Hort. Mal. (1980) 88; Taxon 29 (1980)
606. — N. foetida (WIGHT) SLEUM.; FI.
Males. I, 7 (1971) 55. — Premna nimmo-
niana J. GRAH. Cat. Pl. Bombay (1839)
155:

The basionym antedates that of Stemo-
nurus foetidus WIGHT by six years.
Stemonurus Bl.

Add to Distr.: Lesser Sunda Is. (Flores;
near Paku, 900 m, ScHMuTz 3371, speci-
men in fr.; identified by SLEUMER).

This collection nicely fills a gap in the
generic range and shows again that the Les-
ser Sunda Islands flora fits in with the rest
of Malesia.

Sarcostigma W. & A.

Add to Distr.: Lesser Sunda Islands (Flo-
res).

Sarcostigma paniculata PIERRE.

Add to Distr.: Lesser Sunda Is. (Flores; Pa-
ku, 500 m, ScHmMutTz 4818a).

Juncaceae

Juncus bufonius L.
Add to Distr.: Borneo (Sabah: Mt Kinaba-

4: 214

4: 214

8: 338b

11. G. niuginiense VELDK.

lu), 3300—3760 m alt.

VELDKAMP (Reinwardtia 10, 1982, 25)
noted that the specimens differ in many de-
tails from the common form. I still consider
it an alien brought by tourists.

Add the following species:
5. Juncus nupela VELDK. Blumea 24 (1977)
415.

Distr. Malesia: Central New Guinea
(Star Mts, Tel Basin, 3000 m, VELDKAMP
6369).

Note. The author finds this to be allied
to the northern hemisphere J. balticus
WILLD. and tabulated differences with three
other species, including J. inflexus L. No
developed ovules could be found.

Luzula DC.

R. BRowN and BENTHAM assumed the Aus-
tralasian specimens of Luzula to belong to
a broad concept of L. campestris. In this
they were followed by BuUCHENAU in his
monograph, except that the latter distin-
guished them to represent an outlying varie-
ty, var. australasica. MERRILL (1922) ac-
commodated the Philippine specimens also
in L. campestris sensu lato. BACKER (FI.
Males. I, 4, 1951, 215) identified the Philip-
pine and Papuan specimens as belonging to
the variety.

The precise naming of the Australasian
specimens of this affinity complex from
Australia, Tasmania and New Zealand has
given rise to a confused nomenclature and
species distinction which M. E. JANSEN has
tried to solve (Blumea 24, 1978, 527—532, 1
fig.). For Malesia he distinguished two new
endemic species, L. philippinensis M. E.
JANSEN and L. papuana M. E. JANSEN. In
the key he did not include how they are dis-
tinguished from L. campestris (L.) DC. It
must be left to a future monographic study
of the genus to see whether these two taxa
can be upheld at specific rank.

Labiatae

Leucas lavandulifolia J. E. SM. Change
into:
3. Leucas linifolia (ROTH) SPRENG. Syst.
Veg. ed. 15, 2 (1825) 743; Bru. Lab. Gen.
Sp. (1834) 617, (1835) 744; FosBeRG & Sa-
CHET, Smithson. Contr. Bot. 47 (1981) 25.
— Phlomis linifolia RotH, Nov. Pl. Sp.
Ind. Or. (1821) 260. — L. lavandulaefolia
J. SM. in Rees, Cycl. (1812) Leu 20, nom.
illeg. (superfl.); KENG, Fl. Males. I, 8 (1978)
338.

Note. Smitu’s name is illegitimate as a

1982]

8: 340b

8: 361

93218

9: 234a

G:9293

6: 294

6: 295

Addenda, corrigenda et emendanda

superfluous name since he cited Leonurus
indicus L. as a synonym. Leonurus indicus
can no longer be transferred to Leucas, be-
cause of L. indica R.BR. ex SPRENG., based
on Phlomis indicus L.

Leucas flaccida R.Br. Change into:

6. Leucas decemdentata (WILLD.) J.SM. in
Rees, Cycl. (1812) Leu 20; Draxeg, Ill. Fl.
Ind. Mar. Pac. (1890) 263; FosBERG & Sa-
CHET, Smithson. Contr. Bot. 47 (1981) 25.
— Stachys decemdentata Sou. ex Forst. f.
Prod. (1768) 91, nomen. — Phlomis de-
cemdentata WILLD. Sp. Pl. 3 (1800) 124. —
L. flaccida R.BR. Prod. (1819) 505; KENG,
Fl. Males. I, 8 (1978) 340. — L. stachyoides
SPRENG. Syst. Veg. ed. 15, 2 (1825) 743.
Satureia gracilis (BTH.) Logs.

Add to Distr.: Borneo: Sabah (Headquar-
ters Taman Negara Sabah, c. 1560 m alt.,
R. H. WitemMseE 605, in disturbed places).

Liliaceae

Thysanotus tuberosus R.BR.
In arecent revision of the genus Thysanotus
N. H. BritTan (Brunonia 4, 1981, 67—181)
assigned the Papuan specimens to T. bank-
sii R.BR., a name which since BENTHAM’S
Flora was considered a synonym. In the key
it appears that the minute differential char-
acters either do not hold or are inconsistent
with the descriptions. I cannot accept T.
banksii as a distinct species.
Astelia alpina R.BR.
Add to Distr.: West Central Celebes (Lati-
modjong Range; Mt Rante Mario, summit
zone, very local).

Note. The collection was made by Dr. J.
M. B. SmitH in February 1981 and is a re-
markable addition.

Loganiaceae (LEENHOUTS)

Wood anatomy. See A. M. W. MENNE-
GA in E. & P. Nat. Pfl. Fam. ed. 2, 28b I
(1980) 112-161.
Palynology. See W. Punt in E. & P.
Nat. Pfl. Fam. ed. 2, 28b I (1980) 162—191.
Phytochemistry. See N. G. BisseTin E.
& P. Nat. Pfl. Fam. ed. 2, 28b I (1980)
211-237.
Delimitation and subdivision. See
A. J. M. LEEUWENBERG ef al. in E. & P.
Nat. Pfl. Fam. ed. 2, 28b I (1980) 1—255.
FosBeRG & SACHET (Smithson. Contr.
Bot. 45, 1980, 18—19) suggest the combina-
tion of Loganiaceae-Potalieae with Gentia-
naceae-Tachiineae, preferably as part of
the Gentianaceae, but on rather vague and
superficial arguments. This seems at least
contrary to wood anatomical data (MENNE-
GA, /.c. 158) and to palynology (PUNT, pers.

6: 308a

6: 320a

6: 320b

6: 328a

6: 328b

6: 331a

567
comm., 1980).
Fagraea fragrans Rox.
Add to Distr.: Ceylon, the whole of

Indo-China, SW. New Guinea.

Insert the following species:

8a. Fagraea graciliflora LEENH. Blumea 27
(1981) 209.

Distr. Malesia: Central Celebes (Mt Ro-
roka Timbu; vAN BALGooy 3247, DE VOGEL
5390).

Ecol. Montane ridge forest dominated
by conifers and Fagaceae, at c. 2100 m. Fi.
May.

Note. Though the texture of the flowers
and the distinctly exserted stamens and
style remind of the situation in sect. Cyrto-
phyllum, the new species will have to be in-
cluded in sect. Fagraea which is the most
primitive in the genus (cf. Punt & LEEN-
HouTs, Grana Palynol. 7, 1967, 510—515).
It is closest allied to F. tubulosa Bu. which
is found in Sumatra and Malaya.

In the key given in Fl. Males. I, 6 (1962)
302, the new species keys out under the first
lead of couplet 19 as F. tubulosa BL. from
which it can be distinguished as follows:
19a. Leaves about 2 times as long as wide.
Inflorescences glomerulous, with
bracteolespeeeeaaenee 8. F. tubulosa
Leaves 13.5—22 by 4—6.5 cm, 2.5—4
times as long as wide. Inflorescence a
terminal 3-flowered cyme and in the
axil of the upper leaf-pair either a
cyme or a solitary flower. Bracteoles
absent sevenerieeet 8a. F. graciliflora
Fagraea ridleyi K. & G.

Add to Notes: Though typical specimens of
F. blumei and F. ridleyi are clearly distinct,
fruiting material is not always easy to
name, especially in Borneo. As a whole the
number of flowers and fruits is smaller in
the present species, the pedicel and calyx in
fruit are less densely warty, the calyx is lar-
ger, the nerves are fewer in number and
more prominent beneath, the axillary scales
are less conspicuous, and the dried leaves
are more olive yellow, in contrast to the
usually dark redbrown ones of F. blumei.
Fagraea auriculata Jack.

Add to Distr.: Great Nicobar (cf. Fl. Males.
Bull. no 29, 1976, 2547) and Celebes.
Add to Notes: vAN BaALGooy 3492 from
Central Celebes combines the long-petioled
leaves with small auricles of ssp. parviflora
with the large flowers of ssp. auriculata.
The collection vAN BaLGooy 3678, also
from Central Celebes, represents ssp. bor-
neensis.

Fagraea resinosa LEENH.

Add to description: Tree, 7.5—9 m by 2—20
cm, or climber. Leaves 8—16 cm long,
width from 3.5 cm onwards. Bracteoles 2

19a.

568

FLORA MALESIANA

6: 336

6: 359a

6: 365,
959

6: 371

and 3 cm resp. Calyx 3—3.5 cm high, in
fruit up to 4 cm.

Add to Distr.: Sarawak (4th Div.), N. Bor-
neo (Mt Kinabalu; Sandakan Distr., Telu-
pid).

Add to Ecol.: Altitude up to 1550 m.
Buddleja Houst. ex LINNE.

Add to literature: LEEUWENB. Meded.
Landbouwhogeschool 79-6 (1979) 1—163.
Strychnos axillaris COLEBR.

Add to Distr.: Ceylon.

Neuburgia corynocarpa (A. GRAY) LEENH.
In a detailed study of the specimens of N.
corynocarpa in Papua B. J. Conn (Bruno-
nia 2, 1979, 99—105, 4 fig.) has come to the
conclusion that N. sarcantha (Gitc & BE-
NED.) LEENH. Fl. Males. I, 6 (1962) 366 can-
not well be distinguished at specific level.
Accordingly he reduced this to a variety of
the former, var. sarcantha (GiLG& BENED.)
B. J. Conn, which he keyed out (/.c. 104).
He also described the structure of seedlings.

Distr. Add: Moluccas (Obi) and New
Ireland.

Note. An isotype of N. sarcantha is
found in B (LEDERMANN 13005) and re-
places the neotype with which it completely
agrees.

Geniostoma FORST.

Recently B. J. Conn (Blumea 26, 1980,
245-364, 29 fig.) made a world revision of
the genus resulting in the name change of
one and the creation of a new species. He
also gave a new key to the species of Papua-
sia (incl. Solomons). In using my key (FI.
Males. I, 6, 1962, 371) the two can be ac-
commodated in couplet 3, second lead, af-
ter:
3. Corolla 2—4 mm long.

4. Pistil hairy..... 4. G. antherotrichum
4. Pistil glabrous...... 5. G. leenhoutsii
4. Geniostoma antherotrichum Gitc & BE-
NED. Bot. Jahrb. 54 (1916) 158, f. 2;
LEENH. FI. Males. I, 6 (1962) 371, as syn.
under G. rupestre; B. J. CoNN, Blumea 26
(1980) 317, f. 15. — G. arfakense Kan. &
Hat. Bot. Mag. Tokyo 56 (1942) 163, f. 7;
LEENH. FI. Males. I, 6 (1962) 373.

Note. This species has a variety:
var. archboldianum (MErRR. & PERRY) B. J.
Conn, Blumea 26 (1980) 320, f. 15G. — G.
archboldianum MeErRR. & PERRY, J. Arn.
Arb. 23 (1942) 408.

Distinct by smaller leaves (2-6 by 1—2
cm) and flowers solitary or in triads. A few
collections intermediate with the type vari-
ety.

5. Geniostoma leenhoutsii B. J. Conn, Blu-
mea 26 (1980) 323, f. 17.

Distr. Solomons (type), in Malesia: Pa-
pua New Guinea (Central Distr.: Woitape);
in all 2 specimens known.

4: 265a

6: 962b

5: 195b

5: 429

4: 53a

[ser. I, vol. 93

Note. I doubt whether this new species
will be tenable. — Edit.

Myoporaceae

Myoporum papuanum KRAENZL.

Add to Distr.: Lesser Sunda Is.: W. Timor
(Kie, near Amanubar, C. W. Kooy 1271,
dated 7-2-1981).

Pittosporaceae

Pittosporum pumilum SCHODDE.

Add to literature: STEEN. Blumea 24 (1978)
482.

Add to description: Pedicels 6 mm. Flowers
functionally female, deep purple. Sepals 5,
free, ovate-oblong, blunt, long-hairy, 5 by
2.5 mm. Petals 5(—6), cohering at base, li-
gulate, glabrous, bent outwards, apex
rounded, c. 12 by 2.5 mm. Stamens re-
duced, c. 3 mm. Ovary densely brown-
hairy, ellipsoid, 4 by 2 mm; style glabrous
1 mm. Fruit red to deep-brown.

Add to Distr.: Papua New Guinea, Mt Ke-
nive (Nisbet), 2500 m, LAE 65011; f7. July.

Proteaceae

Macadamia hildebrandii STEEN. Add:

Uses. This endemic tree of Celebes has
been successfully planted as a fire-lane tree
in the very large plantations of Pinus mer-
kusii in the Aek na Uli area, on the north
flank of Toba Lake (N. Sumatra). Thirty
years old trees have the virtue of keeping
dense foliage and branching from the base
upwards.

Rhizophoraceae

The distinction of sterile material of Rhizo-
phora and Bruguiera may give difficulty.
Dr. Dinc Hou found an easy way for iden-
tification (Blumea 10, 1960, 628) by means
of a hand-lens. In Bruguiera the leaf-scars
show 3 distinct, usually horseshoe-shaped
bundles of leaf traces. In Rhizophora there
are several vascular bundles in two rows or
a crescentic pattern. This was recently also
advanced by KENNEALLY c.s. in their work
on Australian mangroves (Nuytsia 2, 1978,
178—180, 1 fig.).

Styracaceae

Styrax crotonoides CLARKE ssp. fraserensis
(Putz & No) STEEN., comb. nov. — S.
fraserensis Putz & No, Mal. For. 40 (1977)
249, f. 1; Tree Fl. Malaya 3 (1978) 263.
A small tree. Average mature leaves
6—10.5 by 2.5—5 cm; nerves 4—6 pairs.

1982]

Addenda, corrigenda et emendanda

569

4: 54b

8. 239b

Fruits roundish, c. 2 cm diam.

Distr. Malesia: Malaya: Fraser’s Hill,
at c. 1300 m.

Notes. I have carefully compared the
descriptive data with those of S. crotonoi-
des, but find no essential differences, and
also that the ones mentioned are slightly
overlapping. In the species the mature
leaves average 8.5—20 by 4—10 cm, nerves
5—9 pairs. Fruit round to ovoid, tending to
have a smallish or indistinct tip at apex,
1.7—3 cm diam. Lowland, 90—300 m.

Obviously a hill race, with smaller leaves.
The shape of the fruit induced me to main-
tain it as a taxon.

Styrax ridleyanum PERK.

Putz & No (Tree Fl. Malaya 3, 1978, 264)
have discarded this from Malaya and find
the specimen YEor KEP 3639 best to place
under the allied S. benzoin DryYAND. be-
cause of the short inflorescence. I must ad-
mit that this is indeed no differentiating
character. I had no opportunity to re-study
this specimen.

However, another one, also with short
inflorescence, exactly tallies with the differ-
ences with S. benzoin as given in the key of
my elaborate treatment (Bull. Jard. Bot.
Btzg. III, 12, 1932, 223): twigs dark brown-
ish; leaves with smaller stellate scales than
in S. benzoin; buds rather broad; petals in-
side appressed-pubescent all over; connec-
tive ditto; stigma 3-lobed, much wider than
the style.

Distr. Malesia: Malaya: K. Trengganu,
Bt Lah, off Sg. Nerus, near Kp. Merjor,
SINCLAIR & Kian SF 40896 (dupl. in L).

Note. I expect that KEP 3639 from Ku-
antan will turn out to belong to S. ridleya-
num, as I studied this in 1932.

Symplocaceae

Insert the following species:
5a. Symplocos columbuli Noort. Blumea 26
(1980) 417, fig.

Distr. Malesia: N. Sumatra, Leuser Re-
serve, in montane moss forest, 2100—2500
m.
Note. In Noortesoom’s key to fruiting
material (Fl. Males. I, 8, 1977, 231) it
comes out in two places because the leaves
are hairy underneath but finally glabrous,
viz. via lead 3a to couplet 8, from the two
species of which it is distinguished by: fruits
narrow-ellipsoid, c. 15 mm long, with c. 10
conspicuous lengthwise ribs. Via lead 3b
one arrives at couplet 25, where the same
characters single it out.

Sofar only known in the fruiting state;
affinity probably with S. atjehensis Noort.
from the same area.

8: 267

8: 61b

4: 131

4: 131b

Symplocos ophirensis CLARKE var. kalien-
sis STONE, Mal. For. 43 (1980) 260, f. 6.

Distr. Malesia: Malaya: Pahang, G.
Ulu Kali, STONE 13965.

Note. According to STONE closest to
var. densereticulata Noot., differing in a
glabrous disk, branches, and fruit, style 5
mm. From all other varieties different by a
rounded leaf base and a convex petiole.

Ulmaceae

Celtis paniculata (ENDL.) PLANCH.

Add to synonymy: Strombosia philippinen-
sis sensu LAM & Hottu. Blumea 5 (1942)
178.

Add to Distr.: Moluccas (Talaud Is.: LAM
3175).

Umbelliferae

Add the following species:

5. Oreomyrrhis plicata MaTHias & Con-
STANCE, J. Arn. Arb. 58 (1977) 190, f. 1—6.
— Fig. 3.

Leaves tufted at apex of a thick, hardly
branched ligneous stem 5—15 cm long,
erect, linear to lanceolate, 1-pinnate, the
sheathing petiole about as long as the blade;
blade 1—4 cm by 1—3 mm, plicate, seg-
ments 5—11, linear, 2—5 mm long, erect,
entire. Peduncles erect, solitary, exceeding
the leaves, 2.5—12 cm long. Flowers pedi-
celled, in a head.

Distr. Malesia: Papua New Guinea (Mt
Suckling, summit of Goé Dendeniwa:
VELDKAMP & STEVENS 5748), 3325—3625 m,
rocky ridges.

Note. Distinctly different from all other
species by the simply pinnate leaves, folded
lengthwise, with closely adpressed, linear,
entire leaf segments.

Replace 2. Apium tenuifolium by:

2. Apium leptophyllum (Perrs.) F.v.M. ex
Bru. Fl. Austr. 3 (1866) 372; Back. &
Baku. f. Fl. Java 2 (1965) 175. — Cnidium
tenuifolium MorNncH. Meth. (1794) 98,
excl. syn. Pimpinella dioica L., nom. illeg.
— Pimpinella leptophylla Pers. Syn. 1
(1805) 324. — Helosciadium leptophyllum
(PerRs.) DC. Mém. Soc. Phys. Geneve 4
(1828) 493; Prod. 4 (1830) 105, cum syn.
numer. — Apium tenuifolium (MOENCH.)
THELL. in Hegi, Ill. Fl. Mitteleur. 5, 2
(1926) 1140, nom. illeg.; BUWALDA, Fl. Ma-
les. I, 4 (1949) 131.

Distr. Southern hemisphere, intro-
duced in various countries: Europe, India,
Taiwan, Pacific Islands (Hawaii: Kauai; Fi-
ji, H. J. Lam 6827); in Malesia: Java (W.:
Lembang; E.: Pudjong, Trétés, Ever-
AARTS, in litt.), Philippines (Luzon: Ba-

FLORA MALESIANA [ser. I, vol. 93

570

, x 10 (Courtesy Journal Arnold Arbo-

, X0.5, b. foliage leaf, nat. size, c. fruiting um-

bel, x 2.5, d. petal, x9, e. intact mature fruit, x7, f. fruit transection

Fig. 3. Oreomyrrhis plicata MATHIAS & CONSTANCE. @. Habit
retum).

1982] Addenda, corrigenda et emendanda 571

guio, PNH 35028, 35842), Papua New Gui-

nea (Western Highlands, Minj Distr., NGF

41801; Morobe Distr., Sattelberg, CLEMENS

1720; Wau: NGF 29146; Goroka: McCKEE

1160). 4: 368 Replace the key to the species of Xyris L. by
Ecol. Fields, wastelands, roadsides, old the following:

walls, 800—1500 m alt.

