FLORA

MALESIANA

SERIES I - SpERMATOPHYTA
Flowering Plants
Vol. 10, part 4

DEDICATION TO BLUME
ADDENDA
 INDEX TO REVISED FAMILIES

Aceraceae
TAXONOMICAL REVISIONS

THE LuESTHER T. MERTZ LIBRARY

THE NEW YORK BOTANICAL GARDEN

 REPUBLIK INDONESIA
REPUBLIC OF INDONESIA
LEMBAGA ILMU PENGETAHUAN INDONESIA (L.I. P. I.)

INDONESIAN INSTITUTE OF SCIENCES

FLORA MALESIANA
BEING
AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA/
INCLUDING KEYS FOR DETERMINATION DIAGNOSTIC DESCRIPTIONS , /

REFERENCES TO THE LITER ATURE SYNONYMY AND DISTRIBUTION

I I /

AND NOTES ON THE ECOLOGY OF

ITS WILD AND COMMONLY CULTIVATED PLANTS

PUBLISHED
UNDER THE AUSPICES OF THE CENTRE FOR RESEARCH
AND DEVELOPMENT IN BIOLOGY BOGOR JAVA AND / /

OF THE RIJKSHERBARIUM LEIDEN NETHERLANDS/ /

PREPARED
ON AN INTERNATIONAL 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 10
SPERMATOPHYTA

GENERAL EDITORS:
Dr. C. G. G. J. VAN STEENIS (f 1986)
Dr. W. J. J. O. DE WILDE

KLUWER ACADEMIC PUBLISHERS

DORDRECHT BOSTON LONDON
/ /

1 9K4 I y«y
Library in Congress Catalog Card Number 72-175112

ISBN 0-7923-0421-7

Published by Kluwer Academic Publishers,

P.O. Box 17, 3300 AA Dordrecht, The Netherlands.

Kluwer Academic Publishers incorporates

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Publication dates

Part 1 20 Jan. 1984

Part 2 21 May 1986
Part 3 19 Oct. 1986
Part 4 3 Aug. 1989

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© 1989 Kluwer Academic Publishers
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 no

v"' CONTENTS

Title-page (3)
Conienis (5)
Dedication by C. G.G.J, van Steenis (7)
Abbreviations and signs (41)

TAXONOMICAL REVISIONS
in alphabetical sequence

Alseuosmiaceae by C. G.G.J, van Steenis 335

Araucariaceae by D.J. de Laubenfels 419
Aristolochiaceae by Ding Hou 53
Chloranthaceae by B. Verdcourt 123
Chrysobalanaceae by G.T. Prance 635
Coniferales by D.J. de Laubenfels 337
Cruciferae by B. Jonsell 541
Ctenolophonaceae by A.M.N, van Hooren & H.P. Nooteboom 629
Cupressaceae by D.J. de Laubenfels 443
Elaeagnaceae by J.F. Veldkamp 151
Ixonanthaceae by R. Kool 621
Linaceae by A.M.N, van Hooren & H.P. Nooteboom 607
Magnoliaceae by H.P. Nooteboom 561
Menispermaceae by L.L. Forman 157
Monimiaceae by W.R. Philipson 255
Olacaceae by H Sleumer
. 1

Opiliaceae by P. Hiepico 31
Pinaceae by D.J. de Laubenfels 447
Podocarpaceae by D.J. de Laubenfels 351
Polygalaceae by R. van der Meijden 455
Sabiaceae by C.F. van Beusel^om & Th.P.M. van de Water 679
Sphenostemonaceae by C. G.G.J, van Steenis 145
Taxaceae by D. J de Laubenfels
. 347
Trimeniaceae by W.R. Philipson 327
Triuridaceae by J. P.M. van Meerendonk 109

ADDENDA
to volumes 4-10

Addenda, corrigenda et emendanda by C. G.G.J, van Steenis el al 717

INDEX
fndex to scientific plant names by E.E. van Nieuwkoop 721
 1

^i^tcsii..^'

l^r\ 11 1.; jj (,. ..: liVUU Y lbiJ\J JJ -L ii l;l-cj^

Dedicated to the memory of

CARL LUDWIG BLUME
 DEDICATION'
Many botanists must have wondered why as yet no volume of Flora Malesiana was dedicated to
the outstanding botanist Carl Ludwig Blume, undisputed pioneer in planning the compilation
of a 'Flora Malesiana'.
The writing of this Dedication would have been greatly facilitated if a full biography of Blume
had been existent, but none is available; there is not even a bibliography of his works. Only recent-
ly, in 1979, two biographical attempts were made, by J. Maclean and by A. den Ouden, but

only for the period 1820-1832; together with other biographical and obituary notes they are here
assembled in Appendix B. I have also compiled a bibliography: Appendix A.*
There are various reasons to account for the lack of data. At Leiden there are, in the Rijksher-
barium archives, only few letters addressed to or written by Blume, and this is also the case for
the University archives. Also Treub (B) in his papers on the history of the Botanic Gardens at
Bogor complained about the lack of correspondence of Blume. The largest source of (official)
letters is contained in the huge 'Rijksarchief at The Hague, but it will require a large, time-

consuming effort to unearth these (D: 5). Blume's large private library was auctioned at Leipzig
in March 1863, soon following his death, by the firm of O.T. W'eigel (B; D: 9).
It has sometimes been suggested that the lack of a full biography - to which Blume was certain-

ly entitled - could be explained by the fact that Blume had few friends (D: 8) and that his contem-

porary colleagues were antagonistic. But this explanation does not really hold, as a biography of
the charismatic Miquel was not written before a century after his death. In the Netherlands the
climate is not favourable for biographies of scientists, at least not in botany (D: 7).
For the reasons given above I have waited a long time to frame a worthy dedication, in the hope
that some historiographer would feel attracted to compose a full biography of Blu.me. In the
absence of this I have ventured to accumulate material myself, recently supported by a study of
Maclean (B) on Blume's years in Java and based on archival research in the 'Rijksarchief, and
an unpublished essay by den Ouden (B) on the same period based on details from several hun-
dreds of letters in the same archives.
To my do not give sufficient attention to personality and motiva-
regret biographers frequently
tions, but confine themselves to an appreciation of achievements. I have tried to form an opinion
about this facet of Blume. From Blume's profuse writings much can be learned about his motiva-
tions and his attitude towards society and people. It stands beyond doubt - and that must soon
have been realized by his contemporaries (E, F) - that in the science of botanical taxonomy Blume
was on a level with the great taxonomists of the previous century. But in the eyes of his close col-
leagues he was an autocratic, dominant, unsympathetic person, and this impression still lingers
around his name and overshadows the singular value of his scientific work. His sharp pen and
especially his fanatical pursuit of a monopolistic position for the Rijksherbarium estranged him
from his surroundings. Goddijn (B: 1931) has pointed this out very well.
.My purpose in composing this dedication is to give a sketch of Blume's life, his work and his

motivations in a detached way. Blume has a right to an impartial judgement; activities and per-
sonalities should be kept well apart. In a few cases, where there is lack of clarity about the inter-
pretation of historical data, I will give Blume the benefit of the doubt.

(1) Shortly May 1976 the author of this Dedication and former Editor of Flora Male-
before his death in
C.G.G.J. van STthsis, finished the text of the manuscript. He had the intention to use this
siana, Professor
biography of Blume to conclude volume 10 of the Flora. Wc wholeheartedly like to carry out his intention
here. - The General Editor.
(2) The documentation here presented is recorded in six appendices: A. Bii'mf's publications annotated;
B. Biographical sources; C. References to cited literature; D. Notes (mostly additional information considered
useful to illustrate the situations under which Blume had to work, his surroundings, personalia, etc.); E.
Eponymy; F. Honorary distinctions and memberships.

The photograph on the opposite page is copied from Rumphia } (1847), Bi.umi silting above his treasures
of the Javanese flora, including Nepenthes, Rafflesta, Rhizanthes, orchids and a rattan, presumably Plec-
lotomia, the picture dating from the lime when he was at the height of his career.

(7)
 Flora Malesiana

Towards the end of the 18th century two earUer attempts to compile a Flora of Malesia were
made, namely by Francisco NoRONa in 1786 and by Louis Deschamps in 1794-1798 (B: van
Steenis & VAN Steenis-Kruseman, 1970; C: van Steenis c.s. 1954). Both attempts were abortive
,

by the unfortunate loss of the material these collectors had made.

In the early 19th century the time had come for the more thorough exploration of the tropical
floras, both in the New and in the Old World. In the Indies it was started by W. Roxburgh and
N. Wallich. In Malesia there had been quite some botanical activity in Raffles' time, notably
by W. Jack in Sumatra and by Th. Horsfield and L.T. Leschenault de la Tour in Java, but
these researches had led only to publications by Jack.
The proper achievement fell to Blume, after the establishment of the Botanic Gardens at Bui-
tenzorg (Bogor) in 1817, where a year later Blume started a research period of seven years which
led to his brilliant scientific career.

Carl Ludwig Blume, born at Brunswick (Germany) on 9 June 1796, was a son of the merchant
Christiaan Nicolaas Ludwig Blume and of Melusine Caroline Sophie Drechsler. His father
died before he was born and his mother died when he was five years old. He was an eager boy
and was attracted by the study of pharmacy. To a high degree he was interested in travel books
of foreign countries, a trend and interest possibly strongly developed in Germany since Hum-
boldt's time, known as the 'Wanderlust', a tendency perpetuated to the present day (D: 10).
Blume's interests were probably directed towards the many unexplored regions of the globe, in-
cluding the tropics. By 1813 he used his heritance to buy clothes and equipment, and enlisted as
a volunteer in the 'Liitzowsche Jagercorps', fighting against the French. Later on he went to the
Netherlands, where on 29 December 1814 the Medical Board of the Dutch Forces appointed him
as a military apothecary of the second class. On 6 April 1815 he was placed with the ambulance
of the second division of the mobile forces in Belgium. He was present at Waterloo. According
to the military Stamboeken (Registers) he was an apothecary of the second class in the hospitals
at Den Helder and Leiden between 1814 and 1817.
Whenin 1815 Prof. S.J. Brugmans was commissioned to bring back the collections of natural
history from Paris to the Netherlands - collections which the French had taken there in 1795
-
Blume was appointed as his assistent.
In some way or other, young Blume enjoyed the support of the Duchess of Braunschweig,
financiallyand otherwise. She fostered his career and had recommended him to Prof. Brugmans
(t 1819), who urged Blume
- who had performed his task excellently - to study natural history
and medicine. Blume followed this advice and took a degree as Doctor of Medicine on 9 July 1817
at Leiden (A: 1817). Shortly before this date, apparently in view of his doctorate, Blume finished

his activities as an apothecary in the hospital at Leiden. On 17 October 1817 he returned in the
service of the hospital as an M.D., after having obtained, on 6 October 1817, the degree of a
health-officer of the second class of the forces and hospitals. On 1 1 January 1818 he was honour-
ably discharged as a surgeon-major and on 28 March 1818 became a health-officer of the second
class of the forces in the Netherlands East Indies. On 28 May 1818 followed the same appointment
for the first class; he worked at Leiden till 17 March 1818.
January 1819, Blume was appointed deputy-director un-
Shortly after his arrival in Java, on 1 1

der C.G.C. Reinwardt in charge of the organization of Education, Medical service, Agriculture,
Arts and Scientific investigation. He was then only 22 years old, but obviously highly esteemed
for his ambition, zeal, knowledge and energy. His initial salary was / 500 annually. He lived in
Reinwardt's house Buitenzorg (Bogor), enlarged for this purpose, in the Botanic Gardens.
at
He married the rich Wilhelmina Nicolasina Cranssen. This marriage was obviously not very
successful. He was divorced in April 1830 in Brussels and he remarried at the end of that month
Johanna Alletta Wilhelmina Waardenburg, by whom he had 7 children.
At that time the Government was much concerned about serious tropical diseases, small-pox,
typhoid, cholera, and in 1820 Reinwardt wrote a detailed report on the state of vaccination in

(8)
 Dedication

the years 1818- 1819. All civil servants were informed of the Government's intention to maintain
and promote vaccination. Blume was provisionally appointed 'Inspector of Vaccine' in 1819. He
informed the Government that it was desirable to use indigenous plants instead of imported
medicines which often lost their value during the long sea-voyage, and the Government requested
him to make proposals.
In the seven years between 1819 and 1826 Blume travelled widely in West and Central Java,
as far east as Rembang, often accompanied by assistants, draughtsmen and interested persons,
collecting plants and also animals; gathering information on all sorts of aspects, including the
medicinal value of certain plants, inspecting epidemics, etc. in short he was engaged in an overall,
;

thorough scientific exploration.

He gathered many duplicates and his herbarium specimens are still in excellent condition. It has
never become clear to me how these early explorers managed to dry and preserve their collections
so well in the everwet tropics under the primitive conditions of the time, trekking from camp to
camp (B: van Steenis-Kruseman, 1950).
In 1821/22 he was in Bantam in the company of the civil servant J.B. Spanoghe; in 1822 he
made a large exploration of Mt Salak in; 1 823 of Mt Gedeh; in 824 he made a large tour of inspec-
1

tion in thecompany of the clerk G.H. Nagel, the gardener W. Kent, and the draughtsman A.
Latour, to many places: Kuripan (near Bogor) with hotwells in limestone then surrounded by
primary forest, Mt Seribu (hills SW. of Jakarta), then to the Krawang region (E. of Jakarta)
eastwards to Indramayu, proceeding to Cheribon, ascending Mts Tjeremai, Tangkuban Prahu,
Burangrang, going as far as Tegal. Furthermore, he explored the then completely forest-clad large
island of Nusa Kambangan (S. Central Java) where he detected Rafflesia. In 1825 he was again
in Rembang (Central Java), but also in Bantam, ascending Mt Parang.
These must have been hectic, creative years in Blume's life. In view of later controversies I have
listed these explorations, which show that Blume covered a considerable part of West and Central
Java, and that his travels partly covered the same habitats which had been visited by Kuhl and
VAN Hasselt, members of the 'Natuurkundige Commissie', but also went beyond these. The
result was of course that the majority of species were collected by both parties.
In all probability Blume studied, analyzed, and described his collections in situ, facilitating
later publication. In addition to all this field work he published scientific reports on many of these
explorations, in part made public in a number of letters which he wrote to the brothers Nees von
EsENBECK, published in the Regensburg journal Flora (A: 1823-1826).
Finally he compiled all this material in the voluminous Bijdragen (A: 1825-1827), containing
the concise treatment of some 700 genera and about 2400 species, belonging to 170 families of
flowering plants. This achievement is colossal, as he had only very few books at his disposal, viz.
Willdenow, Species Plantarum, Persoon, Synopsis, Sprengel, Anleitung and Systema Vegeta-
bilium, DE Jussieu, Families des Plantes, Roxburgh, Flora Indica vol. 1 and W. Jack, Malayan
,

Miscellanies. He had of course also at his disposal the works of Rheede and Rumphius but they
were of hardly any taxonomical use. He mentioned in his Enumeratio that he had seen the plates
of Norona, obviously of a set since lost, but could not have had much profit from them for his
purpose.
writing of the Bijdragen itself was a tremendous task, let alone the research incorporated
The
inthem, a great deal of the genera and species being new to science. This research work has ap-
peared to be of very high quality, testified by the fact that a very large amount of his newly pro-
posed genera still stand and that others, now merged with earlier described ones, were always good
taxa and were later often still recognized as subgenera or sections. A great merit was that Blume

hardly ever failed to recognize their proper affinity and almost always placed them in the proper
family, evidence of his great systematic capacity. In view of the rather primitive state of tropical
botany in his time this deserves great respect. Blume's skills in this field also appeared from a first
attempt to construct a system of affinity for tropical orchids, laid down in the Tabellen en Platen
voor de Javaansche Orchideen (A: 825), issued in part simultaneously with fascicle 6 of the Bij-
1

(9)
 Flora Malesiana

dragen. He complained that he had had no access to contemporary literature on the family by
R. Brown, C.S. Kunth, and L.M.A. du Petit Thouars, which he only received during the print-
ing of his own system of the orchids. This first attempt was much later crowned with his
monograph Flora Javae. Nova Series (A: 1858-1859) of the Orchidaceae, the largest and least
known family of the Malesian flora.
In addition he published in the first five fascicles of the Bijdragen data on the useful and
medicinal plants of the families treated.
Apart from his work with vaccination and his exploration and botanical research work, another
duty had fallen to Blume, when he was in June 1822 appointed director of the Botanic Gardens
at Buitenzorg (Bogor), at Reinwardt's request succeeding him. Reinwardt himself repatriated

in that year. The annual salary was/ 1000. This was a task in itself; besides enriching the garden
with plants he collected during his own travels, he was also in contact with other gardens abroad,
for instance at Mauritius and Calcutta, with the purpose of exchange.
Blume was well aware of the fact that he should attempt to stimulate our consulates in foreign

countries to collect plants or seeds for the garden, a policy which he later also followed when he
was director of the Rijksherbarium. For the Buitenzorg garden he wished to have more Chinese
and Japanese species and to obtain material he wrote to the Dutch consul in Canton and the
representative in Deshima (Japan). A year later, in 1824, he instructed the Dutch in Japan how
to dispatch seeds and plants to Batavia (Jakarta).
Furthermore, Blume sent Javanese and other exotic plants in small baskets to the university
gardens at Leiden, Utrecht and Ghent in the Netherlands and also dispatched seeds to the 'Societe
de Flore' at Brussels.
Blume published the first Catalogus van ... 's-Lands Plantentuin te Buitenzorg (A).
In 1823
were many manuscript names of Reinwardt under the latter's name. Blume himself
In the listing
added several new genera under his own name with valid descriptions. Without doubt Blume was
the botanical 'motor' of this catalogue, Reinwardt having been too much occupied by ad-
ministrative and organizational matters, and besides having been previously occupied by his large
exploration of eastern Malesia. It should be added that Reinwardt's plant-systematical
knowledge was meagre (D: 8).
On 11 June 1822 Blume was also definitively appointed as 'Inspector of Vaccine' and had to
attend to his medical-pharmaceutical duties as well. He reported on the virtues of hotwells in
Krawang (A: 1825, 1839), gained information on the fight against cholera, etc. for which he in-
itiated medical treatment, and paid attention to medicinal plants (A: 1825, 1832). On 12 August
1823 he was appointed commissioner of the civil health service. In short, his duties were manyfold
and his achievements in these years are of tremendous proportion.
In 1824 Blume received permission to extend his research to all Dutch possessions in the East
Indies and was allowed to publish in the journals of Dutch societies. The Government would pay
for the printing of a book on botany, obviously the Bijdragen, with the provision that all
discoveries, observations and prepared specimens would be the exclusive property of the Nether-
lands Government.
In a letter dated 6 August 1825, no. 365, Blume informed the Governor-General about the pro-
posed publication of his large book Flora Javae, pointing out that this was urgent as other persons
who had explored in the Netherlands Indies were already active in having their discoveries printed.
These other persons were obviously the French explorer L.T. Leschenault, the American Th.
Horsfield, and especially the British W. Roxburgh and W. Jack. He said that with the insecure
life in the tropics, when so many fell an early victim to tropical diseases, he felt that he had to

safeguard his research, the result of his extensive field work and observations for science.
Therefore he had decided to publish the very concise Bijdragen in anticipation of the large work
Flora Javae which he had in mind. He mentioned that his own health slightly deteriorated, but
there is no evidence that he was ever seriously ill in Java (D: 11). The Bijdragen were certainly
not merely a striving for priority.

(10)
 Dedication

It was then that years of negotiation started about financing the expensive Flora Javae. For its

elaboration he requested a leave of three to four years in Europe, necessary for the acquisiton of
information which the new literature and the comment of experienced botanists could offer, and
this required visits to some of the famous herbaria in Europe. He offered to stay in Europe on
part of his salary.
September 1825 the Governor-General permitted him a two-years stay in the Netherlands,
In
at one third of his pay. After a frustrated attempt of Blume to ship a large amount of living
material to the Netherlands, and an offer to pay for his own passage, the Government finally
decided by 26 June 1826 to commission Blume for two years leave to the Netherlands on half-pay.
His medical activities and the vaccination were assigned to his colleague Peitch and the botanical
work in the Gardens would be looked after by the gardeners James Hooper and Alexander Zip-
pelius who, together, would be paid from the other half of Blume's salary. These were times of
poor economy in the kingdom.
Blume took with him 29 cases of herbarium material, sailing in the ship 'Christina Bernardina',
destination Brussels, then the capital of the kingdom. He had the good fortune that the ship ar-
rived safely, so many earlier dispatches by shipwreck, for instance several of
having been lost

Reinwardt's. By the end of 1826 Blume arrived in Holland. By far the main part of the collec-
tions were made by himself, minor ones were included, e.g. those made by Reinwardt in Java
and East Malesia (Celebes, Moluccas, Timor), local Javanese collections made by the gardeners
ZiPPELius, Kent and Hooper m the vicinity of Buitenzorg, etc. It should be stressed, however,
that none of the collections of Kuhl and van Hasselt were included, as these were property of
the 'Natuurkundige Commissie'. Later, in 1828, these latter collections were dispatched to the
Museum of Natural History at Leiden by G. van Raalten, who had been taxidermist in the ser-
vice of the 'Natuurkundige Commissie', assisting Kuhl and van Hasselt. Van Raalten was also
a capable draughtsman; he died at Kupang (Timor) in 1829.
Van Breda's archive, now at the 'Hollandsche Maatschappij', Haarlem, contains a partial
abstract of a letter dated 22 July 1825 by G. van Raalten (B: 1825), in which he complains that
Blume - who had inspected the orchids in the Kuhl & van Hasselt herbarium - had noted which
species had been depicted of their collections. He became afraid that Blume's pubUcation would
precede the publication of the Kuhl & van Hasselt plants and found this unfair. He felt extreme-
ly sorry for the misfortunes which befell Kuhl and van Hasselt. This letter was certainly one

of the arguments for later, unjust accusations that Blume stole scientific property. Van Raalten
pointed out that Blume had agreed with van Hasselt to work out the orchids jointly, which
Blume also acknowledged in his Bijdragen; in fact some 27 names have a dual authorship, as I
have elucidated (B: van Steenis, 1980). As a non-botanist van Raalten did not understand that
in such unfortunate situations the dead have no claim unless they left manuscripts.
A testimony that Blume, after his departure from Java, had no access to manuscripts or draw-
ings of Kuhl and van Hasselt is the fact that in the Bogor Library there is - or at least was,
before World War- a book containing drawings of Kuhl & van Hasselt (on Asclepiadaceae,
II

Orchidaceae, is a further proof that Blume did not have these documents (D: 1).
etc.); it

Still, the letter by van Raalten, which was badly understood and interpreted, had influence.

Accusations and slander lead a long life, and are often eagerly reproduced by antagonists. Thus
even Temminck, the director of the Museum of Natural History at Leiden, wrote in 828 - when 1

the Kuhl & van Ha.sselt herbarium was transferred to Blume - that the latter should guarantee
priority to the manuscript names of Kuhl & van Hasselt in publishing, although Temminck must
have been quite well informed about the situation. I regret that Smit (B: 1979) in his essay still

accepted van Raalthn's accusation.

On the arrival ofin Brussels, he reported to I). J. van Hwijck (1786- 1855), administra-
Blume
tor of Education, Arts and Sciences in the Uepartmcnt of the Interior, who was very much im-
pressed by Blume's personality and works. In December 1826 van Ewijck spoke highly of
Blume. praised his diligence and knowledge and declared himself in favour of the /'lora .lavae

(11)
 Flora Malesiana

plans. The Minister contacted his colleague of the Colonies, who in his turn applied to King
Willem I. This was followed by endless discussions who would pay for the publication of Flora

Javae. The result was that Blume received 7000 florins and that the Dutch Government would
buy 50 copies (5 florins for each instalment), the Netherlands Indies' Government would buy 4
copies, and that he was allowed to appoint a draughtsman (Arckenhausen) for a period of four
years. Blume had in mind to publish 250 instalments.
In the meantime Blume pursued his activities in Holland, continued the Bijdragen, and com-
posed a new work under the title Enumeratioplantarum Javae (A: 1827-1828). The treatment
. . .

was more elaborate than that of the Bijdragen. It was published in Leiden. He mentioned on the
title page that he had also used material from Kuhl and van Hasselt, but this is hardly possible

as this came only available to him in 1828 (D: 1).