Xyridaceae (B. HANSEN, Copenhagen)

KEY TO THE SPECIES

1. Median bracts at apex emarginate, margin otherwise entire, grey field 2 by 1.2 mm. Burma, Thailand, Viet-
PEPE Troy fi in AM 5 ec Oe RR eS ai Okie mes coc X. lobbii RENDLE
1. Median bracts not emarginate, although sometimes irregularly torn, if emarginate then margins obviously
lacerate above.
2. Median bracts with upper margin lacerate.
3. Margin of leaves formed by numerous retrorsely pointing, coalescent rows of cells, rough from small tu-
bercles; scape compressed, usually with two strong ridges................-..-05- 1. X. complanata
3. Margin of leaves not so; scape terete, with low ridges.................2...-220000- 2. X. bancana
2. Median bracts not lacerate at margin, rarely with a few dents.
4. Scape terete with 6—15 longitudinal ridges, leaves up to 8 mm wide with short transverse ribs connect-
ingathe lonpitudinal’ OMES eeleycere te Pe oes eae) = shakes vaus «.o)atareta ers eel sks teat otees ot tetenne) oer 3. X. indica
4. Scape terete to compressed, without ridges or with 1—7 ridges, leaves never with transverse ribs.
5. Leaves 6—20 mm wide, inflorescence depressed hemispherical, wider than long; plant extremely robust
4. X. grandis
5. Leaves up to 6, rarely 7.5 mm wide, inflorescence ellipsoid, globular or longer than wide.
6. Median bracts without a greenish or greyish field below apex, at most with a + conspicuous, narrow
0 One ae Be On. eee aCe ONS A arn DAA I at dient ote 5. X. capensis
6. Median bracts with a greyish or greenish field below apex.
7. Leaf sheath with margin + finely ciliate below, oblong-triangular greyish-brown field below apex of
median bracts 3—10 times longer than wide, not protruding upwards............. 6. X. wallichii
7. Leaf sheath not ciliate below, triangular greyish field below apex of median bracts at most 2.5 times
longer than wide.
8. Scape 4-angular with papillate longitudinal ridges; leaves with papillate margins, otherwise smooth
7. X. oligantha
8. Scape terete, without ridges.
9. Lateral sepals with smooth, entire crest.
10. Plants gracile, leaves at most 3 mm wide, often rough by protruding cell walls (when dry), grey
field on median bracts ridge-like protruding towards apex of bract......... 8. X. pauciflora
10. Plants robust, leaves more than 3 mm wide, not rough, grey field on median bracts not protrud-
ing towards apex of bract although convex. Burma, Thailand, Laos, Cambodia, Vietnam
X. intersita MALME
9. Lateral sepals with crest coarsely serrate, often by 1—3 dents only.
11. Plants robust, field on median bracts 1.8—2.6 by 1.4—2.6 mm, not protruding; with subglobose

underground tubers. Thailand, Laos, Cambodia ................-.--+-- X. tuberosa RIDL.
11. Plants gracile, grey field on median bracts 1—2.2 by 0.4—1.5 mm, ridge-like protruding towards
anexofi bract-awithoutstubenrsmer ceeince ae aoe eee nei See eer aoe ee 8. X. pauciflora

4: 369

1. Xyris complanata R.Br. Prod. 1 (1810)
256; v. ROYEN, Fl. Males. I, 4 (1953) 369.
— xX. indica auct., non LINNE: VAHL,
Symb. Bot. 3 (1794) 7, p.p. quoad spec.
KoENIG. — X. anceps auct., non LAMK:
VaHL, En. PI. 2 (1805) 205. — X. elongata
RupGE, Trans. Linn. Soc. 10 (1811) 289. —
X. walkeri KuNTH, En. Pl. 4 (1843) 19. —
X. malaccensis StEuD. Syn. Pl. Glum. 2
(1855) 287.

Distr. Ceylon, India, China (Hainan),
Indochina, Thailand, throughout Malesia
to Australia.

Ecol. Open, wet places on sandy soil,
often among grasses and Sphagnum, from
sea level to 1600 m.

2. Xyris bancana Mig. Fl. Ind. Bat., Suppl.
(1861) 608; v. RoYEN, FI. Males. I, 4 (1953)
369, incl. var. lacerata v. ROYEN, I.c. 370.
— X. ridleyi RENDLE, J. Bot. 37 (1899) 505;
v. RoyeENn, Blumea 7 (1953) 309, incl. var.
penicillata v. ROYEN, Fl. Males. I, 4 (1953)
370, 371. — X. borneensis RENDLE, J. Bot.
37 (1899) 506; v. RoyEN, Fl. Males. I, 4
(1953) 370. — X. chinensis MALME, Svensk
Bot. Tidskr. 21 (1927) 386. — X. glaucella
Ma.ME, Bull. Jard. Bot. Btzg III, 10 (1929)
388. — X. subcomplanata Matme, Bull.
Mus. Hist. Nat. Paris II, 2 (1930) 685. — X.
papuana v. RoyeEN, Blumea 7 (1953) 307;
Fl. Males. I, 4 (1953) 371.

Distr. Thailand, Cambodia, Vietnam,

FLORA MALESIANA

[ser. I, vol. 93

Hong Kong; in Malesia: Sumatra (Banka),
Malaya (Kedah Peak), Borneo, New Gui-
nea (Misool I.; Aru Is.: Trangan) as far east
as Sepik. Fig. 4.

Ecol. On (temporarily) boggy or wet,
invariably acid, sandy soils (kerangas,
sandstone, heath woodland), largely con-
fined to low altitudes below 100 m, but in
Indochina and Malaya at 1100—1300 m and
in West New Guinea on the Star Mts at
1200—1300 m and on white sands in the Ba-
liem Valley at 2000 m, nowhere at altitudes
between. Fig. 4.

3. Xyris indica Ling, Sp. Pl. 1 (1753) 42;

Fig. 4. Distribution of Xyris bancana Miq. The dots provided with an oblique line represent localities above
1000 m altitude.

v. ROYEN, Fl. Males. I, 4 (1953) 373. — X.
robusta Mart. in Wall. Pl. As. Rar. 3
(1832) 30. — X. calocephala Mia. F\. Ind.
Bat. 3 (1857) 527. — X. capito HANCE, J.
Bot. 14 (1876) 262.

Distr. Ceylon, India, Burma, China
(Hainan), Thailand, throughout Malesia to
Australia.

Ecol. On open, swampy places and
along or in ricefields in the lowland, rarely
up to 900 m.

4. Xyris grandis Riwiey, J. Linn. Soc. Bot.
38 (1908) 332; v. RoyeEN, Fl. Males. I, 4
(1953) 372. — X. chlorocephala v. ROYEN,

Addenda, corrigenda et emendanda 573

Blumea 7 (1953) 308; Fl. Males. I, 4 (1953)
372.

Distr. Indochina, Thailand, in Malesia:
Malay Peninsula, Sumatra.

Ecol. In wet places, among mosses over

rocks, 500—2200 m.
5. Xyris capensis THUNB. Prod. Fl. Cap.
(1794) 12; Nusson, Ofvers. Forh. Kongl.
Svenska Vet.-Akad. 48 (1891) 155, mel.
var. nilagirensis (STEUD.) NILSSON ef var.
schoenoides (MART.) Nitsson; v. ROYEN,
Fl. Males. I, 4 (1953) 374, incl. var. — X.
schoenoides Mart. in Wall. Pl. As. Rar. 3
(1832) 30. — X. nilagirensis STEUD. Syn.
Pl. Glum. 2 (1855) 288. — X. melanocepha-
la Mia. Fl. Ind. Bat. 3 (1857) 528. — X. su-
matrana MaAIME, Bull. Jard. Bot. Btzg III,
10 (1929) 391. — X. novoguineensis
Hatus. Tokyo Bot. Mag. 56 (1942) 422. —
X. flabellata vy. RoYEN, Blumea 7 (1953)
308; Fl. Males. I, 4 (1953) 375.

Distr. S. America, S. Africa, India,
Thailand, Indochina, China, throughout
Malesia.

Ecol. In open, swampy places among
sedges on Sphagnum, 600—3300 m.

6. Xyris wallichii KUNTH, En. Pl. 4 (1843)
16. — X. oreophila Rw. J. Fed. Mal. St.
Mus. 7 (1916) 121; v. Royen, Fl. Males. I,
4 (1953) 372. — X. malmei v. RoYEN, Blu-
mea 7 (1953) 307; Fl. Males. I, 4 (1953) 370.

Distr. India (Silhet, Khasya), Burma
(Moulmein), Vietnam (Chapa), Thailand;
in Malesia: Malay Peninsula.

Ecol. In wet, mossy places on rocks,
850—1300 m.

7. Xyris oligantha STEuD. Syn. Pl. Glum. 2
(1855) 288. v. Rovyen, Fl. Males. I, 4 (1954)
599. — X. pauciflora auct., non WILLD.: R.
Br. Prod. 1 (1810) 256, guoad descr.

Distr. Australia (Cape York Peninsu-
la); in Malesia: Aru Is. and New Guinea
(Papua, Western Distr.).

Ecol. In wet, sandy places among sedges

and grasses, up to 60 m.
8. Xyris pauciflora Wir.ip. Phytogr. 1
(1794) 2; v. Royen, Fl. Males. I, 4 (1953)
371. — X. pauciflora var. oryzetorum Mia.
Fl. Ind. Bat. 3 (1857) 529. — X. dajacensis
v. RoyEN, Blumea 7 (1953) 308; Fl. Males.
I, 4 (1953) 372. — X. maritima KoyaMa,
Philip. J. Sc. 84 (1956) 367.

Distr. Ceylon, India, Burma, Thailand,
Indochina, China, Hong Kong, Taiwan,
throughout Malesia to Australia.

Ecol. In open, wet places on sandy soil,
from sea level up to 900 m.

The following species have to be Exclud-
ed:

4: 369b Xyris tuberosa RiwLey, J. Fed. Mal. St.

Mus. 10 (1920) 122.

Distr. So far recorded from Laos, Cam-
bodia and Thailand (type, ‘Mainland
shores of Takuapa’) only.

4: 371b Xyris lobbii RENDLE, J. Bot. 37 (1899) 506,

t. 403, f. 17-24.
Distr. So far recorded from Burma
(type), Thailand and Vietnam only.

‘se ‘ fHLuscbe wap ea aCe Rey « Srna 4 ler, | ad -

—d - =<_ od

poo ai) viper (G1) +) .eetf MM RDE (ETO? eas

ii

-

ca oe ape ne ey aaa

seg) a {tee Lave Sotivihs
we, wiot) Gaye %~ |: ‘é a al »
i) ars. SYep Pan be (be o-s: ; 2 ,

Oa abi lg i
bint Line yas
+ Chis imal, roereher’ iti

INDEX TO SCIENTIFIC PLANT NAMES

compiled by

M.J. VAN STEENIS-KRUSEMAN and E.E. vAN NIEUWKOOP

Families and higher taxa have been entered under their name.

Names of families which have been revised in volumes 4—9 have been entered and are printed in bold type,
so that as far as this is concerned this index is complete for all preceding volumes as well.

Suprageneric epithets have been entered under the family name to which they belong preceded by the indica-
tion of their rank (subfamilies, tribes, efc.).

Infrageneric epithets have been entered immediately under the generic name to which they belong preceded
by the indication of their rank (subgenera, sections, series, efc.).

Infraspecific epithets have been entered under the specific name to which they belong preceded by the indi-
cation of their rank (subspecies, variety, forma, efc.).

Epithets of new names and new combinations have been printed in bold type, synonyms in italics.

Page numbers in bold type denote main treatment; an asterisk behind a page number denotes the presence
of a figure of the concerned taxon; ‘map’ printed behind a page number denotes that a map of the concerned

taxon is present on that page.

Some minor printing errors in plant names have been corrected.
Of synonyms with a double authority, the latter has not always been cited in full. The full authority can

easily be derived from the text.

Acacia 562
Acanthopanax (Decne &
Planch.) H. Witte 2, 4, 7,
9, 103, 554
aculeatus H. Witte 103
malayanus M.R. Henders.
103, 104*, 554
trifoliatus (L.) Voss. (non (L.)
Merr.) 103, 554
Acanthophora Merr. 17
scandens Merr. 19
Aceraceae 4: 3—4, 592; 6: 915;
7: 820; 8: 549; 9: 280
Actinidiaceae s.str. 4: 37—39
Actinophyllum palmatum Bl. ex
Boerl. 99
Adamboe Rheede 558
Agavaceae 191, 192
Agelaea trinervis (Llanos) Merr.
558
Aizoaceae 4: 267—275; 7: 133
Albizia lophantha Bth. 2
‘Alethris’ 230
Aletris L. 189, 191, 192, 230
foliolosa Stapf 230, 231*,
232%
‘foliosa’ 230
japonica Lambert 232
rigida Stapf 230, 231
spicata (Thunb.) Franch. 230,
PRY AIRY.
sumatrana Masam. 230
Alismataceae 5: 317—334; 6
915; 9: 553
Alismatales 199
Allium fragrans Vent. 234
inodorum W. Ait. 234
Allophyllus 275

Alseuosmia A.Cunn. 557
Amaranthaceae 4: 69—98, 593;
5: 554; 6: 915—917; 7:
820; 8: 549
Amaryllidaceae 190, 191
Anacardiaceae 8: 395—548; 9:
553
Anakasia Philipson 1, 7, 9, 89
simplicifolia Philipson 89, 90*
Ancistrocladaceae 4: 8—10; 5:
553; 9: 275, 281
Ancistrocladus Wall. 275
Anguillaria indica (L.) R. Br.
198
Anisoptera Korth. 239, 241,
255, 258, 259, 261, 263,
265, 266, 267, 269, 271,
274, 276, 277, 280, 283,
284, 288, 290, 291, 327,
328 map, 338
sect. Anisoptera 257, 279, 328
sect. Antherotriche Heim 328
sect. Glabrae Heim 279, 328,
335
sect. Pilosae Heim 328
aurea Foxw. 328, 334
bantamensis Hassk. 364
brunnea Foxw. 333
calophylla Perk. 333
cochinchinensis Pierre ex La-
ness. 271, 330
costata Korth. 239, 243, 244,
245, 249, 256, 270, 271,
278, 303, 304, 326*, 328,
330, 331, 333, 334, 354
costata (non Korth.) Rid]. 332
curtisii Dyer ex King 251, 270,
328, 329, 331

(Anisoptera curtisii)
var. Jatifolia King 329

curtisii (non Dyer ex King)
Foxw. 334

curtisii (non Dyer ex King)
Merr. 331

forbesii Brandis 334

glabra Kurz 271, 335

glabra (non Kurz) Pierre 331

grandiflora Brandis 328

grossivenia Sloot. 253, 262*,
328, 330*, 331

guiso DC. 448

kostermansiana Dilmy 334

laevis Ridl. 271, 326*, 327,
328, 335

lanceolata Walp. 333

mangachapoi DC. 365

marginata Korth. 253, 256,
261, 328, 329*

marginatoides Heim 331

megistocarpa Sloot. 328, 331,
332

melanoxylon Hook. f. 341

mindanensis Foxw. 331

mindanensis (non Foxw.)
Wyatt-Smith 328, 335

oblonga Dyer 330

oblonga (non Dyer) F.-Vill.
333

odorata Kurz 361

palembanica Mig. 303

‘parviflora’ 436

parvifolia Warb. 436

plagata Bl. 423

polyandra Bl. 334

reticulata Ashton 253, 328,
333

(575)

576

FLORA MALESIANA

(Anisoptera)

robusta Pierre 331

scaphula (Roxb.) Kurz 243,
244, 271, 328, 335

sp. Brandis 335

sp. Dyer 334

sp. ‘B’ Wyatt-Smith 331

thurifera (Blco) Bl. 241, 243,
244, 249, 255, 256, 278,

326792851595 35277,1 5995

354
ssp. polyandra (Bl.) Ashton
334
ssp. thurifera 333
thurifera (non Bl.) Foxw. 335
thurifera (non Bl.) Ridl. 335
tomentosa Brandis 333
vidaliana Brandis 333
Anisopteroxylon 241
Anisum moluccanum Rumph.
105
Anomopanax Harms 7, 27
arfakensis Gibbs 27
celebicus Harms 27
cumingianus (Presl) Merr. 76
digitata Merr. 27
philippinensis Harms 27
schlechteri Harms 29
variifolius C.T. White 29
versteegii Harms 29
warburgii Harms 27
Anthericum L. 205
Antherotriche Turcz. 327
lanceolata Turcz. 333
Anthoshorea Pierre ex Heim
437, 485
Aphoma Raf. 197
Apium leptophyllum (Pers.)
F.v.M. ex Bth. 569
tenuifolium (Moench.) Thell.
569
Aponogetonaceae 4: 11—12; 5:
553; 7: 213-218
Aporosa minahassae Koord. 491
Avalianlen le 2sr4e 7ee8.u9> al
apoensis Elmer 21
armata (Wall.) Seem. 22
beccarii Ridl. 19
bipinnata Blco 17, 18*, 21
f. inermis Steen. 21
bipinnata Reinw. 22
calyculata Z. & M. 87
capitulata Jungh. & de Vriese
DD
chinensis L. 22
chinensis (non L.) Bl. 17
cochleata Lamk 75
dasyphylla Miq. 17, 19, 20*
var. latifolia Miq. 17
var. strigosa Miq. 17

var. urticifolia (Bl. ex Miq.)

Bakh. f. 17

(Aralia)
decaisneana Hance 22
decomposita Reinw. ex de
Vriese 22
disperma Bl. 86
ferox Miq. 17, 21
f. nana Steen. 22
ferox (Miq.) King 19
filicifolia C. Moore 76
filicifolia Ridl. 21
foliolosa Seem. 22
glauca Merr. 21
glomerulata Bl. 94
guilfoylei (Bull) Cogn. & Mar-
ché 76
hypoleuca Pres| 21
javanica Miq. 17, 19
javanica (non Miq.) F.-Vill.
21
montana Bl. 17, 22
var. acutata Miq. 22
var. crassifolia Bakh. f. 22
naumannii E. Marchal 76, 77
nodosa Bl. 78
palmata Lamk 42
palmata Reinw. ex BI. 99
pendula Blco 78
pinnatifida Jungh. 16
reinwardtiana Steud. 99
scandens (Merr.) Ha 4, 17, 19
thomsonii Seem. 22
tripinnata Blco 73
umbellifera Lamk 35
umbraculifera Roxb. 78
urticifolia Bl. ex Miq. 17
Araliaceae 1—105, 553—554
tribe Aralieae Harms 6, 7, 9
tribe Mackinlayeae Harms 6,
9
tribe Schefflereae Harms 6
Aralidium Mig. 1, 4, 6, 7, 8, 14
dentatum Miq. 16
integrifolium Heine 16
pinnatifidum (Jungh. & de
Vriese) Miq. 15*, 16
Araliopsis Kurz 89
Arbor camphorifera Rumph.
375 pu
Arenga 289
Arthrocnemum Mog. 557
ciliolatum Bunge ex Ung.-
Sternb. 557
indicum (Willd.) Moq. 557
leiostachya (Bth.) Paulsen 557
Arthrophyllum BI. 1, 4, 7, 8, 9,
53, 56 map
ahernianum Merr. 55, 56, 58,
59*
alternifolium Maingay ex
Clarke 67
alternifolium Maingay ex
Ridl. 55, 58*, 64, 67, 553

[ser. I, vol. 93

(Arthrophyllum)

angustifolium Ridl. 55, 56
ashtonii Philipson 55, 56, 58*
blumeanum Z. & M. 60
var. ellipticum (Bl.) Miq. 60
var. oblongatum Miq. 60
var. ovalifolium (Jungh. &
de Vriese) Miq. 60
borneense Baker 60
borneense Merr. 56
cenabrei Merr. 55, 65
ceylanicum Miq. 67
collinum Philipson 55, 59*
congestum Ridl. 60, 63
crassum Philipson 55, 59*, 63
dilatatum Miq. 60
diversifolium Bl. 54*, 55, 58,
59*, 60, 61*, 62*, 64
var. lanceolata Miq. 60
diversifolium (non Bl.) Harms
63
ellipticum Bl. 60
elmeri Merr. 56
engganoense Philipson 55,
59*, 63
havilandii Ridl. 60, 63
Javanicum Bl. 60
kjellbergii Philipson 55, 58*,
65
lancifolium Ridl. 101
macranthum Philipson 55,
59*, 60, 63
maingayi Philipson 55, 57*,
58*, 64, 67
merrilliana Furtado 56
montanum Ridl. 55, 57*, 58*,
64, 65, 553
nitidum Ridl. 65
‘ovalifolium’ 60
ovatifolium Jungh. & de
Vriese 60
ovatum Ridl. 65
pacificum Philipson 55, 60
papyraceum Philipson 55, 64
pinnatum (Lamk) Clarke 64,
67, 76
pinnatum (non Clarke) F.-
Vill. 56
proliferum Philipson 55, 57*
pulgarense Elmer 55, 58*, 65
reticulatum Bl. ex Miq. 67
rubiginosum Ridl. 60, 63
rufosepalum Ridl. 60, 63
sablanense Elmer 56
serratifolium Miq. 68
stonei Ah-Lan Lim 553
trifoliatum Rid. 87

Anthropodium R. Br. 189-192,

202
capillipes 202
neocaledonicum 202
strictum R. Br. 202, 203*

1982]

Index to scientific plant names 577

Asparagopsis Kunth 215
decaisnei Kunth 216
Javanica Kunth 216
schoberioides Kunth 216

Asparagus L. 189-192, 215

cochinchinensis (Lour.) Merr.