Blume dedicated the first volume to the Nees von Esenbecks at Regensburg, with whom he
had early friendly relations for several years. Blume's frequent letters to them on his experiences
in the exploration of Java were published in several volumes of the journal Flora, and he frequent-
ly sent them cryptogams, mosses and fungi; when he returned to Holland in 1826 he stuffed empty

spaces in his cases between his parcels with moss samples, especially hepatics, which enabled
Th.F.L. Nees von Esenbeck to publish on Javanese Hepaticae in 1830. Partly out of courtesy
the latter published a paper on Javanese Fungi, with Blume as co-author (A: 1827). As a matter
of fact Blume extended his interest distinctly to cryptogams, and earlier had already pictured and
studied mosses and fungi himself in the field. This interest did not wane, because in 1841 he readily
agreed with Zollinger to buy lichen collections from Java where Zollinger intended to explore.
The second volume of the Enumeratio, dedicated to W.J. Hooker, consists mostly of descrip-
tions of Pteridophyta; in fact it is the first account of them in Java. It proves Blume's thorough
botanical knowledge, because he was mostly versed in Spermatophyta. Notwithstanding that, this
volume is as complete and its contents as accurate as that of the flowering plants, according to
Hennipman (C: 1979).
When in 1828 Blume's leave came to an end, he requested discharge of his position as chief
of the Civil Health Service. This was granted because he would continue to work on botany and
would not return to Java.
By Royal Decree of 22 June 1828 he was granted from 1 July 1828 onwards an annual salary
of 3000 florins for his services and an annual half-pay of 2000 florins, till he had obtained
another position. Blume had to cede in this same year his immense collection of animals and
insects to the Museum of Natural History at Leiden. As compensation he would receive an an-
nuity (B: GiJZEN, 1938).
The first two parts oi Flora Javae appeared under authority of Blume and
in Brussels, in 1828,
his adjunct. Dr. J.B. Fischer. J.G.S. van Breda Ghent and
(C: 1827-1829), then professor at
by profession a zoologist, would participate, or at least elaborate, the Orchidaceae and Asclepia-
daceae. For this purpose the drawings and descriptions of plants made by Kuhl and van Hasselt
were also given to van Breda.
On 31 March 1829 the Rijksherbarium was founded at Brussels, with Blume as director, with
the title of professor. One of his first actions was to instigate that the Botanic Gardens at Buiten-
zorg should regularly provide consignments of plants to the Rijksherbarium, and furthermore,
that the members of the 'Natuurkundige Commissie' in the Indies should not distribute specimens
to foreign herbaria.
The Rijksherbarium did not long exist at Brussels because of the 1830 rebellion, and was saved
in the nick of time and transported to Leiden by Fischer and von Siebold. This subject has been
fully reported by my wife (C: van Steenis-Kruseman, 1962). Blume himself was not on the spot,
because he was on his honeymoon to Geneva. He combined this tour with the object of inspecting
the Roemer herbarium, which was for sale, to see whether it was worth-while to purchase it for
the Rijksherbarium collections.
The Rijksherbarium, after its transfer to Leiden, was at that time not affiliated to the Universi-

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 Dedication

ty, but was subjected immediately to the Ministry of the Interior. That Ministry drafted an In-
struction for the director, effectivefrom the first of January, 1831 (C: van Dam, 1832).
Blume continued the issue oi Flora Javae. Mid- 1830 35 instalments had been published. Unfor-
tunately, the subscriptions appeared insufficient and money ran out, and the work was temporari-
ly abandoned.
Blume did his best to expand the Rijksherbarium collections on a large scale. Via the Ministry
of Foreign Affairs he urged civil servants abroad and in the colonies to collect plants and make
herbaria. For this purpose he composed a booklet of instruction (of which I have not been able
'

to trace a copy) on how plants should be made into a herbarium, as drying plants in the tropics
brings along difficulties by the moist climate and the often bulky and/or fleshy structure of the
was the problem of frequent insect damage once plants are dried.
material. Moreover, there
With some people Blume succeeded. There is e.g. a large collection of several hundreds of
specimens made by the Dutch consul in Venezuela, J.G. van Landsberge, made in 1842. This
collection is arranged by families, but remains unidentified to the present; it contains many
duplicates. On the whole, however, Blume's urging did not meet with great success.
Blume also approached missionaries to collect plants in their territory, and stimulated phar-
macists to do the same; those whom he tutored Leiden he gave special attention and instruction.
at

Although in this way many people sent overseas were aware of his wishes, the results were very
meagre, as compared for example with the results of F. von Mueller in Australia in his contacts
with missionaries. The latter's success is probably to be ascribed to the fact that he maintained
a very regular correspondence with them and kept them timely informed of results. Besides, von
Mueller lived much closer to them.
In general, the attempt to acquire botanical material by stimulating an interest in the tropical

flora among medical men and other residents in the colony and the collecting of specimens was,
as far as I can judge, not successful either. The endeavour in itself was excellent, but possibly
precocious in the early 19th century.
In addition Blume was engaged in buying collections which were for sale. A curious, significant
example was a collection of Javanese plants offered in 1837 for sale to the Government by the
German physician J.G.H. Kollmann, who was in the service of the Dutch East Indian army. This
offer was referred to Blume who found to his great surprise that this collection contained also
the set of duplicates (about 4000 specimens) which he had conscientiously left at the Botanic
Gardens in Buitenzorg (C: van Steenis-Kruseman, 1950; D: 4).
It should be borne in mind that it was factually impossible for Blume to work on incoming col-

lections without having a large staff of botanists at his disposal. From numerous letters in the
'Rijksarchief it is evident that he pleaded time and again for the appointment of staff officers.
Notwithstanding the esteem he was held in by the Ministry of the Interior and the sympathy of
some high officials, notably van Ewijck, it was of no avail. He could not even attain a permanent
position for his two closest collaborators. Dr. J.B. Fischer and his draughtsman and handy-man
J.C.P. Arckenhausen(C:Griepc.5., 1977; D: 12). Financially the Netherlands were at that time
at low ebb. Blume, moreover, was unfortunate with respect to the few scientific co-operators he
had. Van Hasselt and Flscher met untimely deaths and van Breda took another job.
Members of the 'Natuurkundige Commissie' were entitled to work out the results after seven
it was only P.W. Korthals who per-
years of exploration in the East Indies, but in this category
formed excellent work. Korthals was possibly a modest man, in the shadow of Blume, but his
work in the field and in science was of the same high quality. Korthals would have been an ex-
cellent staff member, but after his retirement he devoted his time to philosophical contemplation.

(1) A. DE Candollf. mentioned (B: 1862) thai Biomi; told him the Netherlands Indies' Government had
ordered, at Blumk's request, thai all physicians in their service should have A. P. di ("ani)oi.ij:'s essay Sur

les propnitis des planles (1816) as a botanical guide.

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 Flora Malesiana

as became evident from Zollinger's diary. Blume cannot be blamed for the fact that Korthals
abandoned botany (B: Zollinger, 1841; D: 8).
H. VAN Hall had worked with Blume on a temporary basis from about 1850, but was only of-
ficially appointed adjunct-director in 1854, the only permanent scientific collaborator Blume was

ever allowed.
Deficiency of technical staff was another drawback; here again attempts to expand failed.
Apart from his draughtsman Arckenhausen technical assistants were few. This must in my opin-
ion be one of the main reasons that hardly any duplicates were distributed in order to exchange
material with foreign institutes to enrich the Rijksherbarium. Foreign colleagues complained that
Blume asked for their material, but seldom gave a return. This greedy and monopolistic attitude
made him unsympathetic. Evil tongues claimed that it was Blume's intention not to distribute
duplicates, as he wanted to prevent new species to be described by others. I cannot believe this
to be correct for in any case he could have distributed duplicates of species already described by
himself. Obviously Blume was not in favour of seeing undescribed species published on dupli-
cates. Not until the 1860s, under Miquel's directorship, numerous duplicates were distributed,
partly unnamed. The same open policy was followed by the Herbarium Bogoriense (with an ex-
ception of selected Javanese collections made by C.A. Backer) and this more generous attitude
was also kept up by Merrill in Manila, and from Miquel onwards by the Rijksherbarium. In
the first place this is done for the greater safety of the collections (in that respect we have but to
think of the disastrous effect of the fires in Berlin and Manila) but also because all research on
Malesian plants must be welcomed, irrespective where and by whom. It is self-evident that in case
of free-for-all descriptions a lot depends on the quality of the collaborators. It is true that not
infrequently mediocre or uncritical collaborators have created more extra work rather than solved
the problems for their successors.
A great inconvenience associated with duplicates of the early Dutch collectors was the fact that
they were not numbered, neither by Blume himself, nor by Korthals, Reinwardt, or others.
Through this, typification is difficult and it is sometimes impossible to know which duplicates
belong to which collection. The more praiseworthy a Teijsmann, who consecutively numbered the
Buitenzorg collections! But then the latter had more personnel. Blume's limited staff was certain-
ly one of the reasons that the numerous collections remained undivided. Whom could he trust to

distribute the unmounted collections in a responsible way?

As already mentioned it was not before the 1860's that Miquel, Blume's successor, instigated
the policy of free distribution of duplicates, but certainly did not do it himself; he had it done
by technical personnel. Without doubt the distribution of duplicates was extremely important as
the result was the acquisition of numerous duplicates in exchange from foreign herbaria in
Europe. It is a pity that this distribution lacked carefulness as regards the labels. Occasionally
specimens with Blume's handwriting were sent elsewhere, for instance to Paris, while specimens
retained at Leiden have labels written by a clerk. This is not seldom a nuisance in connection with
the assignment of the holotype. Sometimes the use of wrongly printed labels is confusing, for in-
stance of Korthals specimens of which sometimes 'Sumatra'-labelled plants are really from SE.
Borneo. At Blume's and Miquel's times most of the Malesian collections were not mounted; this
was only done towards the end of the 19th century.
As the prime botanist he was, Blume's interests were by no means restricted to those of a scien-
tist working in seclusion. He was always keen on the development of the colony towards better

living and status. He stressed the importance of promoting the cultivation of plants not only in
the interest of big enterprise, but he held the opinion that there had to be a balanced situation
for the benefit of all! This comprised also the introduction of new, useful plants. If one reads his
general papers it appears that he had wide interests from his early stay in Java onwards. In the
first five instalments of the Bijdragen he provided families with notes on their useful species. He

wrote a monograph on the peppers (A: 1826) and as early as 1820 he took the initiative to advise
the Government on the importance of cultivating indigo and of importing cochineal, and last but

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 Dedication

not least to import Cinchona, which materialized only three decades later through Hasskarl. In
many papers he advocated more activity in agricultural matters and stressed the importance for
national well-being in commercial, hence financial, aspects, for the Dutch as well as for the native
people.
As a medical man, in his capacity of 'Inspector of Vaccine' and during his
, many travels, Blume
was of course in intimate contact with the Javanese people and he took their welfare as much to
heart as that of the Dutch people; he clearly regarded them all as co-citizens. For example, he
pleaded openly in a letter to theGovernor-General of the Indies (A: 1829) for the desirabihty of
abolishing opium, as he found this a menace for the population; only much later this was
regulated indeed by the Opium Law.
In 1842 Blume founded, together with Ph.F. von Siebold and on the instigation of J. Pierot,
the 'Societe Royale pour I'Encouragement de I'Horticulture dans les Pays-Bas' (Royal Dutch
Society for the Advancement of Horticulture). This was part of his endeavour to make botany
subservient to the general interest of the kingdom and to create a stimulant for new financial and
commercial interests. In a first issue of the above-mentioned 'Societe' (A: 1844) he compiled a
large list of useful plant species. Also later he showed his unfailing devotion by a stimulating paper
on timbers resistant to pile-worm (A: 1859).
Altogether he held enlightened, progressive ideas - not so popular in those days - and in his
opinion the native people ought to have their share of welfare, not in the least because their man-
power was an prosperous colony. In this respect it is significant that he named
essential aspect of a
the genus Santiria after Bapa Santir, an old Sundanese, who accompanied Blume on his explora-
tions of Mt Salak. It was Junghuhn who took this amiss (B: Junghuhn, 1853) and suggested that
Blume was consciously deceptive in pretending to be generous, but really threw a blame on great
botanists and other dignified man who were the only persons entitled to be honoured by eponymy.
In his colonial arrogance Junghuhn called Bapa Santir an inferior person, not more than a sim-
ple 'pakkedrager' (kuli, carrier), whereas in all probability Bapa Santir was an intelligent man
and an outstanding local authority on plants who knew his way in the forest, knew the vernacular
names and uses of forest plants and assisted Blume in many ways. It is testimony of the irony
of fate because in history Junghuhn is reputed to be the pioneer and advocate of a progressive
society of freethinkers, whereas Blume is remembered as a distinctly conservative person, though
all his writings give evidence of a progressive, liberal mentality. It appeared that Blume, mirabile

dictu, was the more enlightened of the two; he was certainly devoid of any racial prejudice.
In 1843 Blume started the journal De Indische Bij, another endeavour to promote an interest
among the Dutch public in the understanding of the colony. Only one volume was issued (1843),
mainly filled with papers by himself and his friend C.F.E. Praetorius, Director of Cultures in
Java, on all kind of subjects, partly political, partly ethnographical, on Borneo and South
Sumatra, and on plant fibres.
Returning to Blume's scientific works: in spite of the untimely abandoning of his Flora Javae,
he set up another large-scale work in the thirties, Rumphia, the scope of which included also other

parts of Malesia. The first fascicle appeared in 1836. It consisted finally of four volumes
(1836-1849). This work was of the same critical standing as Flora Javae, to which it was similar
in size and printing. In a sense it is an attempt towards a Flora Malesiana. Towards the end of

the forties Blume again managed to issue some important parts of Flora Javae, namely the Filices
(instalments 36-39 in 1847 and 40 in 1851) and the Loranihaceae (instalments 41 & 42 in 1851).
How these issues and Rumphia were financed is unknown to me.
The abrupt end of Flora Javae was regrettable and H.C van Hail, professor of botany at
Groningcn, was much concerned about its continuation, which he found of national importance
(C: van Hall. 1856). In a session of the Royal Academy of 28 June 1856 he proposed that this
lofty body might form a committee to approach Blumi; in order to come to a proposal from the
Academy to the Government for further financing Flora Javae; at that time 42 instalments, each
with 6 plates, had been issued. do not know if van Haii 's pleading led to any further action,
I

but it shows that / lora Javae had supporters.

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 Flora Malesiana

After Blume's death there obviously remained illustrated printed material for a continuation
of Flora Javae. These 23 coloured plates, called Planches inedites, mostly represented species of
Loranthaceae and Ericaceae, all provided with analyses. They were offered for sale as a packet
by the firm van der Hoek, Leiden, in 1862 or 1863 (A: 1863; C: van Steenis, 1947).
Towards the end of the forties, when Blume was in his prime, he must have been disappointed
with the untimely discontinuation of the two works on which he had set his heart. Flora Javae

and Rumphia, and the insufficient public interest in his journal De Indische Bij. Moreover, clouds
had gradually gathered round his claim that the Rijksherbarium had the monopoly for housing
and possessing all collections made in the colonies by persons in the pay of the Government. He
based this claim on the Instruction for the Rijksherbarium of 1832. This claim, however, was an
optimistically exaggerated interpretation of art. 10 of this instruction which read (transl.): 'The
Director will attempt to acquire collections, notes and drawings from all civil servants or people
in the pay of the Government through proposals at the proper place and authority' (C: van Dam,

1832). Blume may have had a moral right to claim these collections, but could not refer to a legal
right. His claim was not attended to and this must have been a thorn in his flesh.
Blume opposed the founding of Herbarium Bogoriense by Teijsmann in 1844, claiming that
the latter should send the specimens to the Rijksherbarium, or at least the duplicates, but he found
insufficient understanding with Teijsmann, who foresaw that he would have little profit from this
in the way of a speedy naming of the specimens. Furthermore, Teijsmann's assistant, J.K.

Hasskarl, had assembled a large private herbarium which he took with him on repatriation to
Germany. Then von Siebold's herbarium was elaborated at Munich by Zuccarini (D: 2) where
the types were left. W.H. de Vriese, professor at Amsterdam, had acquired the herbarium of
Splitgerber, made in Surinam, but had not donated this to the Rijksherbarium. Finally,
JuNGHUHN, officially belonging to the medical department in Java, had assembled a very large
herbarium in Java, which Blume could not get into his hands (D: 3). It was purchased by Leiden
University, under the condition that it should not be incorporated in the Rijksherbarium; it was
entrusted to de Vriese. Finally, there was the rising star of tropical botany, F.A.W. Miquel, who
originally published valuable monographs of Piperaceae, Cycadaceae, Casuarinaceae, Melocacti
(partly for de Candolle's Prodromus), and later elaborated various large families in Martius'
Flora Brasiliensis. He became also more and more interested in Asiatic plants, starting with his
Analecta Botanica Indica, published by the Royal Academy. Miquel was a man of immense out-
put and diligent handling of material, with an open mind for collaboration, which he brought in
practice himself. Considering that, if the Junghuhn collection fell into Blume's hands, identifica-
tions would be endlessly retarded, combined with Junghuhn's natural desire that it should be
speedily worked out, de Vriese reasonably entrusted Junghuhn's collection for this purpose to
Miquel. With elaborate support (e.g. Bentham's), the latter indeed published the Plantae
Junghuhnianae. This must have caused immense irritation to Blume, who was constantly on the
barricades defending his institute, stressing again and again that collections made by government
officials with government money ought to be deposited in the Rijksherbarium. This monopoly
also concerned himself. My wife (C: van Steenis-Kruseman, 1979: 51) wrote: 'whatever has been
said to Blume's discredit, one thing is certain, and that is, that he was possibly the only botanist
(and a devoted, not to say inspired one) in his period who had no private herbarium.'
It is ironic but true that Blume's strict monopolistic claims made people reluctant to put their

collections under his care, though Blume was, although not legally, at least morally in his rights.
Even admitting that his claims were correct, it must be said that he should have realized that, if
all these collections had been donated to the Rijksherbarium, he could as a single person never

have mastered them. This would have been necessary, as some people wanted names and iden-
tifications. He should have tried to compromise and initiate collaboration and division of labour,
at least with Miquel and de Vriese, and not sit tight-fisted on propriety of collections. But ob-
viously he could not well adjust himself to the changing conditions of the times and the rise of
capable colleagues in his specialized field. This led to most unfortunate friction and a clash of

(16)
 Dedication

personalities. He offended especially Junghihn in writingwith his sharp pen an acid comment
in Rumphia (1847 or 1849?) on Junghuhn's so-calledLycopodium arboreum which he had 'at
first sight' recognized as belonging to the conifer genus Dacrydium, and Blume renamed Primula
imperialis Jungh. as P. Kuhlii Blume, claiming that Kuhl had found this first and thus had priori-
ty for eponymy, nomenclaturally wrong of course. Junghuhn complained that Blume begrudged
him to describe Acer javanicum and had renamed this wrongly A. niveum, in which Junghuhn
in turn was wrong. In short, about 1850 the fight was on and several very sharp and polemic

papers were published to and fro (D: 13).

The unfortunate result was that Blume became a still more isolated and probably a rather em-
bittered person. Apart from odd fascicles of Flora Javae and Rumphia he had no opportunity
for further great undertakings. He then put himself to proceed with a subject, stipulated in the
1832 Instruction for the Rijksherbarium, namely compiling a catalogue of the collections of that
herbarium. \s this implied identifications, this was not a clerical task for a non-botanist. My wife
mentioned (C: van Steenis-Kruseman, 1979: 35) that the scientific arrangement of the collections
was started by J. Pierot (1831-1840), who was succeeded by J.H. Molkenboer (assisted by C.
Kerbert and Schultes Jr, the son of J. A. Schultes) (1840-1846), and finally by H. van Hall
(1853-1862). Their work of course facilitated Blume's later Museum Botanicum. These helpers
were named 'assistants', only van Hall was designated the title 'conservator'.
In December 1850 Blume had to face an official new Instruction for managing the Rijksher-
barium collections (C: Thorbecke, 1850). This was to meet official complaints by de Vriese,
Junghuhn and von Siebold and especially Miquel, all influential persons, who wanted to bor-
row material, requests only reluctantly given in to by Blume. Miquel wrote to von Schlechten-
dal (B: Stafleu, 1970: 321): 'Es ist mir endlich gelungen, das Reichs Herbarium zu offnen. Nach
einem Befehl der Regierung sind die Samml. aus Borneo, die noch ganz unbearbeitet waren, mir
zur Disposition gestellt.' Blume was ordered to proceed with the catalogue; no unicates were to
be removed from the collections; furthermore, the director had to refrain from publishing
discoveries by still living persons of the former 'NatuurkundigeCommissie', unless with their con-
sent. It must have irritated him considerably that responsibility and authority were restricted.
The catalogue, named Museum Botanicum, was printed in fascicles, all filling one sheet (16
pages), apparently with the intention to publish the fascicles monthly. It consists of two volumes
in which the fascicles of volume one are dated by the year and month. The first volume was dated

from 1849 to 1852 and finished with an index. The second volume was started with a fascicle dated
1852, but fascicles 2-8 are undated, fascicles 9-16 being dated November 1855 to June 1856.
There was no index; this was later composed by myself and Chew Wee Lek (C: van Steenis &
Chew Wee Lek, 1974). As Beumee (B: 1948) and Stafleu & Cowan (B: 1976) have pointed out
there are discrepancies about the dates of publication and this induced the latter towards sug-
gesting that Blume withheld literature from his colleagues (Miquel, Weddell, etc.) and that in
other cases the possibility of antedating cannot be excluded. Miquel (B: 1856) severely criticized
the doubtful datings of the fascicles. It is quite probable that not every fascicle was for sale at
the published date, but sold in lots, and confusion remains. In the absence of well-founded data
regarding the authority which paid for the publication, who arranged the sale, and whether one
could subscribe, we must refrain from further comments (D: 6). Possibly Blume still had a
manuscript for one other fascicle which is known as Metan}>es botaniques (A: 1855). Up till the
present it was assumed not to have been effectively published. This is, however, wrong, as I have
discussed earlier (B: van Steenfs, 1986). The pamphlet was privately published and donated by
Blimi; to his close friends; at least two copies still exist.
The Museum Botanicum is an important, critical work; it contains some attempts towards revi-
sions and, though species and genera from all over the tropics were dealt with, the main text refers
to the Malesian flora. We do not know why the Museum Botanicum was rather abruptly abandon-
ed. It is not unthinkable that Blume wanted to unburden himself from his old love, the Or-
chidaceae, and saw an opportunity to publish this masterly work which he had had constantly in

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 Flora Malesiana

mind since his early Buitenzorg years. This work had been interrupted several times, first when
his collaboration with van Hasselt came to an untimely end by the latter's death, and later by
the early leaving of van Breda. Now it was published as Flora Javae, Nova Series (A:
1858-1859). There is also a French-titled edition, with a preface in French, but otherwise iden-
tical. According to W.E.G. Seemann Blume complained that the Government had not
(B: 1859)

contributed to its Blume, who was a man of means, had taken the risk of
financing; obviously
financing it himself. Besides the excellent works of R. Brown, Lindley and Reichenbach on the
Orchidaceae and the affinities within the family, also Blume's work is very important and of
similar standing,and naturally of special importance for Malesian botany.
Blume, naturalized as a Dutch citizen in 1851, died in Leiden, after a long, painful illness on
February 3, 1862, at the age of 65.
As said before, Blume is through his large oeuvre - including eight important and critical

botanical works of high standard: the Catalogus, the monograph on Piperaceae, the Bijdragen,
the Enumeratio, Flora Javae, Rumphia, Museum Botanicum, and Flora Javae, Nova Series - one
of the great botanists of the former century. A ninth treatise, on cholera in Asia (A: 1831), is
medical.
His creative output is imposing. He distinguished eight new families, to wit, Apostasiaceae
(now mostly judged a subfamily among Orchidaceae), Burmanniaceae, Cardiopteridaceae,
Dipterocarpacaeae, Hernandiaceae, Myricaceae, Sabiaceae, and Schisandraceae. In addition he
described, from Malesia alone, some 300 new genera of which 160 are still used, and 140 are now
in synonymy, either for reasons of nomenclature or for new systematical insights. However, they
were all proper taxa and are still frequently recognized as infrageneric taxa, e.g. Tarrietia and
Campanumoea. Furthermore, he described his genera and species almost always in the proper
families cq. genera, testimony of his systematic vision.
As to his scientific achievement, his talent was soon recognized, both in Holland and abroad
and he was soon made a member of learned societies (F). As usual for members of the 'Leopol-
dina', they should have a cognomen; Blume took for himself the well-chosen name Rumphius
Secundus.
Many generic names (E) and very many species were named after him. We are pleased that the
journal of the Rijksherbarium, Blumea, is named after him.
As an explorer Blume was exemplary in multidisciplinary approach by making observations on
the spot, having a draughtsman with him, interrogating the native people about the uses and ver-
nacular names, collecting insects and other animals, and paying attention to soils, mineral wells,
etc., and by timely reporting about his field research, a good habit which young explorers of the
present day should take more to heart. Through his medical profession he made also observations
about native diseases and tried to cope with these to relieve suffering of the people.
All his endeavours in this field and also his many advices on agricultural and horticultural af-
fairs were focussed on tying up scientific botany and practice for the benefit of society. As such
he was the opposite of the scholar in the ivory tower. His sharp observation power paired with
interest were not confined to botany, as appears from his conclusions on serious contagious
diseasesamong which cholera and typhoid were the most dangerous. As 'Inspector of Vaccine'
he went to Central Java on inspection during a cholera epidemic and observed that the disease
was especially prevalent in the lower lands, and less so in villages in the mountains. He deduced
that cholerawas spread by the polluted water and that the freshwater wells in the mountains were
less contaminated.He prescribed all sorts of simple means for a diet and medicinal substances
from native plants, but in the first place he advised boiling the drinking-water, and optionally ad-
ding some cinnamon in polluted areas. When settled at Leiden Blume published a book on Asiatic
cholera (A: 1831). Shortly after, he attended a congress of naturalists and surgeons at Halle, a
town at that time suffering from a serious epidemic of cholera. He observed that in the rather
isolated 'Franckische Stiftung', a community of some 1800 souls, there was no cholera. These
people were followers of the pietist A.H. Francke, founder of this 'Stiftung' in 1663. To his

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 Dedication

satisfaction he observed that this group of people got its own water from wells through a systein
of tubes several miles outside Halle. In Holland, where at that time cholera also was a serious
disease, he noted that it was rare in the southwestern island province of Zeeland, and he correlated
this with the fact that drinking-water there was mostly rainwater. The next year he wrote a pam-
phlet (A: 1832) on the subject which he had printed in 1000 copies at his own expense. He forward-
ed free copies to all municipalities, stressing that boiling all drinking-water was the simple remedy.
One would expect that the arguments for this cheap advice were immediately accepted, and at least
tested. But his opinion wascompletely overruled by the powerful voice of G.J. Mulder, a chemist
of great influence, who declared that Blume's conclusions were nonsense and that all water from
ditches and canals was fit for drinking and had nothing to do with the dispersion of cholera.
Blume's role looks to me similar to the one of Semmelweiss in Vienna and his fight against
puerperal fever. Thirty years later Blume's conclusions were of course fully accepted.
As a civil servant Blume excelled in activity for the benefit of the country and colony, in pro-
moting the interests of agriculture and horticulture, throughout his life. As a director of the Rijks-
herbarium he did all he could under the circumstances, to raise it to a first-rate institution. As
my wife (C: van Steenis-Kruseman, 1979: 37) put forward, Blume succeeded in greatly enriching
the Rijksherbarium with important standard collections, e.g. Spanoghe (Timor), Korthals (W.
Malesia), Forsten (Celebes), von Siebold, Textor and Burger (Japan), Sieber, Schultes,
Cuming, Persoon, Dozy, and Molkenboer (Bryophytes). Besides this, he acquired large sets of
duplicates from the collections of Wallich, Ecklon & Drege (Cape), and Plantae Preissianae
(Australia). He purchased also several smaller collections from South America.
In the preceding pages I hope to have succeeded in making it clear that the slander of which
Blume was a victim was unfounded and can be defused by factual evidence.

will now proceed with some remarks on Blume's personality and his motives, as an addition
I

to what already may transpire from the precedings pages. Much can be learned about this from
his published papers. A perusal of his personal letters to his colleagues abroad will add probably
more but this falls beyond my capacity. Another source is the opinion of third parties which can
be found, for instance, in biographical papers. However, the latter are mostly an evaluation of
the quantity and quality of achievements and seldom enter into personal facets. Among the
obituaries of Blume only Goddijn (B: 1931) ventilated some well-considered remarks.
Blume was a most intelligent person devoted to science and with a broad outlook, dedicated
to promote the interests of his second fatherland and all its inhabitants. He pleaded for a society
in which everyone, irrespective of race, should benefit from increasing profit. He was antagonistic
to the idea of a 'Cultuurstelsel' and pleaded for a free society.
'

As to his social contacts, it is difficult to ascertain much factual evidence without having access
to his personal correspondence. His family life seems to have been happy and his wife sometimes
shared his stays abroad. In Java he had good friends, e.g. Praetorius, Spanoghe and several
others. As to his contacts with foreign colleagues, Blume apparently often took part in the annual
'Versammlung Deutscher Naturforscher und Aerzte' in Germany.
in his native country he must have had friendly relations, among them the Nees von Esenbecks
at Regensburg. According to Roland (B: 1944) he and his wife paid in September-October 1834
a lengthy visit to Paris where he had many friends (amongst others Decaisne, Brongniart). He
met many prominent personalities, compared material from Java o^ Araceae, Annonaceae, etc.
with Paris collections, bought books, acquired and bought collections and frequently stayed with
J.E. Gay (who had very rich collections) for studying material, often together with A. Moquin-
Tandon, the monographer of Chenupodiaceae. The latter said of Blume (B: Roland, 1944: 74):

(I) In the Netherlands East Indies the system in which ihc local people were forced lo grow various sorts
of crop suitable for the European market (in force mainly in Java, 1828 1890).