189, 216, 217
declinatus L. 217
dubius Decne 216
graminifolius L. 225
lucidus Lindl. 216
var. dolichocladus Merr. &
Rolfe 216
penduliflorus Zipp. 216
racemosus Willd. 190, 216,
217
Astelia Banks & Soland. ex R.
Br. 189-192, 232, 234
map
alpina R. Br. 189, 190, 233*,
567
novoguineensis Krause 235
papuana Skottsb. 190, 233
f. minor Skottsb. 232

Baillonodendron Heim 371
malayanum Heim 373
Balanocarpus Bedd. 273, 276,
388, 415, 561
sect. Microcarpae Heim 415
sect. Pachynocarpoides Heim
415
sect. Sphaerocarpae Heim
391, 409
acuminatus Brandis 475
acuminatus (non Brandis)
Heim 389
bancanus Boerl. 424
anomalus King 421
bracteatus Merr. 414
cagayanensis Foxw. 431
coriaceus Brandis 474
curtisii King 414

Balanophora abbreviata Bl. 554
elongata BI. 554
hansenii Hambali 554
papuana (non Schltr) Soepad-
mo 554
Balanophoraceae 7: 783—805; 8:
549; 9: 554
Banksia dentata 562
Basellaceae 5: 300—304
Batatas crassicaulis Bth. 558
Batidaceae 5: 414—415; 6: 917
Befaria Mutis 275
Betulaceae 5: 207—208; 6: 917
Bignonia chelonoides L. f. 554
colais Ham. ex Dillw. 554
Bignoniaceae 8: 114—186; 9: 554
Bixaceae s.str. 4: 239-241
Boerlagiodendron Harms 3, 5,
Wo Sil, Se
agusanense Elmer 52
barbatum (Becc.) Harms 41
boerlagei (Warb.) Harms 46
borneense (Seem.) Merr. 39
camiguinense Merr. 48
carpophagarum (Becc.) Harms
45
catanduanense Merr. 37
caudatum Merr. 37
celebicum Harms 49
clementis Merr. 52
dinagatense Merr. 34
diversifolium Merr. 52
elegans (Ridl.) Harms 38
eminens (Bull) Merr. 48
fenicis Merr. 48
geelvinkianum (Becc.) Harms
38, 53
helleborinum (Becc.) Harms
39
heterophyllum Merr. 38
humilis Elmer 44
insidiator (Becc.) Harms 45
insigne (Miq.) Harms 53

grandifolius Ridl. ex Sym. 473 lauterbachii Harms 43

heimii King 388

hemsleyanus King 536

latifolius Brandis 475

longiflorus Foxw. 473

maximus King 478

multiflorus Sym. 475

ovalifolius Ridl. 407, 465

ovalifolius (non Ridl.) Foxw.
413

pahangensis Foxw. 475

penangianus King 475

pubescens Ridl. 432

sibogae Boerl. 475

sp. Sloot. ex Merr. 475

sphaerocarpus Heim 411

wrayi King 389

Balanocarpus (non Bedd.) King

388

ledermannii Harms 53

lineare Merr. 39

luzoniense Merr. 52

micranthum Harms 51

mindanaense Merr. 48

moluccanum (Miq.) Bakh. f.
& Ooststr. 42

monticola Harms 53

novoguineense (Scheff.)
Harms 43

pachycephalum Harms 45, 46

palmatum (Zipp. ex Boerl.)
Harms 42

pectinatum Merr. 45

pfeilii (Warb.) Harms 41

pulcherrimum (Vidal) Harms
40

ramosii Merr. 43

(Boerlagiodendron)

sayeri Harms 51
serratifolium Elmer 49
sessiliflorum Laut. 47
sibuyanense Elmer 40
simplicifolia Elmer 34
stenolobum Harms 38
tayabense Merr. 48
teysmannii (Boerl.) Harms 44
tricolor Philipson 51
trilobatum Merr. 51
warburgii Harms 46
yatesti Merr. 38
Bonnetiaceae 281
Brachycyrtis 197
Brachypodandra Gagn. 345
Brassaia 7
Brassaiopsis Decne & Planch. 2,
4,7, 9, 89
cystostyla (Miq.) Seem. 95
elegans Ridl. 91, 92*, 95
glomerulata (BI.) Regel 91, 94
minor Stone 91, 93
ovalifolia Ridl. 93
palmata (Roxb.) Kurz 93
polyacantha (Wall.) Banerjee
91, 93, 94*
simplex (King) Stone 91, 93
speciosa Decne & Planch. 94
sumatrana Ridl. 91, 93
var. sumatrana 91
var. variaefolia Philipson 93
sumatrana X glomerulata 93
Brownlowioideae 268
Bruguiera 568
Buddleja Houst. ex L. 568
Bulbisperma Reinw. ex Bl. 227
ovigera Reinw. ex Bl. 229
Bulbospermum B\. 227
Javanicum Bl. 229
Burmanniaceae 4: 13-26, 592;
5: 553; 7: 820; 9: 554—
555
Burseraceae 5: 209—296, 567; 6:
917—928; 7: 820—822; 9:
246, 555—556
Butomaceae 5: 118—120, 566
Byblidaceae 7: 135—137

Caesalpinioxylon palembangense
241
Caesia R. Br. 189—192, 203
setifera Baker 204*
Caldesia parnassifolia (Bassi ex
L.) Parl. 553
Callitrichaceae 4: 251—252
Calycites 241
Campanulaceae 6: 107-141,
928; 8: 549; 9: 556
Canarium L. 555—556
acutifolium Merr. 556
balsamiferum Willd. 556

578

(Canarium)
batjanense Leenh. 555, 556
cestracion Leenh. 556
decumanum Gaertn. 555
fusco-calycinum Ridl. 555
grandifolium (Ridl.) H.J.
Lam 555
hirsutum 555
oleosum Eng]. 556
rigidum (BI.) Miq. 556
Cannabinaceae 4: 222—223
Cappar(id)aceae 6: 61—105; 7:
822
Caprifoliaceae 4: 175-194; 6:
928—930; 9: 556—557
subfam. Alseuosmoideae 556
Capura hullettii Ridl. 358
Caraipa 281
Carexs Ea 107 19s. 12s" 123%"
125227 Al 29*
subg. Carex 107, 117, 141
sect. Acrarrhenae Fries
Sippe Longebracteatae
Pax 136
sect. Acutae Fries
subsect. Praelonga Kuk.
170
sect. Anomalae (Carey)
Mackenz. 111, 159
sect. Anomalae (non Carey)
Nelmes 157
sect. Borneenses Nelmes 139
sect. Capitellatae Meinsh.
111, 158
sect. Carex 111, 171
sect. Confertiflorae Franch.
ex Ohwi 160
sect. Cruciatae (Clarke)
Nelmes 117
sect. Cryptostachyae (Ohwi)
Nelmes 110, 148
sect. Decorae (Kik.) Ohwi
109, 139
sect. Digitatae Fries
subsect. Radicales Kuk.
153
sect. Dispalatae Ohwi 159
sect. Extensae Fries
ser. Tumidae (Kiik.) Ko-
yama 160
subsect. Baccantes Koya-
ma 136
subsect. Capitellatae
(Meinsh.) Koyama 158
subsect. Rhizopodae
(Ohwi) Koyama 159
sect. Ferrugineae Tuckerm.
ex Bailey 110, 165
sect. Filicinae (Clarke) Nel-
mes 117
sect. Folliculatae Mackenz.
110, 166

FLORA MALESIANA

[ser. I, vol. 93

(Carex)
sect. Frigidae Fries
subsect. Decorae Kuk.
139
subsect. Ferrugineae
(Bailey) Kuk. 165
sect. Graciles Tuckerm. ex
Kuk. 109, 111, 172
sect. Hymenochlaenae Dre-
jer
subsect. Debiles (Carey)
Kuk. 166
subsect. Graciles (Kiik.)
Ktik. 172
sect. Hypolytroides Nelmes
110, 134
sect. Japonicae Franch. 159
sect. Indicae Clarke 117
subsect. Indicae Koyama
117
subsect. Japonicae (Kuk.)
Koyama 132
subsect. Scaposae Kuk.
136
sect. Japonicae Kuk. 109,
132
sect. Lageniformes (Ohwi)
Nelmes 109, 110, 118,
148
sect. Longispicae Clarke
111, 169
sect. Mapaniifoliae Nelmes
& Airy Shaw 109, 136
sect. Mitratae Kuk. 110,
150
sect. Molliculae Ohwi 160
sect. Muricatae Fries
subsect. Multiflorae Kuk.
178
sect. Occlusae Clarke 110,
168
sect. Oligostachyae Clarke
109, 139
sect. Orthocerates Koch
subsect. Folliculatae
(Mackenz.) Koyama 166
subsect. Pseudocypereae
(Bailey) Koyama 167
sect. Paciricae Ohwi 132
sect. Paludosae Bailey 111,
113, 176
subsect. Lacustres Carey
176
sect. Phyllostachyae
Tuckerm. 153
sect. ‘Polystachae’ 136
sect. Polystacheae Clarke
117
ser. Cruciatae Clarke 117
ser. Filicinae Clarke 117
ser. Longispicae Clarke
136, 139

(Carex)
ser. Stramentitiae Clarke
117
sect. Polystachyae
(Tuckerm.) ex Kiik. 110,
136
sect. Praecoces Christ 150
subsect. Cryptostachydeae
Franch. ex Ohwi 148
subsect. Lageniformes
Ohwi 148
sect. Praelongae (Kuk.) Nel-
mes 111, 158, 170
sect. Pseudocypereae
Tuckerm. ex Bailey 111,
167
sect. Radicales (Kiik.) Nel-
mes 110, 111, 153, 156
sect. Rarae Clarke 158
sect. Rhizopodae Ohwi 111,
159
sect. Rhomboidales Kuk.
110, 144
sect. Scabrellae Kiik. 144
sect. Scleriiculmes Nelmes
168
sect. Stramentitiae (Clarke)
Nelmes 117, 157
sect. Surculosae Raym. 110,
144
sect. Sylvaticae (Tuckerm.)
Boott ex Mackenz. 110,
166
sect. Trachychlaenae Drejer
110, 157
sect. Tumidae Kuk. non
Meinsh. 159
sect. Tumidae Meinsh. 176
sect. Unciniaeformes Kik.
subsect. Capitellatae
(Meinsh.) Kuk. 158
sect. Vigneastra Tuckerm.
107—110, 117
sect. Vigneastra (grex) Indi-
cae (Tuckerm.) ex Bailey
117
sect. Vulgares (Aschers.)
Nelmes 171
subg. Indocarex Baill. 117,
141
subg. Primocarex Kuk. 117
subg. Vignea (Beauv.) Clarke
107, 176
sect. Capituligerae Kuk.
subsect. Divisae Christ ex
Kuk. 176
sect. Divisae (Christ) Kuk.
111, 176
sect. Elongatae Kunth 111,
180
sect. Elongatae Kunth sensu
Kuk. 179

1982] Index to scientific plant names 579
(Carex) (Carex) (Carex)
sect. Heleonastes Kunth breviceps Kuk. 144 ceylanica Boeck.
111, 181 var. recurvirostris Kuk. 144 var. saturata Kuk. 124

sect. Multiflorae (Carey)
Bailey 111, 178
sect. Muricatae Fries
subsect. Paniculatae
Kunth ex Kiik. 177
sect. Paniculatae (Kunth)
Meinsh. 111, 177
sect. Stellulatae Kunth 111,
179, 180
sect. Vignea Beauv.
subsect. Multiflorae
Carey 178
sect. Vigneastra 182
abacta L.H. Bailey 166
acaulis d’Urv. 153
acrophila S.T. Blake 172
acuta (a) L. 117, 171
alopecuroides D. Don 113,
114, 160
var. ‘chlorostachya’ 160
var. chlorostachys (D. Don)
Clarke 160
alta Boott 180, 181
anomocarya Nelmes 112, 144,
145*
appendiculata (Trautv.) Kiik.
172
appressa R. Br. 108, 116, 177
var. virgata (Sol. ex Boott)
Kuk. 177
arenaria L. 176
arenicola F. Schmidt 177
arnottiana Nees ex Drejer
182
arridens Clarke 140, 141
asperinervis Koyama 172
atjehensis Kuk. 166
‘atjehiensis’ 166
atrosanguinea Nelmes 142
baccans Nees 115, 137*, 162*,
180, 183
var. nigra O. K. 137
var. siccifructus Clarke 137
basilata Ohwi 180
bengalensis Roxb. 120
var. scaberrima Boeck. 122
var. virgata Boeck. 122
bilateralis Hayata 108, 117,
163*, 172, 174
blepharolepis Nelmes 118
borbonica Lamk 183
borneensis Clarke 139, 140,
141
var. clemensii (Ktik.) Kiik.
f. angustifrons Kiik. 141
brachyathera Ohwi 108, 113,
163*, 165
var. brevispiculosa Koyama
165

breviculmis R. Br. 108, 109,
112, 114, 150
ssp. royleana Kik. 151
var. kingiana Kiik. 151
var. breviculmis 151
var. montivaga (S.T. Blake)
Noot. 151
var. perciliata Ktik. 151
var. perciliata (non Kuk.)
Ridl. 165
breviglumis Ridl. 140, 141
brevis S.T. Blake 151
breviscapa C.B. Clarke 112,
148, 561
brizopyrum Kunze 180, 181
brownii Tuckerm. 108, 113,
114, 161
ssp. brownii 111, 161
ssp. transversa (Boott) Kern
163*
var. dissociata Koyama 163
var. transversa (Boott) Kuk.
ex Matsum. 161, 163
var. viridis Boeck. 161
brownii x transversa 163
brunnea Thunb. 108, 117,
140, 163*, 173
ssp. meyenii (Nees) Koyama
174
var. dolichocarpa Nelmes
WB5 se!
var. meyenii (Nees) Koyama
173% Wi
var. subteinogyna Kik. 173,
174
brunnea (non Thunb.) Nelmes
175
buennemeijeri Nelmes 122
bukaénsis Palla 144
bulbostylis Kuk. 151
var. ciliato-marginata Kuk.
151
var. hispidula Kuk. 151
buruensis Nelmes 173, 174
caespitosa (non L.) R. Br. 172
canescens L. 182
canescens (non L.) Boott 181
var. B 181
capillacea Boott 108, 112,
158, 161*
var. capillacea 159
var. major Nelmes 159
var. nana Franch. 159
var. sachalinensis (F.
Schmidt) Ohwi 158, 159
capitellata Boiss. & Bal. 158
celebica Kuk. 112, 139
ceramica Nelmes 122
cernua Boott 182

chinensis Retz. 144
chlorostachys D. Don 160
cirrhulosa Nees 115, 117
cladostachya Wahlenb. 182
clarkeana Kiik. 122
clemensii Kuk. 141
commixta Steud. 115, 118, 130
compacta Poir. 184
composita (non Boott) Clarke
138
concolor Nees 155
condensata Nees 120
confertiflora Boott 165
conorrhyncha Nelmes 151
constricta S.T. Blake 139
continua Clarke 120
continua (non Clarke) S.T.
Blake 128
continua (non Clarke) Clarke
122
contracta Boeck. 132
courtallensis Nees ex Boott
155, 156
var. angustifolia Boott 155
craspedotricha Nelmes 180,
181
cruciata Wahlenb. 109, 116,
118, 120, 182
var. condensata Ridl. 120
var. cruciata 120, 131*
var. rafflesiana (Boott)
Noot. 120, 122, 131*
cryptostachys Brongn. 114,
118, 148
cumingiana Steud. 144
cumingii (Boott) ex Clarke
128
curta Gooden. 108, 114, 181
curtisii Ridl. 149
curvirostris Kunze 137
‘cyrtostachys’ 148
decora Boott
var. losirensis Kuk. 143
densiflora Pres] 117
depauperata 132
dietrichiae Boeck. 126
dimorpholepis Steud. 182
divisa Huds. 176
divulsa Stokes
var. javanica Nelmes 182
dolichocarpa C.A. Mey. ex
Krecz. 167
dolichostachya Hayata 108,
112, 151
doniana Spreng. 160
var. cacuminis Nelmes 160
drepanorhyncha Franch. 166
duriuscula C.A. Mey. 108,
116, 176

580

FLORA MALESIANA

(Carex)
echinata Murr. 108, 114,
162*, 179, 180
eggytera Steud. 151
elibates Nelmes 157, 158
elmeri Kiik. 146
elongata L. 180
eminens Nees 138
eremostachys S.T. Blake 112,
159
erythrolepis Kuk. 182
euphlebia S.T. Blake 169
exploratorum Nelmes 169
eymae Nelmes 140, 141
fallax Steud. 178
var. franchetiana Ohwi 179
fascicularis Soland. ex Boott
167
ferruginea Scop. 165
fibrata Boott ex Vidal 117
filicina Nees 116, 122, 182
f. saturata Kuk. 124
var. angustifolia Nelmes
122
var. ciliata O. K. 122
var. laevis O. K. 122
var. saturata (Clarke) Kuk.
122
var. zipelii Nelmes 122
finitima Boott 108, 113, 114,
166
fleckeri Nelmes 124
floribunda Boeck. 138
foliosissima (non F. Schmidt)
Franch. 152
folliculata L. 166
formosensis Lév. & Van. 108,
112, 148, 152
fuirenoides Gaudich. 118, 126
var. cirrhulosa Kiik. 117
fuirenoides (non Gaudich.?)
Clarke 126
fuirenoides (non Gaudich.?)
F.-Vill. 117
furusei Koyama 163
fusiformis Nees 166
var. borneensis (Clarke)
Kuk. 141
var. enervosa Kiik. 166
fusiformis (non Nees) Stapf
141
gajonum Nelmes 179, 180
galactolepis Nelmes 120, 122
gaudichaudiana Kunth 108,
114, 117, 171
var. humilior Kiik. 171
var. thunbergii (Steud.)
Kuk. 172
gembolensis Clarke 132
var. crebra Nelmes 132
var. timorensis Clarke 122
gibbsiae Rendle 142

(Carex)

gracilis R. Br. 172, 173
gracilispica Hayata 112, 149
graeffeana Boeck. 114, 162*,
169
var. samoensis Nelmes 169
haenkeana Pres| 182
harlandii Boott 146
f. longibracteata Gross 144
var. angustior Kuk. ex
Back. 144
harlandii (non Boott) Merr. &
Chun 144
hattoriana Nakai & Tuyama
173
hatusimana Ohwi 150
havilandii Clarke 143
hebecarpa C.A. Mey.
var. lachnosperma Clarke
168
var. /igulata (non Kik.)
Back. 168
var. maubertiana Franch.
168
heleonastes Ehrh. 181
helferi Boeck. 108, 114, 136
hexasticha Reinw. ex Miq. 170
hispida Willd. 157
hispidangula Koyama 156
horsfieldii Boott 108, 116,
124, 126, 131*
var. major Nelmes 124
horsfieldii (non Boott) Miq.

118
hypolytroides Rid]. 112, 115,
131*, 134
hypolytroides Gross & Mattf.
134

hypolytropsis Gross 134
hypsophila Miq. 143
var. havilandii (Clarke)
Kuk. 143
var. verticillata (Zoll. &
Mor.) Kuk. 143
imbricata Kuk. 180
impunctata Boott 132
indica L. 108, 116, 117, 118,
126, 131*, 133*
var. fissilis (Boott) Kiik.
126
var. laetebrunnea Clarke
126
var. milnei Boott ex Clarke
126
ischnostachya Steud. 164
jackiana Boott 113, 114, 146
ssp. jackiana 146, 147
ssp. parciflora (Boott) Kuk.
146
var. B 146
var. breviculmis Thw. 148
var. minor Clarke 146, 147

[ser. I, vol. 93

(Carex)

var. tumens Kuk. 146, 151
japonica Thunb. 160
Japonica (non Thunb.) Boott

160

ssp. subtransversa (Clarke)
Koyama 160

var. chlorostachys Kuk. ex
Matsum. 160

var. mesogyna Kuk. 160

var. minor Boott 160
Javanica Boeck. 137
kanehirae Ohwi 173, 174
kemiriensis Nelmes 171
kerrii Nelmes 171
kinabaluensis Stapf 140, 141
kuntzeana Boeck. 138
lacerans Kuk. 172
lachnosperma Nees 168
lamprochlamys S.T. Blake

116, 126, 131*

var. diplocolea Nelmes 128
lateralis Kuk. 113, 146, 147
lenticularis D. Don 170
leucantha Arn. ex Boott 155
leuchochlora Bunge 151
leucostachys Ridl. 140, 141
ligata Boott

var. formosensis (Lév. &

Van.) Kiik. 152

var. nexa Kuk. 152
ligata (non Boott) Ridl. 149
lindleyana Nees 182
loheri Clarke 113, 147

f. grandimascula Kuk. 147
‘longebracteata’ 138
longibracteata Steud. 138, 139

f. angustifolia Kuk. 138

f. distans Kuk. 138

var. gigantea Kuk. 138

var. major Miq. 138
longipes D. Don 108, 117,

174, 175*

var. ramosa Kuk. 174
longispica Boeck. 155
lutchuensis Ohwi 149
macrothyrsa Miq. non So-

land. ex Boott 122
maculata Boott 108, 122, 157,
171
f. humilior Kuk. 157
var. neurochlamys Kuk.
157, 158
var. sanguineo-squamata
Kuk. 157, 158
madoerensis Clarke 156
matlaccensis C.B. Clarke 108,
115, 153, 157
malayana Nelmes 149
manca Boott

var. contigua Gross 144

mapaniifolia Ridl. 136

1982] Index to scientific plant names 581
(Carex) (Carex) (Carex)
maubertiana Boott 112, 162*, var. minor Kiik. 165 pterocaulos Nelmes 156
168 olivacea (non Boott) Ktik. 163 pullei Nelmes 143

maximowiczii Miq. 171
megacarpa Koyama 173, 174
melanophora S.T. Blake 130
merrillii Kuk. 143
meyenii Nees 173, 174
michauxiana Boeck. 108, 112,
162*, 166
f. asiatica (Hultén) Akiya-
ma 167
ssp. asiatica Hultén 167
var. asiatica (Hultén) Ohwi
166
mitrata Franch. 150, 153
monopleura Krecz. 181
montivaga S.T. Blake 151
moritzii Steud. 126
moupinensis Franch. 134
multiflora Muehl. ex Willd.
178
multifolia Ohwi 152
muricata L. 182
muricata (non L.) Ohwi 180
myosurus Nees 115, 136, 138
var. celebica Nelmes 138
var. eminens (Nees) Clarke
138, 139
var. floribunda (Boeck.)
Kuk. 139
neoguineensis Clarke 122, 124
neurochlamys F.v.M. 157, 158
nigra (L.) Reich. 172
‘nikkoensis’ 132
nikoensis Franch. & Savat.
132, 134
nilagirica Steud. 122
nipposinica Ohwi 161
nodiflora Boeck. 115, 128
nodiflora (non Boeck.) Kuk.
140
notha 170
nubigena D. Don 108, 117,
178*
var. fallax Clarke 178
var. fallax Kuk. 178
nubigena (non D. Don) Kuk.
179
oblonga Nelmes 122
oedorrhampha Nelmes 113,
163, 165, 561
var. arfakiana Ohwi 163
var. microcarya Nelmes
163, 164
oligostachya Nees 115, 144
olivacea Boott 113, 164*
ssp. confertiflora (Boott)
Koyama 165
ssp. Olivacea 164
ssp. recurvisaccus (Koyama)
Koyama 165

var. altissima Kuk. 163
omiana Franch. & Sav. 180
var. perileia Koyama 179
pairaei F. Schultz
var. javanica Nelmes 182
palawanensis Ktik. 115, 149
paludosa Gooden. 176
pandanus Ohwi 169
paniculata L. 170, 177
paniculata (non L.) Benth.
177
papuana Nelmes 128
pentacarpa Boeck. 122
perakensis C.B. Clarke 109,
114, 115, 140
var. borneensis (Clarke)
Noot. 141, 162*
var. perakensis 141, 142,
162*
var. vansteenisii (Kuk.)
Noot. 141, 142
perciliata (Kiik.) Nelmes 151
perileia S.T. Blake 179, 180
petecticalis Nelmes 171
Phacelostachys Nelmes 143
var. losirensis (Kuk.) Nel-
mes 143
‘phacodes’ 170
phacota Spreng. 113, 117,
158, 163*, 170, 171
phacota (non Spreng.) Kik.
171
Dhilippinensis Nelmes 169
platycarpa Hochst. ex Steud.
170
plesiocephala Turr. 156
pocilliformis Boott 153
praelonga Clarke 170
pruinosa Boott 113, 158,
163*, 170
f. submutica O. K. 170
f. tristigmatosa Back. 157,
170, 171
ssp. mMaximowiczii (Miq.)
Kuk. 171
var. aristata O. K. 170
pruinosa (non Boott) Kiik.
157
pseudocyperus L. 108, 113,
167
var. fascicularis (Soland. ex
Boott) Boott 163*, 167

var. haenkeana (Presl) Ktik.

182
var. pseudocyperus 168

pseudocyperus (non L.) R. Br.

167
pseudorivulorum Kuk. 140,
141

punctata 170
pycnothyrsos Kiik. 122
radicales Boott 153
rafflesiana Boott 122
var. continua Kuk. 122
var. continua (non (Clarke)
Kuk.) Kiik. 128
var. macrothyrsa Kuk. 122
var. minor Hochr. 122
var. scaberrima Kuk. 122
var. tenuior Clarke 122
var. virgata Nelmes 122
ramosii Kuk. 112, 115, 155,
157
rara Boott 159
ssp. capillacea Kiik. 158
var. nana Kuk. 159
rara (non Boott) Stapf 158
rechingeri Palla 169
‘recurvirostra’ 137
recurvisaccus Koyama 165
remota L. 114, 180
ssp. alta (Boott) Kuk. 180,
181
var. brizopyrum Kuk. 180
ssp. remota 181
ssp. rochebrunii (Franch. &
Sav.) Kuk 181
var. rochebrunii Clarke 180,
181
remotiflora Hayata 166
repanda Clarke
var. implumis Clarke 120, 122
rhizomatosa Steud. 144
var. impunctata (Boott)
Ktik. 132, 144
var. aristulata Kuk. 144
rhizopoda Maxim. 159
rhynchachaenium Clarke 108,
112, 150
riparia Curt. 176
riparia (R. Br.) Poir. 184
rivulorum Ridl. 140, 141
rochebrunii Franch. & Sav. 181
rostrata Michx. 166
royleana Nees 151
rugata (non Ohwi) Nelmes 151
rugulosa Kiik. 176
samoensis Boeck. 157
sarawaketensis Kiik. 116, 122,
128
var. brevirostris Kuk. 122
var. glabrinux 124
var. minor 128
‘satsumensis’ 134
saturata Clarke 124
satzumensis Franch. & Sav.
108, 115, 132
scaberrima Clarke 122

582

FLORA MALESIANA

(Carex)
scabrata Schwein. 160
scabriculmis Ohwi 175
scabrifolia Steud. 182
sclerioides Ridl. 124
secta Boott 170
semiglomerata Kuk. 122
setulifolia Nelmes 140
simplicissima F.v.M. 158
smitinandii Raym. 118
sociata Boott 152
sp. 113
sp. 176
sp. Nelmes 176
spadiceo-vaginata Ohwi 173,
174
spathaceo-bracteata Kuk. 172
spathulata Nelmes 171
spatiosa Boott 118
var. bogorensis Clarke 118
speciosa Kunth 108, 111, 115,
154*, 155
var. abscondita Kuk. 155
var. angustifolia (Boott)
Kuk. 155, 156
var. angustifolia Raym. 155
var. courtallensis Kuk. 155
var. minor Boeck. 155
spiculata Boott 138
spongocrepis Nelmes 120
spongoneura Nelmes 122
Squamata Krecz. 180
stellulata Gooden. 179
stenophylla Wahlenb. 176,
177
var. duriuscula Trautv. 176
stenura Nelmes 155, 156
stramentitia Boott ex Boeck.
108, 116, 130, 131*
striata R. Br. 161
subfilicina Ohwi 182
sublateralis Koyama 147
subteinogyna Ohwi 172, 174
subtransversa Clarke 160, 161
sumatrensis Clarke 143
sylvatica Huds. 166
tartarea Ridl. 143
tatsutakensis Hayata 147
teinogyna Boott 108, 117,
174, 175
var. scabriculmis Kuk. 175
teinogyna (non Boott) Back.

‘teiogyna’ 175

teres Boott 114, 163*, 171
var. spathulata Kuk. 171

thorelii Camus 156

thunbergii Steud. 172

timorensis (Clarke) Nelmes

122
tonkinensis Franch. 140
transversa Boott 163

(Carex transversa)
var. dissociata Franch. &
Sav. 163
tricephala Boeck. 108, 112,
115, 156
tricholoma S.T. Blake 151
tricuspidata Kuk. 165
var. brevispiculosa Kuk.
165
var. minor Kuk. 165
tristachya Thunb. 112, 114,
152
var. pocilliformis (Boott)
Kuk. 153
var. pseudopocilliformis
Gross 149
var. tristachya 153
tumida Boott 163
turrita C.B. Clarke 109, 114,
142
var. merrillii (KUk.) Noot.
143
var. turrita 142, 162*
turrita (non Clarke) Kiik. 143
typhoides Bory 183
tyttholepis Nelmes 128
uda Maxim.
var. sachalinensis F.
Schmidt 159
vacua Boott ex Boeck. 120,
122
valida Nees 120
vansteenisii Kuk. 140, 142
var. brevispiculosa Kuk. 142
verticillata Z. & M. 109, 112,
140, 143, 174
var. havilandii (Clarke)
Nelmes 143
var. Jutescens Nelmes 143
vesiculosa Boott 108, 116,
130, 131*
var. congesta Kiik. 132
var. latifolia Kik. 132
var. paniculata Clarke 130
virgata Miq. 122
virgata Soland. ex Boott 177
vulcanica Elmer 128
vulgaris Franch.
var. gaudichaudiana Boott
172
vulgaris (non Franch.) F.v.M.
172
wahuensis C.A. Mey. 182
walkeri Arn. ex Boott
var. turrita (Clarke) Kuk.
142
walkeri (non Boott) Boeck.
143
walkeri (non Arn. ex Boott)
Kuk. 137
wightiana Nees
var. perakensis Kuk. 140

[ser. I, vol. 93

(Carex)
xestogyne Nelmes 122
Caryolobis indica Gaertn. 552
Celastraceae 6: 227-291,
389—421, 930—932; 9: 557
Celastrus hindsii Bth. 557
monospermoides Loes. 557
Celtis paniculata (Endl.) Planch.
569
Centrolepidaceae 5: 421—427
Cephaloschefflera 2
Ceratophyllaceae 4: 41—42
Chenopodiaceae 4: 99—106, 594;
6: 932; 8: 549; 9: 557
tribe Salicornieae 557
Chlaenaceae 268, 275, 282
Chlamysporum Salisb. 211
chrysantherum (F.v.M.) O. K.
211
tuberosum (R. Br.) O. K. 213
Chloopsis Bl. 225
acaulis Bl. 226
caulescens Bl. 226
Chlorophytum Ker-Gawl. 189,
191, 192, 205
glaucum 206
laxiflorum Baker 205
laxum R. Br. 205
f. javanicum (Hassk.) Back.
205, 206
longissimum Ridl. 206
malayense Ridl. 205, 206
nimmonii 206
orchidastrum s.s. 206
orchidastrum (non Lindl.)
Ridl. 206
Cladium scleroides F.v.M. 187
Clethraceae 7: 139—150
Clusiaceae 275
Cnidium tenuifolium Moench.
569
Cochlospermaceae 4: 61—63
Columniferae (order) 281
Combretaceae 4: 533—589; 5:
564; 6: 932—933; 7: 823
Compositae 5
Compsoa D. Don 195
Connaraceae 5: 495—541; 6:
933—936; 7: 823; 8: 549;
9: 557-558
Connarus L. 558
euphlebius Merr. 558
impressinervis Stone 558
monocarpus L.
ssp. malayensis Leenh. 558
Convallaria japonica L. f. 227
Convolvulaceae 4: 388—512,
599; 5: 558; 6: 936-941;
7: 823; 9: 558
Copaifera palustris (Sym.) de
Wit 552
Cordyline 191

1982]

Cornaceae 8: 85—97; 9: 4, 6, 16
Cornales 5

Corynocarpaceae 4: 262—264; 5:

557; 6: 941
Cotylelobiopsis Heim 275
beccariana Heim 552
Cotylelobium Pierre 239, 259,
263, 265—268, 270, 274,
276, 277, 278, 280, 284,
288—291, 327, 338, 340,
342 map, 347, 561
asperum Sloot. 343
beccarianum Heim 341
beccarii Pierre 341
burckii (Heim) Heim 253,
256, 341, 342, 343*
flavum Pierre 342
flavum (non Pierre) Ridl. 345
harmandii Heim 341
hopeifolium Heim 482
lanceolatum Craib 341, 343*
leucocarpum Sloot. 342
lewisianum (Trim. ex Hook.
f.) Ashton 277, 278
malayanum Sloot. 253, 256,
343
melanoxylon (Hook. f.) Pierre
243, 253, 275, 340*, 341,
342*, 343, 552, 561
philippinense Heim ex Brandis
365
Crassula scutellaria Burm. f. 75
Crassulaceae 4: 197—202; 9:
558—560
Crypteroniaceae 8: 187—204
Cyperaceae 7: 435—753, 823; 9:
107—187, 560—561
subfam. Cyperoideae 186
tribe Hypolytreae 186, 187
Cyperus compactus Retz. 561
esculentus L. 561

Dacrycarpus imbricatus 2
Dacryodes longifolia (King) H.J.
Lam 555
var. longifolia 555
multijuga Leenh. 556
rugosa (BI.) H.J. Lam 555
var. rugosa 555
Dammara selanica Rumph. 505
Datiscaceae 4: 382—387; 7: 823
Delarbrea Vieill. 2, 7, 9, 24
collina Vieill. 23*; 24 map
lauterbachii Harms 24
paradoxa Vieill. 25
paradoxa (non Vieill.) Britten
24
sp. Hemsl. 24
Dendropanax Decne & Planch.
2, 3, 9, 101, 102 map
borneensis (Philipson) Merr.
100*, 101, 102

Index to scientific plant names 583

(Dendropanax)

lancifolius (Ridl.) Ridl. 101

maingayi King 101

parviflorus (non (Champ.)
Bth.) Clarke 101

(Dimorphanthera)

wrightiana (Koord.) J.J.S. 563
Dioticarpus Dunn 391, 415
Dipsacaceae 4: 290-292; 5: 557
Diptera 3

Deplanchea bancana (Scheffer)
Steen. 554
glabra (Steen.) Steen. 554
Dianella Lamk 189—193, 206
albiflora Hall. f. 207 (erron.
cox)
austro-caledonica Seem. 209
bambusifolia Hall. f. 207
bancana Miq. 207, 209
caerulea (non Sims) Merr. 207
carinata Hall. f. 207
celebica (Bl.) Kunth 209

ensata (Thunb.) R.J. Henders.

208
ensifolia (L.) DC. 207
flabellata Hall. f. 207
javanica (BI.) Kunth 207,
208*, 209
f. alba Schlittl. 209
f. rubra Schlittl. 209
f. stenophylla Schlittl. 209
ledermannii Krause 208
levis (non R. Br.) C.T. White
208, 209
monophylla Hall. f. 207
montana Bl. 207, 209
monticola Krause 208
nemorosa Lamk 207, 209
odorata (Rumph.) BI. 207,
209
parviflora Ridl. 208
parviflora Zipp. ex Hall. f.
207
pullei Krause 208
revoluta (non R. Br.) Schauer
207
robusta Elmer 207
serrulata Hall. f. 207
sparsiflora Schlittl. 208, 209
Dichapetalaceae 5: 305—316,
567; 6: 941-943; 7: 823
Dichopogon Kunth 202
strictus (R. Br.) Baker 202
Dilleniacea ? nervosa Wall. 548
Dimorphanthera
sect. Pachyanthae 563
albida P.F. Stevens 563
amblyornidis (Becc.) F.v.M.
var. steinii Stevens 563
apoana (Merr.) Schltr
var. mindanaensis (Merr.)
Stevens 563
ingens (Sleum.) Stevens 563
mindanaensis Merr. 563
napuensis P.F. Stevens 563
Steinii Sleum. 563
wisselensis P.F. Stevens 563

Dipterocarpaceae 237—552, 240

map, 561—562
subfam. Dipterocarpoideae
237, 240 map, 242, 266—
270, 273, 274, 279, 282,
283, 561
tribe Dipterocarpeae 275,
276, 277, 281, 290
tribe Shoreae 276, 280, 290
subfam. Monotoideae 237,
240 map, 241, 266, 268,
270, 274, 276, 278, 282,
283
subfam. Pakaraimoideae 237,
240 map, 266, 268, 270,
276, 278, 282

Dipterocarpophyllum humei Se-

ward 242
zeraibense Seward 242

Dipterocarpoxylon 241, 242

africanum Bancroft 242
annamense Colani 241

Dipterocarpus Gaertn. f. 239,

241, 242, 245, 246, 247,
258, 259, 261, 263, 265—
269, 271, 273—280, 283,
289, 290, 291, 294*, 297
map, 561

sect. Alati Dyer 291, 293

sect. Angulati Dyer 291, 293

sect. Plicati Dyer 291, 293

sect. Sphaerales Dyer 291, 293

sect. Tuberculati Dyer 291,
293

acutangulus Vesque 294*, 297,
322

affinis Brandis 301

alatus Roxb. 247, 249, 269,
27S

alatus (non Roxb.) Foxw. 318,
325

angulatus Dyer 320

angustialatus Heim 303

angustifolius W. & A. 321

appendiculatus Scheff. 324

appendiculatus (non Scheff.)
Dyer 322

applanatus Sloot. 295, 310

apterus Foxw. 247, 293, 312,
561

artocarpifolius Pierre ex
Laness. 271, 321

balsamiferus Bl. 307

bancanus Burck 303

basilanicus Foxw. 324

baudii Korth. 243, 271, 293,
303

584

(Dipterocarpus)
beccarianus Vesque 311
beccarii Dyer 311
var. glabrata Dyer 311
blancoi Bl. 317
borneensis Sloot. 253, 256,
293, 295, 319
caudatus Foxw. 293, 305
ssp. caudatus 305
ssp. penangianus (Foxw.)
Ashton 305
caudiferus Merr. 293, 299
chartaceus Sym. 293, 304
cinereus Sloot. 295, 316
concavus Foxw. 297, 315,
322, 323, 326
confertus Sloot. 295, 315,
323, 326
confertus (non Sloot.) Sloot.
321.322
conformis sensu Ashton 321,
323
conformis Sloot. 278, 295,
321
ssp. borneensis Ashton 321
ssp. conformis 321, 322
coriaceus Sloot. 251, 253, 297,
324
cornutus Dyer 271, 294*, 295,
311, 312
costatus Gaertn. f. 243, 244,
256, 271, 293, 295, 321,
325
costulatus Sloot. 293, 295, 310
crinitus Dyer 293, 299, 300*,
304
cuneatus Foxw. 305
cuspidatus Ashton 297, 324
dryobalanops Steud. 375
duperreana Pierre 303
dyeri Pierre 243, 295, 315
dyeri (non Pierre) Foxw. 324
elongatus Korth. 256, 271,
293, 294*, 295, 304, 310,
312, 561
eurynchoides Scheff. 324
eurynchus Miq. 297, 322, 324,
325
exalatus Sloot. ex Wood 309
fagineus Vesque 251, 293,
295, 315, 316, 320
fulvus Bl. 302
fusiformis Ashton 295, 319
geniculatus sensu Ashton 321
geniculatus Vesque 295, 320
ssp. geniculatus 320
ssp. grandis Ashton 255,
321
gibbosus Sloot. 311
glabrigemmatus Ashton 295,
318
glandulosus Thw. 321, 325

FLORA MALESIANA

(Dipterocarpus)

globosus Vesque 253, 293,
295, 310, 311, 322
gracilis Bl. 239, 243, 244, 245,
256, 293, 301, 304, 307,
308, 325, 326
grandiflorus (Blco) Blco 243,
256, 294*, 295, 310, 317,
326
griffithii Miq. 317
guiso Blco 448
hasseltii Bl. 239, 243, 274,
293, 301, 306*, 307*
helicopteryx Sloot. 322
hirtus Vesque 299
hispidus Thw. 304
hispidus (non Thw.) F.-Vill. 303
humeratus Sloot. 268, 295,
311, 312
insignis Thw. 322
insularis Hance 321
intricatus Dyer 243, 271
junghuhnii Becc. 375
kerrii King 239, 243, 256, 293,
305
kunstleri King 271, 293, 294*,
295, 301, 309, 310
kutaianus Sloot. 299
lamellatus Hook. f. 295, 312
lampongus Scheff. 307
lasiopodus Perk. 301
lemeslei Vesque 321
littoralis Bl. 243, 293, 309
lowii Hook. f. 295, 313, 314*
macrocarpus Vesque 308
macrorrhinus Sloot. 299
malaanonan Blco 383
mangachapoi Blco 364
marginatus Korth. 302
mayapis Blco 333
microcarpus Zipp. ex Miq.
334
motleyanus Hook. f. 317
mundus Sloot. 294*, 295, 319
nobilis Dyer 323
nudus Vesque 282, 295, 320
obconicus Foxw. 305
oblongifolius Bl. 246, 253,
DS Os iilin295 aos
obtusifolius Teysm. ex Miq.
243, 255, 271, 278, 293,
304
var. glabricalyx Smitinand
304
var. subnudus Ryan & Kerr
304
var. vestitus Smitinand 304
ochraceus Meijer 255, 297,
325
orbicularis Foxw. 293, 326
pachyphyllus Meijer 294*,
295, 313

[ser. I, vol. 93

(Dipterocarpus)

palembanicus Ashton 319
palembanicus Sloot. 295, 318
ssp. borneensis Ashton 319
ssp. palembanicus 318
palosapis Blco 517
parallelus Korth. ex Burck 330
parviflora Zipp. 334
parvifolius Heim 321
penangianus Foxw. 305
pentagonus DC. 307
pentapterus Dyer 320
perakensis Ashton 297, 325
perturbinatus Foxw. 305
philippinensis Foxw. 243, 297,
325
pilosus Roxb. 301
pilosus (non Roxb.) Brandis
303
pilosus (non Roxb.) F.-Vill.
301
plagatus Blco 423
polyspermus Blco 514
prismaticus Dyer 316
pseudofagineus Foxw. 316
pterygocalyx Scheff. 318
pubescens K. & V. 308
pulcherrimus Ridl. 317
punctulatus Pierre 304
quinquegonus BI. 307
retusus Bl. 239, 243, 244, 245,
255, 293, 294*, 308, 309
retusus (non Bl.) Ridl. 311
rigidus Rid]. 253, 295, 310,
311
rotundifolius Foxw. 293, 304
sarawakensis (Browne) Sloot.
253, 271, 297, 323
schmidtii Heim 303
scortechinii King 303
semivestitus Sloot. 295, 316
skinneri King 303
skinneri (non King) Ridl. 305
var. hirtus Ridl. 303, 304
sp. Ashton 318
spanoghei Bl. 308
var. cordata Burck 308
speciosus Brandis 271, 309,
310
stellatus Vesque 282, 297, 323,
326
ssp. parvus Ashton 323
ssp. stellatus 323
stenopterus Vesque 317
subalpinus Foxw. 307, 308
sublamellatus Foxw. 293, 297,
322
tampurau Korth. 306
tampurau (non Korth.) Burck
299
tawaensis Sloot. 322
tempehes Sloot. 247, 293, 297