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 Flora Malesiana

'Je suis sorti avec M. Blume dont j'aime beaucoup la figure gracieuse, la gatte et la vitalite vrai-
ment meridionale.'
The fact that so many honours befell him (F) indicates that he must have enjoyed the sympathy
of many persons abroad who took the initiative to make the proposal. In political circles in
Holland he certainly was also appreciated; the fact that he did not succeed in building up a staff
of collaborators for which he pleaded in vain for two decades, can be ascribed to the rather poor
economic situation of the kingdom, unfavourable for creating permanent scientific positions.
I believe that the later strenuous relations with his Leiden scientific contemporaries must, to
a large part, be ascribed to feelings of envy towards his great capacities by the autocratic von
SiEBOLD and Junghuhn, the mediocre de Vriese and the frustrated Reinwardt and Hasskarl,
who all eagerly grasped any opportunity to damage his image. In this they were in a way assisted
by Blume's rigid, autocratic personality.
Unfortunately it is difficult to obtain more impartial contemporaneous information from
neutral, disinterested parties. Among the rather neutral sources there is one, from the Swiss
Heinrich Zollinger, who wrote an extensive diary which is now deposited in the Central Library
atZurich (B: Zollinger, 1841). The part of this diary relating to Zollinger's stay at Leiden, Oc-
tober to December 1841 was typed out and generously put at my disposal by Prof. Dr. H. Wan-
,

ner, Zurich.
Zollinger, at the suggestion of A. de Candolle, was considering a botanical-zoological ex-
ploration of Java and wanted subscriptions from biologists, authorities, and institutes for his
endeavour. After having obtained some in Switzerland, France, and Belgium, he came to Hol-
land, in 1841, where Miquel gave him some hope. Withhis letters of recommendation he tried
to obtain subscriptions from the Rijksherbarium and from the National Museum of Natural
History at Leiden. Above all, he sollicited free transport for himself and his equipment to Java
from the Dutch authorities as a contribution to his future work in the colony. In his diary Zol-
linger gave his free opinion on several scientists he visited (Reinwardt, Temminck, Schlegel,
DE Vriese, Ammann, Splitgerber, Schwaner, von Siebold, Korthals) (D: 8). He paid visits
to Blume and noted about him (B: Zollinger, 1841: 25): 'Blume ist ein kleines, elegantes,
vornehmes, lebhaftes Mannchen, das sich auf verschiedene Weise ein grosses Vermogen und eine
grosse Reputation erworben hat. Er war sehr freundlich und zuvorkommend, gab mir Rathe aller
Art. Ob nun im Herzen es anders aussieht, warum er so gegen mich ist, weiss ich nicht. Ich will
das Beste denken und auf meiner Huth sein'; I.e. 29: 'Er schwatzte mir freundlich vor, wie bis
jetzt noch kein Privatunternehmen wie meines auf Java, gelungen. Wie ich dort nichts neues mehr
finden werde, besonders im Westen; ich miisse mich zeitig nach einer Anstellung umsehen. Aus
dem Ganzen schien mir hervorzugehen dass er mich ganz abzuhalten oder fiir den hollandische
Dienst zu gewinnen sucht; denn auf beide Weise kommt nichts in fremde Hande, oder im letzteren
alles zuerst in die seinen'; I.e. 31 (summarized in English): von Siebold suggests that Blume is

a rather tough person and reckons that Zollinger will anyway send him plants, obviously
alluding to Blume's refusal to subscribe to a set of Zollinger's plants; I.e. 33: Blume subscribed
to buy Lichenes from Java and offered him an iron trunk. He spent another evening in Blume's
beautiful house, with a large library, but the trunk did not turn up. 'Blume hat fiinf hiibsche
Kinder und eine hochgebildete Frau. Er zeigte mir seine Rumphia und andere Sachen, die auf Java
bezug haben. Wir sprachen meist von Indien. Ich soil 3 Kisten (lebende) Pflanzen miterhalten'
(obviously for the Botanic Gardens at Buitenzorg). At the advice of Blume he went to Mr. Ar-
RiENS, a high official at The Hague, who suggested an audience with the Minister of the Navy,
but Zollinger had no success; all he got was a permission to collect in the colony, antiquities ex-
cepted. In passing, Zollinger followed Blume's advice and sollicited to be attached to the
Botanic Gardens at Buitenzorg, but there was no vacancy at that time. Thus, Zollinger had not
much success at Leiden, as far as botany was concerned. It remains guesswork whether Blume
could have achieved more for him if he had backed him up.
Summing up my impression of Blume's personality, it appears that he was not a social,

(20)
 Dedication

amicable person, but self-centered and keeping aloof; also conscious of his capacities and dignity
but lacking flexibility. However, his motives were honest, and this becomes clear from scanning
his own writings and other literature, if judged against the background of his time and
cir-

cumstances. It is true that he had a sharp pen and in defending the rights and interests of the Rijks-
herbarium his acid reprimanding of Junghuhn, no less a dominant authority than himself, un-
necessarily hurt personally, which, to say the least, led to a severe estrangement.
However, the slander to which Blume became a victim is unjustified, and may well have been
induced by jealousy of his brilliant scientific achievements and envy of his monopoHstic position
at the Rijksherbarium. In my view Blume was an enlightened scientist, whose image
may hereby
be restored.

C. G.G.J. VAN Steenis

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 Flora Malesiana

Appendix A - Blume's publications

1817 - Dissertatio inauguralis medica, de Arsenica et Ratione qua in Animalia agit. Leiden. 49
pp.
With verses by D.J. Veegens, a friend, and Prof. S.J. Brugmans.

1821 - Minerale wateren van Tjipannas en Tjiradjas.

Bataviaasche Courant, 15 Sept. 1821.
Repr. in Indisch Magazijn, Tweede Twaalftal, no 1/2, 1845: 162-166.
Contains a chemical analysis, obviously made by Blume himself, of the mineral contents
of these waters.

1822 - Gedachten op eene reize door het Zuid-Oostelijk gedeeUe der Residentie Bantam.
Bataviaasche Courant, 16 Febr. to 30 Nov. 1822.
Repr. in Indisch Magazijn, Tweede Twaalftal, nos 3/4, 1845: 1-36.
Report of trip, describing the history, anthropology, ethnography and poHtics of the Badui
people in SW. Java. No botany involved.

1 822 - Beschrijving van de heilige graven der Badoeis in het Zuid-Oostelijk gedeelte der Residen-
tie Bantam.
This appeared as the chapter 'Mengelingen' in the Bataviaasche Courant, nos 7, 8, 10, 13,

27-29, and 32, 16 Febr. to 30 Nov. 1822.

Repr. in Indisch Magazijn, Tweede Twaalftal, nos 3/4, 1845: 1-36.
An ethnographical description of the Badui people in SW. Java, their sacred graves, etc.
In the library of the Institute Taal-, Land- & Volkenkunde, Leiden University, there is a 85
pp. manuscript (H 75) with the title 'Gedachten op eene reize, in de maanden December en
Januari jl., in het zuidoostelijke gedeelte der Residentie Bantam gedaan. Getrokken uit de
Javaansche Couranten van J 822,' which is probably copied literatim.

1 823 - Catalogus van eenige der merkwaardigste zoo in- als uitheemsche gewassen, te vinden in

's-Lands Plantentuin te Buitenzorg. Batavia. 112 pp., 1 pi.

Several new genera and species. Many nomina nuda under Reinv^ardt's name.
Repr. in Arnold Arboretum 1946.

1823 - Beschrijving van eenige gewassen, waargenomen op eenen togt naar den Salak in den
jaare 1822.
Verhand. Batav. Genootschap van K. & W. 9: 129-202.
Mostly descriptions of plants {Magnoliaceae, Loranthaceae, Dipterocarpus, Cedrela, Piper,
etc.).

1823 - Letter to Nees von Esenbeck. Flora 6: 713-716.

Report on a planned trip in Java.

1823 - Bijdrage tot de kennis onzer Javaansche eiken.

Verhand. Batav. Genootschap van K. & W. 9: 203-223, 6 pi.

Account of Quercus in Java (incl. also Lithocarpus).

1823 - (with C.G. Nees von Esenbeck) Pugillus plantarum Javanicarum, e Cryptogamicarum
variis ordinibus selectus.
Nova Acta Acad. Caes. Leop. -Carol. 11 (1): 117-138, pi. 12 & 13.
Descriptions of Pteridophytes, the new species under dual authorship 'Nees & Bl.'.
(22)
 Dedication

1 824 - Letter to Nees von Esenbeck: Veber die Vegetation des Berges Gedee aufder Insel Java.
Flora 7: 289-295.
Extract from a larger paper in Dutch, see below (1825). Sketches on the exploration of Mt
Gedeh made together with the hortulanus Kent. Blume did not ascend Mt Pangrango.

1824 - Epidemie onder de buffets.

Bata\iaasche Courant, 10 Jan. 1824: 'Verslag van den kommissaris van den burgerlijk
geneeskundigen Dienst in Nederlandsch Indie C.L. Blume.'
See also: Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 91-94.
Epidemic disease among the buffaloes.

1825 - Letter to the Governor-General, dated 8 Dec. 1824, published in the Bataviaasche
Courant, 12 Jan. 1825.
Report on Blume's discovery of Rafflesia in Nusa Kambangan I. (S. Java), the first

discovery of the genus in Java. He did not name it here.

1825 - Bestijging van den berg Tjerimai, gewoonlijk genoemd Tjerme, in de Residentie Cheri-
bon.
Bataviaasche Courant, 2 Febr. 1825.
Repr. in Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 102-116.
Report of a trip from Krawang eastwards to Panarukan, Linggadjati, culminating in the as-
cent of Mt Tjeremai, with many botanical data on plants encountered.

1825 - Over de gesteldheid van het gebergte Gedeh.

Verhand. Batav. Genootschap van K. & W. 10: 55-104.
Lively topographicaland botanical description of an ascent of Mt Gedeh from Bogor via
Puntjak, along Megamendung, Tjibeureum and Kandangbadak through the crater and
along the Alun-Alun to the summit. Blume did not ascend Mt Pangrango, and thus missed
Primula imperialis.

1 825 - Inlandsche middelen tegen diarrheen.

Bataviaasche Courant, 23 Febr. 1825.
Native recipes against diarrhoea.
See also: Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 116.

1825 - Tabellen en Platen voor de Javaansche Orchideen. Batavia. 5 tab., 16 pi. Folio.

Famous exposition of a system of the Javanese orchids and their affinities; 73 spp. depicted
in detail. Issued with the Bijdragen (1825-1827) part 6.

1825 - (with C.G. Nees von Esenbeck & C.G.C. Reinwardt) Hepaticae Javanicae editae con-
junctis studiis et opera.
Nova Acta Acad. Caes. Leop. -Carol. 12: 181-238, 409-417.
Account of hepatics in Java.

1825 - lets over de planten onder den naam van Paima. bij de Hindostaners en de Javanen
bekend.
Bataviaasche Courant, 9 March 1825.
Repr. in Indisch Maga/ijn, Tweede Twaalftal, no 3/4, 1845: 179-183.
A note on plants known under the vernacular name 'patma' = Rafflesia).
(

(23)
 Flora Malesiana

1825 - Korte beschrijving van de Pat ma der Javanen.

Bataviaasche Courant, 23 March 1825 (22 pp., in L).
Repr. in Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 183-194.
Short description of the 'patma' (-Rafflesia) of the Javanese.

1825 - Die Patma-Pflanze der Indier und Javanesen und Beschreibung einer neu entdeckten
Blume auf der Insel Noesa Kambangan, die an Grosse alle bis dahin bekannt gewesenen
ubertrifft.
Liter. Wochenbl. der Borsenhalle, Hamburg, no 29: 454-462. Repr. in L.
As the preceding.

1825 - Beitrage zur Kenntnis von Bantam, dem westlichsten Bezirk auf Java.
Hertha II: 227-257.
Not seen. Probably similar to entries in 1822.

1825 - Letter to Th.F.L. Nees von Esenbeck: Reise von Batavia nach Krawang in der Preanger
Regentschaft. Flora 8 (2): 577-585.
Report of journey from Batavia to Krawang.

1 825 - Etwas Uber die Rhizantheae, eine neue Pflanzenfamilie, und die Gattung Rafflesia insbe-
sondere.
Flora 8 (2): 609-624.

1825 - Letter to Th.F.L. Nees von Esenbeck: Ueber Pflanzen der Gegend von Batavia. Flora
8 (2): 676-680.
Flora of the vicinity of Batavia.

1825 - Letter to the Governor-General, dated 20 Nov., on the flowering of a new species of a
new genus of Araceae with a very large inflorescence, obviously Amorphophallus cam-
panulatus, in the Botanic Garden, with reference to Tacca phallifera Rumph.
Bataviaasche Courant, 23 Nov. 1825.

1825-1827 - Bijdragen tot de Flora van Nederlandsch Indie. 17 fascicles, 1169 pp.
For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 236.
In all 107 families are treated, in which 700 genera and over 2300 species were incorporated.
There are many new genera and very many new species, all described in concise Latin. In
the first 5 fascicles each family has also a paragraph with notes on its useful plants.
On p. 265 Blume mentioned that his plan was to treat the orchids together with van
Hasselt; 27 species out of the 296 were jointly described. Through van Hasselt's early
death this joint venture was frustrated.
There is a typed Index to the names in L.
Data on useful plants mentioned in fascicle 1 were copied in Alg. Konst- en Letterbode 1826-
1: 26-29, 37-41.

1826 - Monographie der Oost-Indische pepersoorten.

Verhand. Batav. Genootschap van K. & W. 11: 139-245, 6 pi., 41 fig.

Monography of Netherlands-Indian species of Piper.

1826 - De Tacca Culat van Rumphius wedergevonden. Mededeeling van de waarnemingen van
C.L. Blume.
Alg. Konst- en Letterbode 1826-1: 333-334.

(24)
 Dedication

Report about Blume's recollection of a Rumphian aroid in the island of Nusa Kambangan,
S. Ja\a: Amorphophallus campanulatus.

1826 - Letter to Nees von Esenbeck: Bruchstucke einer Reise auf der Insel Java. Flora 9 (2):
417-426, 433-441.
Report on a trip in NW. Java, including also an ascent of Mt Tjeremai.

1827 - (with Th.F.L. Nees von Esenbeck) Fungi Javanici.

Nova Acta Acad. Caes. Leop. -Carol. 13 (1): 9-22, pi. 2-7.

1827 - Over een nieuw plantengeslacht, de Brugmansia, uit de natuurlijke familie der Rhizan-
theae.
In: H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen 2: 419-423.
Brugmansia, a new genus of the Rafflesiaceae.

1 827 - Observations sur le structure des poivres.

Ann. Sc. Nat. 12: 216-224.
Extract in French of the monograph of Piper (1826).

1827 - Bijdrage tot de kennis van het landschap Bantam, in het westelijk gedeelte van Java, etc.
Cybele (Tijdschr. Bevordering Land- en Volkenkunde) VI' stuk: 1-36.
Contribution to the knowledge of Bantam, West Java. Almost literatim reproduced under
the same title in Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 1-36.

1 827 - Over de siaat der indigo-teelt.

In: P. VAN Griethuizen, Over de staat der indigo-teelt. De Nederl. Hermes, Tijdschr.

Koophandel, Zeevaart en Nijverheid 2, no 10: 40-42.

Brief information and references on cultivation of indigo.

1827-1828 - Enumeratio plantarum Javae et insularum adjacentium minus cognitarum vel

novarum ex herbariis Reinwardtii, Hasseltii, Kuhlii, Blumei, etc. Leiden. 2 vols. 278 pp.
Description of some families of Angiosperms and the Pteridophytes. Properly a continua-
tion of the Bijdragen (1825-1827), although in more detail and with longer descriptions.
Repr. Den Haag 1830, Amsterdam 1968.

1 828 - Het Duizend-Gebergte (Goenong SeribuJ.

In: G.H. Nagel, Schetsen uit mijne Javaansche portefeuille; Javaansche tafereelen: 69-75
(in L).
Remarks on the landscape of the 'Thousand Hills', in the plain SW of Jakarta. Also a brief
description of the limestone hills Kuripan, SW of Bogor, famous for their hotsprings, which
yielded several plants not found anywhere else, amongst them a Cycas sp.

1828-1851 - Flora Javae nee non insularum adjacentium. Brussels. 3 vols.

Three sumptuous folio volumes, with analyses, plates, and descriptions in great detail. The
authorship is partly ascribed to his assistant Dr. J.B. Fischer, who was his 'adjutore'. The
preface is probably most interesting, but being not in sufficient command of the Latin
language, cannot evaluate it.
1

For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, (1976) 236. I

23 Planches in^dites were for sale in probably 1863 (see also C: van Steenis, 1947).

1829 Letter to the Governor-Cicncral. Algcmccn Handelsblad of April Isi, no 26.

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 Flora Malesiana

On the occasion of the appointment of Governor-General van den Bosch; on the impor-
tance of stimulating cultures for the general welfare, commerce, and the benefit of the com-
mon people. Blume pleaded for the gradual abandoning cq. restriction of the use of opium.

1831 - Reistogte naar Buitenzorg, het Duizend-Gebergte, Koeripan en in de omstreken van

Batavia, 1824; door een ambtenaar.
Recensent (de Recensenten) XXIV, 2: 427-442, ^dl-AlX.
This contributionis not written by Blume himself, but by one of the civil servants accom-
panying him, A. ZipPELius or A. Latour, on a trip to the hills W
of Bogor. By Blume
himself also described in the entry of 1828, Het Duizend-Gebergte.
Contains no scientific observations.

1831 - Ueber einige Ostindische, und besonders Javanische Melastomataceen.

Flora 15 (2): 465-527.
A thorough study of the family Melastomataceae in which Blume described 12 new genera,
all standing to the present day, mainly based on species described in the Bijdragen
(1825-1827).

1831 - Over eenige Oost-Indische, byzonder Javaansche, Melastomataceae.

H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen
In: 6: 211-268.
The same as the preceding entry.

1831 - Eenige woorden over de redding van Herbarium door Dr. J.B. Fischer.
het Rijks
Alg. Konst- en Letterbode no 23, 10 June: 356-359 & no 24, 17 June: 314-311 (in L).
Detailson the transfer of the Rijksherbarium from Brussels to Holland by Dr. J.B. Fischer.

1831 - Over de Asiatische cholera, uit eigene waarnemingen en echte stukken. C.G. Sulpke,
Amsterdam, viii + 203 pp. (In University Library at U).
Historical account and personal experience with cholera in the Netherlands Indies, exten-
sively documented; measures taken by the government to cope with this disastrous illness.

1 832 - Vruchten mijner ondervinding in het afweren en genezen der cholera. Amsterdam. 3 1 pp.
(in L).

A most interesting paper prescribing how to deal with patients suffering from cholera, in
Java caWedfebris endemica bataviae. Recipes for external and internal use. Prescribing the
boiling of drinking-water. Paper printed in 1000 copies at the author's expense, distributed
freely to boards of municipalities in the Netherlands.

1832 - Beschrijving van Calamus draco Willd., etc.

In: H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen 7: 115-129.
Extensive Latin description of a rattan from S. Sumatra collected by his friend C.F.E.
Praetorius.

1832 - Uittreksel uit eenen brief van den Heer J.B. Spanoghe aan den hoogleeraar C.L. Blume.
Alg. Konst- en Letterbode 1832-1: 356-361.
Notes on the situation in Bima (Sumbawa), with biographical notes on Spanoghe by Blume.
Plant list of Bima.

1834 - Observationes de genere Helicia Lour.

Ann. Sc. Nat. ser. II, 1 (1) Bot.: 211-220.
Review of the genus, with new species.

(26)
 Dedication

1834 - Eenige waarnemingen omtrent de Culilawan boom van Rumphius.

Tijdschr. Natuurlijke Geschiedenis en Physiologie 1: 45-64, t. 2.
Repr. in: Wiegman, Archiv Naturgeschichte 1 (1835) 1 16-126, and in: Jahrb. Pharm. Berlin
35 (1835)9-29.
On Cinnamomum described by Rumphius.

1834 - Eenige opmerkingen over de natuurlijke rangschikking van Rohdea, Tupistra en

mede de beschrijving eener nieuwe soort van dit laatste geslacht.
Aspidistra, als
Tijdschr. Natuurlijke Geschiedenis en Physiologie 1: 67-85, pi. 3 & 4.
Botanical relations between three genera, and description of a new species of Aspidistra.

1834 - De novis quibusdam plantarum familiis expositio et olim jam expositarum enumeratio.
Tijdschr. Natuurlijke Geschiedenis en Physiologie 1: 131-162.
Repr. in Ann. Sc. Nat. sr. II, 2 Bot.: 89-106.
A preprint was issued in 1833, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976)
2367.
Description of a number of newly proposed families, Apostasiaceae, etc., with a few new
species.

1835 - Neesia, genus plantarum javanicum repertum, descriptum et figura illustratum.

Nova Acta Acad. Caes. Leop. -Carol. 17 (1): 73-84, pi. 6.
A new genus of Bombacaceae named after Th.Fr.L. Nees von Esenbeck.

1835-1848 - Rumphia, sive commentationes botanicae imprimis de plantis Indiae orientalis,

tum penitus incognitis turn quae in libris Rheedii, Rumphii, Roxburghii, Wallichii, aliorum,
recensentur. Leiden, Amsterdam. 4 vols. Folio.
For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 238.
Conditions for sale were mentioned in Ann. Sc. Nat. ser. II, 4 (1835) 318.
JuNGHUHN mentions that part of the work was elaborated and illustrated by J. Decaisne
and the Latin was supervised by D.J. Veegens, a friend of Blume.

1837 - Levensbyzonderheden van Franz Junghuhn.

Alg. Konst- en Letterbode 1837-11: 277, footnote.
Biographical notes on F. Junghuhn.

1837 - Levensbyzonderheden van Dr A. Fritze.

Alg. Konst- en Letterbode 1837-11: 277, footnote.
Biographical notes on A. Fritze, Inspector of Physicians and benefactor of Junghuhn.

1837 - Naschrift op den brief van Junghuhn uit Djocjakarta.

Alg. Konst- en Letterbode 1837-11: 278-280.
Appendix to a letter of Junghuhn.

1838 - Revue des palmiers de I'archipel des Indes orientates.

Bull. Sc. Phys. & Natur. en Neerlande no 9: 61-67.

Repr. in Ann. Sc. Nat. ser. 11, 10 Bot.: 369-377.

1838 - Miquetia, genus novum plantarum javanicarum.

Bull. Sc. Phys. & Natur. en Neerlande no 13: 93-95.
Repr. in Ann. Sc. Nat. sdr. II. 10 Bot.: 255-256.

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 Flora Malesiana

Description of a new genus of Icacinaceae, named after Miquel, then director of the Rotter-
dam Botanical Garden.

1838 - (transl.) Advertisement for sustaining the edition of Flora Javae, Rumphia, etc.

Alg. Konst- en Letterbode 1838-11: 322, 401.

1839 - Beschrijving der minerale bronnen, welke nabij Tjiratjas in de Residence Krawang
warden gevonden.
Tijdschr. Ned. -Indie 2 (1): 451-455.
Description of mineral wells near Tjiratjas in Krawang, E of Jakarta.

1843 - Levensbyzonderheden over Th.St. Raffles.

De Indische Bij 1: 49, footnote.
Praise of Raffles' humane government.

1843 - Engeland's staatkunde omtrent China.

De Indische Bij 1: 61-77.
To stimulate the necessity of increasing naval power in Netherlands-Indian waters and ex-
tend commercial relations with Japan. In a footnote on p. 76 Blume refers again to the
necessity of regulating the trade in opium.

1843 - Toelichting aangaande de nasporingen op Borneo van G. Miiller.

De Indische Bij 1: 103-176.
On the geography, anthropology, commercial situation etc. of W. Borneo, from cor-
respondence with G. MiJLLER. In a footnote on p. 104 Blume reveals the bad management
of the Governor-General Daendels, and he praises Raffles for his humane, unselfish ad-
ministration.

1843 - Bladvulling.
De Indische Bij 1: 320.
An occasional note on common social progress, whereby also the native people should pros-
per. Private property of land by non-natives is discouraged. Native rule should not be under-
mined. Adat should be maintained.

1843 - Over een Nederlandsch Gezantschap in Japan.

De Indische Bij 1: 479-480.
Importance of a Netherlands Embassy in Japan.

1843 - Over eenige Oost-Indische planten welke eene uitmuntende vezelstof opleveren, en
Gedachten over het nut van dergelijke kulturen tot opbeuring van de buiten Java gelegene
etablissementen.
De Indische Bij 1: 481-509.
On the importance of fibres, from ramie, cotton and Musa; tissues provided by Blume were
examined.

1844 - (with P.P. VON Siebold) Ontwerp tot oprigting van de Koninklijke Nederlandsche
Maatschappij tot Aanmoediging van den Tuinbouw.
Jaarb. Ned. Mij. Aanmoed. Tuinbouw over 1844: iii-iv.
Tentative rules for the newly erected society.

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 Dedication

1844 - Over het nut der invoering van vreemde gewassen en de laatste pogingen om daardoor
den tuinbouw hier te lande op le beuren.
Jaarb. Ned. Mij. Aanmoed. Tuinbouw over 1844: 41-88.
On the use of importing exotic plants for horticulture in the Netherlands.

1 844 - Naamlijst van Oost-Indische en bepaaldelijk Javaansche gewassen, etc.

Jaarb. Ned. Mij. Aanmoed. Tuinbouw over 1844: 88-90, t. 1-4 (col.).
Unsigned, but attributed to Blume.

1 844 - Ueber das Lycopodium arboreum Jungh.

Amtlicher Bericht uber die Versammlung Deutscher Naturforscher and Aerzte Abt. 2, 22:

85-89.
Identified as Dacrydium cf. elatum Wall, on type material shown to him by W.H. de
Vriese. In Rumphia 3 (1849) 219, 221 Blume later added sour remarks.

1 844 - Ueber ein Surrogat des Chinesischen Thees.

Amtlicher Bericht iiber die Versammlung Deutscher Naturforscher und Aerzte Abt. 2, 22:

90-92.
Made public in a session of the Society at Bremen, 23 Sept. 1844. As Prof. G.J. Mulder
had shown the alkaloid theine is the same as caffeine, Blume suggested that tea could be
made from dried leaves of coffee.

1845 - De Koffij-thee.
Astrea, Tydschr. van Schoone Kunsten, Wetenschap en Letteren 1: 285.
Same as preceding.

1 845 - Minerale wateren van TJipannas en Tjiradjas. Opmerkingen nopens de bruikbaarheid van
dien le TJipannas (Preanger Reg.), beschrijving en scheikundig onderzoek van dien te
Tjiradjas (Krawang).
Indisch Magazijn, Tweede Twaalftal, no 1/2: 162-166.
Reprint of an article published in the Bataviaasche Courant of 15 Sept. 1821.

1845 - Gedachten op eene reis door het zuidoostelijk gedeelte der Residentie Bantam.
Indisch Magazijn, Tweede Twaalftal, no 3/4: 1-36.
Account of his experience on a trip through SE. Bantam in W. Java. Account of the Badui
people. Reprint of an article published in 1822.