1982]

(Dipterocarpus)
teres Steud. 375
thurifer Blco 333
tonkinensis A. Chev. 308
trinervis Bl. 308
var. canescens Bl. 308
var. elegans Bl. 308
trinervis (non Bl.) Foxw. 307
tuberculatus Roxb. 243, 269,
AYA), PAN PAKS
turbinatus Gaertn. f. 239,
271, 304
turbinatus (non Gaertn. f.) F.-
Vill. 333
undulatus Vesque 313
ursinus Sloot. 311
validus Bl. 293, 301, 302*,
561
validus (non Bl.) Brandis 311
vanderhoevenii K. & V. 303
velutina Vidal 303
verbeekianus 241
vernicifluus Blco 302
verrucosus Foxw. ex Sloot.
293, 296*, 297, 298*
vestitus Wall. 304
warburgii Brandis 271, 301
woodii Merr. 301
zeylanicus Thw. 293
Disporopsis Hance 189, 191,
193, 219
fusco-picta Hance 219, 220*
Disporum Salisb. 189, 190, 191,
193, 217
calcaratum D. Don 219, 220
cantoniense (Lour.) Merr. 218*
chinense (Ker-Gawl.) O. K.
219
horsfieldii D. Don 218
leschenaultianum D. Don 218
luzoniense Merr. 219
multiflorum (Bl.) D. Don 219
pullum Salisb. 218
var. multiflorum Ridl. 218
pullum (non Salisb.) Merr.
219
Doona Thw. 269, 274, 275, 277,
436, 552
Javanica Burck 421
micrantha Burck 421
multiflora Burck 475
odorata (Roxb.) Burck 421
Dracaena 191, 192
ensata Thunb. 207
ensifolia L. 207
graminifolia (L.) L. 225
Drapiezia Bl. 217
multiflora Bl. 218
var. albiflora Zoll. 218

Drosera L. 562

banksii R. Br. ex DC. 562
peltata J.E. Smith 562

Index to scientific plant names 585

Droseraceae 4: 377—381; 5: 557;
6: 943; 9: 562
Dryobalanops Gaertn. f. 237,
239, 241, 242, 246, 259,
261, 263, 265—269, 271,
274-282, 284, 288-291,
371, 373 map, 391
abnormis Sloot. 373
aromatica Gaertn. f. 246, 253,
256, 271, 273, 289, 371,
Sy, SE, BoB, SUG,
Sie
beccariana Ridl. 373, 375
beccarii Dyer 289, 372, 375
camphora Colebr. 375
fusca Sloot. 253, 256, 372,
377
hallii Korth. 330
javanica 241
kayanensis Becc. 374
keithii Sym. 372, 373
lanceolata Burck 245, 289,
372, 374
neglectus Korth. ex Burck 421
oblongifolia Dyer 247, 271,
279, 372, 373, 374, 379
ssp. oblongifolia 373
ssp. occidentalis Ashton 373
oblongifolia (non Dyer)
Wyatt-Smith 378
oiocarpa Sloot. ex Heyne 375
oocarpa Sloot. 377
ovalifolia 1.H. Burkill 373
rappa Becc. 251, 253, 256,
372, 377, 378*
schefferi Hance 350
sericea Korth. 420
spectabilis 241
vriesti Becc. 375
Dryobalanoxylon 241
Duvaliella Heim 292
problematica Heim 316
Dyerella Heim 340

Elaeocarpaceae 275, 281, 282
Elaeocarpus 275
Elaeogene Miq. 345, 349
sumatrana Miq. 351
Elatinaceae 4: 203—206
Elatineae 275
Eleutherococcus Maxim. 553
malayanus (M.R. Henders.)
Stone 554
trifoliatus (L.) S.Y. Hu 554
Elisma natans 182
Ellipanthus beccarii Pierre 558
var. beccarii 558
Engelhardtia selanica Bl. 505
Epacridaecae 6: 422—444, 943
Eremolaena boinensis 282
Eremopanax Baill. 53

Ericaceae 6: 469—914; 7: 827; 8:
549; 9: 562-563
Eriophorum comosum (Wall.)
Nees 187
filamentosum Boeck. 187
Erythroxylaceae 5: 543—552; 8:
549
Eschweileria Zipp. ex Boerl. 31
barbata (Becc.) Boerl. 41
boerlagei Warb. 46
carpophagarum (Becc.) Boerl.
45
elegans Ridl. 38
gawadensis Baker f. 51
geelvinkiana (Becc.) Boerl. 38
helleborina (Becc.) Boerl. 39
insidiatrix (Becc.) Boerl. 45
insignis (Miq.) Boerl. 52
novoguineensis (Scheff.)
Boerl. 43
palmata Zipp. ex Boerl. 42
pfeilii Warb. 41
pulcherrima (Vidal) Boerl. 40
teysmannii Boerl. 44
Euaraliopsis Hutch. 91
palmata (Roxb.) Hutch. 94
sumatrana (Ridl.) Hutch. 91
Euphoria malaanonan Blco 448
vel Nephelium Blco 448
Eupteron Miq. 7, 72, 78
nodosa (Bl.) Miq. 78
Eustrephus 191, 193
celebicus (Bl.) D. Dietr. 209
Javanicus (Bl.) D. Dietr. 209
Exocarya Benth. 186
montivaga Domin 187
scleroides (F.v.M.) Benth.
185*, 187

Fagaceae 7: 265—403; 8: 549; 9:
253, 255, 563
Fagara ? avicennae Lamk 105
Fagraea
sect. Cyrtophyllum 567
sect. Fagraea 567
auriculata Jack 567
ssp. auriculata 567
ssp. borneensis 567
ssp. parviflora 567
blumei 567
fragrans Roxb. 567
gracilliflora Leenh. 567
resinosa Leenh. 567
ridleyi K. & G. 567
tubulosa Bl. 567
Fatsia 3
Fernandoa macroloba (Miq.)
Steen. 554
Fimbristylis eragrostis (Nees)
Hance 560
fusca (Nees) Clarke 560
malayana Ohwi 155

586

Flacourtiaceae 5: 1-106, 565; 6:
943—944; 7: 827; 9:
563—564

Flagellaria indica L. 195

Flagellariaceae 4: 245—250; 5:
557; 9: 564

Flueggea Rich. 225

wallichiana Kunth 226

‘Fluggea’ 225

Folium polypi mas (et femina?)
Rumph. 42

Fritillaria cantoniensis Lour. 218

Funckia Willd. 232

Gahnia javanica 179
Gastonia Comm. ex Lamk 1, 2,
4, 5, 7, 8, 9, 67, 68 map
boridiana Harms 70
eupteronoides T. & B. 68
papuana Mig. 68
‘sasuroides’ 35
saururoides Roxb. 35
serratifolia (Miq.) Philipson
66*, 68 map
simplicifolia Zipp. ex Seem.
33

spectabilis (Harms) Philipson
2, 3, 8, 68 map, 69*, 70*
sundaica (Miq.) Baill. 99
winkleri Harms 68
Geitonoplesium 191, 193
Gelibia Hutch. 72, 77
branderhorstii (Harms)
Hutch. 77
elegans (C. Moore & F.v.M.)
Hutch. 77
Geniostoma Forst. 568
antherotrichum Gilg & Bened.
568
var. archboldianum (Merr.
& Perry) B.J. Conn 568
archboldianum Merr. & Perry
568
arfakense Kan. & Hat. 568
leenhoutsii B.J. Conn 568
rupestre 568
Gentianaceae 567
tribe Tachiineae 567
Geomitra Becc. 554, 555
clavigera Becc. 555
Geraniaceae 6: 445—449; 9:
565—566
Geranium L. 565
ardjunense Z. & M. 565
balgooyi Veldk. 565
editum Veldk. 565
frigidurbis Moerman 565
homeanum Veldk. 565
hyperacrion Veldk. 565
lacustre Veldk. 565
leptodactylon Veldk. 565
monticola Ridl. 565

FLORA MALESIANA

(Geranium)
nepalense Sweet 565
niuginiense Veldk. 566
papuanum Ridl. 565
subcompositum Veldk. 565
whartonianum Veldk. 565
wilhelminae Veldk. 565
Gilibertia Ruiz & Pav. 101
borneensis Philipson 102
maingayi Philipson 101
saururoides DC. 35
Gladiolus odoratus indicus
Rumph. 207
Gloriosa L. 189, 190, 191, 193
superba L. 190, 193, 194*,
195*
virescens Lindl. 195
Gnetaceae 4: 336—347; 6:
944—949
Gomphandra capitulata (Jungh.
& de Vriese) Becc. 22
Gonocaryum Miq. 566
macrophyllum (BI.) Sleum.
566
Goodeniaceae 5: 335—344; 6:
949—952; 7: 827; 9: 566
Grewia 281
Griselinia 6, 16
Griseliniaceae 16
Guttiferae 275, 280, 281, 282
Guttiferales 275, 281, 282
Gymnosiphon BI. 554
aphyllus Bl. 554

Haemodoraceae 5: 111-113
Halongia Jeanplong 211
purpurea Jeanplong 211
Haloragaceae 7: 239—263, 828
Halosarcia P.G. Wilson 557
indica (Willd.) P.G. Wilson
557
ssp. indica 557
ssp. leiostachya (Bth.) P.G.
Wilson 557
Hamamelidaceae 5: 363—379; 6:
952
Hancea Pierre 391, 397
beccariana Pierre 407
‘beccarii’ 407
cernua Pierre 398
dryobalanoides Pierre 403
griffithii Pierre 406
mengerawan Pierre 400
micrantha Pierre 401
microptera Pierre 407
pierrei Pierre 405
Hanguana major Shaw 564
malayana (Jack) Merr. 235,
564
var. anthelminthica (BI.)
Bakh. f. 564
Haploclathra 281

[ser. I, vol. 93

Haplophragma macrolobum
(Miq.) Steen. 554
Harmsiopanax Warb. 1, 3, 4, 6,

7, 8,9
aculeatus (Bl.) Warb. ex
Boerl35 1125
harmsil K. Sch. 3, 11
ingens Philipson 3, 11, 13*
ssp. ingens 10*, 13, 14
ssp. moniliformis Philipson
14
Hedera 2, 4, 101
australiana F.v.M. 83
disperma (Bl.) DC. 87
glomerulata (Bl.) DC. 94
nodosa (Bl.) Hassk. 78
polyacantha Wall. 93
umbelliferum (Lamk) DC. 35
Hederopsis Clarke 7, 86
maingayi Clarke 87
major Tidl. 87
Heimiatoma Pierre 292
Helmholtzia novoguineensis
(Krause) Skottsb. 235
Helosciadium /eptophyllum
(Pers.) DC. 569
Heteromorpha 6
‘Heterophragma macrolobium’
554
Heterosmilax 191, 192
Homalium Jacq. 563
acutissimum Gilg 564
bismarckense Craven 564
caput-avis Craven 564
dentrecasteauxense Craven
564
foetidum (Roxb.) Bth. 564
maneauense Craven 564
reductum Craven 564
streimannii Craven 564
subcordatum Craven 564
tatambense Sleum. 564
velutinum Craven 564
Homalostachys sinensis Boeck.
134
Hopea Roxb. 237, 239, 244,
245, 246, 255, 257, 258,
259, 261, 263, 265—269,
271, 273, 275—280, 283,
284, 288, 289, 291, 327,
388, 391, 398 map
sect. Dryobalanoides Miq.
257, 280, 281, 392*, 397,
406, 410*
subsect. Dryobalanoides
392, 393, 397
subsect. Sphaerocarpae
(Heim) Ashton 393, 394,
409, 429
sect. Euhopea Miq. 415
sect. Hancea (Pierre) Heim
391, 397

1982]

(Hopea)
sect. Hopea Ashton 277, 280,
388, 406, 415
subsect. Hopea 257, 388,
393, 395, 415, 416*, 418,
429
subsect. Pierrea (Heim)
Ashton 259, 393, 396,
418, 429, 430*, 562
sect. Petalandra Heim 415
Bracteata group Sym. 409
Euhopea group Sym. 415
Nervosa group 429
Pierrea group Sym. 429
acuminata Merr. 243, 394*,
395, 420
acuminata (non Merr.) Ash-
ton 423
aequalis Ashton 394, 409
albescens Ridl. 483
albescens (non Ridl.) Foxw.
421
altocollina Ashton 394, 409
andersonii Ashton 396, 425
ssp. andersonii 425
ssp. basalticola Ashton 426
anomala Foxw. 421
apiculata Sym. 396, 430*, 431
aptera Ashton 253, 395, 418
argentea Meijer 398
aspera de Vriese 548
auriculata Foxw. 395, 413
avellanea Heim 405
balangeran Korth. 509
bancana (Boerl.) Sloot. 396,
424
basilanica Foxw. 396, 425
beccariana Burck 243, 251,
269, 271, 390*, 392*,
393*, 404, 406*, 407
beccariana (non Burck) Sym.
404
bilitonensis Ashton 243, 253,
396, 430*, 433
borneensis Heim 403
brachyptera (Foxw.) Sloot.
395, 414
bracteata Burck 395, 410*,
414, 415
bullatifolia Ashton 396, 433,
434
cagayanensis (Foxw.) Sloot.
243, 396, 431
celebica Burck 243, 244, 396,
427
celebica (non Burck) Diels 419
celtidifolia Kosterm. 395, 406,
415
centipeda Ashton 253, 395,
416*, 423
cernua T. & B. 253, 255, 393,
398, 400

Index to scientific plant names

(Hopea)

coriacea Burck 253, 393, 398

curtisii King 421

dalingdingan Gutierrez 403

dasyrrhachis Sloot. 253, 395,
417, 424

dasyrrhachis (non Sloot.) Ash-
ton 423

dealbata Hance 418

decandra Buch.-Ham. ex
Wight 423

depressinerva Ashton 395, 420

diversifolia Miq. 426

diversifolia (non Miq.) Scheff.
421

dolosa Sloot. 427

dryobalanoides Miq. 264*,
286*, 390*, 393, 402, 403,
404

dyeri Heim 392*, 394, 407

enicosanthoides Ashton 251,
396, 436

fagifolia Miq. 241, 421

faginea Wall. 360

ferrea Laness. 243, 253, 256,
267, 392, 395, 416*, 421

ferruginea Parijs 392*, 393,
404

floribunda Wall. 493

fluvialis Ashton 256, 393, 399

forbesii (Brandis) Sloot. 243,
DAA~-2535 255; 2565395;
416*, 417

foxworthyi Elmer 393, 397,
403

foxworthyi (non Elmer) Foxw.
403

garangbuaya Ashton 399

glabra W. & A. 271

gabrifolia C.T. White 243,
256, 396, 428

glaucescens Sym. 396, 429

globosa Brandis 421

glutinosa Elmer 397

gracilis Miq. 436

grandiflora Wall. 360

gratissima Wall. 487

gregaria Sloot. 243, 244, 396,
428

griffithii Kurz 243, 256, 394,
397, 406

var. pedicellata Brandis

406, 408

grisea Brandis 482

hasskarliana Heim 421

heimiana Brandis 483

helferi (Dyer) Brandis 243,
395, 418

hongayanensis Tard. 429

hopeifolia (Heim) Sloot. 483

inexpectata Ashton 394, 405

intermedia King 407

587

(Hopea)

intermedia (non King) Brandis
408
intermedia (non King) Foxw.
404
iriana Sloot. 243, 255, 259,
396, 428
javanica (Burck) Heim 421
johorensis Sym. 393, 404
kelantanensis Sym. 399
kerangasensis Ashton 253,
393, 401
laevifolia Parijs 461
lanceolata de Vriese 451
latifolia Sym. 269, 271, 393,
404, 562
laxa Sym. 433
linggensis Boerl. 456
longiflora Brandis 473
longifolia (non Dyer) Merr.
473
longirostrata Ashton 393, 399
lowii Dyer ex Brandis 421
lowii (non Dyer) Foxw. 424
macrophylla de Vriese 523
malibato Foxw. 243, 393, 398,
402*, 403, 404*
maquilingensis Foxw. 420
maranti Miq. 540
megacarpa Ashton 396, 426
mengerawan Mig. 393, 400*
mengerawan (non Miq.) Bran-
dis 408
mesuoides Ashton 395, 413
micrantha Hook. f. 253, 393,
401
micrantha (non Hook. f.)
Burck 407
micrantha (non Hook. f.)
Dyer 404
micrantha (non Hook. f.)
Foxw. 399
micrantha (non Hook. f.)
Hance 405
micrantha (non (Hassk.)
Hook. f.) Heim 421
micrantha (non Hook. f.)
King 403, 408
microcarpa Heim 398
microptera Dyer ex Brandis
407
mindanensis Foxw. 396, 435
minima Sym. 414
var. penangiana Sym. 414
var. perakensis Sym. 414
minutiflora Fischer 421
montana Sym. 395, 413
multiflora (Burck) Brandis
475
var. venosa Boerl. 475
multiflora (non Brandis)
Foxw. 421, 426

[ser. I, vol. 93

588 FLORA MALESIANA
(Hopea) (Hopea)
myrtifolia Miq. 392*, 394, pubescens Ridl. 271, 393, 397
408 quisumbingiana Gutierrez 393,
myrtifolia (non Miq.) Heyne 398
404 resinosa Sym. 433

nabirensis Sloot. 428

nervosa King 271, 394, 397,
409, 410*, 412*

nervosa (non King) Foxw. 411

nicholsoni Heim 407

nigra Burck 395, 411

nodosa Sloot. 396, 427

novoguineensis Sloot. 243,
395, 419, 428

nutans Rid]. 259, 269, 271,
396, 424

oblongifolia Dyer 429

odorata Roxb. 243, 256, 263,
269, 270, 271, 393, 395,
422*

odorata (non Roxb.) Foxw.
423

odorata (non Roxb.) Hance
421

odorata (non Roxb.) Vidal
434

ovalifolia Boerl. 449

ovalifolia (non Boerl.) Foxw.
451

ovoidea Ashton 396, 426

pachycarpa (Heim) Sym. 391,
396, 415, 432*, 434*

papuana Diels 395, 419

parviflora Bedd. 259, 423

parvifolia (Warb.) Sloot. 436

paucinervis Parijs 396, 431

pedicellata (Brandis) Sym.
243, 394, 408

pentanervia Sym. ex Wood
253, 255, 396, 425

philippinensis Dyer 243, 396,
430*, 434

Dhilippinensis (non Dyer) Ash-
ton 435

pierrei Hance 243, 256, 394,
403, 405, 562

pierrei (non Hance) Brandis
407

pierrei (non Hance) Foxw.
397, 403, 404

pierrei (non Hance) Ridl. 406,
407, 408

plagata (Blco) Vidal 243, 388,
391, 395, 416*, 423, 424,
426, 562

plagata (non (Blco) Vidal)
Foxw. 426

plagata (non Vidal) Sym. 426

polyalthioides Sym. 396, 431,
433

pterygota Ashton 253, 396,
434

reticulata Tardieu 429
rudiformis Ashton 394, 409
samarensis Gutierrez 396, 427
sangal Korth. 241, 243, 271,
288*, 395, 416*, 420, 421
sarawakensis Heim 403
scabra Ashton 395, 419
scabra Buch.-Ham. 419
scaphula Roxb. 335
selanica W. & A. 505
semicuneata Sym. 396, 426
seminis de Vriese 451
sericea (Korth.) Bl. 420
siamensis Heim 408
similis Sloot. 395, 417
singkawang Miq. 537
siranda Miq. 436, 562
sp. Sloot. ex Merr. 426
sp. ‘“Gyam’ Foxw. 423
sp. nov. Sym. 429
sp. nov. aff. H. pachycarpa
(non Sym.) Meijer &
Wood 409
sphaerocarpa (Heim) Ashton
395, 411
splendida de Vriese 522
squamata Turcz. 517
suava Wall. ex DC. 466
subalata Sym. 263, 269, 270,
271, 393, 394*, 395, 413
subalata (non Sym.) Ashton
413
sublanceolata Sym. 394, 397,
409, 411
sulcata Sym. 393, 399
sumatrana King ex Gilg 436
tangili Bl. 514
tenuinervula Ashton 396, 435
treubii Heim 253, 394, 406,
410*
ultima Ashton 243, 395, 418
vaccinifolia Ridl. ex Ashton
253, 395, 410*, 414
vasta Wall. ex DC. 423
vesquei Heim 253, 393, 401
wightiana Wall. ex W. & A.
263, 271
woodiana Gutierrez 403
wyatt-smithil Wood ex Ashton
396, 429, 430*
Hopeoides Cretzoiu 327, 335
scaphula Cretzoiu 335
Hopeoxylon 241
‘Hoppea’ 391
Horsfieldia Bl. ex DC. 9
aculeata (Bl.) DC. 11
peltata Bth. 11

Hydrocaryaceae 4: 43—44
Hydrocharitaceae 5: 381—413,
569; 6: 952; 7: 828; 9: 566
Hydromystria G. Meyer 566
laevigata (H. & B. ex Willd.)
Diaz-Miranda & Philcox
566
Hydrophyllaceae 4: 207—209
Hypericaceae 8: 1—29
Hypolytrum 134, 186, 187
humile (Steud.) Boeck. 136
nemorum (Vahl) Spreng. 235
Hypoxis spicata Thunb. 232

Icacinaceae 7: 1—87, 828; 9: 566
Indokingia Hemsl. 67
Iphigenia Kunth 189, 190, 191,
193, 197
indica (L.) A. Gray ex Kunth
190, 198*
Ipomoea campanulata L. 558
carnea Jacq.
ssp. fistulosa (Mart. ex
Choisy, in DC.) D.
Austin 558
crassicaulis (Bth.) B.L. Robin-
son 558
fistulosa Mart. ex Choisy in
DC. 558
illustris (Clarke) Prain 558
macrantha R. & S. 558
tuba (Schlechtend.) G. Don
558
Iridaceae 8: 77—84
Irvingia F.v.M. 72
australiana (F.v.M.) F.v.M.
84
Isauxis (Arn.) Reichb. 345, 349
Isoptera Scheff. ex Burck 276,
436, 447
borneensis Scheff. ex Burck
247, 451
borneensis (non Scheff. ex
Burck) King 453
burckii Boerl. 448
dasyrrhachis Sloot. ex den
Berger & Endert 417
seminis Burk. 451
sumatrana Sloot. ex Thore-
naar 453

Juglandaceae 6: 143—154, 953
Juncaceae 4: 210—215; 5: 557; 6:
953; 9: 566
Juncus balticus Willd. 566
bufonius L. 566
inflexus L. 566
nupela Veldk. 566
Juncaginaceae 4: 57; 5: 554

Kalopanax resectum Miq. 91, 93
sumatranum Migq. 91

1982]

Kielmayera 281
Kissodendron Seem. 7, 72, 81
australianum (F.v.M.) Seem.
83, 84
australianum (non (F.v.M.)
Seem.) Boerl. 81
bipinnatum Gibbs 81
Klotzschia 5
Kobresia 134

Labiatae 8: 301—394; 9:
566-567

Lauraceae 280

Laurineae d. Pterigiae 275

Laxmannia sessiliflora Decne
235

Leeaceae 7: 755—782

Leguminosae 557

Lemnaceae 7: 219—237

Lentibulariaceae 8: 275—300

Leonurus indicus L. 567

Lepironia 187

Lethea Norona 217

Leucas decemdentata (Willd.) J.