1845 - Fragment uit een Dagboek gehouden op eene reis over Java. Bestijging van den berg
TJerimai, gewoonlijk genoemd Tjerme, in de Residentie Cheribon.
Indisch Magazijn, Tweede Twaalftal, no 3/4: 102-116.
Report on an exploration of Ml Tjeremai, above Cheribon. Reprint of an article published
in 1825.

1845 - Over iniandsche middelen tegen diarrhoe.

Indisch Magazijn, Tweede Twaalftal, no 3/4: 1 16.

Indigenous recipes against diarrhoea. Copied from the paper published in 1825.

1845 - De patma van Noesa Kambangan.

Indisch Magazijn, Tweede Twaalftal. no 3/4: 179-194 (in L)-

Reprint of an article published in the Bataviaasche Courant of 9 & 23 March 1825, in which
he described his finding of Rafflesia Nusa Kambangan
in I. (S. Central Java) and claimed
this to be the largest flower, superseding Nelumbntm.
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 Flora Malesiana

1846 - An article in the 'Handelsblad'.

In this article Blume advised to hold expositions of colonial products from the East and
West Indies in the Netherlands from time to time.

1849-1856- Museum botanicum Lugduno-Batavum sive stirpium exoticarum, novarum vel

minus cognitarum ex vivis out siccis brevis expositio et descriptio. Leiden. 2 vols.
Appeared in dated parts each of 16 pp. In all, 1: 396 pp., 60 fig.; II: 256 pp., 58 fig. The
second volume was not finished and had no index.
For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, (1976) 240. 1

A most important work, being a scientific catalogue of the Rijksherbarium collections,

hence containing descriptions and treatments of plants from all over the world.
An Index to volume 2 was prepared by C. G.G.J, van Steenis & Chew Wee Lek at the
Rijksherbarium in 1974 (see Appendix C).

1850 - Antwoord aan den Heer W.H. De Vriese.

Alg. Konst- en Letterbode 1850-11: 99-109, 114-123. Repr. 34 pp. in L.
Blume defends his criticism on the identity of Lycopodium arboreum and the reduction of
Pinusmerkusii, and the right of the Rijksherbarium to be the depository of collections made
by civil servants.

1850 - Opheldering van de inlichtingen van den Heer Fr. Junghuhn.

Alg. Konst- en Letterbode 1850-11: 258-261, 274-279. Repr. 19 pp. in L.
On Junghuhn's collection and the right of the Rijksherbarium as the proper public deposi-
tory of botanical collections.

1852 - Copy of a letter to J.G. Baud, Minister of the Colonies, dated 14 March 1840, 'nopens
de bereiding van thee uit koffie-bladeren, met aanbeveling tot het nemen van proeven in het
groot op Java zelf.
Natuurk. Tijdschr. Ned. Indie 3: 122-126.
Proposal to prepare tea from coffee leaves and suggesting experiments with this on a large
scale in Java.
There are two other entries on the subject in 1844 & 1845; see also Astrea 1 (1851) 256.

1855 - Melanges botaniques. 8°. No 1, 1 Aug. 1855: 1-8; no 2, 1 Sept. 1855: 9-12. Facsimile
inTaxon 35 (1986) 274-285.
Until June 1985 assumed not to have been published; see Stafleu & Cowan, Taxonomic
literature, ed. 2, 1 (1976) 241.
The new names etc. in the Melanges were validated by Walpers in his Annales 4 (1857)
6A1-6AA and a rather large extract was published in Flora 41 (1858) 254-256.
L. Vogelenzang, librarian of the Rijksherbarium, found in Vesque's bibliography of
J. Decaisne (C: 1883) that the latter had a copy of the Melanges in his library, now incor-

porated in the Bibliotheque Nationale at Paris. H. Heine located another copy in the
Bibliotheque Central of the Museum d'Histoire Naturelle at Paris which had belonged to
the library of A.Th. Brongniart. The original copy mentioned in Flora is still not located.
It was probably dedicated to Nees von Esenbeck.

The pamphlet was not for sale, but it was effectively published and at least two copies exist.
Both Paris copies were autographed to Blume's close friends. He may have sent more copies
to other botanists with whom he was befriended. Obviously Blume published it at his own
expense and the reason for this is unknown. He could have published it in his Museum
Botanicum Lugdunum-Batavum
The first numero of the Melanges contains a discussion on paper-making by the Sino-

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 Dedication

Japanese and three species are described of Broussonetia (2 new). Furthermore there is a sec-
tion 'synonymic de quelques plantes peu connues', concerning speciesand genera of Cuno-
niaceae, Saxifragaceae, Rosaceae, Guttiferae (Cratoxylon), Dipterocarpaceae, Ulmaceae,
Moraceae, and Nepenthes. Numero 2 contains Chrysobalanaceae and Rosaceae (Pygeum)
(B: VAN Steenis, 1986).

1858 - Bijdrage tot de kennis der Oost-Indische Orchideen en het maaksel (de organisatie) van
hare bevruchiingswerktuigen.
Versl. & Meded. Kon. Ned. Akad. Wetensch., Amsterdam 7: 100-115, 2 pi.

Interpretation of the orchidaceous flower, with special regard to Apostasiaceae.

1858(-1859) - Flora Javae et insularum adjacentium. Nova Series. Leiden, pp. 8 + 6+ 162, 66
col. pi.
Also edited with a French title, see below.
A sumptuous work in which Blume summarized his large knowledge on orchids in which
he had great insight since he wrote the Bijdragen (1825-1827).

1 858(- 1 859) - Collection des Orchidees les plus remarquables de I'Archipel Indien et du Japon.
The French-titled version of the Flora Javae, Nova Series.
For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 240.

1859 - (with A.H. van der Boon Mesch) Geschikte materialen uit de Overzeesche bezittingen
voor het vervaardigen van papier.
Report about useful materials from overseas territories suitable to manufacture paper.

1859 - Vanda suaveolens Bl.

Ann. Hort. Bot. ou Fl. Jard. Pays-Bas 2: 1-2, 1 col. pi.

1859 - Over eenige Oost-Indische houtsoorten in verband met de verwoestingen door den paal-
worm of andere schelpdieren hier te lande en elders aangerigt.
Versl. & Meded. Kon. Ned. Akad. Wetensch. 9: 25-49. Repr. 25 pp. in L.

A scholarly review of timbers resistant against teredo and other molluscs, in which Blume
summarized experience onwards of Rumphius and collected data from all kinds of sources,
indicating valuable species to be used in sea harbours.

1860 - De houtteelt verbonden met den landbouw.

Tijdschr. Ned. Mij. ter Bevordering van de Nijverheid 23: 1-29.
Cultivation of timber species in relation to agriculture.

1861 - Monographie des Anoectochilus, Goodyera et genres voisins, les plus remarquables de
du Japon.
I'archipel Indien et
Belg. Hon. 11: 369-378, 1 pi.

Extract from Flora Javae, Nova Series (1858-1859).

1863 - Flora Javae. Planches in^dites.

23 coloured folio plates of Javanese plants with names and analyses. These were probably
intended for further instalments of the Flora Javae, but remained without text.

On the back of some plates an advertisement was printed by a booksellers firm in Leiden;
herein Blumf.'s works were offered for sale, as a packet, probably one or (wo years after
his death.
I have distributed a few copies to some herbaria, with a nolo, in November 1947 (C: 1947).
Further particulars I published in Blumea 6 (1948) 263.
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 Flora Malesiana

Excluded

1 823 (April) - Herinnering aan acht merkwaardige dagen van mijn /even, op een uitstapje naar de
top van de Gounong (berg) Gedu.
This concerns a 16 pp. manuscript which has wrongly been attributed to Blume. It was writ-
ten by a party following Blume's trail to the lower part of the crater of Mt Gedeh above
Tjibodas. It is preserved in the library of the Instituut van Taal-, Land- en Volkenkunde,
Leiden University (H 338).

Appendix B — Biographical sources

Aa, A.J. VAN DER. 1878. Biographisch woordenboek der Nederlanden. Bijvoegsel: 34-35; ibid.:
111-115. - A concise biography.
Anonymous. 1827-1856. Algemeene Konst- en Letterbode 1827-11: 137; 1829-1: 227; 1831-1: 50,
359; I833-I: 429; 1838-11: 290; 1851-1: 257; 1853-1: 193, 305; 1855: 118; 1856: 57.
Anonymous. 1: 228. - Blume was in Berlin and offered (obviously at a meet-
1853. Bonplandia
Boehmeria tenacissima Bl. which he said had a great durability and could possibly
ing) fibres of
be of importance for the navy. He was then presented to the King of Prussia. On the fibres of
this Boehmeria he pubUshed in the Melanges botaniques (A, 1855).
Anonymous. 1855. Bonplandia 3: 155. - Here it was reported that Reinwardt sold his library
for Dfl. 20,000. His herbarium was donated to the University herbarium of Leiden, on the con-
it should not be incorporated in the Rijksherbarium.
dition that
N.B. In the 'Instruction' of 1832 (see C: van Dam) it had been officially decreed that the
University herbarium was to be merged with the Rijksherbarium!
Anonymous. 1858. Flora 41: 254-256. - Extract review oi Melanges botaniques.
Anonymous. 1862. Leidsch Dagblad, 5 Febr. 1862, no 598. Repr. of 3 pp. in L. - Formal
obituary.
Anonymous. 1862. Bonplandia 10: 47. - Obituary note.
Anonymous. 1862. Botanische Zeitung 20: 56. - Obituary note.
Anonymous. 1862. Proceedings Linnean Society of London 1862: xcvi-xcviii. - Obituary note.
Anonymous. 1862 or 1863. Annuaire de 1' Academic de Paris. - Obituary note (not seen).
Anonymous. 1875. Allgemeine Deutsche Biographic 2: 746-747. - Short biography.
Anonymous. 1875. Album Studiosorum Lugdunum Batavum 1575-1875, column 1243. - Short
biography.
Anonymous. 1930. Nieuw Nederlandsch Biographisch Woordenboek 8: 132-133. - Short
biography.
Backer, C.A. 1936. Verklarend woordenboek van wetenschappelijke plantennamen: 70. - Brief
biography.
Baillon, H.E. 1877. Dictionnaire de Botanique 1: 433.
Beumee, J.G.B. 1948. C.L. Blume, Museum Botanicum. Fl. Males. Bull, no 3: 69-70. - On the
dates of publication.
Boerlage, J.G. 1896. Botanische literatuur. Encyclopaedic van Nederlandsch-Indie ed. 1, 1: 210,
272-273, 280.
Bretschneider, E. 1898. History of European botanical discoveries in China: 308-309. London.
- Brief biography; Blume illustrated some Chinese plants.
BuRDET, H.M. 1972. Cartulae ad botanicorum graphicem. Candollea 27: 327-328.
Candolle, a. DE.1862. Memoires et souvenirs de A. P. de Candolle: 150, 383, 412.
1880. Phytographie: 318. - Praises the excellent figures in Blume's Museum Botanicum.

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 Dedication

CoLENBRANDER, H.T. 1926. Kolonialc Geschiedenis 3: 111.

Danser. B.H. 1938. Who can give further information about the dates of publication of Blume's
Flora Javae? Chron. Bot. 454-455.
4:

1939. The publication dates of Blume's Flora Javae. Blumea 3: 203-211.

GoDDiJN, W.A. 1931. 's-Rijks Herbarium 1830-1930. Meded. Rijksherb. 62b: 1-53. - Rather
extensive biographical notes.
Museum van Natuurlijke Historie, 1820-1915: 100-101. Rotterdam.
GiJZEN, A. 1938. 's-Rijks
- On Blume's zoological contributions to the Leiden Museum.
Hall, H.C. van. 1862. C.L. Blume. De Nederl. Spectator, 22 Febr. 1862, no 8: 57-59. -
Biographical data; rather extensive (in L).

Hasskarl, J.K. 1850. Antwoord aan den heer C.L. Blume, wegens onderscheidene te mijnen aan-
zien geuite beschuldigingen, vervat in zijn antwoord aan den heer W.H. de Vriese, Leiden 1850.
Alg. Konst- en Letterbode 1850. Repr. of 16 pp. in L. - Hasskarl defending his rights to have
a private herbarium.
Jacobs, M. 1980. C.L. Blume (1796-1862). Fl. Males. Bull, no 33: 3362-3363.
Jansen, p. & W.H. Wachter. 1941. Ned. Kruidk. Arch. 51: 343. - Biographical references.
JuNGHUHN, aan C.L. Blume vanuit Djocjakarta, 2 Febr. 1837. Alg. Konst- en Let-
F. 1837. Brief
terbode 1837-11: 275-277.
1850. Inlichtingen aangeboden aan het publiek over zeker geschrift van den heer C.L.
Blume, en antwoord aan dien Heer. Alg. Konst- en Letterbode 1850, no 41. Repr. 9 pp. in L.
- Self-defense in keeping his private herbarium.
1850. Vervolg der inlichtingen aangeboden aan het publiek over een geschrift van den heer
C.L. Blume. Alg. Konst- en Letterbode 1850. Repr. 29 pp. in L. - Polemics with Blume.
1851. Een woord over den Sambinoer-boom van Sumatra, betrekkelijk deszelfs botanische
bepaling. Ned. Kruidk. Arch. 2: 2-16. - On Blume's reduction of Junghuhn's Lycopodium
arboreum to Dacrydium.
inwendigebouw, 1: 183-186. 2nd Dutch ed.
1853. Java, zijnegedaante, zijn plantentooi, en
Kalkman, C. 1979. The Rijksherbarium, Blumea 25: 13-26, especially p. 14.
past and present.
KosTER, J.Th. Facsimile handwritings of Blume. - Unpublished (in L).
Lasegue, A. 1845. Musee botaniquede M. Benjamin Delessert: 268,293,307,315,346,347,506,
535, 562.
Leenhouts, P.W. 1980. Het Botanisch Kabinet te Franeker: 34.
Lintum, C. te. 1913. Een eeuw van vooruitgang, 1813-1913. Zwolle (not seen). - Blume was
far ahead of his time in having found the solution of the combat against cholera by the simple
boiling of drinking-water.
Maclean, J. 1979. Carl Ludwig Blume and the Netherlands East Indies. Janus 66: 15-29.
-
Period 1820-1831; valuable biographical essay. Maclean traced many letters in the Colonial
Archives of the 'Rijksarchief, The Hague.
Miouel, F.A.W. 1856. Review of Blume, Museum Botanicum. Bot. Zeit. 14: 185-188, 540-541.
- MiQUEL complained severely about Blume's antedating issues of the Museum Botanicum and
his attempts to withhold information from his colleagues.
Guden, a. den. 1979. C.L. Blume, pcriodc 1826-1832. Unpublished essay, made under supervi-
sion of Dr. P. Smit, Biohistorical Institute, Utrecht. - A thorough account, largely based on
official letters and documents of the period mentioned, as present in the 'Rijksarchief, The
Hague.
Pritzel. G.A. 1872. Thesaurus litcraturac boianicae: 29. - Blumi's selected bibliography.
Pi;lle, a. a. 1917. Botanische literaluur. lincyciopacdic van Nedcrlandsch Indie cd. 2, vol. I:

317,394-395; ibid. 1919. Vol. 4: 422.

Raalten, G. van. 1825. Unpublished letter to J.G.S. van Breda (?). - Erroneous accusation that
Blume stole property or information from Kuml & Van Hasshlt (in L).
koiANi>. M. 1944. Alfred Moquin-Tandon. Un naturalistc ^ Paris sous Louis-Philippe. Journal

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 Flora Malesiana

d'un voyage inedit (1834). Paris, Mercure de France ed. 3: 351 pp. — Historically a most in-
teresting booklet full of biographical data of French botanists. Blume paid a prolonged stay
to Paris in Sept. /Oct. 1834.
RoMER, L.S.A.M. VON. 1921. Historische Schetsen. Batavia. 335 pp., 109 pi.; a very brief
obituary on p. 193. - It is most peculiar that in the brief history of cholera (pp. 232-238) the
author, himself a physician, makes no mention at all of Blume's important work on the sub-
ject.
ScHOUTE, D. 1937. Occidental therapeutics in the Netherlands East Indies during three centuries
of Netherlands settlement. Publication of the Netherlands Indies Health Service: 114-119. -
Cited the governmental regulations and instructions for the native chiefs, extension of the vac-
cination, etc. Some of these might have actually been written by Blume, who was chief of vac-
cination and later even chief of the medical service.
Seemann, B. 1863. Journ. Bot. 1: 64. - Short obituary.
Seemann, W.E.G. 1859. Bonplandia 7: 52-53. - Blume complained that the Netherlands
Government did not contribute funds towards the publication of the Flora Javae, Nova Series,
and that this was printed at his own expense. Seemann had received the volume, or at least first
sheets of it, on 3 Nov. 1858. He criticizes Blume for having given too little attention to the works
of LiNDLEY and Reichenbach.
SiRKS, M.J. 1915. Indisch Natuuronderzoek: 109-112, portr. Amsterdam. - Brief biographical
notes.
Smit, p. 1979. The Rijksherbarium and the scientific and social conditions which influenced its

foundation. Blumea 25: 5-11. - In this excellent essay on the foundation of the Rijksher-
barium Smit erroneously mentioned (p. 9) that Blume transferred the Kuhl & Van Hasselt
specimens to Leiden in 1826.
& R.J.Ch.V. ter Laage (eds.). 1970. Essays in biohistory. Regnum Vegetabile 71.
Stafleu, F.A. 1966. Wentia 16: 28-31. - In an excellent biography of Miquel some notes on
Blume.
1970. The Miquel-Schlechtendal correspondence. A picture of European botany, 1836-
1866. In: P. Smit &
R.J.Ch.V. ter Laage, Essays in biohistory. Regnum Vegetabile 71:
295-341. - Many data on Blume and his works. Page 307: Decaisne made several drawings
for Rumphia. Page 324: Junghuhn sold his herbarium to the University of Leiden on the condi-
tion that it should not be incorporated in the Rijksherbarium. Page 326: Reference to Miquel,
who was glad that in February 1851 a new, more 'liberal' Instruction for the Rijksherbarium
was issued by the Government. Page 33 1 Reference to Miquel's complaint about the irregulari-
:

ties with the dates of Museum Botanicum. Page 334: Reference to the difficulty in choice of

a successor of Blume.
1978. Flora Malesiana I, 8: (7)-(16). - Dedication to the memory of F.A.W. Miquel, con-
taining some notes on Blume.
& R.S. Cowan. 1976. Taxonomic literature. Ed. 2, vol. 1 234-241 (Regnum Vegetabile 94).
:

Steenis, C.G.G.J. van. 1941. Natuurwet. Tijdschr. Ned. Ind. 101: 216. - The Planches inedites
appeared at least before 1883.
1948. On the date of publication of Blume's Planches inedites. Blumea 6: 263.
1979. The Rijksherbarium and its contribution to the knowledge of the tropical Asiatic flora.
Blumea 57-77, especially pp. 60-62. - Blume's endeavours.
25:
The publication of Blume's Tabellen en Platen voor de Javaansche Orchideeen.
1980.
Miscellaneous Papers Landbouwhogeschool, Wageningen 19: 289-291.
1986. Blume's Melanges botaniques effectively published, 1855. Taxon 35: 272-273; fac-
simile of the Melanges: 274-285.
&
M.J. VAN Steenis-Kruseman. 1970. The plates of Javanese plants of Francisco Norona,
with a revised evaluation of his generic names. In: P. Smit & R.J.Ch.V. ter Laage: Essays in
biohistory. Regnum Vegetabile 71: 353. - Blume has seen Norona's plates in Java, as well as

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 Dedication

Reinwardt. Incidentally Blume mentioned a few Norona names in the synonymy of his
works.
Steenis-Kruseman, M.J. van. 1950. Carl Ludwig Blume. Flora Malesiana I, 1: 64-66, 600,
portr. - Brief personalia; account of Blume's travels and publications.
1979. Directorate of C.L. Blume. Blumea 25: 35-39.
Treub, M. 1889. Geschiedenis van 's-Lands Plantentuin te Buitenzorg. Meded. 's Lands Planten-
tuin 6: 1-79. Batavia. - History of the Botanic Gardens, Bogor, from 1817 till 1844.
1892. Korte geschiedenis van 's-Lands Plantentuin te Buitenzorg: 7-9, portr. - Short
history, as above.
Ule, Witty. Geschichte der Kaiserlichen Leopoldinisch-Carolinischen Akademie der Natur-
forscher 1852-1882. No. 1071 (not seen).
Veth, P.J. 1884. Ontdekkers en onderzoekers: 45-149. Leiden. - Mostly on Reinwardt; por-
trait of Blu.me.

Vos, C. DE. 1888. Korte schets van de geschiedenis der plantkunde etc.: 91-92. Bolsward.
Vriese, W.H. de. 1851. Naschrift (to Junghuhn's paper). Ned. Kruidk. Arch. 2: 13-17 (in L).
- Defending Junghuhn.
1851. Teregtwijzing van C.L. Blume's naamsverwarring. Alg. Konst-en Letterbode 1850-11:
35-38. Repr. of 4 pp. in L. - On the reduction of Pinus merkusii to P. finlaysoniana.
\\ EiGEL, T.O. 1863 (Jan.). Verzeichniss der nachgelassenen Bibliothek von C.L. Blume. Leipzig.

III-VI + 81 pp. - With portrait of Blume.

Winkler-Prins, C. 1949. Encyclopaedic ed. 6, 4: 374.
Wit, H.C.D. de. 1949. 47. Blume. Flora Malesiana I, 4: civ-cv. - A brief account of Blume's
life; discussion of achievements and main publications.
- Typed copies of letters to and
1950. History of Malesian botany. 29 pp., unpublished.
from Blume, partly relating to herbarium Reinwardt, but largely official letters on the sale
and distribution of Flora Javae and Ruinphia. Several derived from the 'Rijksarchief, The
Hague.
Zollinger, H. 1841. Tagebuch(ined.), 5 Oct.-31 Dec. - Unpublished diary of Zollinger; typed
copy by H. Wanner in L.

Appendix C - References to cited literature

Arckenhausen, J.C.P. See his biography by H.-G. Griep et al., vide infra.
Breda, J.G.S. van. 1827-1829. Genera et species Orchidearum et Asclepiadarum quas in
itinerere per insulam Java collegerunt Dr. H. Kuhl et Dr. J.C. van Hasselt. Ghent. Folio. 15
fol. & 15 tab. col.
Da.m, van. 13 Febr. 1832. Ontwerp van eene instructie voor den Directeur van het Rijksherbarium
(Ministry of the Interior, 5th Div., No. 254 - Concept of an Instruction to the Director of the
Rijksherbarium).
Directions for the director in 14 articles: how to manage the collections, the accommodation,
and availability to other botanists, loans, the making of a catalogue of the col-
the facilities for
lections, exchange of duplicates, desirability of acquiring collections from civil servants, the fu-
sion of the University Herbarium with that of the Rijksherbarium, the order that the director
writes an annual report on the important accessions, and that proposals of the director had to
go via the Curators of the University.
A particularly ticklish point was stipulated in art. 10, in which the director was prohibited to
publish on discoveries of still living persons and explorers without their consent.
The Instruction was approved by the Minister of the Interior and was stipulated to be effective
from January 1st, 1831.

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 Flora Malesiana

Griep, H.-G., H. Ullrich &G. Wagenitz. 1977. Johann Christian Arckenhausen (1784-1855).
In H. Ulrich (ed.), Goslarer Kiinstler und Kunsthandwerker 1: 1-32, illust. (D, 12).
Hall, H.C. van. 1856. Voorstel omtrent de voortzetting van de uitgave der Flora Javae. In W.H.
DE Vriese: Tuinbouwflora 3: 365-366.
Hennipman, E. 1979. The collections of Pteridophytes at the Rijksherbarium. Blumea 25:
103-106.
Reinwardt, C.G.C. 1826. Nova plantarum indicarum genera. Syll. Plant. Ratisb. 2: 1-15.
1 Ueber den Charakter der Vegetation auf den Inseln des Indischen Archipels. Ein Vor-
828.
trag. Kon. Akad. Wiss. Berlin: 1-18.
Steenis, C. G.G.J. VAN. 1947. Introduction to the Planches inedites Flora Javae (mimeographed).
- Pamphlet, consisting of a coloured folio plate of Blume's Planches inedites with at the back
an advertisement for the sale of Blume's works, probably from 862 or 1 863 Copies were sent 1 .

to some selected European libraries.

& Chew Wee Lek. 1974. Index to C.L. Blume, Museum Botanicum Lugduno-Batavum,
vol. 2, 1856-1857. Leiden. 24 pp.
M.J. VAN Steenis-Kruseman & C.A. Backer. 1954. Louis Auguste Deschamps. Bull. Brit.
,

Mus. Nat. Hist., Hist. ser. 1, no 2: 51-68, pi. 13 (a reproduction of the drawing Deschamps
made of Rafflesia).
Steenis-Kruseman, M.J. van. 1950. KoUmann's collection of Javan plants. Bull. Jard. Bot. Btzg
ser. Ill, 18: 463-466.
1962. Contributions to the history of botany and exploration in Malaysia. 8. Heinrich Burger
(71806-1858), explorer in Japan and Sumatra. 9. The transfer of the Rijksherbarium from
Brussels to Holland in 1830. Blumea 11: 495-505; 505-508, photo. 1

1979. The collections of the Rijksherbarium. Blumea 25: 29-56.

Thorbecke, J.R. 11 Nov. 1850. Instructie voor den Directeur van het Rijks-Herbarium te Leijden
(Ministry of the Interior, 5th Div., No. 254 - Instruction fortheDirector of the Rijksherbarium
at Leijden). 22 pp. (in L).

Instruction to replace that of 1832 (see under C: van Dam), consisting of 28 articles. New
stipulations were: the director should be present on the first three days of the week; not more
than one family of plants can be borrowed by a single person; the director is prohibited to use
data from the still living members of the former 'Natuurkundige Commissie' without their per-
mission; he is not allowed to have a private collection; as to exchange, priority has to be given
to Dutch botanists and onwards of December 1st, 1850.
institutes, effective
A most was in art. 18: anybody could claim to receive duplicates from the
peculiar stipulation
overseas territories (the names of which had already been printed and the plants described) even
when nothing was offered in exchange. So it has happened recently that, in cleaning a school
somewhere in Holland, a set of Javanese sheets was found, obviously claimed by a former en-
thusiastic teacher who had, it seems, no employ for it.
Vesque, J. 1883. Catalogue de la Bibliotheque de feu M. J. Decaisne. Avec une notice biogra-
phique par M. le Dr. Ed. Bornet. Paris. Libraire de la Bibliotheque Nationale: 13. - Listing
under no 56: 'Blume, Melanges botaniques (Premier et deuxieme numero). Leyde, 1855, br.
in-8, de 12 pp. -Envoi autogr. de I'auteur a M. Decaisne.
Vriese, W.H. de. 1858. Reinwardt's Reize naar het Oostelijk gedeelte van den Indischen Archipel
in het jaar 1821 etc. Amsterdam.

Appendix D - Notes

1)- Later it was said that Blume misused the collections and manuscripts of A. Zippelius, a
gardener of the Botanic Gardens at Buitenzorg (Bogor), who made a long exploration trip to the
Moluccas, SW. New Guinea, and Timor, where he died.