Sm. 567
flaccida R. Br. 567
indica R. Br. ex Spreng. 567
lavandulaefolia J. Sm. 566
lavandulifolia J.E. Sm. 566
linifolia (Roth) Spreng. 566
Sstachyoides Spreng. 567
Leucopogon juniperinum 270
Ligustrum 3
Liliaceae 189—235, 567
subfam. Aletroideae 191
tribe Narthecieae 191
subfam. Allioideae 191
subfam. Asparagoideae 191
tribe Asparageae 191
tribe Aspidistreae 191, 220,
Pepa
tribe Convallarieae 191
tribe Polygonatae 191
subfam. Asphodeloideae 191
tribe Asphodeleae 191
tribe Asphodelinae 191
tribe Dianell(in)eae 191
tribe Johnsonieae 191
tribe Lomandreae 191
subfam. Dracaenoideae 191
tribe Dracaeneae 191
tribe Milliganieae 191
subfam. Lilioideae 191
tribe Tulipeae 191
subfam. Luzuriagoideae 191
subfam. Melanthoideae 191
tribe Anguillariaceae 191
tribe Iphigenieae 191
tribe Petrosavieae 191
tribe Tricyrtae 191, 197
tribe Tricyrtideae 191, 197
tribe Uvularieae 191

Index to scientific plant names

(Liliaceae)
subfam. Mondoideae 191
tribe Ophiopogoneae 191
tribe Peliosantheae 191
subfam. Smilacoideae 191
Liliales 199
Lilium L. 189—192, 214
longiflorum Thunb. 214, 215
‘longifolium’ 214
‘philippense’ 215
philippinense Baker 214, 215
Limnobium Rich. ex Steud. 566
laevigata 566
stoloniferum 566
Linaria alpina 182
Lipocarpha chinensis (Osb.)
Kern 560
Liriope Lour. 189—192, 223
brachyphylla Merr. 230, 231
graminifolia (L.) Baker 225,
228*
muscari (Decne) L.H. Bailey
225
muscari (non (Decne) L.H.
Bailey) Hatus. 225
spicata Lour. 225
Litsea palustris 499
Lobelia 556
victoriensis v. Royen 556
Loganiaceae 6: 293-387, 953,
960; 7: 828—829; 9:
567-568
tribe Potalieae 567
Lomandra 187, 191, 192
Lophira Banks ex Gaertn. 275
Lophiraceae 275
Lophopyxidaceae 7: 89—91
Lourya Baill. 227
campanulata Baill. 229
Luzula DC. 566
campestris (L.) DC. 566
var. australasica 566
papuana M.E. Jansen 566
philippinensis M.E. Jansen
566
Luzuriaga 191
Lythraceae 275

Macadamia hildebrandii Steen.
568
Machaerina rubiginosa 187
Mackinlaya F.v.M. 1, 6, 7, 9, 27
amplifolia Hemsl. 27
brassii Philipson 29
celebica (Harms) Philipson 4,
DIS LBS
digitata (Merr.) Philipson 29
klossii Philipson 29
radiata Philipson 27, 28, 29
schlechteri (Harms) Philipson
4, 27, 29
subulata Philipson 29

589

(Mackinlaya)
versteegii (Harms) Philipson
29
warburgii (Harms) Philipson

Macropanax Mig. 2, 7, 9, 38, 86
concinnus Mig. 86, 87
cyrtostylum Miq. 95
dispermus (Bl.) O. K. 86
floribunda Mig. 87
glomerulatum (Bl.) Miq. 95
maingayi (Clarke) Philipson
86, 87, 88*

oreophilus Miq. 87

sp. Vidal 56

undulatus (Wall. ex G. Don)
Seem. 87

Mahurea 281

‘Majau’ Durant 492

Malpighiaceae 5: 125—145, 566;

6: 960

Malvaceae 275, 280

Malvales 274, 280, 281, 282

Mapania Aubl. 560
cuspidata (Miq.) Uittien 560
macrocephala (Gaudich.) K.

Sch. 560

Marcgraviaceae 275

Marila 281

Marquesia Gilg 237, 240 map,

274, 279, 282
excelsa R.E. Fr. 282
Melanthium cochinchinense
Lour. 216
indicum L. 198
Memecylanthes Gilg & Schltr
556

Meryta 89
colorata F.M. Bailey 105
spathipedunculata Philipson

34
Mesua acuminatissima (Merr.)
Kosterm. 370

Meta-aletris Masam. 230
rigida (Stapf) Masam. 230
sumatrana (Masam.) Masam.

230

Metanarthecium Maxim. 230

brachyphyllum (Merr.) Ma-
sam. 230

Methonica Tourn. ex Crantz 193
superba (L.) Crantz 195

Miyoshia Makino 199
sakurii Makino 200

Miyoshiaceae 199

Miyoshiales (order) 199

Mocanera Blco 292, 327
grandiflora Blco 317
guiso Blco 448
malaanonan Blco 383
mangachapoi Blco 364
mayapis Blco 333

590

FLORA MALESIANA

(Mocanera)
plagata Blco 423
polysperma Blco 514
thurifera Blco 333
verniciflua Blco 292, 302
Mondo Adans. 225
graminifolia (L.) Koidz. 225
japonicum (L.f.) Farwell 227
malayanum (Ridl.) Farwell
226
Monoporandra Thw. 263, 275,
279
Monotes A. DC. 237, 240 map,
241, 261, 263, 268, 274,
D275 276s 279, 28, 282;
284, 338
africanus 275
Moringaceae 4: 45—46; 5: 554;
6: 960
Mormoraphis Jack ex Wall. 53
sumatrana Jack ex Wall. 60
Myodocarpus 6

Myoporaceae 4: 265—266; 9: 568

Myoporum papuanum Kraenzl.
568

Myricaceae 4: 276—279
Myrrhidendron 6

Najadaceae 6: 157-171
Nanolirion Bth. 204
Neisandra Rafin. 391, 415
Neobalanocarpus Ashton 239,
246, 263, 265, 266, 269,
271, 214, 276; 278; 288,
289, 291, 388
heimii (King) Ashton 267,
2695 27105 QING 2765 217,
388*, 389*
Neolourya Rodriguez 227
Neuburgia corynocarpa (A.
Gray) Leenh. 568
var. sarcantha (Gilg & Be-
ned.) B.J. Conn 568
sarcantha (Gilg & Bened.)
Leenh. 568
Neuwiedia curtisii Rolfe 223
singapureana (Baker) Rolfe
223
zollingeri Rchb.
var. singapureana (Baker)
de Vogel 223
Nolina javanica Hassk. 205
Nothapodytes foetida (Wight)
Sleum. 566
nimmoniana (J. Grah.) Mab-
berley 566
Nothofagus Bl. 563

sect. Calucechinus (Hombr. &

Jacq.) Krasser 563
subsect. Calucechinus 563
sect. Calusparassus (Hombr.
& Jacq.) Krasser 563

(Nothofagus)
sect. Nothofagus 563
subsect. Antarctica Steen.
563
subsect. Nothofagus 563

subsect. Quadripartitae Steen.

563
betuloides (Mirb.) Oerst. 563
Nothopanax Miq. 72
anisum Miq. 105
cochleatum (Lamk) Miq. 75
crispatum (Bull) Merr. 76
cumingii (Presl) Seem. 76
elegans (C. Moore & F.v.M.)
Seems i
fruticosum (L.) Miq. 73
var. plumatum (Hort.)
Merr. 73

var. victoriae (Hort.) Merr.

73
guilfoylei (Cogn. & Marche)
Merr. 76
macgillivrayi Seem. 74
obtusum (Bl.) Miq. 73
ornatum (Bull) Merr. 76
pinnatum (Lamk) Miq. 76
scutellarium (Burm. f.) Merr.
75
tricochleatum Miq. 75
zippelianum (Miq.) Seem. 81
Nothoscordum Kunth 190, 191,
192, 234
fragrans Kunth 234
inodorum (W. Ait.) G. Nich-

ols. 192, 234
Nyctaginaceae 6: 450—468; 7:
829

Nyssaceae 4: 29-31

Ochnaceae 7: 97—119; 9: 275
Oleoxylon Roxb. 292
Onagraceae 8: 98—113
Ophiopogon Ker-Gawl.
189—192, 225
acaulis (Bl.) Ridl. 226
caulescens (Bl.) Back. 224*,
226, 228*
‘gauliscens’ 226
intermedius D. Don 226, 227
var. macranthum Ridl. 226
japonicus (L. f.) Ker-Gawl.
226. 220228
japonicus (non (L. f.) Ker-
Gawl.) Koord. 226
malayanus Ridl. 226
merrillii Masam. 227
prolifera Lindl. 226
sp. 228*
‘spicata’ 228*

[ser. I, vol. 93

Ophiopogon (non Ker-Gawl.)
Kunth 223
Oplopanax 2
Oreobolus ambiguus Kuk. &
Steen. 561
ktikenthalii Steen. 561
Oreomyrrhis plicata Mathias &
Constance 569, 570*
Oreopanax 2
Oroxylum indicum (L.) Kurz 67
Osmoxylon Mig. 1, 3, 4, 5, 7, 8,
9, 31, 34 map
amboinense Miq. 33, 35
articulatum Philipson 32, 35
barbatum Becc. 32, 41, 42
boerlagei (Warb.) Philipson
33, 44, 46*
borneense Seem. 32, 33, 39
camiguinense (Merr.) Philip-
son 33, 48, 52
carpophagarum Becc. 45, 46
catanduanense (Merr.) Philip-

son 32, 37, 38
caudatum (Merr.) Philipson
323i)

celebicum Philipson 33, 49

cumingii Seem. 51

dinagatense (Merr.) Philipson
32, 34

eminens (Bull) Philipson 33,
40, 48, 49

fenicis (Merr.) Philipson 33,
48

geelvinkianum Becc. 32, 38,
39

globulare Philipson 32, 40, 47,
48

helleborinum Becc. 39

heterophyllum (Merr.) Philip-
son 32, 37, 38

humile (Elmer) Philipson 33,
44, 52

insidiator Becc. 33, 45

insigne (Miq.) Becc. 33, 52

kostermansii Philipson 32, 41

lanceolatum Philipson 32, 35,
36*

lineare (Merr.) Philipson 32,
38, 39, 553

luzoniense (Merr.) Philipson
32, 33, 48, 52

masarangense Philipson 33, 45

micranthum (Harms) Philip-
son’33, 50*:/ Sis 55

miquelii Boerl. 32, 33

moluccanum (Miq.) Becc. 42

novoguineense (Scheff.) Becc.
30*, 33, 43*

oblongifolium Philipson 32,

spicatus (Lour.) Ker-Gawl. 225
wallichianus (Kunth) Hook. f.
226

33, 37
palmatum (Lamk) Philipson
32, 41, 42, 43

1982] Index to scientific plant names 591

(Osmoxylon) Pakaraimaea Maguire & Ashton (Parashorea)

pectinatum (Merr.) Philipson
33, 45, 48, 52

pfeilii (Warb.) Philipson 32,
41

pulcherrimum Vidal ex F.-
Vill. 32, 40, 49

ramosii (Merr.) Philipson 32,
43

serratifolium (Elmer) Philip-
son 33, 49

sessiliflorum (Laut.) Philipson
33, 40, 41, 47, 51

simplicifolium (Elmer) Philip-
son 32, 34

soelaense Philipson 32, 40, 47,
48

spathipedunculatum (Philip-
son) Philipson 32, 34

talaudense Philipson 33, 47,
49

teysmannii (Boerl.) Philipson
33, 44

trilobatum (Merr.) Philipson
33845. '515°52

umbelliferum (Lamk) Merr.
32°35

yatesii (Merr.) Philipson 32,
38

zippelianum (Miq.) Becc.
42

Otophora hullettii Ridl. 358
Oxalidaceae 7: 151—178, 829

Pachychlamys (Dyer ex King)

Dyer ex Ridl. 437, 504
beccarianus Dyer ex Brandis
513
beccarianus (non Dyer ex
Brandis) Ridl. 537
brachypterus Dyer ex Brandis
513
‘ghysbertiana’ 523
gysbertsiana Ridl. 523
hemsleyanus Ridl. 536
thiseltonii Rid]. 537

Pachydiscus Gilg & Schltr 556
Pachynocarpus Hook. f. 345,

349
grandiflorus Ridl. 349
ridleyanus Anders. 352
ridleyanus (non Anders.) Ridl.
352
ruminatus Brandis 352
Stapfianus King 349
umbonatus Hook. f. 349
umbonatus (non Hook. f.)
Ridl. 350, 352
verrucosus Heim 349
verrucosus Ridl. 350
wallichii King 349, 351

237, 263, 266, 268, 279,
282, 283, 561
Palmervandenbroekia Gibbs 7,
72, 84
papuana Gibbs 85
Panax 2
subg. Acanthopanax Decne &
Planch. 103, 554
aculeatus Ait. 103
anisum DC. 105
armatus Wall. ex G. Don 22
bandanense Zipp. ex Span. 76
cochleatum (Lamk) DC. 75
conchifolium Roxb. 75
crispatum Bull 76
cumingiana (Presl) Rolfe 76
elegans C. Moore & F.v.M.
77
fruticosum L. 73
macgillivraei (Seem.) Bth. 74
murrayi (non F.v.M.) F.v.M.
79
obtusum Bl. 73
ornatum Bull 76
palmatum Roxb. 93
pinnatum Lamk 46, 76
rumphii Hassk. 75
scutellarioides Reinw. ex BI.
75
secundum Schult. 76
zippelianum Miq. 81
Pandorea pandorana (Andr.)
Steen. 554
Papaveraceae 5: 114—117
Papaya silvestris Rumph. 78
Parahopea Heim 437, 485
balangeran Heim 509
Paramapania 187
Parashorea Kurz 239, 245, 259,
263, 266, 267, 269, 271,
276, 277, 278, 280, 281,
283, 284, 289—292, 379,
381 map
aptera Sloot. 380, 381
aptera (non Sloot.) Foxw. 380
balangeran (Korth.) Merr. 509
densiflora Sloot. & Sym. 271,
380
globosa Sym. 380, 381
longisperma (Roxb.) Kurz 481
lucida (Miq.) Kurz 380, 383,

387
lucida (non (Miq.) Kurz)
Sloot. 383

macrophylla Wyatt-Smith ex
Ashton 258, 379, 380,
382*
malaanonan (Blco) Merr. 243,
288, 380, 381 map, 383,
384*, 385*, 387
var. tomentella Sym. 385

parvifolia Wyatt-Smith ex
Ashton 380, 382

plicata Brandis 383

poilanei Tard. 383

smythiesii Wyatt-Smith ex
Ashton 267, 380, 382, 387

sp. Browne 382

sp. Thomas 385

stellata Kurz 243, 380, 383

stellata (non Kurz) Sloot. 387

tomentella (Sym.) Meijer 380,
385, 386*, 387

warburgii Brandis 387

Paratropia cumingiana Presl 76
nodosa (Bl.) DC. 78

Passifloraceae 7: 405—434, 829

Pedaliaceae 4: 216—221; 5: 557;

7: 829

Peekeliopanax Harms 7, 67
spectabilis Harms 70

Peliosanthes Andr. 189—192,

227
albida Baker 229
campanulata (Baill.) Rodri-
guez 229
graminea Ridl. 229
grandifolia Ridl. 229
humilis Andr. 229
hypogyna Ridl. 229
javanica (Bl.) Dietr. 229
lurida Ridl. 229
mantegazziana (Pamp.)
Pamp. 228
monticola Ridl. 229
parviflora Ridl. 229
sessiliflora Ridl. 229
stellaris Ridl. 229
sumatrensis Ridl. 229
teta Andr. 228
ssp. humilis (Andr.) Jessop
228*, 229 map
ssp. teta 228, 229 map
var. angustifolia Ridl. 229
var. mantegazziana Pamp.
228
tonkinensis Wang & Tang 229
violacea Wall. ex Baker 229
viridis Ridl. 229

Pentace 281
Pentacme A. DC. 269, 275, 436,

465
contorta (Vidal) Merr. &
Rolfe 466
malayana King 466
mindanensis Foxw. 467
paucinervis Brandis 467
siamensis (Miq.) Kurz 272, 466
var. laevis Pierre 466
var. suavis (DC.) Pierre 466
suavis A. DC. 466
tomentosa Craib 466

592

Pentapanax Seem. 2, 3, 9, 25
elegans Koord. 25, 26*
var. elegans 25
var. puberula (sphalm.) 25
var. pubescens Koord. 25
Pentaphragmataceae 4: 517—528
Pentaphylacaceae 5: 121—124,
566
Pentaphylax auryoides 554
Periomphale Baill. 556
papuana Steen. 557
Perissandra Gagn. 345
Petalandra Hassk. 276, 391, 415
micrantha Hassk. 421
Petasula Noronha 95
Petrosavia Becc. 189—192, 198
sakuraii (Makino) J.J. Smith
ex Steen. 199, 200*
sinil (Krause) Krause 199
stellaris Becc. 199, 200*
Petrosaviaceae 191, 199
Philesiaceae 191, 193
Philydraceae 4: 5—7; 7: 829
Phlomis decemdentata Willd.
567
indicus L. 567
linifolia Roth 566
Phytolaccaceae 4: 228—232; 5:
557
Pierrea Heim non Hance 429
pachycarpa Heim 432
penangiana Heim ex Brandis
389
Pierreocarpus Dunn 391
Pierreocarpus Ridl. ex Sym. 429
pachycarpa Ridl. ex Sym. 433
‘Piliosanthes’ 227
Pimpinella dioica L. 569
leptophylla Pers. 569
Pinus merkusii 568
Pittosporaceae 5: 345—362; 6:
960—963; 7: 829—830; 9:
5, 568
Pittosporum pumilum Schodde
568
Platanaceae 2
Pleomele 192
Pleranda 5, 7
Plumbaginaceae 4: 107—112
Podostemaceae 4: 65—68; 6:
962—964
Polemoniaceae 4: 195—196
Polygonum 263
Polyscias J.R. & G. Forster 1,
22 U5 i Os Pg Cols TP TE!
map, 89
sect. Eupteron (Miq.) Philip-
son 7, 73, 78
sect. Gelibia (Hutch.) Philip-
son 73, 77
sect. Kissodendron (Seem.)
Philipson 73, 81

FLORA MALESIANA

(Polyscias)

sect. Palmervandenbroekia
(Gibbs) Philipson 73, 80,
84

sect. Polyscias 8, 72, 73

acuminata Vidal 79

anisum Harms 105

australiana (F.v.M.) Philipson
81, 83

var. disperma (F.v.M.)