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 Dedication

Surely Zippelius made a most important collection, but he left no manuscripts at L; we only
have a box full of old provisional labels. As a matter of fact, P. Bleeker found in the archives
of the 'Natuurkundige Vereeniging' at Batavia manuscripts and notes of Zippelius that were of-
fered to Blume about 1850, under the condition that the latter should publish them. Blume never
replied to this. In fact this request came two decades too late, as Blume had worked on Zippelius's
material (received through the intermediary of J.B. Spanoghe in ± 1830/31) and published this
earlier in Rumphia and in the Museum Botanicum. Blume honoured Zippelius by naming the
Piperaceous genus Zippelia after him. (See also the footnote under D: 4.)
Blume has also been accused of having left at Bogor no duplicate specimens of the collections
he took 10 the Netherlands, but this is untrue (see C: van Steenis-Kruseman, 1950; and D: 4).
As to the Kuhl & van Hasselt collections: they did not add many novelties to what Blume
himself had collected. The sites where he travelled covered most of theirs, and even far beyond
eastwards. Besides, the Kuhl & van Hasselt collections came only in Blume's hands in 1828
when he had already published his Bijdragen (1825-1827) and Enumeratio (1827-1828). And as
late as 1844 van Breda offered him a packet of notes written by Kuhl and van Hasselt, when
the main part of Flora Javae (1828-1851) had already been published.

2) - Ph.F. von Siebold, a most meritorious scientist, withheld his collections from Blume. Most
of VON Siebold's botanical collections were not made by himself, but by Burger, Textor, Keiske
and others (see C: van Steenis-Kruseman, 1962). Von Siebold also was a dominating, ambitious
person. The Flora Japonica was authored by 'Siebold & Zuccarini', but the latter, professor at
Munich, was the proper author responsible for the research. Von Siebold hardly had any claim
towards being a botanical taxonomist. As Burger belonged to the 'Natuurkundige Commissie',
their herbarium should properly go to the Rijksherbarium. Though Burger's share in the under-
taking was very large - he also wrote a large manuscript on Japanese fishes - von Siebold later
refused to support Burger's second appointment to the 'Natuurkundige Commissie' for the ex-
ploration of W. Sumatra, because the latter would not be sufficiently endorsed with scientific
knowledge (/.c. 501), a most ungracious and unjust gesture.
VoN Siebold claimed later to have been the saviour of the Rijksherbarium in 1830, whereas his
sole purpose was to get back specimens collected during his internment by the Japanese in
Deshima (I.e. 501). Whatever the great merits of von Siebold may have been, these facts throw
a distinct shadow on his honesty and tolerance regarding other people.

3) - F. W. Junghuhn was a physician of the army since 1835, but his superior, A.E. Fritze, per-
840 he was charged with making investiga-
mitted him to devote himself to the study of nature. In 1

tions in the Batak Lands, W. Sumatra. After his return to Java Junghuhn was appointed a
member of the 'Natuurkundige Commissie' (1845-1848). Through h\s Reisers dureh 7Gvoand Die
Battaldnder auf Sumatra it became clear that Junghuhn had amassed a great herbarium, and
Blume claimed this for the Rijksherbarium. Junghuhn refused, which caused Blume's irritation.
As Junghuhn was no taxonomist and had made errors in precursory papers (amongst others with
Lycopodium arboreum), Blume's sharp remarks on this led to a strong mutual animosity between
him and Junghuhn.

4) - According to my wife (C: van Steenis-Kruseman, 1950), G.H.J. Kollmann was a German
senior physician, in the service of the Dutch East Indian army and stationed at Buitenzorg (Bogor)
in 1821 - 1835, on leave in Europe in 1835-1837. In 1837 he offered the Dutch government a col-
lection of Javanese plants for sale. ' His letter and material were designated to Blume, who, to

(I) About the contents of the collection which came in Koilmann's hands more can be found in J. Macii-an,
Scieniiarum Historia 15 (2), 1973. 12
1 I I.V They comprised zoological collections as well as ethnographical

ones besides the herbarium specimens. According to Kollmann they were acquired at auctions (presumably
in Java) and contained not only Blumi collections but also Zippflius plants (M.J. van Steenis-Kruseman).

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 Flora Malesiana

his surprise, found that this was the set of duplicates (more than 4000) of his collection he
painstakingly left at Buitenzorg when returning to Holland. Kollmann himself never collected.
Obviously the collection had been stored somewhere in the annexes of the Palace at Buitenzorg,
adjoining the Botanic Gardens. The curator of the Gardens, James Hooper, was subordinated
to the Intendant of the Palace. In some way or other Kollmann appropriated this collection. The
rumour that Blume did not leave duplicates at Buitenzorg appears fully untrue.
he never Why
alluded in print to the curious way in which the Bogor duplicate collection came into his hands,
can only be guessed at (D: 14). He was either loyal to Kollmann, with whom he had friendly rela-
tions, or he found it unnecessary to justify himself. Anyway it shows his loyalty to the Buitenzorg
Gardens.

5) - Both J. Maclean and A. den Ouden (B: 1979) have searched in the 'Rijksarchief, The
Hague, where all official correspondence by Blume is kept. For a proper biography the period
1830-1862 should also be covered. Moreover, personal letters will be kept in the archives of
several botanical institutes as Blume had contacts with many botanists.

6) - It is quite possible that, as soon as Blume had finished the text for a fascicle of Museum
Botanicum, he sent it to the printers and assumed it then to be effectively published. In his splen-
did isolation, surrounded by envious, hostile colleagues and antagonists, Blume did not care
about their interests. Leiden was at that time a centre where nobody did care about collaboration
or sympathy, each staff member promoting self-interests; a most unfortunate situation.

7) - The number of extensive biographies of prominent Dutch botanists is small. I know off-hand
only those of C.G.C. Reinwardt, Hugo de Vries, W. Beijerinck, F. Junghuhn, J. P. Lotsy,
F.A.W. MiQUEL, and H.J. Lam. Such biographical studies require much time, and also historical-
minded people to compose them. If one should like to have a posthumous biography made, it is
best, in my opinion, to write an autobiography; one ought to think timely of this.

8) - The diary of H. Zollinger contains notes on his stay at Leiden in 1841, with interesting per-
sonal information on members of the biological circle at Leiden. Amongst others about the com-
plaints ofReinwardt that Blume did not give him sufficient honour and published all novelties
under his own name. But C.A.L.M. Schwaner, a German geologist and member of the
'Natuurkundige Commissie', said that this was due to the fact that Reinwardt did not publish
himself, even not his own report on the exploration in East Malesia, and that Reinwardt's
reasons for not publishing was that he was afraid not to come up to the expectations the botanical
public had of him. As a matter of fact, the lecture Reinwardt held for this select public, the 'Ver-
sammlung Deutsche Naturforscher und Aerzte' on 20 September 1828 about the vegetation of
Malesia, was not exciting, but mediocre (C: Reinwardt, 1828). The same holds for his paper
Nova plantarum indicarum genera; many genera were assigned to wrong families and several
others had been described before. Reinwardt's creative efforts lay mainly in the organization
of botany and cultures in Java, not in research. His report on the exploration of the Moluccas
was after his death pubhshed in 1858 by W.H. de Vriese (C: 1858), together with a biography.
Another fact Zollinger mentioned was that it was not due to Blume that P. Korthals aban-
doned botany. Korthals told Zollinger at the time the first was working out his most important,
meticulous observations, that botany was an inferior branch of science as compared with
philosophical and etymological studies, which he found more interesting and scholarly.

9)- According to Weigel's catalogue (B: Weigel, 863), Blume had a very large library, the total
1

number of entries being 2123, largely concerning botany (1527 entries). It is peculiar that Blume's
works are only represented by 9 items. None of his publications on useful and medicinal plants
were represented.

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 Dedication

10) - As
a matter of fact, the majority of biologists, physicians, and explorers in the early part
of century concerned with the biology of the Indies were scientists with the German nationali-
last

ty or of German descent, e.g., Arckenhausen, Blume, Burger, J.B. Fischer, Hasskarl,

JUNGHUHN, KUHL, MaCKLOT, SaL. MuLLER, ReINWARDT, VON ROSENBERG, SCHLEGEL,
ScHWANER, VON SiEBOLD, ZiPPELius. Also in South American and African botanical pioneer ex-
ploration Germans played a prominent role in the former century.

11) - As to his health, Blume withstood illnesses obviously rather well, probably because he ap-
plied his own devices, drinking boiled water, etc. He was reported to suffer of fever during his
trip to Rembang (see A: 1828). In 1826 (Java) Blume complained of illness. In Holland he was
rather seriously ill about 1829. Early 1850 he suffered of laryngitis.

12) - H.-C. Griep c.s. Arckenhausen reproduced a letter

(C: 1977) in their biography of J.C.P.
(in the 'Rijksarchief, The Hague) from Blume The Hague (d.d.
to the Minister of the Interior at
27 Dec. 832), in which he pleads for the second time for a permanent position of Arckenhausen.
1

Blume mentioned that he had 500 drawings, mostly from Latour, made in Java. These drawings
1

were sketches which should be made ready for reproduction in Flora Javae and often needed to
be supplemented by details (from herbarium material). Arckenhausen could manage to prepare
7 or 8 drawings monthly. As the publication of Flora Javae at Brussels needed monthly 12 draw-
ings for the two instalments, Blume had attracted a certain Mr. Vivien as draughtsman (in 1827)
and .Mr. Sixtus (in 1828) for keeping pace. Vivien disappeared in 1829 and he was replaced by
Arckenhausen. The Minister was of the opinion that Arckenhausen should be paid from the
Flora Javae project funds. The latter worked for Blume at least until 1832, possibly longer. After

repatriation to Germany Arckenhausen remained draughtsman in Goslar, drawing all kinds of

plants and animals, mostly for Krebs, Naturgeschichte. After Arckenhausen's death (1855) his
estate was sold in 1862, among which 134 plates of Flora Javae. In the library of the Natur-
wissenschaftliche Verein, in volume 19 (portfolio), 190 plates of Blume's work are preserved, of
which some unfinished sketches. Whether they are originals or printed copies, and whether there
are unpublished drawings among them, has still to be examined. Plates by Arckenhausen are
reproduced too in Rumphia, volumes 1-3.
In Java J.Th. Bik was another artist, originally in the service of Reinwardt, who drew for
Blu.me.

13) - Why the polemic papers between Blume and Junghuhn, de Vriese, and others (see B)
started as late as 1850 is unclear, because Blume had already in 1844 (see A) reduced Lycopodium
arboreum - the subject of controversy. Blume's denigrating words accompanying the reduction
were published by him in Rumphia (3: 219, 221) and these gave offence to Junghuhn and de
Vriese. Stafleu & Cowan (Taxonomic literature, ed. 2, vol. 1, 1976) gave 1847 as date for this
part of Rumphia, but it might be that 1849 fits better (as mentioned by Lorentz, c/. Flora Male-
siana I, 4: clxxii, and also accepted by de Wit).

14) - Why Blume more openly and publicly is not clear. It is of course
did not defend himself
rumours without published evidence. He was clearly not a very
a fact that one cannot well oppose
militant personality. Blume took action only twice: first, when he revealed the transfer of the
Rijksherbarium from Brussels and gave honour to Flscher (A: 1831); and second, in defending
himself against Junghuhn (A: 1850). For the rest he satisfied himself by writing explanatory let-
ters. Though convinced of his view on the cause of cholera, he did not officially oppose Mulder
in public. In all these matters am inclined to believe Blume felt below his dignity to expose
I it

himself.

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 Flora Malesiana

Appendix E - Eponymy

Blumia C.G.D. Nees 1823, nom. rejic. /^Magnolia L.).

Blumia K.P.J. Sprengel 1826 ( = Saurauia Willd.).
Blumea H.G.L. Reichb. 1828 ( ^ \eesta Bll-me).
Blumea A. P. DC. 1833.
Blumeodendron Kl'KZ 18"3.
Blumella van Tieghem 1895 ^ Elytranthe Blu )ae ~ Macrosolen Blume).
(

Blumeopsis Gacsep 1920. .

The journal Blumea, official botanical journal of the Rijksherbarium; vol. 1, \934-hodie.

Epithets for species, blumei, blumii, etc. , are too numerous to enumerate here.

Appendix F - Honoran distinctions and memberships

1829 (31 March): Ridder (Knigh:) m de Orde .an de .Nederlandse Leeuw; the Netherlands.
1851: Legion d'Honneur; France.
1851: Preussische Rothe .Adler-Ordens, 3. Klasse; Prussia.
1853: Knight Cross of the Albrechis Order of Sachsen; Saxony.
1853: Large golden medal for merits from the King of Belgium.

1822: Council member Bata\iaasch Genootschap van Kunsien en Wetenschappen, Baiavia; Neth-
erlands Indies.
1825 (6 Febr.j: Corresponding member of the .Maatschappij van Landbouw en Kruidkunde; the

182" -.e Koninklijk Instituui van Wetenschappen, Letterkunde en SchooneKunsten

...- ._... . iiherlands Royal Academy); the .Netherlands.
1827 (29 June): Member of the Provinciaals Utrechtsch Genootschap voor Kunsten en
Wetenschappen, Utrecht; the Netherlands.
1829 (7 Jan.): Member of the Konigliche Botanische Gesellschaft zu Regensburg; Bavaria, Ger-
many.
1833: Member of the HoUandsche Maatschappij van \\etenschappen, Haarlem; the Netherlands.
1845: Doctor honoris causa and .Matheseos magister of Leiden University; the Netherlands.
1851 corresponding member of the Institut de France, Paris; France.
(7 April): Foreign
1853 (May): Ordinary member of the Kaiserliche Akademie fur Naturkunde, .Moscow; Russia.
1855: Honorary member of the '.Maatschappij ter Bevordering der Geneeskunde', Baden; Ger-
many.
1855 (31 March): Member of the Koninklijke Akademie van Wetenschappen, Amsterdam; the
Netherlands.
1856 (10 Oct.): Member of the Royal Academy of Sciences, Stockholm; Sweden.

Member of:
Caesarea Leopoldino-Carolina Academia Naturae Curiosorum, Bonn; Germany. Cognomen:
Rumphius secundus.
Linnean Society of London; England.
Societas Caesarea Naturae Curiosorum .Mosquensis, Moscow; Russia.
Societas .Medico-Botanica Londinensis, London; England.
Natuurkundige Vereeniging van Nederlandsch-Indie, Batavia; Netherlands Indies.

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 ABBREVIATIONS AND SIGNS
ace. = according ex audi. = ex auciores; according to authors
Ak. Bis. = Aklan Bisaya (Philip, language) excl. =exclusus (masc); excluding, exclusive of
Alf. Cel. = Alfurese Celebes (language) ex descr. = known to the author only from the de-
alt. = altitude scription
Anat. = Anatomy . /. (before a plant name) = forma; form
.Ap. = .Apayao (Philip, language) /. (after a personal name) =filius; the son
app. = appendix, appendices f . (in citations) = figure
appr. = approximate fam. = family
Apr. = .April Feb(r). = February
Arch. = .Archipelago fide = according to
atl. = atlas fig. = figure
auci. div. = auciores diversi; various authors f]. =flore, floret (floruit); (with) fiower, flowering
auct(t). mat. = auciores malayenses; authors dealing For. Serv. = Forest Service
with Malesian flora fr. =fructu, fructescit; (with) fruit, fruiting
- auciores plures; several authors
auctflj. plur. Fr. (after a vernacular name) = French
.Aug. = August
. G. = Gunung (Malay); mountain
Bag. = Bagobo (Philip, language) Gad. = Gaddang (Philip, language)
basionym = original name of the type specimen; its gen. = genus; genus
epithet remains permanently attached to the taxon genus delendum = genus to be rejected
which is typified by it provided it is of the same Germ. = German
rank. geronl. = Old World
Bg. = Buginese (language) haud = noi, not at all
Brk. = Bikol (Philip, language) holotype = the specimen on which the original de-
Bil. = Bila-an (Philip, language) scription was actually based or so designated by
Bill. = Billiton the original author
Bis. = Bisaya (Philip, language) homonym = a name which duplicates the name of an
Bon. = Bont6k (Philip, language) earlier described taxon (of the same rank) but
Born. = Borneo which is based on a different type species or type

Bt = Bukit; mountain specimen; all later homonyms are nomenclaturally

Bug. = Buginese (language) illegitimate, unless conserved
Buic. = Bukidnon (Philip, language) I. = Island
c. = circiier, about ibfid). = ibidem; the same, in the same place
C. Bis. = Cebu Bisaya (Philip, language) Ibn. = Ibanag (Philip, language)
cf. - confer, compare ic. = icon, icones; plate, plates
Chab. = Chabecano (Philip, language) ic. inedit. = icon ineditum, icones inedila; inedited
citations = see references plate(s)
cm = centimetre id. =idem; the same
c.n. =see comb, now i.e. =id est; that is
comb, now =combinaiio nova; new combination If. = Ifugao (Philip, language)
CS = cross-section or transversal section of an organ Ig.= Igorot (Philip, language)
c.s. =cum with collaboration
suis; Ilg.= Ilong6t (Philip, language)
cum fig. = including the figure Ilk. = Iloko (Philip, language)
cur. =curanie: edited by in adnol. = in adnotatione; in note, in annotation
D (after a vernacular name) = Dutch incl. =inclusus (masc); including, inclusive(ly)
Daj. = Dyak (language) indet. = indetermined
d.b.h. = diameter at breast height Indr. = Indragiri (in Central Sumatra)
D.E.I. = Dutch East Indies inedit. =inediius (masc); inedited
descr. added behind a reference = means that this in herb. = in herbario; in the herbarium
contains a valid description in lilt. = in lilteris; communicated by letter
diam. = diameter in sched. = in schedula; on a herbarium sheet
Distr. (as an item) = Distribution in sice. =in sicco; in a dried state
Distr. (with a geographical name) = District in syn. = in synonymis; in synonymy
ditio = the same, see do Is. = Islands
Div. = Division, or Divide Is. (after a vernacular name) = Isinai (Philip, lan-
div. =diversus (masc); various guage)
do = diiio (Ital.); the same Ism. = Isamal (Philip, language)
Dum. = Dumagal (Philip. language) isotype = a duplicate of the holotype; in arboreous
dupl. = duplicate plants isotypes have often been collected from a
E = east (after degrees: eastern longitude) single tree, shrub, or liana from which the holo-
E (after a vernacular name) = English type was also derived
Ecol. = Ecology Iv. = Ivatan (Philip, language)
cd. = edited; edition; editor J(av). = Javanese (language)
e.g. = exempli gratia; for example Jan. = January
elah. = elaboravit; revised Jr = Junior
emfend). = emendavit; emended KIg. = Kalinga (Philip, language)
em(erg). cd. = emergency edition Kul. = Kuliman (Philip, language)
Engl. = English Kuy. = Kuy6non (Philip, language)
etc.. dc. =el cetera; and (the) other things Lamp. = Lampong Districts (in S. Sumatra)

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 Flora Malesiana

Lan. = Lanao (Philip, language) nom. rejdc.) = nomen rejiciendum; name rejected by
lang.= language the International Rules of Botanical Nomenclature
/.c. =loco cilalo; compare reference nom. seminudum = a name which is provided with
lectotype = the specimen selected a posteriori from some unessential notes or details which cannot be
the authentic elements on which the taxon was considered to represent a sufficient description
based when no holotype was designated or when which is, according to the International Rules of
the holotype is lost Botanical Nomenclature, compulsory for valid
livr. = livraison, part publication of the name of a taxon
lice. =l.c. (plur.) nom. suhnudum = nomen seminudum
LS = longitudinal or lengthwise section of an organ nom. superfl. = a name superfluous when it was pub-
m = metre lished; in most cases it is a name based on the same
M = Malay (language) type as an other earlier specific name
Mag. = Magindanao (Philip, language) non followed by author's name and year, not placed
Mak. = Makassar, Macassar (in SW. Celebes) in parentheses, and put at the end of a citation =
Mai. = Malay(an) means that this author has published the same
Mai. Pen. = Malay Peninsula name mentioned in the citation independently.
Mand. = Mandaya (Philip, language) These names (combinations) are therefore homo-
Mang. = Mangyan (Philip, language) nyms.
Mar. = March line 5-4 from bottom. The same can
Compare 56b
Mbo = Man6bo (Philip, language) happen with generic names.
Md. = Madurese (language) {non followed by abbreviation of author's name) be-
Minangk. = Minangkabau (a Sumatran language) fore a reference (citation) headed by an other
min. part. =pro minore parte; for the smaller part author's name = means that the second author has
mm = milimetre misinterpreted the taxon of the first author.
Mng. = Mangguangan (Philip, language) Compare 419a under species 47 the synonym H.
p.
Morph. = Morphology celebica. Diels misapplied the name H. celebica as
ms(c), MS(S) = manuscript(s) earlier described by Burck.
Mt(s) = Mount(ains) non = non aliorum; not of other authors
al.
n. = numero; number non = not seen by the author
vidi
N = North (after degrees: northern latitude); or New nov. =nova (femin.); new (species, variety, etc.)
{e.g. in N. Guinea) Nov. = November
NE. = northeast n.s. = new series
nee = not n. sp. = nova species; new species
rieerl. - Netherlands, Netherlands edition n. (sp.) prov. = nomen (specificum) provisorium;
Neg. = Negrito (Philip, language) provisional new (specific) name
N.E.I. = Netherlands East Indies n.v. = non vidi; not seen
neotype = the specimen designated to serve as no- NW. = northwest
menclatural type when no authentic specimens Oct. = October
have existed or when they have been lost; a neotype op.cit. =opere citato; in the work cited
retains its status as the new type as long as no auth- p. =pagina; page
entic elements are recovered and as long as it can P. = Pulau, Pulu (in Malay); Island
be shown to be satisfactory in accordance with the Pal(emb.) = Palembang
original description or figure of the taxon Pamp. = Pampangan (Philip, language)
N.G. = New Guinea Pang. = Pangasinan (Philip, language)
N.I. = Netherlands Indies paratype = a specimen cited with the original descrip-
no = numero; number tion other than the holotype
nom. =nomen; name (on\y) = nomen nudum part. alt. = for the other part
nom. al. = nomen aliorum; name used by other P. Bis. = Panay Bisaya (Philip, language)
authors P.I. = Philippine Islands
nom. alt(ern). = nomen alternativum; alternative pi. = plate
name plurim. = plurimus; most
nom. cons(erv). = nomen conservandum, nomina p.p. =pro parte; partly
conservanda; generic name(s) conserved by the In- pr. max. p. =pro maxima parte; for the greater part
ternational Rules of Botanical Nomenclature pro = as far as is concerned
nom. fam. cons. = nomen familiarum conservan- prob. =probabiliter; probably
dum; conserved family name prop. = propositus; proposed
nom. gen. cons. = see nomen conservandum Prov. = Province
nom. gen. cons. prop. = nomen genericum conser- pr.p. =pro parte; partly
vandum propositum; generic name proposed for pt = part
conservation quae est = which is
nom. illeg(it). = nomen illegitimum; illegitimate 9«ooc^ basionym, syn., specimina, etc. =as far as the
name basionym, synonym(s), specimen(s), etc. are con-
nom. leg(it). = nomen legitimum; legitimate name cerned
nom. nov. = nomen novum; new name references = see for abbreviations the list in vol. 5,
nom. nud. = nomen nudum; name published without pp. cxlv-clxv
description and without reference to previous pub- Res. = Residency or Reserve
lications resp. = respective(ly)

(42)
 Abbreviations and signs

S = south (after degrees: southern latitude) taxon = each entity throughout the hierarchic ranks
S (after a vernacular name) = Sundanese (language) of the plant kingdom which can be described and
Sbl. = Sambali (Philip, language) discriminated from other taxa of the same rank
SE. = southeast Taxon. = Taxonomy
sec. = secus; according to Tg = Tandjung (Malay); cape
sect. =sectio; section Ting. = Tinggian (Philip, language)
sens. ampl. (ampliss.) = sensu amplo (amplissimo); Tir. = Tirurai (Philip, language)
in a wider sense, in the widest sense transl. = translated
sens. lai. =sensu laio; in a wide sense type = each taxon above the rank of a species is typi-
sens. sir. (sirictiss.) = sensu siricto (sirictissimo); in by a type belonging to a lower rank, for in-
fied
the narrow sense, in the narrowest sense stance a family by a genus, a genus in its turn by
Sept. = September a species; a species or infraspecific taxon is typified
seq., seqq. =sequens, sequeniia; the following by a specimen. Thenameof ata.xon is nomenclatu-
ser. = series rally permanently attached to its type; from this it
=sensu lato; in a wide sense
S.I. cannot be inferred that the type always represents
S.-L. Bis.= Samar-Leyte Bisaya (Philip, language) botanically the most typical or average structure
Sml. = Samal (Philip, language) found in the circumscription of the taxon.
s.n.=sine numero: (specimen) without the collec- type specimen = the specimen or other element to
number
tor's which the name of a species or infraspecific taxon
Sp. = Spanish (language) is (nomenclaturally) permanently attached; botan-

sp(ec). = species; species ically a type specimen is a random specimen on

specim. = specimen(s) which the name was based by description. There-
sphalm. = sphalmaie; by error, erroneous fore, it does not need to represent the average or
spp. = species; species (plural) most typical representative of a population. See
Sr = Senior holotype, isotype, lectotype, syntype, paratype,
5.5.= see sens. sir. and neotype
ssp. = subspecies; subspecies typ. excl. = typo excluso; type excluded
s.sir. = see sens. str. typ. incl. = typo incluso; type included
Stat, now = status nova; proposed in a new rank typus = see type and type specimen
Sub. = Subanum (Philip, language) var. = varietas; variety
subgfen). = subgenus; subgenus var. nov. = varietas nova; new variety
subseci. =subsectio; subsection Vern. = Vernacular
subsp. = subspecies; subspecies vide = see
Sul. = language)
Siilu (Philip, viz. = videlicet; namely
Sum. E.C. = Sumatra East Coast vol. = volume
Sum. W.C. = Sumatra West Coast W = west (after degrees: western longitude)
Suppl. = Supplement Yak. = Yakan (Philip, language)
SW. = southwest ± = about
syn.= synonymum; synonym & = and
synonyms = the names of taxa which have been re- = diameter
ferred to an earlier described taxon of the same (^ =male (flower, etc.)
rank and with which they have been united on $ = female (flower, etc.)
taxonomical grounds or which are bound together 5, -bisexual (flower)
<^

nomenclaturally (cJ) = dioecious with unisexual flowers

(9)
syniypes = the specimens used by the original author (c??) = monoecious with unisexual flowers
when no holotype was designed or more specimens (<? 5) = polygamous
were simultaneously designated as type (??) = polygamous
t. = tabula; plate oo = many
Tag. = Tagalog (Philip, language) > = more than (in size, number, etc.)
Tagb. = Tagbaniia (Philip, language) < = less than (size, number, etc.)
Tagk. = Tagaka-6lo (Philip, language) X 2/5 = 2/5 of natural size
Tapan. = Tapanuli (in NW. Sumatra) X montana = means that the epithet montana is that
of a hybrid

(43)
 CHRYSOBALANACEAE (G.T. Prance, Kew)^

Trees or shrubs (or rarely suffrutices outside Malesia). Leaves simple, alternate,
often coriaceous, glabrous or with an indumentum on undersurface, margin en-
petioles often with 2 lateral glands. Stipules 2, minute and caducous to large
tire;

and persistent, usually linear-lanceolate. Inflorescence racemose, paniculate or

cymose; flowers bracteate and usually bibracteolate; bracts and bracteoles small
and caducous or larger and enclosing flower or groups of flowers and persistent.
Flowers actinomorphic to zygomorphic, hermaphrodite or rarely polygamous,
markedly perigynous. Receptacle campanulate to cylindrical or rarely flattened
cupuliforum, often gibbous at base; calyx lobes 5, imbricate, often unequal,
erect or reflexed. Petals 5 (absent in some Neotropical species), inserted on mar-
gin of disk, commonly unequal, imbricate, deciduous, rarely clawed. Stamens
indefinite, 2-60 (to 300 in Neotropics), inserted on margin of the disk, in a com-
plete circle or unilateral, all fertile or some without anthers and often reduced
to small tooth-like staminodes; filaments filiform, free or ligulately connate,
short and included to long and far exserted; anthers small, 2-locular, longitudi-
nally dehiscent, glabrous or rarely pubescent. Ovary basically of three carpels
but usually with only one developed, the other two aborted or vestigial, various-
ly attached to (the base, middle or mouth oO receptacle, usually sessile or with
short gynophore, pubescent or villous; ovary unilocular with two ovules or bi-

locular with one ovule in each locule. Ovules erect, with micropyle at base
(epitropous). Style filiform, basally attached; stigma 3-lobed or truncate. Fruit
a fleshy or dry drupe of varied size, interior often densely hairy; endocarp much
varied, thick or thin, fibrous or bony, often with a special mechanism for seed-
ling escape. Seed erect, exalbuminous, the testa membraneous; cotyledons
amygdaloid, plano-convex, fleshy, sometimes ruminate. Germination hypogeal
with the first leaves opposite or alternate or epigeal with opposite first leaves.