Philipson 84

belensis Philipson 78, 80

bipinnata (Gibbs) Philipson 81

borneensis Philipson 78, 80

branderhorstii Harms 77

caroli Harms 81, 83

cibaria White & Francis ex
Lane-Poole 29

clemensiana Harms 79

cumingiana (Presl) F.-Vill. 64,
13) 14515; 16377

cumingii (Presl) Harms 76

disperma Blco 86

elegans (C. Moore & F.v.M.)
Harms 3, 77

filicifolia (C. Moore) Bailey 76

floribunda Elmer 79

florosa Philipson 78, 79

forbesii Baker f. 79

fraxinifolia Philipson non
Harms 80

fruticosa (L.) Harms 73, 77

var. plumata (Hort.) Bailey

73

gjellerupii Harms 83

grandifolia Volkens 74

guilfoylei (Cogn. & Marché)
L.H. Bailey 73, 76

jacobsii Philipson 84, 85

javanica K. & V. 73, 77

joskei L.S. Gibbs 86

ledermannii Harms 3, 7, 78,
79, 80

macgillivrayi (Seem.) Harms
73, 74

murrayi (F.v.M.) Harms 80

nodosa (BI.) Seem. 7, 71*, 78,
79

obtusa (Bl.) Harms 73

‘palawanensis’ 80

palmervandenbroekii Bernardi
84, 85

papuana (Miq.) Seem. 68

philipsonii Bernardi 78, 80

pinnata J.R. & G. Forster 77

roemeriana Harms 86

royenii Philipson 81, 83, 84

rumphiana Harms 76

schultzei Harms 81*, 82*, 83,
84

scutellaria (Burm. f.) Fosb.
13); 15. 7)

[ser. I, vol. 93

(Polyscias)
sleumeri Philipson 84
sorongensis Gibbs 76
sp. C.T. White 74
tricochleata (Miq.) Fosb. 75
verticillata Stone 73, 75
vogelkopensis Philipson 84,

85

zippeliana (Miq.) Valeton 81
Pontederiaceae 4: 255—261; 5:
557; 6: 964
Portulacaceae 7: 121—123
Premna nimmoniana J. Grah.
566
Primocarex Kuk. 117
Primulaceae 6: 173—192, 964
Proteaceae 5: 147—206, 566; 6:
965; 7: 830; 9: 568
Protolirion Ridl. 198
miyoshia-sakuraii Makino
200
paradoxum Ridl. 199
sakurati (Makino) Dandy 200
sinii Krause 199
Pseudobrassaiopsis Banerjee 91
polyacantha (Wall.) Banerjee
94
Pseudopanax 2, 86
Pseudosandalum O. K. 31
amboinense Rumph. 35
miquelii (Boerl.) O. K. 33
umbelliferum (Lamk) O. K.
35
‘Pseudosantalum’ 31
Pseudosindora palustrus Sym.
21S S52
Pteranthera Bl. 345, 359
mangachapoi DC. 365
sinensis Bl. 365
Pterigium Correa 275, 292, 371
costatum (Gaertn. f.) Correa
292, 321
teres Correa 371, 375
‘Pterygium’ 292
Punicaceae 4: 226—227

Radermachera glandulosa (BI.)
Miq. 554
Reaumuriaceae 275
Restionaceae 5: 416—420, 569
Retinodendron Korth. 345, 349
bancanum King 350
kunstleri King 350
moluccanum Heim 353
pallidum King 358
pauciflorum Korth. 351
rassak Korth. 353
scortechinii King 357
Retinodendropsis Heim 345
aspera Heim 354
Rhipogonum 191, 192
Rhizophora 568

1982]

Rhizophoraceae 5: 429—493; 6:
965—967; 8: 550-551; 9:
568
Rhodea ftupistra Schult. 223
Rhododendron subg. Rhododen-
dron
sect. Vireya (Bl.) Copel. f.
subsect. Euvireya Copel. f.
ser. Buxifolia Sleum. 562
anagalliflorum Wernh. 563
atjehense Sleum. 563
burtii P.J.B. Woods 562
capellae P. Kores 562
frey-wysslingii J.J.S. 562
irroratum Franch. 563
ssp. kontumense (Sleum.)
Chamberlain 563
nervulosum Sleum.
var. exuberans Sleum. 563
pulleanum Koord. 562
roseiflorum P.F. Stevens 562
rubineiflorum Craven 562
ruttenii J.J.S. 562
scortechinil K. & G. 562
stenophyllum Hook. f. 563
vinkii Sleum. 562
Rhuacophila Bl. 207
celebica Bl. 209
javanica Bl. 209
Richetia Heim 275, 437, 468
acuminata Heim 475
coriacea Heim 474
latifolia Heim 475
oblongifolia Heim 475
penangiana Heim 475
Ridleyinda O. K. 437, 447
borneensis O. K. 451
Romnalda 191, 192
Rosaceae 557
Rourea prainiana Talbot 558
Roureopsis asplenifolia Schel-
lenb. 558
Ryparosa javanica (Bl.) Kurz
563

Salacia kalahiensis Korth. 557

Salicaceae 5: 107—110

Salicornia /eiostachya Bth. 557

Salvadoraceae 4: 224—225

‘Sama rupa chengal’ Foxw. 426

Sambucus javanica Bl. 556

Sandersonia 197

Santiria grandiflora Kalkman
555

Sapindaceae 275

Sapotaceae 246

Sarcolaenaceae 268, 275, 282,
283

Sarcospermataceae 4: 32—34; 6:
967

Sarcostigma W. & A. 566

paniculata Pierre 566

Index to scientific plant names

593

Satureia gracilis (Bth.) Loes. 567
Saul Roxb. ex W. & A. 436, 447
iallarea Roxb. 493
Saururaceae 4: 47—48
Scaevola sericea Vahl 566
taccada (Gaertn.) Roxb. 566
Scaphula Parker 327, 335
glabra Parker 335
Schefflera 1—5, 7, 8, 9, 38
sect. Brassaia 3, 4
subg. Agalma 4
actinophylla (Endl.) Harms 3,
8

altigena Frodin 3
aromatica (Bl.) Harms 4
chimbuensis Frodin 3
eriocephala Harms 4
falcata Philipson 3
gemma Frodin 3
longifolia (Bl.) Vig. 8
nervosa (King) Vig. 2, 4
odorata (Blco) Merr. & Rolfe
73
pagiophylla Harms 3
rugosa (Bl.) Harms 2, 3
singularis B.C. Stone 3
stahliana (Warb.) Frodin 3
stolleana Harms 4
straminea Frodin 3
thaumasiantha Harms 3
Scheffleropsis 7
Schelhammera R. Br. 189, 190,
191, 193, 200
multiflora R. Br. 201*
pedunculata 201
undulata 201
Schelhammeria 200
Schellhammera 200
Scheuchzeria palustris 182
Scheuchzeriaceae 199
Schoenoxiphium 134
Schoutenia 268, 276, 281, 282
Schubertia Bl. 9
aculeata Bl. 11
Sciadophyllum palmatum Bl. 99
Scirpodendron 187
Scleria sinensis H. Pfeiff. 134
ustulata F.M. Bailey 187
Scutellaria prima Rumph. 75
secunda Rumph. 64
secunda angustifolia Rumph.
76
secunda latifolia Rumph. 75
tertia Rumph. 73
Scyphostegiaceae 5: 297—299; 6:
967; 7: 830
Sedum L. 558
ambiflorum R.T. Clausen 559
ambiflorum (non R.T. Clau-
sen) Backer 560
australe Merr. 559
australe (non Merr.) Steen.
560

(Sedum)
erythrospermum Hayata 559
ssp. australe Merr. 558,
559*, 560
ssp. erythrospermum 559
formosanum N.E. Brown 558,
559
mariae R.-Hamet 559
parvisepalum Yamamoto
ssp. parvisepalum 560
ssp. philippinense H. Ohba
559*, 560
Seidlia Kostel. 345, 349
‘Semayor’ Durant 505
Shorea Roxb. ex Gaertn. f. 239,
241, 243—247, 258, 259,
261, 263, 265, 266, 268,
269, 271, 273—280, 283,
284, 287*, 288—291, 353,
379, 388, 393, 436, 448
map, 562
sect. Anthoshorea Heim 258,
265. 260) 267 2 279
288, 436, 437, 442, 485,
486*, 488*, 499, 503
sect. Brachypterae Heim 265,
266, 267, 277, 280, 287,
438, 443, 499, 503, 521,
S28), SY, S2Y/, Se)!
subsect. Brachypterae 438,
443, 486*, 504, 506*,
508*
subsect. Smithiana Ashton
438, 443, 503, 504*, 562
sect. Doona 255, 265, 266,
269, 274, 552
sect. Eushorea Brandis 445
sect. Hopeoides Heim 436,
485
sect. Isoptera Scheff. ex
Burck 436
sect. Isoptera (Scheff. ex
Burck) Foxw. 447
ciliata subgroup Sym. 447
sect. Mutica Brandis 247, 258,
266, 267, 277, 279, 280,
287, 436, 439, 444, 503,
527*, 529, 551
subsect. Auriculatae Ashton
439, 444, 472, 528*, 529,

562
subsect. Mutica 258, 439,
445, 534, 552

subsect. ‘Muticae’ 534
sect. Neohopea Ashton 277,
436, 437, 440, 467, 468*
sect. Ovalis Ashton 266, 279,
280, 287, 436, 439, 445,
547*
sect. ‘Pachycarpa’ 521
sect. Pachycarpae Heim 246,
258, 265, 266, 267, 277—

594

(Shorea sect. Pachycarpae)
280, 287, 288, 436, 439,
444, 520*, 521, 522*
sect. Pachychlamys Dyer ex
King 504
sect. Pentacme (A. DC.) Ash-
ton 265, 266, 267, 269,
277, 436, 437, 440, 465,
467*
sect. Pinanga Brandis 436,
521
sect. Richetioides Heim 245,
258, 259, 265, 266, 267,
277, 279, 280, 288, 289,
436, 437, 440, 468
subsect. Polyandrae Ashton
437, 440, 469, 470*
subsect. Richetioides 437,
440, 470, 472*
sect. Rubella Ashton 266, 267,
277, 287, 436, 438, 442,
498*, 500*, 502, 503, 521,
529, 547
sect. Shorea 257, 258, 265,
266, 267, 277, 280, 288,
437, 439, 445, 466, 468,
491
subsect. Barbata Sym. ex
Ashton 437, 440, 461,
462*
subsect. Shorea 437, 439,
446*, 447, 461
subg. Anthoshorea (Heim)
Meijer 485
subg. Richetia (Heim) Meijer
436, 468
subg. Rubroshorea Meijer
266, 436, 534
Balau group Sym. 445
Barbata group Sym. 461
Meranti Damar Hitam group
Sym. 468
Meranti Pa’ang group Sym.
485
Red Meranti group Sym. 499,
503, 529, 547
ovalis subgroup 547
parvifolia subgroup 529
pauciflora subgroup 503
Richetia group Sym. 468
acuminata Dyer 271, 445, 529,
535
acuminatissima Sym. 441, 482
acuta Ashton 253, 444, 533*
agamii Ashton 244, 263, 270,
272, 442, 485, 497
ssp. agamii 497
ssp. diminuta Ashton 497
agsaboensis W.L. Stern 545
alba Ridl. 465
albida Sym. 241, 247, 251,
253, 254*, 256, 259, 288,

FLORA MALESIANA

(Shorea albida)

442, 498*, 499, S501*,
502*, 509, 562
almon Foxw. 252*, 443, 503,
507
almon (non Foxw.) Browne
540
alutacea Ashton 441, 472, 481
amplexicaulis Ashton 444,
521, 524, 525*, 526, 528
andulensis Ashton 443, 514
angustifolia Ashton 441, 472*,
478, 479
angustiloba Foxw. 447
aptera Burck 513
argentifolia Sym. 272, 445,
539
asahii Ashton 279, 440, 462*,
463, 465
assamica Dyer 239, 243, 244,
272, 442, 485, 491
ssp. assamica 491
ssp. globifera (Ridl.) Sym.
272, 490*, 491, 562
ssp. koordersii (Brandis)
Sym. 244, 491, 492
ssp. philippinensis (Brandis)
Sym. 244, 491, 492, 561
ssp. philippinensis (non
Sym.) Browne 497
astrostricta Scort. ex Foxw.
540
astylosa Foxw. 440, 460, 562
atrinervosa Sym. 440, 446*,
457
attopoensis Pierre 493
auriculata Scort. ex Foxw. 532
bailloni Heim 532
bakeriana Heim 523
bakoensis Ashton 441, 479
balangeran (Korth.) Burck
251, 253, 256, 443, 486*,
499, 503, 509
var. angustifolia Boerl. 509
var. binnendijkii Boerl. 489
balangeran (non Burck) Vidal
458
balangeroides Boerl. 538
balanocarpoides Sym. 279,
441, 475
barbata Brandis 465
beccariana Burck 255, 444,
522*, 525, 526
beccarii Dyer ex Brandis 526
bentongensis Foxw. 442, 494
biawak Ashton 279, 440, 446,
462*, 464
blumutensis Foxw. 441, 480
borneensis Pierre 451
brachyptera Heim 513
bracteata Pierre ex Laness.
466

[ser.. I, volae2

(Shorea)

bracteolata Dyer 272, 442,
485, 496
brunnescens Ashton 439,
446*, 449
bullata Ashton 444, 517
cambodiana Pierre 494
camphorifera Roxb. 375
carapae Ashton 255, 551
chaiana Ashton 261, 441, 477
chrysophylla Ridl. 525
ciliata King 439, 450, 460
ciliata (non King) Foxw. 460
ciliata (non King) Ridl. 461,
465
cinerea Fischer 383
cochinchinensis Pierre 493
var. oligoneura Boerl. 538
var. saigonensis Guérin 493
collaris Sloot. 441, 477
collina Ridl. 439, 447, 458
collina (non Ridl.) Foxw. 453,
457
compressa Burck 272, 526
confusa Ashton 442, 497
conica (Sloot.) 441, 479
contorta Vidal 243, 244, 272,
440, 466, 467*
cordata Ashton 442, 495
coriacea Burck 253, 255, 443,
509, 510
costata Presl 321
costata (non (Correa) Presl)
King 450
crassa Ashton 253, 440, 457
crassifolia Ridl. 494
cristata Brandis 508, 509
curtisii Dyer ex King 245—249,
250*, 251, 272, 403; 445;
541, 542*, 543*
ssp. curtisii 541, 562
ssp. grandis Ashton 543,
562
cuspidata Ashton 441, 472,
483
dasyphylla Foxw. 546, 551
dasyrrhachis Sloot. ex Endert
417
dealbata Foxw. 253, 442, 485,
562
discolor Heim 533
dispar Ashton 443, 500*, 502
dolichocarpa Sloot. 475
domatiosa Ashton 440, 460
dryobalanoides Dyer ex Bran-
dis 484
dyeri (non Thw. ex Trim.)
Heim 508
elliptica Burck 442, 498*, 500
elliptica (non Burck) Sym. 455
exelliptica Meijer 439, 455
eximia (Miq.) Scheff. 548

1982]

Index to scientific plant names

(Shorea eximia)
var. angustifolia Burck 548
eximia (non (Miq.) Scheff.)
Foxw. 507
faguetiana Heim 441, 472, 484
faguetioides Ashton 441, 480
falcifera Dyer ex Brandis 243,
253, 260*, 439, 456, 461
falciferoides Foxw. 438*, 440,
458, 467
ssp. falciferoides 458
ssp. glaucescens (Meijer)
Ashton 458, 459
fallax Meijer 444, 503, 509,
516
farinosa C.E.C. Fischer 243,
442, 487
ferruginea Dyer ex Brandis
444, 532, 533
flava Meijer 456
flaviflora Wood ex Ashton
255, 444, 508, 518
flemmichii Sym. 253, 443, 507
floribunda (Wall.) Kurz ex
Dyer 493
floribunda (non Kurz) F.-Vill.
517
foraminifera Ashton 253, 445,
534, 538
forbesii King ex Brandis 417,
S37
foveolata Scort. ex Foxw. 496
foxworthyi Sym. 439, 453
franchetiana Heim 526
furfuracea Miq. 540, 551
furfuracea (non Miq.) Brandis
548
furfuracea (non Miq.) Rolfe
507
fusca Burck 551
gardneri (Thw.) Ashton 272
geniculata Sym. ex Ashton
253, 263, 279, 439, 451
gentilis Parijs 546
gibbosa Brandis 441, 472,
482, 483, 484
gisok Foxw. 458, 459
glauca King 251, 272, 440,
461
glauca (non King) Browne 456
glaucescens Meijer 459
globifera (Ridl.) Sym. 491
grandiflora Brandis 536
gratissima (Wall. ex Kurz)
Dyer 243, 253, 442, 487,
491
gratissima (non (Wall. ex
Kurz) Dyer) Foxw. 489
guiso (Blco) Bl. 239, 241, 243,
244, 253, 256, 272, 439,
447, 562
gysbertsiana Burck 523

595

(Shorea gysbertsiana)

var. scabra Burck 523, 526
harmandii Pierre ex Laness.
493
aff. harmandii (non Pierre)
Foxw. 492
havilandii Brandis 253, 262*,
439, 448
helferi Kurz 418
hemsleyana (King) King ex
Foxw. 261, 288, 445, 536
ssp. grandiflora (Brandis)
Ashton 536
ssp. hemsleyana 536
hemsleyana (non King) Foxw.
536
henryana Pierre 243, 442,
486*, 489
hopeifolia (Heim) Sym. 441,
472, 482
hypochra Hance 243, 244,
256, 272, 442, 485, 494
aff. hypochra (non Hance)
Browne 485
hypoleuca Meijer 440, 446*,
459, 460
iliasii Ashton 441, 472, 480
inaequilateralis Sym. 251, 253,
356, 358, 443, 505, 507
inappendiculata Burck 439,
455
inappendiculata (non Burck)
Sym. ex Desch 453, 457
induplicata Sloot. 253, 441,
476
isoptera Ashton 277, 440,
468*, 469*
javanica K. & V. 288, 442,
492, 493
johorensis Foxw. 443, 503,
508*, 513
kalunti Merr. 483
koordersii Brandis 491
kuantanensis Ashton 440, 472
kudatensis Wood ex Meijer
441, 472, 483, 562
kunstleri King 444, 503, 519
laccifera Heyne ex Wall. 493
ladiana Ashton 253, 440,
462*, 463
laevifolia Endert 461
laevis Ridl. 243, 440, 461
lamellata Foxw. 442, 493
lamellata (non Foxw.) Ashton
492
laurifolia Wall. 493
laxa Sloot. 253, 261, 440, 474
lepidota (Korth.) Bl. 445, 537
lepidota (non Bl.) Foxw. 514
leprosula Miq. 246, 248, 249,
259, 267, 272, 445, 540,
541, 542

(Shorea)

leprosula (non Miq.) Boerl.
541
leprosula x curtisii 270
leptoclados Sym. 513
leptoderma Meijer 439, 450
longestipulata Tard. 489
longiflora (Brandis) Sym. 440,
472*, 473, 474*
longipetala Foxw. 448
longisperma Roxb. 255, 441,
477, 481
lucida Miq. 387
lumutensis Sym. 439, 453, 456
lunduensis Ashton 440, 458
macrantha Brandis 253, 288,
445, 527*, 536
macrobalanos Ashton 440,
473
macrophylla (de Vriese) Ash-
ton 246, 247, 272, 444,
522*, 523, 528, 562
macroptera Dyer 444, 472,
527455295, 932
ssp. baillonii (Heim) Ashton
532
ssp. macroptera 272, 532,
583
ssp. macropterifolia Ashton
532, 533
ssp. sandakanensis (Sym.)
Ashton 532
macroptera Sloot. 532
malaanonan Bl. 383
malibato Foxw. 440, 460
mangachapoi Bl. 365
maranti Burck 540
maritima Pierre 494
martiniana Scheff. 272, 522
materialis Rid]. 251, 253, 256,
439, 456, 461
maxima (King) Sym. 441,
472*, 478
maxwelliana King 248, 272,
279, 440, 462*, 464*, 465
mayapis Bl. 333
meadiana Sym. 449
mecistopteryx Ridl. 272, 444,
521, 525
megistocarpa Foxw. 538
mekongensis Pierre ex Laness.
466
micans Ashton 440, 463, 465
mindanensis Foxw. 487
mollis 303
monticola Ashton 255, 444,
503, 519
montigena Sloot. 243, 244,
267, 442, 485, 487, 489
mujongensis Ashton 441, 472,
484
multiflora (Burck) Sym. 248%,

596

(Shorea multiflora)
272, 279, 441, 470, 472,
475
myrionerva Sym. ex Ashton
444, 528*, 529, 530*, 531*
nebulosa Meijer 509
negrosensis Foxw. 243, 443,
499, 502
nervosa Kurz 330
nitens Miq. 538
obovoidea Sloot. 441, 477
obscura Meijer 440, 457
obtusa Wall. 243, 270, 272,
447
var. kohchangensis Heim
448
var. subevenis Boerl. 494
ochracea Sym. 442, 492
ochrophloia Strugnell ex Sym.
439, 447
oleosa Meijer 516
ovalis (Korth.) Bl. 259, 263,
269, 272, 445, 547*, 548*,
550
ssp. Ovalis 548*, 549*, 551
ssp. sarawakensis Ashton
548, 549
ssp. sericea (Dyer) Ashton
269, 270, 272, 551
ovata Dyer ex Brandis 251,
255, 267, 403, 445, 544*,
545
pachyphylla Ridl. ex Sym.
D293 443s Sl 51i2*
Ppahangensis Foxw. 494
palembanica Miq. 247, 272,
288, 443, 513
pallida Foxw. 492
pallidifolia Ashton 253, 551
palosapis (Blco) Merr. 243,
444, 509, 517
paludosa Foxw. 538
palustris Ridl. 541
parvifolia Dyer 246, 249, 259,
263, 272, 278, 445, 502,
534, 535, 546
ssp. parvifolia 546, 547, 562
ssp. velutinata Ashton 502,
547
parvifolia (non Dyer) King
545
‘parvistipula’ 508
parvistipulata Heim 443, 503,
508
ssp. albifolia Ashton 509,
516
ssp. nebulosa (Meijer)
Ashton 509
ssp. parvistipulata 508
patoiensis Ashton 441, 476
pauciflora King 272, 443, 503,
511, 545