An extensive review of the generic limits of the family has been published: G.T. Prance &
F. White, The genera of Chrysobalanaceae: a study in practical and theoretical taxonomy and
its relevance to evolutionary biology, Phil. Trans. Roy. Soc. London 320 (1988) 1-184. This con-
taxonomic history, morphology, anatomy, pollen, ecology and distribution
tains full details of
of the family. A condensed version of these subjects is given here. Details of the Neotropical
members of the family are given in: G.T. Prance, Chrysobalanaceae, Flora Neotropica 9 (1972)
1-410. The African members of the family were treated in: F. White, The taxonomy, ecology
and chorology of African Chrysohalanaceae (excluding Acioa), Bull. Jard. Bot. Nat. Belg. 46
(1976) 265-350.

Distribution. Pantropical with 456 species in 17 genera; 365 species in the Neotropics, 57
in Africa, and 34 in Asia, Malesia and the Pacific.
Seven genera are native to the Flora Malesiana region and one species of an eighth genus, Chry-
sobalanus, from Africa and South America, has naturalized in Malesia and Fiji and is therefore
included in this treatment. All four tribes of Chrysobalanaceae are represented in the region. The
genera treated here fall into the following tribes of Pranc i- & White:
Tribe Chrysobalaneae: Chrysobalunus, I.icania, Paraslemon.

(I) Drawings made by Bobbi Angcll. David Woolcott, KirMcn Tind, and Julia Lokcn; David Johnson
assisted with ihc distribution maps.

(635)
636 Flora Malesiana [ser. I, vol. lO"*

Tribe Parinarieae: Hunga, Parinari.

Tribe Couepieae: Maranthes.
Tribe Hirtelleae: A tuna, Kostermanthus.
The genera Atuna, Hunga, Kostermanthus, and Parastemon are confined to the Malesian and
Pacific region. Licania predominantly a Neotropical genus (186 species there) with a single
is

species in West Africa and three in Malesia. Parinari is a pantropical genus with almost equal rep-
resentation in all three major regions of the tropics, and Maranthes is predominantly an African
genus with one abundant and widespread species in Malesia and the Pacific and a single closely
related species in Central America.
Morphology. All species of Chrysobalanaceae are woody and most are trees or treelets. All
are leptocaul. Several, including species of Atuna, Kostermanthus, Licania (Neotropical), Magni-
stipula (African), Maranthes and Parinari, exceed a height of 30 m and are important constituents
of the upper forest canopy or are emergents. Six African and Neotropical species belonging to
Licania, Magnistipula and Parinari are geoxylic suffrutices with massive woody underground
parts, but rather exiguous aerial shoots which are capable of only limited upward growth and a
similar form occurs in Parinari nonda in Australia.
In their architecture and growth-dynamics those Chrysobalanaceae that have been studied ex-
hibit the model of Troll. This has been demonstrated only in African and Neotropical species.
Herbarium specimens of Atuna show a distinct pattern of branching which is difficult to
describe except in terms of development based on the living plant.
Buttresses are normally absent but frequently well-developed in some species of Parinari and
Atuna, for example, P. canarioides, P. costata, P. oblongifolia, A. cordata and A. excelsa, and
the trunk of some species of Parinari, e.g. P. parva and P. gigantea is often fluted at the base.
The leaves, which are simple and spirally inserted, are frequently arranged distichously. Most
species have stiff, coriaceous, evergreen leaves which contain abundant silica inclusions.
Stipules are nearly always present but are sometimes small and caducous. In some Neotropical
species of Parinari the stipules reach a length of 7 cm, they are up to 4 cm in Parinari parva. In
Atuna they are prominently keeled, a unique feature in the family.
The lamina is entire, in all Malesian species. In nearly all species of Parinari, and a few
Neotropical species of Licania, the veins on the lower surface are extremely prominent and form
a dense network occupying more than half of the leaf surface so that the stomata are confined
to relatively small sunken crypts which are densely filled with short curly hairs.
Foliar glands occur in most, possibly all, species. They
secrete nectar which is eaten by ants,
and function chiefly on young leaves. On mature leaves of herbarium specimens they are not
always clearly visible. The structure and distribution on the leaf of the glands varies greatly from
genus to genus and provides characters of considerable taxonomic importance. Small discoid
glands occur in various places on the lower surface or margins of the lamina in Parastemon. There
are larger, sometimes ill-defined, glandular areas towards the base of the lamina in Maranthes.
In Parinari conspicuous glands occur on the petiole.
The inflorescence is very variable. In Chrysobalanus the few-flowered inflorescence is a short
raceme of cymules or is cymose throughout, or is a false raceme or a subsessile fascicle. In
Parastemon the inflorescence is a simple or branched raceme. Hunga and Malesian species of
Licania have simple or branched racemes of usually congested cymules. More complex mixed in-
florescences with cymose ultimate units are found in Kostermanthus and Parinari, and the inflo-
rescence of Maranthes is corymbose.
Since the inflorescence is usually cymose, at least in part, a distinction between bract and
bracteole cannot always be drawn. Bracts and bracteoles are usually small but in nearly all species
of Parinari they are relatively large and enclose small groups of developing flowers.
In most species the flowers appear to be bisexual, but future field work may show that this is
not always so. Parastemon urophyllus is said to be polygamodioecious.
Floral symmetry varies from almost completely actinomorphic, apart from the lateral style, in
1989] Chrysobalanaceae (Prance) 637

Chrysobalanus, Parastemon, and most species of Licania to strongly zygomorphic in Kosterman-

thus. Aciinomorphic flowers are patelliform or shallowly cupuliform, and zygomorphic flowers
usually have a long receptacle-tube, but in Kostennanthus the strongly zygomorphic flowers have
a very short receptacle. In the Chrysobalanaceae the receptacle-surface is always lined with nectar-
secreting tissue, which sometimes, Maranthes corymbosa, almost completely fills the tube.
as in
In most genera the entrance to the receptacle tube is blocked by long straight retrorse hairs, but
these are lacking in Kostermanthus. In Parastemon the nectariferous lining of the receptacle is
freely exposed.
There are always five, completely free, slightly to strongly imbricate sepals which vary from
subequal in Chrysobalanus to markedly unequal in Kostermanthus. In most genera they are acute
or subacute but in Kostermanthus and Maranthes they are suborbicular and deeply concave.
Petals are present in all Malesian species but absent in many Neotropical species of Licania.
There are always five. They are mostly caducous. In shape they vary from linear-spathulate
(Chrysobalanus) to orbicular. They are usually subequal, but in Kostermanthus they are very un-
equal in shape and size and are strongly unguiculate.
Stamens vary in number from two
in Parastemon urophyllus to 40 in Maranthes. In Chrysoba-

lanus, most species of Licania, Parastemon versteeghii, and Maranthes they form a complete or
almost complete circle round the entrance to the flower and all or most are fertile. Otherwise the
fertile stamens are inserted unilaterally opposite the carpel. Staminodes are frequently present op-

posite the style. In several genera the filaments appear to be united at the base, but it is sometimes
difficult to decide whether this represents true union or whether the filaments are free but inserted
on a development of a receptacular rim. In Maranthes the stamens are inserted in two or more
rows on the outer surface of what appears to be a receptacular annulus. In length the filaments
vary from much shorter than the calyx, as in Hunga, Parastemon and some species of Licania,
to very much longer in Maranthes. In Kostermanthus the filaments are united for at least half of
their length to form a conspicuous ligule.
The gynoecium fundamentally is composed of three carpels which are free except for the
gynobasic style. In most species there is only one functional carpel, though one or two small
rudimentary carpels can sometimes be seen. Due to the development of a false dissepiment the
ovary is bilocular in Hunga, Parinari, and A tuna.
The fruit is basically a drupe but there is considerable variation in detail, apparently associated
with dispersal and germination. In Chrysobalanus, Parastemon and Hunga the endocarp has a
smooth surface and is sharply differentiated from the mesocarp. In the other genera the differen-
tiation is less well-defined. In Chrysobalanus and Hunga, seedling escape is effected by means of
longitudinal lines of weakness. In Parastemon and Maranthes two large lateral plates fall away
permitting the seedling to emerge. In Parinari there are two small basal 'plugs' or obturators. All
other genera seem to lack specialized means of seedlings escape.
In Chrysobalanus, Licania, Parastemon, Parinari, and A tuna, germination is cryptocotylar,
whereas in Maranthes it is phanerocotylar.
Vegetative Anatomy. - Leaf anatomy. Indumentum, if present, consisting of long unicel-
lular hairs. Variously positioned glands (cxtrafloral nectaries) with slender upright epidermal
secretory cells commonly present. Wax present as platelets (Fhhrlnbach & Barthi.ott, 1988).
Siomata mostly paracytic, confined to the lower leaf surface. Upper epidermis often composed
of tall cells; with mucilaginous inner walls in some species. Hypodermis often present. Mesophyll
entirely composed of palisade-like cells, more rarely dorsivcntral and differentiated into palisade
and spongy tissue. Astcrosclcrcids occasionally present in mesophyll. Veins mostly with scleren-
chyma sheaths including sclercids with U-shaped wall thickenings, sometimes vertically Iranscur-
rent. Midrib and distal end of petiole with a closed vascular cylinder, with or without additional
adaxial or medullary collateral bundles. Silica bodies and silicified cell walls common, especially
in epidermis.
Young stem. Cork arising superficially. Pcricyclic sclercnchyma ring composed of fibres and
638 Flora Malesiana [ser. I, vol. 10"^

sclereids with U-shaped wall thickenings. Secondary phloem occasionally with secretory (tannin?)
cells. Sieve tube plastids of the S-type (Behnke, 1984). Silica bodies often present in pericycle,
phloem and xylem rays, and in pith.
IVood anatomy. Growth rings absent or, if present, defined by differences in the spacing of
tangential parenchyma bands. Vessels diffuse, often in a weakly oblique pattern, (almost) exclu-
sively solitary, tending to be of two distinct sizes, the larger ones very wide (200-300 [im). Vessel
perforations simple. Tyloses often present in heartwood, sclerotic in some species. Vessel-ray pit-
ting including elongate horizontal or oblique to almost vertical pits with strongly reduced borders,
often unilaterally compound. Fibres often thick-walled, with distinctly bordered pits throughout
the tangential walls, and in the radial walls often confined to fibre-ray contacts (fibre-tracheids);
in contact with vessels often less thick-walled and with biseriate bordered pits ( == vasicentric
tracheids). Parenchyma in fine uniseriate or locally bi(-tri)-seriate, regular or irregular wavy tan-
gential bands. Parenchyma strands typically long, of up to 16 cells. Some axial parenchyma cells
with spiral thickenings in Atuna p.p., Licania, Maranthes p.p., and Kostermanthus (Jer Welle,
1975). Rays predominantly uniseriate, but in some taxa also biseriate, typically weakly heteroge-
neous with (often weakly) procumbent central cells and one row of square to upright marginal
cells (Kribs type III), sometimes homogeneous and composed of procumbent cells only. Silica

bodies universally present in ray cells, more rarely in axial parenchyma cells. Rhomboidal crystals
in chambered axial parenchyma cells noted in Parastemon.

Taxonomic notes based on vegetative anatomy. The above general anatomical description is
based on the literature (for leaf and young stem anatomy mainly Kuster, 1897, as abstracted by
SoLEREDER, 1899; and Prance, 1972, and Prance & White, 1988, for wood anatomy from many
sources), amplified with original observations on slides present in the Rijksherbarium at Leiden.
A number of anatomical characters may prove to be of considerable taxonomic significance at
the genus or species level (mucilaginous leaf epidermis, distribution of silica grains in leaves,
young stem, and wood, vascular pattern and sclerenchyma support of leaf veins and petiole, fibre
and sclereid distribution pattern of the mature bark (Roth, 1981), spiral thickenings in axial
parenchyma cells of the wood, ray width and histology, etc.). However, for the Malesian Chryso-
balanaceae their diagnostic value remains largely untested. On the whole the Chrysobalanaceae
are anatomically rather homogeneous, and as repeatedly emphasized, quite distinct from the
Rosaceae. Anatomically Chrysobalanaceae are also distinct from the numerous families to which
they have been compared in the search for closest relatives.
References: Behnke, Ann. Missouri Bot. Gard. 71 (1984) 824-831; Desch, Manual of
Malayan Timbers 2 (1954) 474-485; Burgess, Timbers of Sabah (1966) 434-436; Fehrenbach
& Barthlott, Bot. Jahrb. 109 (1988) 407-428; Furuno, Anatomy of Papua New Guinea Wood
(Continued), Res. Report of Foreign Wood 8, ShimaneUniv. (1979); Hayashi c.5., Micrographic
Atlas of Southeast Asian Timber, Kyoto Univ. (1973); Lecomte, Les bois de I'lndochine (1926)
59-61; Metcalfe & Chalk, Anatomy of the Dicotyledons 1 (1950) 550-553; Moll &
Janssonius, Mikrographiedes Holzes der auf Java vorkommenden Baumarten 3 (1914)222-230;
Prance, Flora Neotropica 9 (1972) 1-19; Prance & White, Phil. Trans. Roy. Soc. Lond. B 320
(1988) 1-184; Roth, Encycl. Plant Anatomy 9, 3 (1981) 286-295, 402-403; Solereder,
Systematische Anatomic der Dicotyledonen (1899) 341-351; Ter Welle, Acta Bot. Neerl. 24
(1975) 397-405; lAWA Bulletin 1976/2 (1976) 19-29; Ter Welle & Detienne, Flora of the
Guianas A 85 (1986) 109-126. - P. Baas.
Palynology. The pollen of Chrysobalanaceae is very uniform, but is different from that of
Rosaceae. It is of little value for distinguishing between the genera of Chrysobalanaceae or for
arranging them in groups.
Most species have grains with three furrows, but some species have three or four; there are no
special features except occasional equatorial constrictions. With light microscopy the pores are
indistinct, and in some The grains are usually distinctly triangular
species are difficult to observe.
in shape in polar view, except when four-furrowed; they are elliptical to circular in equatorial view
1989] Chrysobalanaceae (Prance) 639

and are oblate-spheroidal, prolate-spheroidal or subprolate in shape as indicated by the ratio:

polar length x 100, divided by the equatorial length = 85-150. The size is very variable from one
genus to another; the polar area is usually small, sometimes medium, but never large. The exine
is medium to rather thick with very little patterning; it is usually scabrous to verrucose, but never
striate.

The pollen of Chrysobalanaceae and Rosaceae is similar but readily distinguishable. The
former is markedly triangular in polar view in the expanded grain, whereas in Rosaceae it is never
more than weakly triangular. Most Rosaceae have more distinctive pores, and many have more
patterning on the wall. A feature that occurs frequently in the Rosaceae is a distinct wedge-shaped
protrusion from the middle of the furrow, obvious in polar view, which does not occur in Chryso-
balanaceae.
Erdtman (1952) states 'pollen morphological objections cannot be raised against regarding the
Chrysobalanaceae diS a separate family.' Our own study o{ Rosaceae X)o\\tr\ {sensu lato) confirmed
that three main types of pollen occur: the Rosaceae sensu stricto, the Chrysobalanaceae and the ,

Seuradoideae types (Prance, 1963). The differences between pollen of Chrysobalanaceae and
Rosaceae are, however, comparatively small. By contrast, the pollen of the Tropaeolaceae,
Geraniaceae, Limnanthaceae, Linaceae, Polygalaceae, and Sapindaceae, families which various
phylogenists (Hallier, 1923; Bonne, 1926; Hauman, 1951; Gutzwiller, 1961) have suggested
are closely related to Chrysobalanaceae, is very different. Pollen morphology thus provides
reasons for keeping the Chrysobalanaceae near to the Rosaceae in the Rosales, and not for remov-
ing it to the Geraniales or Sapindales.
The pollen of Chrysobalanaceae is so uniform that it does not provide good generic characters.
Kostermanthus heteropetala is distinct from all other Chrysobalanaceae examined, including
Daciyladenia (Africa) and Acioa (America) with which it shares a staminal ligule, in having three
swellings on each of the triangular sides of the grain in polar view. Apart from Kostermanthus
no other genus is clearly definable on pollen characters.
References: Bonne, C. R. Hebd. Seanc. Acad. Sci. Paris 182 (1926) 1404-1406; Erdtman,
Pollen morphology and plant taxonomy, Angiosperms (1952) 380-383; Gutzwiller, Bot. Jahrb.
81 (1961) 1-49; Hallier, Beih. Bot. Centralbl. 39(1923) 1-178; Hauman, Bull. Jard. Bot. Etat
Brux. 21 (1951) 167-198; Prance, A taxonomic study of the Chrysobalanaceae. Thesis, Oxford
(1963).
Phytochemistry. Chemical knowledge about the family Chrysobalanaceae is still scanty.
Hegnauer (1973) treated it as Chrysobalanoideae sub Rosaceae. Chrysobalanaceae are note-
worthy for their tendency to accumulate silica (SiOz) in leaves and in the wood where usually every
ray cell contains one globular silica inclusion. Leaf flavonoid patterns are dominated by the
flavonols quercetin and kaempferol; some taxa also have myricetin. Proanthocyanidins (formerly
called leucoanthocyanidins), i.e. condensed tannins, were demonstrated to be present in leaves of

few species of Chrysobalanus, Licania, and Parinari, but galli- and ellagitannins have not yet been
traced in the family. The recent flavonoid investigation of 21 species of f*ormcrr/(CoRADiN, Gian-
NASi & Prance, 1985) resulted in the identification of a number of 3-glycosides of kaempferol,
quercetin and myricetin, and showed restriction of myricetin glycosides to four African species;
dihydroquercetin('taxifolin')-3-glycosides were noticeable only in Asian Parinari insularum from
the Pacific islands and vicenin-like C-glycoflavones only in a few African populations of P. ex-
celsa. Myricetin was also observed in leaves of Licania macrophylla which besides has much con-
densed tannins in all parts, saponins in leaf, pericarp, seed, and stem and root bark; alkaloids
in stem and root bark (Grenand, Morhtti & Jacquhmin, 1987). Cyanogenic glycosides which
are characteristic of a number of Rosaceous taxa have not been traced in Chrysobalanaceae
hitherto. The most noteworthy chemical character known from the family at present is the fatty
acid pattern of their seed triglycerides; conjugated iricnoicand tetraenoic CiK-acids such as alpha-
claeostearic and parinaric acids are present as major fatty acids in seed oils of species of Chrysoba-
lanus, Licania, and Parinari s. I. (i.e. including Atuna, Maranthes and the African Neocarya).
640 Flora Malesiana [ser. I, vol. 10'*

This character, however, which Hnks Chrysobalanaceae biochemically with Prunoideae (same
type of seed oils in some Prunus s.l. species) seems not to be universal in the family. According
to Jones & Earle (1966) seed kernels of a species of Couepia (Central & South America) con-
tained an oil without conjugated unsaturation. Still is known from the chemistry of this
too Httle
taxon to allow a sound chemotaxonomic discussion.
References: Coradin, Giannasi &
Prance, Brittonia 37 (1985) 169-178; Grenand, Moretti
& Jacquemin, Pharmacopees Guyane, ed. Orstom, Paris (1987); Hegnauer,
traditionelles en
Chemotaxonomic der Pflanzen 6 (1973) 84-130; Jones & Earle, Econ. Bot. 20 (1966) 137;
Prance & White, Phil. Trans. Roy. Soc. London B 320 (1988) 28-29. - R. Hegnauer.
Dispersal. The fruits of Chrysobalanaceae are very uniform in basic structure but remark-
ably diverse in functional detail. Despite their uniformity they have become adapted to a wide
range of dispersal agents, sometimes within a single genus or species; however, few species have
been studied in the field.

Chrysobalanus icaco ssp. icaco is dispersed by ocean currents, and also by bats, rodents and
monkeys, and possibly by birds; C. cuspidatus is said to be dispersed by birds.
Some Neotropical species of Licania are bat-dispersed, whereas the fruits of several South
American riverine species float and are also eaten by fish; those of the African species L. elaeo-
sperma are also transported by water. The Malesian species L. splendens is dispersed by the fruit
pigeon Ducula aenea.
Various species of Parinari are known to be dispersed by bats, elephants, baboons and other
primates, a scatter-hoarding squirrel, fruit pigeons, rheas, emus, agoutis and fish. Species of
Couepia, Licania and Parinari are frequently eaten by bats in the Neotropics.
Maranthes corymbosa is dispersed by birds, most notable hornbills and fruit pigeons, and, at
least for short distances, by a scatter-hoarding squirrel. The fruits of some African species are
eaten by monkeys which are possibly mainly destructive.
Atuna is dispersed by ocean currents and a scatter-hoarding squirrel and possibly by wild pigs.
Uses. Members of the Chrysobalanaceae are used by the local people everywhere, for
building, fuel, charcoal and in folk medicine. The fruits and seeds of some species are highly
esteemed, and others are eaten in times of scarcity; some are used in the preparation of alcohoUc
beverages. At present, Chrysobalanaceae are only of local importance commercially, but, with
improved communications and technology, their potential as a source of construction timber,
fruits, and edible and industrial oils appears to be promising.
The Malesian standard timber name for various genera of Chrysobalanaceae is merbatu.
Edible fruits and seeds. Chrysobalanus icaco is tinned and bottled in syrup and sold in Colom-
bia and Venezuela under the name Icacos. The fruit of several Neotropical species of Couepia and
Parinari are eaten. In Amboina a dish called Koku koku is prepared from the mashed seeds of
Atuna excelsa mixed with raw or fried small fish, ginger, onions, chillies and lime juice.
Wood. Despite the large supplies of Chrysobalanaceae wood potentially available, commercial
sawn timber is produced only in relatively small amounts. This is because its high silica content
blunts even tungsten-tipped saws. Because the wood of many species is resistant to marine borers,
it is used throughout the tropics for piers and other marine constructions.

Caulking and waterproofing agent. In the Solomon Islands the principal use oi Atuna excelsa
sensu lato is for caulking the seams of plank-built canoes. The seeds, which are known as 'putty
nut' are pounded to a putty-like consistency. After application the putty hardens and darkens,
but if exposed too long to the sun it cracks, so canoes drawn up on the beach are often kept in
the shade of sheds. In the central and south-eastern Solomons it is used for setting shell inlay in
wood bowls, figures and other articles. The north-western Solomon Islanders also use it for water-
proofing bottles made from gourds. In the Admiralty Islands (Manus) coiled baskets are coated
with it to make them waterproof (B.A.L. Cranstone, in litt., 14 June 1983).
History oi Parinari. The taxonomic history of Parinari is complex. At least some species
of all Malesian genera except Chrysobalanus, and Parastemon have at one time or another been
placed in Parinari.
1989] Chrysobalanaceae (Prance) 641

Aiuna and Maranthes have been included in Parinari. Despite their considerable
All species of
differencesfrom Parinari sensu stricto in virtually all other respects, these genera have one feature
in common - a bilocular ovary. It was the adoption of this character as a generic criterion,
especially by Bentham (1849), that led to the increasingly artificial nature oi Parinari. As Parinari
became more and more heterogeneous even some species with unilocular ovaries were included,
for example, the species now placed in Kostermanthus.
which was based on P. campestris and P. montana from
In the original description of Parinari,
French Guiana, Aublet (1775) mentioned the bilocular ovary, but he does not appear to have at-
tached much importance to it.
De Jussieu (1789), who brought all previously described genera of Chrysobalanaceae together
for the first time, knew some of them only from the original descriptions and illustrations. His
implication that Parinari differs from the other genera principally in its bilocular ovary seems to
have laid the foundations for the subsequent confused history of the group.
De Jussieu was the first to extend the concept of Parinari to another continent by citing in
synonymy two manuscript names of Adanson from Senegal, Mampata and Neou. The former
was subsequently described as P. excelsa and the latter as P. macrophylla by Sabine.
The following year, in his Prodromus, De Candolle (1825), who only knew the four species
mentioned above, divided Parinari into two sections. Section Petrocarya (correctly section
Parinari) was based on a superfluous generic name which Schreber (1789) substituted for the
earlier Parinari. It included Aublet's original species. Section Neocarya was based on P.
senegalensis DC. [now Neocarya macrophylla (Sabine) Prance], but P. excelsa was associated
with it, probably because its type-description is inadequate to characterize it properly. Parinari
macrophylla is not mentioned by De Candolle. He was also apparently unaware of the first true
Parinari to be described from Asia, P. sumatrana Benth., which had been described by Jack in
the illegitimate genus Petrocarya in 1822. De Candolle indirectly emphasized the importance of
the bilocular ovary of Parinari by describing the ovary of all other genera as unilocular.
During the first half of the nineteenth century, in addition to Neocarya macrophylla, a few

other species, which belong to other genera, were described in Parinari or its illegitimate synonym
Petrocarya, because of their bilocular ovary. Thus Jack ( 1 822) described Petrocarya excelsa (now
Atuna excelsa), and Bentham (1840) published Parinari coriacea (now Exellodendron coriacea),
but it was Bentham's treatment of Parinari in Hooker's Niger Flora (1849) that firmly establish-
ed Parinari as an artificial genus.
Whereas workers had implied that the bilocular ovary is a diagnostic character of
earlier
Parinari, Bentham referred to the spurious dissepiment which separates the ovules as 'the essen-
tial character.' Bentham divided Parinari into three sections as follows:
Section Petrocarya (correctly Parinari) included the African species P. excelsa and P.
1:

curatellifolia, all the known American species including P. coriacea (now Exellodendron cor-
iaceum), and, with some doubt, three species Bentham had not seen himself, namely P.
sumatrana Benth. (a true Parinari), P. glaberrima Hassk. (now Atuna excelsa) and P. scabra
Hassk. (now Atuna scabra).
Section 2: Sarcostegia Benth. included two new species, P. polyandra (now Maranthes polyan-
dra) and P. griffiihiana (now Maranthes corymbosa), and, with some doubt, also P. jackiana
Benth. (based on Petrocarya excelsa, now Atuna excelsa) which Bentham had not examined.
Section 3: Neocarya DC. contained P. macrophylla (now Neocarya macrophylla) and its
synonym P. senegalensis.
Bentham's circumscription of Parinari was probably much wider than he imagined, largely
because of the inclusion of the Asian species he only knew from the literature. He appears to have
adopted it with some reservation. Parinari polyandra has c. 40 fertile stamens and Bentham men-
tions that this, in conjunction with the glandular leaves and fleshy 'calyx', might 'suggest the es-
tablishment of a distinct genus.' He clearly believed that the stamen number of Parinari varies
more or less continuously, but the evidence he cites is partly on the species he had not studied.
642 Flora Malesiana [ser. I, vol. 10"^

Bentham's circumscription of Parinari included five genera which are now regarded as distinct,
namely, in additon to Parinari itself, Atuna Rafin., Exellodendron Prance, Maranthes Blume

and Neocarya Prance. Two of these from Malesia had enjoyed a brief period of generic recogni-
tion. Thus, Maranthes was described by Blume in 1825, but three years later he transferred the
type species to his illegitimate Exitelea. A tuna was described by Rafinesque in 838, but remained 1

disregarded for more than 100 years, though one of its species was independently described by
Hasskarl in 1842 as the type of his new genus Cyclandrophora. It appears that Hasskarl had
little faith in his new genus for he united it with Parinari within a year of its publication, although
it has little in common with the latter, other than the bilocular ovary.
Since Bentham (1849) nearly all species of Chrysobalanaceae with false dissepiment (and even
some without) were automatically placed in Parinari regardless of any other consideration.
As new species now placed in Exellodendron, Maranthes and Atuna were described they were
all placed in Parinari. Likewise, equally disparate elements which are now placed in Bafodeya

Prance, Hunga Pancher ex Prance and Kostermanthus Prance joined the assemblage.

KEY TO THE GENERA

based on flowering material

1. Stamens free, not united into a ligule; petals not clawed, ovary uni- or bilocular.
2. Ovary unilocular, inserted at or near base of receptacle.
3. Inflorescence a panicle of cymules; fertile stamens 7-26.

'Stamens 15-26, the filaments hairy, exserted

4. 1. Chrysobalanus

Stamens 7-10, the filaments glabrous, included

4. 2. Licania

3. Inflorescence racemose; fertile stamens 2 or 5 3. Parastemon

2. Ovary bilocular, inserted at mouth or midway up receptacle.

5. Fertile stamens 6-8(-9), markedly unilateral, the filaments equal or not exceeding the calyx lobes.

6. Lower leaf surface glabrous or lanate, with stomatal cavities; bracteoles not enclosing small groups of

flowers; inflorescence a panicle of cymules; ovary inserted midway up receptacle 4. Hunga

6. Lower leaf surface usually areolate with stomatal cavities; bracteoles enclosing small groups of flowers;

inflorescence a much-branched panicle; ovary inserted at mouth of receptacle 5. Parinari

5. Fertile stamens 10-50, usually inserted around complete circle; the filaments far exserted beyond calyx

lobes.
7. Stamens 10-25; inflorescence little branched panicles, or racemes 6. Atuna

7. Stamens c. 45; inflorescence much-branched corymbose panicles 7. Maranthes

1. Stamens united into a strap-shaped ligule; the 2 anterior petals unguiculate and enveloping the ligule in
buds; ovary unilocular 8. Kostermanthus

KEY TO THE GENERA

based on fruiting material

1 . Epicarp crustaceous-verrucose; mesocarp thick, hard, fibrous; endocarp breaking up irregularly on germi-
nation; cotyledons at least slightly ruminate.
2. Stamens free to base (can often be seen persistent around base of young fruit). Cotyledons ruminate
6. Atuna
Stamens united into a unilateral ligule. Cotyledons only slightly ruminate
2. 8. Kostermanthus
1. Epicarp smooth and glabrous or distinctly lenticellate but not crustaceous; if lenticellate then endocarp

opening by a pair of basal stoppers to allow seedling escape. Cotyledons not ruminate.
3. Epicarp lenticellate; opening by a pair of basal stoppers to allow seedling escape, always thick and woody;

fruit bilocular, but often only one loculus developing seed 5. Parinari
3. Epicarp glabrous and smooth without lenticels; opening by lateral plates, longitudinal lines or no special
mechanism of seedling escape; fruit uni- or bilocular.
4. Endocarp opening by a pair of lateral plates to allow seedling to escape; endocarp thick and woody or

thin and bony.

5. Fruit unilocular; endocarp very thin, bony 3. Parastemon
5. Fruit bilocular; endocarp thick, woody 7. Maranthes
1989] Chrysobalanaceae (Prance) 643

4. Endocarp not opening by lateral plates, usually opening longitudinally; endocarp thin and bony.
6. Fruit usually bilocular, 1.5-5 cm long, sometimes sagittate with a distinct stipe, not ridged
4. Hunga
6. Fruit unilocular, either 1-1.3 cm long, ellipsoid or 2-5 cm long and ridged.
7. Fruit ridged. Leaves orbicular 1. Chnsobalanus
7. Fruit smooth, not ridged. Leaves oblong to elliptic 2. Licania

1. CHRVSOBALANUS

Linn. Sp. PI. 1 (1753) 513; DC. Prod. 2 (1825) 525; Hook./, in Benth. &
Hook./., Gen. PI. 1 (1865) 606; Hook./ in Mart., Fl. Bras. 14 (2) (1867) 7;
Prance, Fl. Neotrop. 9 (1972) 14. - Fig. 1.
Shrubs or small trees. Stipules small, connate-axillary, caducous. Leaves
glabrous on both surfaces, without stomatal crypts. Petioles eglandular. Inflo-
rescence terminal or a.xillary cymules or a panicle of cymules. Bracts and brac-
teoles to 2 mm long, eglandular, not enclosing groups of flower buds. Flowers
hermaphrodite. Receptacle campanulate, symmetric, tomentose on exterior and
interior; calyx lobes 5, acute, equal. Petals 5, longer than calyx lobes, not
clawed. Stamens 15-26, on margin of disk; filaments hairy,
all fertile, inserted
united in groups for half length, exserted. Ovary inserted at base of receptacle,
densely pilose; carpel unilocular, with 2 ovules. Style pubescent. Fruit a small
fleshy drupe, epicarp smooth and ridged, endocarp hard, thin, glabrous on in-
terior, with 4-8 prominent longitudinal ridges which correspond to lines of
fracture for seedling escape.

D i s t r . Three species, one in West Africa and the Neotropics, two confined to the Neotropics. One species
naturalized in Malesia and Fiji.

Uses. Edible fruit. The shrub is used for the stabilization of dunes.

1. Chr>$obalanus icaco Linn. Sp. PI. (1753) 513; 1 tomentose on exterior and interior. Calyx lobes
rical,

Browne, Nat. Hist. Jamaica (1756) 250; Jacq. Sel. rounded to acute, tomentellous on both surfaces.
Stirp. Am. Hist. (1763) 155; DC. Prod. 2(1825)525; Pe/o/s white, glabrous, exserted. S/flrne'n5 15-26, the
Hook./, in Mart., Fl. Bras. 14(2) (1867) 7; Prance, filaments joined for up to half of length in small
Fl. Neotrop. 9 (1972) 15; Smith, Fl. Vit. Nov. 3 groups, densely hairy, exserted. Ovary at base of
(1985) 50. - Fig. 1. receptacle, pilose. Fruit ovate to obovate, 2-5 cm
Shrub or small tree to 5 m tall, the branches gla- long; epicarp smooth with longitudinal ridges;
brous and lenticellate. S//pM/« 1-3 mm long, cadu- mesocarp thin and fleshy; endocarp thin, hard,
cous. Leaves orbicular to ovate-elliptic, 2-8 by 2-6 ridged on exterior.
cm, retuse, rounded or with short blunt acumen at Distr. Neotropics, mainly in coastal areas; West
apex, subcuneate at base, glabrous on both surfaces; & Central Africa, naturalized in Fiji, cultivated in

petioles 2-4 mm. Inflorescences small terminal and Vietnam; in Malesia cultivated in Singapore where it

axillary cymules or panicles of cymules, the rachis has escaped and naturalized. Fig. ID.
and branches grey-puberulous. Flowers 4-6 mm Ecol. Dunes, beaches and coastal scrub,
long. /?ecep/flc/^ campanulatc-cupuliform, symmct- Uses. Edible fruit.
644 Flora Malesiana [ser. I, vol. 10^

2mm

1 cm

Fig. 1. Chrysobalanus icaco Linn. A. Detail of flower; B. habit; C. fruit; D. distribution in Malesia.
1989] Chrysobalanaceae (Prance) 645

Fig. 2. Licania splendens (KoRTH.) Prance. A. Habit, xO.5; B. flower, x9; C. flower section, x9; D. fruit,

X 1 (A-C Elmer 20916, D Ramos & Convocar 83437).

2. LICANIA
AuBL. Hist. PI. Guiane Fr. 1 (1775) 1 19, t. 45; DC. Prod. 2 (1825) 527; Hook.
/. in Benth. & Hook./. Gen. PI. 1 Ann. Naturh. Mus. Wien
(1865) 606; Fritsch,
4 (1889) 33; Focke in E. & P. Nat. Pn. Fam. 3, 3 (1891) 58; Prance, F1.
Neotrop. 9 (1972) 21; Prance & Whitm. Tree Fl. Malaya 2 (1973) 328; White,
Bull. Jard. Bot. Nat. Belg. 46 (1976) 280; Prance, Brittonia 31 (1979) 94.
-
Moquilea Aubl. Hist. PI. Guiane Fr. (1775) 521, t. 208; DC. Prod.
1 2 (1825)
526; Hook./, in Benth. & Hook./, Gen. PI. 1 (1865) 606; Focke in E. & P.
Nat. PH. Fam. 3, 3 (1891) 58. - Dahuronia Scop. Introd. (1777) 217, nom. illeg.
- Hedycrea Schreb. in Linn., Gen. PI. ed. 8, (1789) 160, nom. illeg. - 1

Angelesia Korth. Ned. Kruidk. Arch. 3 (1854) 384; Boerl. Handl. Fl. Ned.
Ind. (1890) 424; Burk. Diet. (1935) 159; Corner, Wayside Trees (1940) 526;
1

Hutch. Gen. Flow. PI. (1964) 191. - Trichocarya Miq. Fl. Ind. Bat. 1,
1 1

(1855) 358; ibid. 6 (1858) 1084, p.p. quoad T. splendens tantum. - Geobalanus
Small, Fl. Miami (1913) 80; Hutch. Gen. Flow. PI. (1964) 191. - Coccomelia 1

Ridley, J. Str. Br. Roy. As. Soc. n. 82 (1920) 183; Fl. Mal. Pen. 1 (1922)
671. - AfrolicaniaMiLDBK. Notizbl. Bot. Gart. Bcrlin-Dahlem 8 (1921) 483. -
Fig. 2.
Small to large trees. Stipules small, free, caducous. L^crv^^ glabrous on both
surfaces, without stomatal crypts. Petioles eglandular. Inflorescence a panicle
of cymules. Bracts and bracteoles to 1.5 mm
long, membraneous, eglandular,
not enclosing groups of flower buds. Flowers hermaphrodite. Receptacle cam-
panulate, slightly asymmetric, tomcntose on exterior, tomentosc within; calyx
646 Flora Malesiana [ser. I, vol. 10^

lobes 5, acute, unequal. Petals 5, small, not exceeding the calyx lobes, not
clawed. Stamens 7-10, all fertile, inserted on margin of disk; filaments gla-
brous, included, slightly united at base. Ovary inserted at or near base of recep-
tacle, piloseon exterior; carpel unilocular, with 2 ovules. Style pubescent at
base, the stigma capitate. Fruit a small, fleshy drupe,narrowed to a shortly stip-
itate base; epicarp smooth, not ridged, glabrous, not lenticellate; mesocarp thin,
fleshy; endocarp thin, hard, bony, breaking up in longitudinal lines during ger-
mination, tomentose within.

Dist r About 180 species in the Neotropics, one species in West Africa; three species in Malesia from the
.

Malay Peninsula to New Guinea and the Philippines, but not in the Lesser Sunda Islands.
Uses. The timber is strong and durable and resistant to marine borers. It is hard to work because of silica.
Note. The description above is for the Malesian element of Licania; the genus is much more variable in
the Neotropics. The three Asian species are placed in subgenus Angelesia by Prance & White, Phil. Trans.
Roy. Soc. London 320 (1988) 94.

KEY TO THE SPECIES

1. Fruit 1-1.3 cm long, ellipsoid, not narrowed towards base or apex.

2. Leaves oblong, the apices distinctly acuminate; inflorescence rachis densely puberulousl. L. splendens
2. Leaves elliptic to oblong-elliptic, the apices acute to rounded and emarginate; inflorescence rachis sparsely
puberulous 2. L. palawanensis
1 . Fruit 2.5-5 cm long, narrowed at apex and base, fusiform 3. L. fusicarpa

1. Licania splendens (Korth.) Prance, FI. Neotrop. unilocular, pilose on exterior. Fruit ellipsoid, 1-1.3
9 (1972) 172. - Angelesia splendens Korth. Ned. cm long; epicarp smooth, glabrous; mesocarp thin,
Kruidk. Arch. 3 (1854) 384; Boerl. & Koord. Ic. fleshy; endocarp thin, hard, bony, breaking open by
Bog. 1, 4 (1901) 59, t. 96; Merr. Philip. J. Sc. 10 longitudinal lines of weakness, tomentose within.
(1915) Bot. 307; Enum. Philip. PI. 2 (1923) 236; Cor- Distr. Thailand; in Malesia: Sumatra, Malay
ner, Wayside Trees (1940) 526; Browne, For. Trees Peninsula, W. Java, Borneo, Philippines. Fig. 3.
Sarawak & Brunei (1955) 307. - Licania angelesia Ecol. Commonest in forest, including diptero-
Blume, Melang. Bot. 2 (1855) 358. - Chrysobalanus carp forest, on hill slopes and ridges, but wide-rang-
splendens Korth. ex Miq. F1. Ind. Bat. 1, 1 (1855) ing in peat swamp, freshwater swamp forest, on sea-
358, in syn. - Parinarium fragile Teijsm. & Binn. shores, and in rocky places; 0-400(-800) m altitude.
Cat. Hort. Bog. (1866) 253, nom. nud. - Parinarium Uses. The timber is strong, durable and resistant
nitidum Hook. /. Fl. Brit. India 2 (1878) 310. - to marine borers and is used for saltwater piles, rail-
Ferolia nitida {Hook, f.) Ridley, J. Str. Br. Roy. As. road ties, etc. However, it is extremely hard to work
Soc. n. 82 (1920) 183; Fl. Mai. Pen. 1 (1922) 671. - and requires special tools because of silica. The fruit
Parinarium philippinense Elmer, Leafl. Philip. Bot. is edible but is not widely used.
10(1939)3809. - Fig. 2. Vern. Malay Peninsula: champrai, medang
Tree to 25 m tall, the young branches sparsely Ian- merah, m. puteh, membatu, mempadang, merbatu
ate,soon glabrous. Stipules linear-lanceolate, to 3 kechil; Borneo: piasau-piasau, Kedayan, gandulong,
mm long, caducous. Leaves 4-11 by 1.8-4.2 cm, Dusun, tampaluan, Sabah, sampaluan, Brunei,
oblong, usually acuminate at apex, cuneate at base, buku-buku, bunga, djentihan burung, mauhi,
glabrous beneath; petioles 2-5 mm, canaliculate, Kalimantan; Philippines: taguilom bay; amayan, ba-
glabrous when mature. Inflorescence terminal and lik, D.Bis., dagingan, dagingdingan, S.L.Bis.,
axillary panicles of cymules, 1.5-14 cm long, the gapas, maralibus, Tagb.
rachis and branches grey-puberulous. Flowers c. 2
mm long. Receptacle campanulate, slightly swollen
to one side, grey-tomentellous on exterior, tomen- 2. Licania palawanensis Prance, Brittonia 31 (1979)
tose within; pedicels c. 1 mm long. Calyx lobes acute, 94.
tomentellous on both surfaces. Petals pubescent on Shrub, young branches sparsely puberulous soon
exterior.Stamens 7-10, slightly unilateral, the fila- becoming glabrous. Stipules lanceolate, 1-2 mm
ments glabrous. Ovary at or near base of receptacle. long, glabrous, caducous. Leaves 3-6 by 1.4-3 cm.
1989] Chrysobalanaceae (Prance) 647

Fig. 3. Distribution of Licania splendens (Korth.) Prance (dots) and L. palawanensis Prance (triangles).

elliptic to oblong elliptic, rounded to acute at apex, Tree to 7 m tall, young branches puberulous, gla-

emarginate, subcuneate at base, glabrous beneath; brescent, with small prominent round lenticels. Stip-
petioles 1-3 mm long, c. 1.5 mm wide, lanate be- ules not seen. Leaves 5-10 by 1-4.5 cm, charta-
coming glabrous with age, rugose. Inflorescences ceous, oblong to elliptic, acute to bluntly acuminate
panicles of cymules, 3-4 cm long, the rachis and at apex, cuneate at base, glabrous and glossy on both
branches sparsely puberulous. Flowers c. 2 long. mm surfaces, decurrent onto petiole; petioles 2-3 mm
Receptacle campanulate, slightly swollen to one side, long, rugose, puberulous becoming glabrous with
grey-tomentellous on exterior, tomentose within; age. Inflorescences terminal and axillary panicles of
pedicels c. 1 mm long. Calyx lobes acute, tomentel- cymules, few-flowered, the rachis and branches
lous on exterior, puberulous within. Petals puberu- sparsely puberulous. Flowers c. 2 mm long. Recep-

lous on exterior. Stamens around com-

7, inserted taclecampanulate, grey-tomentellous on exterior,
plete circle, the filaments glabrous. Ovary inserted at tomentose within; pedicels 2-3 long. Calyx mm
base of receptacle, lanate-pilose, unilocular. Fruit lobes acute, narrow, tomentellous on exterior, pu-
(immature) epicarp smooth, glabrous;
ellipsoid, berulous within. Petals not seen. Stamens persistent
mcsocarp endocarp thin, hard, bony, tomen-
thin; beneath young fruit, 0.5-1 mm
long, connate at
tose within, breaking open by longitudinal lines of base. Fruit (2.5-)3-5 cm long, narrowly spindle-
weakness. shaped, narrowed at apex to a tip 2-3 mm long, nar-

Distr. Malesia: Philippines (Palawan). Fig. 3. rowed at base in stipe 5-10 mm long; epicarp
Ecol. Confined to ultrabasic rock formation; smooth, glabrous; mesocarp thin; endocarp hard,
0-3(X) m altitude, including sea-shore forest. bony, c. i mm thick, densely lanate within, without
lines of dehiscence.
V I.icania fuMcarpa (KosTkRM.) Prance, Brittonia Dislr. Malesia: E. Papua New Guinea (Milne
19 (1987) 3W). Hunga fusicarpa Kostfrm. Rcin- Bay Prov., [-erguson I., Morobc Prov.). Fig. 5.
wardtia 10(1985) 123. Ecol. Coastal rain-forest, 0-300 m altitude.
648 Flora Malesiana [ser. I, vol. 10^

F\g. 4. Paraslemon urophyllus (V^ ALL. ex A.DC.) A.DC. A. Hab'n, x 0.5; B. flower, x 10; C. fruit, xl;D.
flower section, x 10 (A, B Sinclair 39504, C, D Sinclair 3319).

3. PARASTEMON

A.DC. Ann. Sci. Nat. Bot. (1842) 208; Miq. F1. Ind. Bat. 1, 1 (1855)
ser. 2, 18

359; Hook./, in Benth. & Hook./., Gen. PI. 1 (1865) 607; Fl. Brit. India 2 (1878)
312; BoERL. Handl. Fl. Ned. Ind. 1 (1890) 426; Focke in E. & P. Nat. Pfl. Fam.
3, 3 (1891) 60; Merr. Philip. J. Sc. 10 (1915) Bot. 307; Ridley, Fl. Mai. Pen.
1 (1922) 672; Merr. & Perry, J. Arn. Arb. 21 (1940) 197; Corner, Wayside
Trees (1940) 526; Hutch. Gen. Flow. PI. 1 (1964) 193. - Diemenia Korth.
Ned. Kruidk. Arch. 3 (1854) 388; Boerl. Handl. Fl. Ned. Ind. 1 (1890) 425. -
Trichocarya Miq. Fl. Ind. Bat. 1, 1 (1855) 357, p.p. - Fig. 4.
Tree or shrub. Stipules small and triangular, caducous. Leaves glabrous on
both surfaces, without stomatal cavities, with 2 small discoid glands at base of
lamina; petioles eglandular. Inflorescence an axillary or rarely terminal simple
or sparsely branched raceme. Bracts and bracteoles small, eglandular, not en-
closing groups of flower buds. Flowers hermaphrodite or polygamo-dioecious.
Receptacle patelliform or shallowly cupuliform, shortly hairy within; calyx
lobes 5, acute, subequal. Petals 5, not exceeding calyx lobes, not clawed. Sta-
mens either 5 and all fertile or 2 fertile with 3 staminodes; the filaments
glabrous, shorter than the calyx lobes. Ovary centrally inserted at base of recep-
1989] Chrysobalanaceae (Prance) 649

tacle, glabrous or densely hairy on exterior; carpel unilocular, with 2 ovules.

Style filiform, puberulous towards the base, with 3 large undivided lobes at apex
or 1 obscure lobe and 2 large, sometimes deeply divided lobes. Fruit a small
drupe to c. 1.5 cm or c. 3 cm long; epicarp smooth, not lenticellate; endocarp
thin, hard, bony, smooth on exterior, glabrous within; with 2 large lateral plates
which break away on germination to allow seedling escape.

Distr. Three species; Nicobar Islands; in Malesia: Malay Peninsula, Sumatra, Borneo, Moluccas, New
Guinea, Admiralty Is.

KEY TO THE SPECIES

1. Fruit 1-1.5 mm long. Primary veins of leaves 8-11 pairs.

2. Receptacle shallowly cup-shaped, glabrous on exterior; fertile stamens 2; style 3-lobed at apex. Inflores-
cence glabrous 1. P. urophyllus
2. Receptacle saucer-shaped, puberulous on exterior; fertile stamens 5; style with one obscure and two large
bifid apical lobes. Inflorescence villous 2. P. versleeghii

1. Fruit 2.3-3.5 mm long. Primary veins of leaves 5-6 pairs 3. P. grandifnictus

1. Paraslemon urophyllus (Wall, ex A. DC.) A. DC. thin, hard; endocarp thin, hard, bony, glabrous
Ann. Sci. Nat. Bot. ser. 2, 18 (1842) 208; MiQ. Fl. within, opening by 2 lateral plates.
Ind. Bat. 1, (1855) 359; Boerl. & Koord. Ic. Bog.
1 Distr. Nicobar Islands; in Malesia: Malay Penin-
1, 4 (1901) 61, t. 97; Ridley, Fl. Mai. Pen. 1 (1922) sula, Sumatra, Borneo. Fig. 5.
672; BuRK. Diet. (1935) 1693; Corner, Wayside Ecol. Characteristic of peat swamp forest where
Trees (1940) 526; Browne, For. Trees Sarawak & it is a common large tree, but wide ranging into

Brunei (1955) 308; Kochum. & Wyatt-Smith, Mai. shorter, more open scrub forest.
For. Rec. 17 (1964); Prance & Whitm. Tree Fl. Uses. The wood is hard to use because of the
.Malaya 2 (1973) 331. - Embelia urophyl/a [Wall. silica content, but it is used locally for general con-
Cat. (1830) n. 2309, nom. nud.] ex A. DC. Trans. struction, posts, and as firewood.
Linn. Soc. 17 (1837) 131. - Diemenia racemosa Vern Malay
. Peninsula: kelat, k. pasir, k. puteh,
(KoRTH.) MiQ. Fl. Ind. Bat. 1, 1 (1855) 358. - nylas; Sumatra: galam tabanga, kayu gelang, malas,
Licania diemenia Blu.me, Melang. Bot. 2 (1855) 10; meriawak; Borneo: mandailas, Brunei, Besaya, sem-
Hassk. Flora 41 (1858) 256, nom. illeg. - palawan, Brunei, tempalawan, Bajau, mengilas,
Paraslemon spicatus Ridley, J. Str. Br. Roy. As. ngilas padang, obah, Sarawak.
Soc. n. 75 (1917) 29. - Fig. 4. Notes. The only record of this species from Java
Tree to 40 m tall, or shrub, the young branches (Blume s.n., L) is very dubious since the collector's
glabrous, the trunk often buttressed. Stipules name was added later. It is probably either misla-
triangular, c. 1 mm long, caducous. Leaves thinly co- belled or from cultivated material. The only dif-
riaceous, narrowly oblong, 2.5-8 by 1.4-2.5 cm, ference given between P. spicatus and P. urophyllus
cuspidate acuminate at apex, the tip 5-15 mm, is that the former is a shrub with sessile flowers.
cuneaic at base; midrib plane above, prominulous Some forms of P. urophyllus have extremely short
beneath; primary veins 8- 1 1 pairs; petioles 4-5 mm pedicels and most sessile-flowered individuals are
long, canaliculate, glabrous. Inflorescence of ax- recorded as being small trees. There is thus no reason
illary and rarely terminal racemes or occasionally to maintain P. spicatus as a distinct species.
slightly branched, 4- 14 cm long, the rachis glabrous.
Flowers polygamo-dioecious, c. 1 .5 mm long. Recep- 2. Paraslemon versleeghii Merr. & Pi:rrv, J. Arn.
tacle broadlycupuliform to flattened sauccr-shapcd, Arb. 21 (1940) 197.
glabrous on cxicrior, lomcntosc within; pedicels up Tree to 40 m tall, the young branches sparsely
to 2 mm long. Calyx lobes acute, glabrous
on exte- puberulous, soon glabrous. Stipules triangular, c. I
rior. Petals 5.Stamens 2 fertile and 3 sterile stami- mm long, caducous. Leaves thinly coriaceous, nar-
nodcs opposite. Ovary inserted at base of receptacle, rowly oblong. 5-9.5 by 1.8-3.7 cm, cuspidate acu-
pilose on exterior, unilocular. Style pilose at base, minate at apex, the tip 7-15 long, cuncaie at mm
glabrous above, the stigma trifid. fruit ellipsoid, base; midrib plane above, prominulous beneath;
1-1.5 cm long; epicarp smooth, glabrous; mcsocarp primary veins 8-12 pairs, inconspicuous, slightly
650 Flora Malesiana [ser. I, vol. 104

"'^^

(Kosterm.) Prance (diamond), Parastemon urophyllus (Wall, ex

Fig. 5. Distribution of Licania fusicarpa
A. DC.) A. DC. (dots), P. grandifructus Prance (squares), and P. versteeghii Merr. & Perry (triangles).

prominulous beneath; petioles 3-7 mm

long, terete, 3. Parastemon grandifructus Prance, Brittonia 39
glabrescent. Inflorescence of axillary and terminal (1987) 366.
racemes, 2-9 cm long, the rachis sparsely villous. Tree to 30 m tall, the young branches glabrous, the
Flowers hermaphrodite, c. 1.5 mm
long; pedicels trunk lightly buttressed to1 m. Stipules caducous
1.5-3 mm, sparsely villous. Receptacle broadly (not Leaves coriaceous, narrowly oblong,
seen).
cupuliform-flattened, sparsely villous on exterior, 5-8.5 by 1.8-3.2 cm, with long cuspidate acumen at
tomentose within. Calyx lobes acute, with hirsute apex, the tip 10-16 mm long, cuneate at base,
margins. Petals 5, with hirsute margins. Stamens 5, glabrous on both surfaces; midrib prominent above,
all fertile, opposite the petals in a complete circle. prominulous or plane beneath; primary veins 5-6
Ovary inserted at base of receptacle, glabrous on ex- pairs,prominulous above, plane beneath; petioles
terior except at base, unilocular. Style pilose at base, 5-8 mm long, glabrous, slightly canaliculate, slight-
glabrous above, the stigma with two large apical ly swollen at base. Inflorescence of axillary and ter-
lobes, the third reduced or missing. Fruit ellipsoid, c. minal racemes, the rachis glabrous. Flowers seen on-
1.6 cm long; epicarp smooth, glabrous; mesocarp ly in fruiting specimens. Calyx lobes 5, acute,
thin, hard; endocarp thin, hard, bony, glabrous glabrous on exterior, glabrous within except for a
within, opening by two lateral plates. few hairs around base. Receptacle glabrous on ex-
Distr. Malesia: Moluccas (Morotai), New below
terior in fruiting condition. Style persistent
Guinea, and Admiralty Is. Fig. 5. fruits, the stigma bifid or trifid. Fruit ellipsoid,
E c o Usually in well-drained hill forest up to 700 m
1 . 2.3-3.5 by 1.3-1.5 cm, epicarp smooth, glabrous;
altitude, but also reported from secondary forest, mesocarp thin, 0.25 mm; endocarp thin, hard, bony,
swampy (Campnosperma) forestand even beach 0.25 mm thick, glabrous within, opening by 2 lateral
forest. plates 1.9-2 cm long.
Vern. New Guinea: mangu, Tobelo, noeng, Distr. Malesia: Borneo (Sarawak, Sabah). Fig.
Irian, gorsauw. Tor, gwarsau, Wainlag, sirebo, 5.
sisero, Kemtoek, sosopi, Japen, telek, Mooi, sino- Ecol. Upland white sand areas.
ree, Biak. Vern. Borneo: ngilas, Ihan, praus, Dyak.