FLORA MALESIANA

(Shorea)

peltata Sym. 261, 440, 473

peltata (non Sym.) Browne
474

Dhilippinensis Brandis 492

pierreana Heim 449

pierrei Hance 448

pilosa Ashton 444, 521, 525

pinanga Scheff. 272, 278, 444,
520*, 522, 524, 525, 526

pinangiana Wall. 360

plagata Foxw. 545

platycarpa Heim 251, 253,
256, 259, 445, 527*, 541,
562

platyclados Sloot. ex Foxw.
246, 255, 272, 443, 515,
562

polita Vidal 243, 442, 485,
487, 488*, 491

polyandra Ashton 253, 261,
440, 470*, 471*

polysperma (Blco) Merr. 241,
243, 252*, 443, 467, 514

polysperma (non Merr.) Keith
513

praestans Ashton 444, 524

pubistyla Ashton 444, 516,
S8*

purpurea DC. 551

quadrinervis Sloot. 258, 445,
529, 534, 535*

resina-negra Foxw. 481

resinosa Foxw. 263, 269, 272,
442, 485, 496

reticulata (non Thw. ex Dyer)
F.-Vill. 423

retinodes Sloot. 253, 256, 442,
485, 495

retusa Meijer 253, 445, 534,
537, 543

revoluta Ashton 253, 255,
445, 534, 543

richetia Sym. 440, 472*, 474

ridleyana King 483, 484

rigida Brandis 551

robusta Gaertn. f. 243, 259,
263, 272, 274, 282, 447

var. schmidtii Heim 448

robusta (non Gaertn. f.) F.-
Vill. 448

rogersiana Raizada & Smiti-
nand 461

rotundifolia Ashton 444,
520*, 524

roxburghii G. Don 239, 243,
253, 256, 261, 263, 265,
272, 442, 485, 486*, 493

rubella Ashton 238*, 253,
442, 500*

rubra Ashton 255, 445, 534,
545

[ser. I, vol. 93

(Shorea)

rugosa Heim 253, 445, 540,
5511
rugosa (non Heim) Foxw. 517
rugosa (non Heim) Sym. 539
var. uliginosa Sym. 539
sagittata Ashton 444, 531
saigonensis Pierre 493
sandakanensis Sym. 532
scaberrima Burck 262*, 288,
443, 506*, 515, 551
scabrida Sym. 253, 445, 534,
537, 543
schefferiana Hance 451
scrobiculata Burck 439, 446*,
449, 450
scrobiculata (non Burck)
Foxw. 448
scutalata King 546
selanica (Lamk) Bl. 243, 244,
256, 443, 503, 505
var. latifolia Bl. 505
var. obtusa Burck S05
seminis (de Vriese) Sloot. 247,
253, 256, 288, 439, 451,
453, 460
sericea Dyer 551
sericeiflora Fischer & Hutch.
489
siamensis Miq. 243, 253, 265,
272, 278, 304, 440, 466
singkawang (Miq.) Miq. 272,
288, 445, 536
ssp. scabrosa Ashton 537
ssp. singkawang 537
slootenii Wood ex Ashton
444, 527*, 529
smithiana Sym. 245, 272, 443,
503, 504*, 562
sororia Sloot. 491
sp. Browne 455, 530
sp. Foxw. 539
sp. Sloot. ex Merr. 503
sp. B. Sym. 543
Sp. Gs sym. 2593/7
sp. nov. Sloot. 505
sp. nov. Sym. ex Desch 449
splendida (de Vriese) Ashton
QAT, 272, 4445 52522
squamata Benth. & Hook. f.
Sul7/
squamata (non Benth. &
Hook. f.) Brandis 508
squamata (non Benth. &
Hook. f.) Browne 516
stellata Dyer 383
stenoptera Burck 246, 253,
261), 272, 444; 521; 523;
524
stipulosa Burck 538
suavis (non Wall. ex DC.)
Pierre ex Lanessan 466

1982]

(Shorea)
subcylindrica Sloot. 441, 476
sublacunosa Scheff. 548
submontana Sym. 439, 450
subpeltata Miq. 387
sumatrana (Sloot. ex Thor.)
Sym. 247, 256, 272, 288,
439, 447, 452*, 453, 454*,
460
superba Sym. 256*, 440, 459,
562
symingtonii Wood 442, 495
talura Roxb. 493
talura (non Roxb.) F.-Vill.
514
tenuiramulosa Ashton 255,
441, 478
teysmanniana Dyer ex Brandis
251, 253, 256, 267, 445,
538
teysmanniana (non Dyer ex
Brandis) Ashton 538
teysmanniana (non Dyer)
Foxw. 515
thiseltonii (King) 537
tomentosa (non Miq.) Pierre
466
trapezifolia (Thw.) Ashton
272
uliginosa Foxw. 251, 253, 256,
445, 539
utilis King 465
vandekoppelii Parijs 493
var. grandifolia Parijs 493
venulosa Wood ex Meijer 253,
255, 443, 503, 506*, 510
verruculosa Dyer ex Brandis
540
vidaliana Brandis 448
virescens Parijs 442, 492
virescens (non Parijs) Ashton
497
vulgaris Pierre ex Laness.
448
waltoni Wood ex Meijer 443,
510
warburgii Gilg 514
warburgii (non Gilg) Perk.
448
xanthophylla Sym. 441, 479
Shoreoxylon 241
Simaroubaceae 6: 193—226,
968—972
Smilacaceae 191, 192
Smilax 191, 192
Sonneratiaceae 4: 280—289, 513;
5: 557; 6: 973
Sparganiaceae 4: 233-234
Sphenocleaceae 4: 27-28
Spondias malayana Kosterm.
553
pinnata (L. f.) Kurz 553

Index to scientific plant names

Stachys decemdentata Sol. ex
Forst. f. 567
Stackhousiaceae 4: 35—36
Staphyleaceae 6: 49—59
Stemonoporus Thw. 255, 261,
263, 265—269, 273, 275,
276, 278, 279, 280, 283,
284, 338
lanceolata 262*
Stemonurus Bl. 566
foetidus Wight 566
Sterculiaceae 275, 280
Stereospermum colais (Ham. ex
Dillw.) Mabberley 554
personatum (Hassk.) Chatter-
jee 554
Stictocardia 558
Streptopus multiflorus (Bl.) D.
Dietr. 218
Strombosia philippinensis sensu
Lam & Holth. 569
Strychnos axillaris Colebr. 568
Stylidiaceae 4: 529-532; 5: 564;
6: 976
Stypandra 191
Styracaceae 4: 49-56; 6: 976; 9:
568-569
Styrax benzoin Dryand. 569
crotonoides Clarke 569

ssp. fraserensis (Putz & Ng)

Steen. 568
fraserensis Putz & Ng 568
ridleyanum Perk. 569

Sunaptea Griff. 278, 345, 359
astrotricha Pierre 361
borneensis Heim 363
bureavi Heim 365
dyeri Pierre 361
faginea Pierre 361
odorata Griff. 360
teysmannianii Heim 360
urbanii Heim 363

Symplocaceae 8: 205—274; 9:

569

Symplocos atjehensis Noot.

569
columbuli Noot. 569
ophirensis Clarke

var. densereticulata Noot.
569
var. kaliensis Stone 569

‘Synaptea’ Kurz 345, 359
bantamensis Kurz 364
cinerea Ridl. 360
cuspidata Ridl. 367
grandiflora Kurz 361
lancaviensis Ridl. 360
lowii Ridl. 369
maingayi Ridl. 367, 369
nitens Ridl. 367
perakensis Ridl. 367
reticulata Ridl. 365

597

Taccaceae 7: 806—819
Tamaricaceae 275
Ternstroemiaceae 275, 280
Teta Roxb. 227
viridiflora Roxb. 228
Tetraplasandra (non A. Gray)
Mia. 7, 67
koordersii Harms 68
paucidens Miq. 68
philippinensis Merr. 68
solomonensis Philipson 68
Textoria Miq. 101
Theaceae 280, 281
Thismia Griffith 554
sect. Sarcosiphon (Bl.) Jonker
555
clavigera (Becc.) F.v.M. 555*
Thymelaeaceae 4: 349—365; 6:
1—48, 976; 7: 830
Thysanotus R. Br. 189—192,
211, 567
banksii R. Br. 567
chinensis Bth. 189, 190, 210*,
211
chrysantherus F.v.M. 211
siamensis Ridl. 211
tuberosus R. Br. 210%, 211,
213, 567
var. parviflora Bth. 213
Tieghemopanax elegans (C.
Moore & F.v.M.) Vig. 77
macgillivrayi Viguier 74
Tiliaceae 268, 274, 276, 280,
281, 282
Trapaceae 4: 43—44; 6: 982
Trevesia Visiani 1, 4, 7, 9, 91, 95
arborea Merr. 97, 99
barbata (Becc.) O. K. 41
beccarii Boerl. 97, 99
burckii Boerl. 96*, 97
carpophagarum (Becc.) O. K.
45

cheirantha (Clarke) O. K. 97
eminens Bull 48
geelvinkiana (Becc.) O. K. 38
helleborina (Becc.) O. K. 39
insidiator (Becc.) O. K. 45
insignis Miq. 52
moluccana Miq. 42
novo-guineensis Scheff. 43
palmata (DC.) Vis. 97
var. cheirantha Clarke 97
var. insignis Clarke 52
pulcherrima (Vidal) O. K. 40
rufo-setosa Ridl. 97
sundaica Mig. 93, 97, 98*, 99
var. glomerata K. & V. 99
teysmannii (Boerl.) O. K. 44
zippeliana Miq. 42
Tricalistra Rid]. 189, 191, 192,
220
ochracea Rid]. 221, 222*

598

Tricoryne R. Br. 189, 190, 191,
193, 213
platyptera Rchb. 212*, 214
pterocaulon Baker 214
Tricyrtis Wall. 189, 191, 193,
195
imeldae Gutierrez 196*, 197
stolonifera 197
Trigoniaceae 4: 58—60
Trigonobalanus Forman 563
doichangensis 563
excelsa nov.sp. 563
Triuridaceae 199
Triuridales (order) 199
Tupidanthus 5, 7
Tupistra Ker-Gawl. 189, 191,
192, 221
gracilis 223
grandis Ridl. 221, 222*, 223
nutans Wall. 223
perakensis Nichols. 223
singapureana (Wall.) Baker
223
squalida 221, 223
violacea Rid]. 221
Tupistra sensu Hutch. 220
Turneraceae 4: 235—238
Typhaceae 4: 242—244; 6: 982

Ulmaceae 8: 31—76, 551; 9: 569
Umbellales 4, 5
Umbelliferae 4: 113—140, 595;
5: 555; 6: 983—984; 7:
830-832; 9: 5, 6, 7, 11,
27, 569-571
subfam. Apioideae 6
subfam. Hydrocotyloideae 5
subfam. Saniculoideae 5
Uncinia Pers. 107, 109, 183 map
subgen. ‘Eu-Uncinia’ 184
subgen. Pseudocarex 184
subgen. Uncinia 184
sect. Platyandra 184
sect. Uncinia 184
compacta R. Br. 184
var. alpina Noot. 185*, 186
var. compacta 186
var. nervosa Clarke 184,
185*, 186
filiformis Boott 184
kingii Boott 184
nervosa Boott 184, 186
ohwiana Koyama 184
riparia R. Br. 184, 186
var. stolonifera Kuk. &
Steen. 184
riparia (non R. Br.) Ohwi 184
rupestris Raoul 184
var. capillacea Kiik. 184
sclerophylla Nelmes 184
subtrigona Nelmes 184

FLORA MALESIANA

[fser. I, vol. 93

Unjala bifida Reinw. ex de
Vriese 42
Unona selanica DC. 505
Upuna Sym. 239, 259, 261, 263,
266, 267, 269, 272, 274,
276, 277, 279, 280, 281,
284, 288, 290, 291, 337
borneensis Sym. 253, 262*,
PP SES Soo, BOs
339*
Uvularia chinensis Ker-Gawl.
218
multiflora (Bl.) Kunth 219

Vaccinium altiterrae Veldk. 563
oranjense J.J.S. 563
Valerianaceae 4: 253—254
Vateria Arn. 345
sect. Isauxis Arn. 345, 349
Vateria L. 261, 263, 265, 266,
269, 273, 275, 276, 278,
279, 280, 283
indica L. 263, 275
rassak Walp. 353
Vateriopsis 239, 263, 276, 278,
279, 283, 284
Vatica L. 239, 245, 249, 253,
255, 258, 259; 261,263,
265—269, 272, 275, 276,
Dili 21D 280s 28283),
284, 288-291, 338, 341,
345, 350 map
sect. Euvatica B. & H. 359
sect. Isauxis (Arn.) B. & H.
345, 349
sect. Pachynocarpus (Hook.
f.) Burck 345, 349
sect. Retinodendron (Korth.)
Burck 349
sect. Sunaptea (Griff.) Burck
266, 279, 341, 345, 348,
359, 370
sect. Vatica Ashton 266, 267,
278, 279, 280, 346, 347,
349
subg. Isauxis (Arn.) Brandis
345, 349
subg. Retinodendron (Korth.)
Brandis 345, 349
subg. Synaptea (Griff.) Bran-
dis 345, 359
acrocarpa Sloot. 349
aerea Sloot. 361
albiramis Sloot. 347, 355, 358
aperta Sloot. 369
apteranthera Blco 364
astrotricha Hance 361
badiifolia Ashton 348, 367
bancana Scheff. 350

bantamensis (Hassk.) B. & H.

ex Miq. 243, 348, 364*

(Vatica)

bantamensis (non B. & H. ex
Miq.) Ashton 367
beccariana Heim 341
beccarii Dyer ex Brandis 363
bella Sloot. 347, 352
blancoana Elmer 349
borneensis Burck 348, 363
brevipes Ashton 348, 366
brunigii Ashton 348, 362
burckii Heim 342
bureavi Heim 365
cauliflora Ashton 347, 370
celebensis Brandis 353
celebica Koord. ex Sym. 491
celebica Sloot. 353
chartacea Ashton 347, 351
‘chinensis’ 364
cinerea King 243, 256, 272,
348, 360
compressa Ashton 348, 361
congesta Ashton 348, 362
coriacea Ashton 253, 348,
360, 362
cupularis Sloot. 349
curtisii King 361
cuspidata (Ridl.) Sym. 348,
367
cuspidata (non Ridl.) Browne
367
dulitensis Sym. 255, 347, 356
dyeri Pierre ex Laness. 361
elliptica Foxw. 347, 369
endertii Sloot. 348, 366
eximia Miq. 548
faginea 360, 361
flavida Foxw. 279, 348, 359
flavovirens Sloot. 243, 244,
348, 366
fleuryana Tardieu 361
forbesiana Burck 351
furfuracea Burck 356
glabrata Ashton 347, 370
globosa Ashton 348, 358
grandiflora (Wall. ex DC.)
Dyer 272, 361
granulata Sloot. 347, 354
ssp. granulata 354
ssp. sabaensis Ashton 255,
354
harmandii Heim 341
havilandii Brandis 268, 347,
351
helferi Dyer 418
heteroptera Sym. 255, 348, 359
hullettii (Ridl.) Ashton 348,
358
imbricata Sloot. 370
javanica Sloot. 348, 362
ssp. javanica 362
ssp. scaphifolia (Kosterm.)
Ashton 362

1982]

Index to scientific plant names

599

(Vatica)

kelsalli Ridl. 352

kunstleri Brandis 350

laccifera (Wall. ex DC.) W. &
A. 493

lamponga Burck 351

lanceaefolia Bl. 241, 359

lankaviensis Ridl. 360

laurifolia (Wall.) Steud. 493

lepidota Korth. 538

leucocarpa Foxw. ex den Ber-
ger & Endert 342

lobata Foxw. 279, 348, 358

lowii King 243, 348, 369

/owii (non King emend. Sym.)
King 369

lutea Ridl. 350

macroptera Sloot. ex Thoren.

369

maingayi Dyer 346*, 348,
368*, 369

maingayi (non Dyer) Sloot.
367

mangachapoi Blco 243, 251,
348, 364, 370

ssp. mangachapoi 364, 562
ssp. obtusifolia (Elmer)
Ashton 365
maritima Sloot. 348, 359
melanoxylon Benth. & Hook.
f. ex Miq.
var. recta Heim 341
micrantha Sloot. 348, 366
mindanensis Foxw. 361
moluccana Burck 353
nitens King 348, 367
oblongifolia Hook. f. 347,
35555355, 561
ssp. crassilobata Ashton
253, 355, 356
ssp. elliptifolia Ashton 253,
355, 356
ssp. multinervosa Ashton
355, 356
ssp. oblongifolia 355*, 356
ssp. selakoensis Ashton 255,
356
obovata Sloot. 348, 363
obtusa Burck 347, 349, 352,
370
obtusa (Wall.) Steud. 370
obtusifolia Elmer 365
odorata (Griff.) Sym. 243,
272, 348, 360, 361
ssp. mindanensis (Foxw.)
Ashton 255, 361, 365
ssp. odorata 272, 360, 361
ovalifolia Ridl. 352
ovalis Korth. 548
pachyphylla Merr. 243, 348,
363, 365
pallida Dyer 279, 348, 358

(Vatica)
papuana Dyer ex Hemsl. 273,
353

parvifolia Ashton 253, 348,
365
patula Sym. 365
pauciflora (Korth.) Bl. 247,
256, 273, 347, 351, 370
pauciflora Walp. 351
pedicellata Brandis 347, 356
pentandra Ashton 347, 370
perakensis King 348, 367, 369
philastreana Pierre 359
ramiflora Sloot. 349, 354, 355
rassak (Korth.) Bl. 243, 244,
DAG; 247, 25551250, 2710)
273, 347, 349, 353, 354*
var. subcordata Bl. 353
reticulata King 365
reticulata (Thw.) A. DC. 365
ridleyana Brandis 347, 352
rotata Ashton 347, 357
ruminata Burck 351
rynchocarpa Ashton 348, 365
sarawakensis Heim 261, 347,
354, 358, 561, 562
scaphifolia Kosterm. 362
scaphula (Roxb.) Dyer 335
scaphula (non Dyer) F.-Vill.
365
schefferi Brandis 350
schouteniana Scheff. 350
schumanniana Gilg 353
scortechinii (King) Brandis
347, 357
simalurensis Sloot. 351
sinensis (non Gmel.) Blco 364
soepadmoi Ashton 347, 352
songa Sloot. 367
sorsogonensis Foxw. 361
sp. Ashton 365
stapfiana (King) Sloot. 273,
347, 349
stapfiana (non Sloot.) Browne
349
stipulata Ridl. 358
stipulosa Miq. 538
subcordata (Bl.) Hall. f. 353
sublacunosa Miq. 548
sumatrana Sloot. 352
teysmanniana Burck 348, 360
thorelii Pierre 352
tonkinensis (Chevalier) Tar-
dieu 361
trigyna Griff. 317
umbonata (Hook. f.) Burck

247, 256, 270, 344*, 346*,

347, 349, 350, 352, 354,
370, 561

ssp. acrocarpa (Sloot.) Ash-
ton 349, 561

ssp. umbonata 349, 561

(Vatica)
urbanii Heim 363
venulosa BI. 253, 261, 347,
350, 351, 370
ssp. simalurensis (Sloot.)
Ashton 351
ssp. venulosa 350
verrucosa Burck 349
vinosa Ashton 279, 347, 357
wallichii Dyer 247, 273, 351
whitfordii Foxw. 365
zollingeriana DC. 351
Vatica (non L.) Dyer 327
Veratronia malayana (Jack)
Miq. 235
Veratrum malayanum Jack 235
Vignea Beauv. 176
Violaceae 7: 179—212, 831
Vitis vinifera 273

Wahlenbergia albomarginata
Hook. f. 556
papuana v. Royen 556
Wardenia King 7, 89, 93
simplex King 93
Woburnia 241

Xanthorrhoeaceae 191, 192
Xerotes arenaria R. Br. 235
filamentosa A. Cunn. 187
longifolia R. Br. 235
leucocephala R. Br. 187
Xylonymus Kalkman 557
versteeghii Kalkman 557
Xyridaceae 4: 366—376, 598; 5:
557; 9: 571-573
Xyris L. 571
anceps auct., non Lamk 571
bancana Miq. 571, 572 map
var. lacerata v. Royen 571
borneensis Rendle 571
calocephala Miq. 572
capensis Thunb. 571, 572
var. nilagirensis (Steud.)
Nilsson 573
var. schoenoides (Mart.)
Nilsson 573
capito Hance 572
chinensis Malme 571
chlorocephala v. Royen 572
complanata R. Br. 571
dajacensis v. Royen 573
elongata Rudge 571
flabellata v. Royen 573
glaucella Malme 571
grandis Ridley 571, 572
indica L. 571, 572
indica auct., non Linné 571
intersita Malme 571
lobbii Rendle 571, 573
malaccensis Steud. 571
malmei v. Royen 573

600

(Xyris)
maritima Koyama 573
melanocephala Miq. 573
nilagirensis Steud. 573
novoguineensis Hatus. 573
oligantha Steud. 571, 573
oreophila Ridl. 573
papuana v. Royen 571
pauciflora Willd. 571, 573

var. oryzetorum Miq. 573

FLORA MALESIANA

[ser. I, vol. 93

(Xyris)

pauciflora auct., non Willd.
573
ridleyi Rendle 571
var. penicillata v. Royen
Syl
robusta Mart. 572
schoenoides Mart. 573
subcomplanata Malme 571
sumatrana Malme 573

(Xyris)
tuberosa Ridley 571, 573
walkeri Kunth 571
wallichii Kunth 571, 573

Zanthoxylum avicennae (Lamk)
DEIAN105
trifoliatum L. 103
Zygophyllaceae 4: 64

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