4. HUNGA
Pancher e-v Prance, Brittonia 31 (1979) 79; FI. Nouv. Caled. et Dep. 12 (1983)
106. - Fig. 6, 8.
Shrubs or small trees. Stipules lanceolate and persistent (absent or very early
1989] Chrysobalanaceae (Prance) 651

caducous in New Caledonian species). Leaves usually glabrous on both surfaces

(lanate beneath in some New Caledonian species), with a pair of, often obscure,
marginal glands towards the base, without stomatal cavities; petioles eglan-
dular. Inflorescence a few-flowered terminal or axially raceme of cymules.
Bracts and bracteoles small, persistent, not enclosing the flowers in small
groups. Flowers hermaphrodite, slightly zygomorphic. Receptacle campanu-
late, slightly asymmetric, shortly puberulous on exterior, densely pubescent
within. Calyx lobes 5, acute. Petals 5, small, not exceeding calyx lobes, not
clawed. Stamens 5-9, not exceeding calyx lobes, unilateral with 3-7 stami-
nodes opposite. Ovary inserted midway up receptacle, densely hairy on exterior;
carpel bilocular with one ovule in each loculus. Style truncate but distinctly 3-
lobed at apex. Fruit small, fleshy, bilocular or often with one loculus underde-
veloped; epicarp smooth, not ridged, not lenticellate; mesocarp thin, fleshy; en-
docarp thin, hard, bony, with a smooth surface, interior very hairy, with 4-6
longitudinal lines of weakness which allow the seedling to escape.

Distr . There are 1 1 species, 8 of which occur in New Caledonia and the Loyalty Is., 3 in Malesia: Papua
New Guinea.

KEY TO THE SPECIES

1. Inflorescence branches glabrescent; flowers glabrescent on exterior. Leaves with conspicuous anastomos-
ing venation, oblong-elliptic to elliptic, 4-8.5 cm broad.
2. Leaves elliptic, 7.5-8.5 cm broad 1. H. novoguineensis
2. Leaves oblong-elliptic, 4-6.5 cm broad 2. H. papuana
1 . Inflorescence branches lanate to puberulous; flowers pubescent on exterior. Leaf with venation not con-
spicuously anastomosing, oblong-lanceolate, 2-3.7 cm broad 3. H. longifolia

1. Hunga novoguineensis Prance, Brittonia 31 2. Hunga papuana (Baker/.) Prance, Brittonia 31

(1979) 88. - Fig. 6 G, H. (1979) 88. - A ngelesia papuana Baker/. J. Bot. 61,
Tree 4 m tall, the young branches puberulous, Suppl. (1923) 13. - Fig. 6 A-F.
soon glabrous, lenticellate. Stipules lanceolate, pu- Small tree, the young branches lanate, soon gla-
berulous, c. 5 mm long, persistent. Leaves coria- brous. Stipules lanceolate, 3-6 mm long, puberu-
ceous, elliptic, 15-19 by 7.5-8.5 cm, glabrous on lous, persistent.Leaves coriaceous, oblong-elliptic,
both surfaces, apex acuminate, the acumen 8-10 10-19 by 4-6.5 cm, finely acuminate at apex, the tip
mm long, subcuneate at base; primary veins 11-14 8-16 mm, rounded to subcuneate at base, glabrous
pairs,anastomosing 4 mm away from margins, pro- on both surfaces; primary veins 9-13 pairs, promin-
minulous above, prominent beneath; petioles 5-6 ulous above, prominent beneath, conspicuously ana-
mm long, puberulous soon becoming glabrous, stomosing 5 mm from margin; petioles 2-4 mm
slightly canaliculate, eglandular. Inflorescence of long, shallowly canaliculate, sparsely puberulous
terminal and axillary panicles, the rachis and bran- when young, soon glabrescent, rugose. Inflores-
ches puberulous. Bracts and bracteoles 1-2 mm cences of terminal and axillary panicles, 3-10 cm
long, puberulous, persistent. /^/ow^ri not seen. Frw;7 long, the rachis and branches sparsely lanate-
sagiltalc pyriform, c. 3.5 cm long, the upper part puberulous when young. Bracts and bracteoles
triangular,2-2.5 cm long, the base with a stipe 6- 10 0.5-2 mm long, sparsely puberulous-glabresccnt on
mm long; epicarp glabrous, smooth, mesocarp thin, both surfaces. I'lowers 1.5-2 mm long. Receptacle
fleshy; endocarp thin, hard, bony, lanale within. campanulatc, glabrous externally, tomentosc within.
Pistr. Malesia: Papua New Cjuinca, two collcc- Calyx lobes glabrous on both faces except for cilio-
tions, from Morobc and Milne Bay Prov. Fig. 7. late margins. Petals glabrous. Stamens c. 7,
Ecol. Oak forest on slopes, at c. 800 m altitude. unilateral with toothed staminodcs opposite. Ovary
 Flora Malesiana [ser. I, vol. 10^
652

Fig. 6. Hunga papuana (Baker f.) Prance. A. Habit, xO.5; B. flower, x7; C. flower section, x9; D. petal,
X xO.5; F. ovary section, x 15. - H. novoguineensis Prance. G. Young fruit section, x 1; H.
15; E. fruit,
habit, xO.5 (A-C Forbes 504, £)-F Womersley NGF 19307, G, H Hartley 12645).
1989] Chrysobalanaceae (Prance) 653
654 Flora Malesiana [ser. I, vol. 10^

age, slightly rugose. Inflorescences axillary and ter- forciliate margins. Stamens6-8, unilateral with 3-5
minal panicles of cymules 1.5-6 cm long, the rachis short staminodes opposite them. Ovary bilocular, in-

and branches appressed lanate when young, becom- sertedmidway up receptacle tube, pilose on exterior.
ing puberulous. Bracts and bracteoles 1-3.5 mm S/^e pilose at base. Fru/7 not seen.
long, sparsely puberulous-giabrescent on both sur- Distr. Malesia: Papua New Guinea (Misima I.),
faces. Flowers 2-2.5 mm long. Receptacle cam- known from a single collection. Fig. 7.
panulate, swollen slightly to one side, lanate-tomen- Ecol Rain-forest on N. slope, at 300 m altitude.
.

tose on exterior, tomentose within. Calyx lobes Fl. July.

pubescent on both surfaces. Petals glabrous except

5. PARINARI

AuBL. Hist. PI. Guiane Fr. 1 Hauman, Bull. Jard. Bot. Brux. 21
(1775) 204;
(1951) 184, quoad subg. Euparinari tantum; Backer & Bakh./. Fl. Java 1
(1964) 521, p.p.; Hutch. Gen. Flow. PI. 1 (1964) 192, p.p. excl. syn. Maranthes
etc.; KosTERM. Reinwardtia 7 (1965) 7, excl. syn. Thelira, Ferolia, Mampata et
Neou; Prance, Fl. Neotrop. 9 (1972) 178; Prance & Whitm. Tree Fl. Malaya
2 (1973) 332; White, Bull. Jard. Bot. Nat. Belg. 46 (1976) 310; Distr. PI. Afr.
10 (1976) 327; Fl. Zamb. 4 (1978) 36; Prance, Fl. Venez. 4 (1982) 325; Smith,
Fl. Vit. Nov. 3 (1985) 44. - Dugortia Scop. Introd. (1777) 217, nom. illeg. -
Pahnarium Juss. Gen. PI. (1789) 342; Lamk, Encycl. Meth. Bot. 5 (1804) 17;
St.Hil. Expos. Fam. 2 (1804) 194, p.p.; R.Br, in Tuckey, Nar. Exped. Riv.
Zaire Cong. (1818) 433; Steud. Nom. (1821) 591; DC. Prod. 2 (1825) 526; Pom.
Diet. Sci. 37 (1825) 544; Bartl. Ord. Nat. (1830) 406; G.Don, Gen. Syst. 2
(1832) 478; Meissn. Gen. (1836/42) 102; Benth. Hook. J. Bot. 2 (1840) 211,
218; Endl. Gen. (1840) 1252, n. 6411; Benth. in Hook., Niger Fl. (1849) 333;
MiQ. Stirp. Surin. Select. 2 (1850) 7; Blume, Mus. Bot. Lugd.-Bat. 2 (1852) 94;
Melang. Bot. 2 (1855) 10; Miq. Fl. Ind. Bat. 1, 1 (1855) 352; ibid. (1858) 1084;
C.MuELL. in Walp., Ann. 4 (1857) 644; Miq. Suppl. Sumatra (1860) 306;
Benth. Fl. Austr. 2 (1864) 426; Hook. /. in Benth. & Hook./., Gen. PI. 1
(1865) 607; Miq. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 237; Hook./, in Mart.,
Fl. Bras. 14 (2) (1867) 49; Baill. Hist. PI. 2 (1869) 435, 482; Kurz, For. Fl. Bur-
ma 1 (1877) 432; Hook./. Fl. Brit. India 2 (1878) 308; Fritsch, Ann. Naturh.
Mus. Wien 4 (1889) 33; Boerl. Handl. Fl. Ned. Ind. 1 (1890) 421, 424; Focke
in E.& P. Nat. Pfl. Fam. 3, 3 (1891) 60; King, J. As. Soc. Beng. 66 (1897) 276;
K. & V. Bijdr. 5 (1900) 332; Bailey, Queensl. Fl. 2 (1900) 524; Brandis, Indian
Trees (1906) 278; Backer, Schoolfl. Java 1 (1911) 445; Ridley, Fl. Mai. Pen.
1 (1922) 666;Merr. Enum. Philip. Flow. PI. 2 (1923) 235; Burk. Diet. (1935)
1693; Corner, Wayside Trees (1940) 527. - Petrocarya Schreb. in Linn. Gen.
PI. ed. 8, 1 (1789) 245, nom. superfl. - Pahnarium sect. Petrocarya DC. Prod.
2 (1825) 526; Benth. in Hook., Niger Fl. (1849) 335, p.p. excl. P. glaberrima
et P. scabra. - Parinarium sect. Neocarya DC. Prod. 2 (1825) 526, p.p. guoadl
P. excelsum. - Balantium Desv. ^atBuch.-Ham. Prod. PI. Ind. Occ. (1825) 34.
- Parinarium subg. Petrocarya (DC.) Miq. Fl. Ind. Bat. 1, 1 (1855) 352. -,
1989] Chrysobalanaceae (Prance) 655

Lepidocarpa Korth. Ned. Kruidk. Arch. 3 (1854) 385. - Ferolia O.Kuntze,

Rev. Gen. PI. 1 (1891) 216, p.p. (non Ferolia Aubl.). - Fig. 15.
Small or large trees or rarely shrubs. Stipules small to large, persistent or ca-
ducous. Leaves usually with stomatal crypts filled with pubescence on lower sur-
face or rarely glabrous, or lanate pubescent without crypts. Petioles usually with
2 circular glands above. Inflorescence a many-flowered complex cyme or cy-
mose and bracteoles eglandular, usually concealing flower buds
panicle. Bracts
individually and in small groups. Flowers hermaphrodite. Receptacle subcam-
pulate to cupuliform, slightly swollen to one side, tomentose on both surfaces;
calyx lobes 5, deltate, acute, densely hairy on both surfaces. Petals 5, as long
as or shorter than sepals, caducous. Stamens 6-10, unilateral, the filaments
glabrous, included, with c. 6 minute staminodes opposite. Ovary inserted on up-
per half of receptacle tube below mouth, pilose on exterior; carpel bilocular with
1 ovule in each loculus; style arcuate, included. Fruit a fleshy drupe; epicarp ver-
rucose; endocarp thick, with a rough fibrous surface, with 2 basal obturators
for seedling escape.

Disir. Pantropical with 18 species in the Neotropics, 6 in Africa and 15 in tropical Asia (P. anamensis),
Malesia, the Pacific region (P. insularam) and northern Queensland, Australia; in Malesia 13 species.
Uses. The fruit of several species are edible, but little-used.
Note. Since inflorescences and flowers are uniform in the Malesian region, the species are difficult to
separate; a key containing all 15 Australasian species, based on leaf characters only, is given here.

KEY TO THE SPECIES

1 . Stomatal crypts absent from leaf underside; leaf underside glabrous or with a persistent lanate pubescence
and then with large persistent stipules 7-40 mm.
2. Leaf undersurface glabrous. Stipules small and caducous.

3. Leaves elliptic to oblong or obovate-elliptic, 9.5-20.5 by 4.5-8.5 cm; primary veins 11-16 pairs. Pan-

icles large and silvery pubescent 1. P. argenleo-sericea

3. Leaves ovate, 5-9 by 2-4.5 cm; primary veins 7-11 pairs. Panicles small and subsericeous brown

pubescent 2. P. canarioides

2. Leaf undersurface densely lanate pubescent, but when removed no stomatal crypts present; stipules large
and persistent, 7-40 mm
long, 3-5 cm broad at base.
4. Leaves oblong-lanceolate, 5-18 cm long on flowering branches, thickly coriaceous, base cuneate

3. P. elmeri

4. Leaves elliptic to oblong, 1 1 -28 cm long, chartaceous, base rounded 4. P. parva

1. Stomatal crypts present on leaf underside; leaf undersurface lanate or at least pubescent in crypts; stipules
usually small, or if larger then early caducous.
5. Leaf lower surface with a series of small glands along lower part of margin. Calyx broadly cupuliform.
6. Leaf apex acute or rounded but not acuminate; primary veins 10-17 pairs. Young branches with small,
almost plane icnticels. Low tree of savanna, savanna forest and forest margins 5. P. nonda

6. Leaf apex acuminate, acumen 3- 10 mm long; primary veins 16-22 pairs; young branches with extreme-
prominent large lenticels; large tree of rain-forest and hills
ly 6. P. papuana
5. Leaf lower surface without marginal glands on lower part. Calyx usually campanulate.

7. Primary leaf veins 20-33 pairs (16-26 pairs in P. costata ssp. polyneura).

8. Petioles 9-17 mm long 7. P. oblonttifolia

8. Petioles 3-6 mm long.
9. Leaves chartaceous, primary veins prominulous above, 16-26 pairs
13c. P. costata ssp. polyneura
9. Leaves coriaceous, primary veins impressed for upper portion, 20-28 pairs 8. P. gigantea
656 Flora Malesiana [ser. I, vol. 10^

7. Primary leaf veins 20 pairs or fewer.

10. Petioles 10-20 mm
long; leaves with or without a metallic sheen above.
II. Leaves with metallic sheen above; petioles 14-20 mm
long; leaves 4-9 cm broad. Borneo
9. P. metallica

11. Leaves without metallic sheen; petioles 10-12 mm long; leaves 6.5-12 cm broad. New Guinea
10. P. prancei

10. Petioles 3-10 mm long; leaves without metallic sheen.

12. Leaves rigidly coriaceous, often broadest well below mid point; midrib and often primary veins
impressed on upper surface
lightly 11. P. rigida

12. Leaves thinly coriaceous or chartaceous, usually broadest at or above middle (except in P. in-
sularum); midrib and primary veins usually plane or prominulous.
13. Leaves ovate, with long thin acumen, tapering from well below mid point; midrib impressed above.
Plants of Pacific Islands (Fiji, Tonga, Samoa, Wallis Is.) P. insularum
13. Leaves elliptic to oblong-lanceolate, tapering from middle or above; midrib usually plane or pro-
minulous. Plants of Sunda shelf.
14. Inflorescence predominantly axillary. Leaves broadly elliptic 12. P. sumatrana
14. Inflorescence terminal and subterminal. Leaves elliptic to narrowly oblong.
15. Leaf apex rounded to acute. Thailand and Indochina P. anamensis
15. Leaf apex acuminate. Burma, Malay Peninsula, Indonesia and the Philippines 13. P. costata

1. Parinari argenteo-sericea Kosterm. Reinwardtia 7

(1965)47, f. 1; 158.
Trees to 35 m tall; the young branches glabrous,
prominently lenticellate. SZ/pw/e^ lanceolate, to 8 mm
long, tomentose on exterior, early caducous. Leaves
chartaceous, oblong, elliptic to subovate-elliptic,
9.5-20.5 by 4.5-8.5 cm, glabrous on both surfaces,
without stomatal cavities beneath, usually 2 glands
beneath at base near junction with midrib, acute to
shortly acuminate at apex, the tip 7-10 mm long,
rounded at base; midrib lightly impressed above ex-
cept near base, prominent beneath; primary veins
11-16 pairs, plane above, prominent beneath, erect-
patent; petioles 5-9 mm long, eglandular, glabrous,
rugulose. Inflorescence a lax, much branched, ter-
Fig. 9. Distribution of Parinari argenteo-sericea
minal panicle 9-15 cm and branches
long, the rachis
Kosterm. (stars) and P. canarioides Kosterm.
densely grey sericeous-tomentose; bracts and brac-
(dots).
teoles ovate, acute, densely tomentellous on exterior,
glabrous within except near apex, caducous. Recep-
tacle campanulate, markedly gibbous, densely grey
mm long; pedicels 1-3
tomentellous on exterior, 2-3 2. Parinari canarioides Kosterm. New & Crit. Mai.
mm long. Calyx lobes 2-3 mm long, narrowly ovate, PI. 3 (1955) 25, t. 12 (For. Dept. Bur. of Planning,
densely grey tomentose on exterior, tomentellous Bogor, Indonesia); Reinwardtia 7 (1965) 159,/. 2.
within. Petals spathulate, 2 mm long, caducous. Fer- Trees to 60 m tall; trunk buttressed to 2.5 m high;
tile stamens 7-8, base forming a conspicuous fused young branches sparsely puberulous, glabrescent,
ring with opposite tooth-like staminodes. Ovary lenticellate. Stipules linear, acute to 5 mm, hirsute,
densely pilose. Style pilose, stigma truncate. Fruit early caducous, present on very young leaves only.
ovoid, 7-8 by 4.5-5.5 cm, exocarp densely len- Leaves chartaceous, ovate, 5-9 by 2-4.3 cm, gla-
ticellate; mesocarp thin, fleshy; endocarp extremely brous on both surfaces when mature, without sto-
hard and thick (1 -8 cm thick), woody, granular, and matal crypts beneath, acuminate at apex, the tip
very irregularly ridged, with 2 small loculi in centre, 5-12 mm long, rounded to subcordate at base;
densely lanate within. midrib lightly impressed above, prominent beneath,
Distr. Malesia: North Borneo (Sabah). Fig. 9. sparsely pubescent when young; primary veins 7-11
Ecol. Hillside forest, to 100 m altitude; forest pairs, plane to prominulous above, prominent be-
along rivers. neath, arcuate; petioles 3-7 mm, glabrous when
Vern. Berangan, Malay. mature, eglandular or with small rather inconspicu-
1989] Chrysobalanaceae (Prance) 657

ous central glands. /n/Zore^cence^ dense-flowered ax- glandular, but glands often obscured. Inflorescences
illary panicles to 4.5 cm long, the rachis and branches of raceme-like reduced terminal and axillary panicles
tomentose; bracts and bracteoles persistent, ovate, or cymules, 1.7-3 cm long, the rachis and branches
puberulous on e.xterior, caducous. Receptacle cam- densely brown tomentose; bracts and bracteoles
panulate, 3 mm long, tomentose on exterior; pedicels large, 2 mm long, ovate, persistent. Receptacle con-
1-2 mm long; calyx lobes elliptic, concave, c. 2 mm, ical, gibbous, to 3 mm long, brown-lanate on exte-
acute, sparsely puberulous on exterior, densely to- rior, pedicels 0.5-2 mm long. Calyx-lobes ovate,
mentellous on interior. Petals elliptic, obtuse, 2 mm, acute, 2-3 mm long, lanate on exterior. Petals white,
tapered to base. Fertile stamens 7-8. Fruit ellipsoid, oblong-ovate, 2-3 mm long, narrowed to base. Fer-
3.5-5 by 1 .5-2.5 cm; epicarp densely to sparsely len- tile stamens 7-9 with tooth-like staminodes oppo-
ticellate; mesocarp fleshy, 1 mm thick; endocarp 5 site. Fruit oblong-ellipsoid, 6.7 by 3.7 cm; epicarp
mm thick, hard, marbled, densely lanate within. sparingly lenticellate.
Distr. Malesia: Sumatra, Borneo, Sulawesi, Distr. Malesia: Malay Peninsula, Borneo (Sara-
Philippines (Palawan). Fig. 9. wak, Brunei, Sabah, NE. Kalimantan), Philippines
Ecol. Forest extending up to 800 m altitude. (Mindanao). Fig. 10.
Uses. The timber is much used, but of poor quali- Ecol. Lowland and hill forest to 900 m, including
ty. Fruit edible, also eaten by pigs. areas on ultrabasic rock.
Uses. The wood is used for supports of Iban long
3. Parinari elmeri Merr. Univ. Calif. Publ. Bot. 15 houses.
(1929) 92; Kosterm. Reinwardtia 7 (1965) 161, f. 4; Vern. Borneo: resak, Iban.
Prance & Whitm. Tree Fl. Malaya 2 (1973) 335.
Trees to 32 m, without buttresses; the young bran- 4. Parinari parva Kosterm. Reinwardtia 7 (1965) 52,
ches densely tomentellous, glabrescent, obscurely f . 5; 162; Prance & Whitm. Tree Fl. Malaya 2 (1973)
lenticellate. Stipules lanceolate, acute, to 18 long mm 335.
by 3 mm broad at base, lateral, tomentellous, persis- Tree to 15 m tall, bole often fluted at base, without
tent. Leaves oblong to oblong-lanceolate, 5-18 by buttresses; the young branches densely tomentellous,
1 .5-7 cm, chartaceous to thinly coriaceous, glabrous glabrescent, conspicuously lenticellate. Stipules lan-
above, densely lanate pubescent beneath, without ceolate, lateral, 13-37 mm long, up to 5 mm broad
stomatal cavities; acuminate at apex, the tip 5-13 at base, persistent, conspicuously reticulate and
mm long, subcuneate at base; midrib plane or slight- densely tomentose on exterior. Leaves chartaceous,
ly impressed and pubescent above when young, oblong to elliptic, 11-28 by 5.5-11 cm, glabrous
prominent beneath; primary veins 14-21 pairs, above, densely lanate-arachnoid pubescent beneath,
prominent beneath, curved at margin; secondary the pubescence completely obscuring reticulate ner-
nerves more or less parallel forming ladder-like vation, but without stomatal crypts; finely acumi-
reticulation; petioles 1.5-6 mm long, tomentellous. nate at apex, the tip 3-13 mm
long, rounded to sub-
cordate at base; midrib plane and pubescent above,
prominent beneath; primary veins 15-23 pairs, ar-

cuate and anastomosing at margin, plane above,

prominent beneath, pilose; petioles 5-8 mm, densely
pale brown pilose, with 2-3 extremely prominent
glands. Inflorescence of short little-branched termi-
nal and axillary panicles to 5 cm long, sometimes
borne on young woody branches, the rachis and
branches densely pale-brown tomentellous; bracts
and bracteoles ovate, acute, mm long, persistent.
to 5
Receptacle campanulate, 3 mm long, pale brown
tomentose on exterior; calyx lobes acute, ovate-
lanceolate, 1-1.5 mm long, densely tomentose.
Petals white. Fruit ellipsoid to narrowly ellipsoid, to
10 cm long by 3 cm broad; epicarp densely len-
ticellate, ridged when dry; mesocarp thin and fleshy;
endocarp thick, hard.
•"====^5;, Distr. Malesia: Malay Peninsula (Kelantan,
Trengganu, Pahang, Johorc), Sumatra, Borneo. Fig.
10.

Fig. 10. Distribution of Pormor/e/fner/ Merr. (stars) Ecol . Mostly on ridge tops and hillsides to 750 m
and P. parva Kosterm. (dots). altitude.
658 Flora Malesiana [ser. I, vol. lO^

5. Parinari nonda F.v.M. ex Benth. F1. Austr. 2

(1864)426; BANKSi&SoL.Bot. Cook's Voy. 1 (1900)
t. 92; Bailey, Queensl. Fl. 2 (1900) 524; Compreh.

Cat. Queensl. Pi. (1913) 167; Pulle, Nova Guinea,

Bot. 8, 2 (1910) 367; Kosterm. Reinwardtia 7 (1965)
170, f. 1 la, excl. syn. P. papuanum et P. salomonen-
se. - Ferolia nonda (F.v.M. ex Benth.) O. Ktze,
Rev. Gen. PI. 1 (1891) 216.
Trees to 15 m
tall, without buttresses, the young

branches sparsely puberulous, soon glabrous, with

small prominulous ienticels. Stipules lanceolate,
membraneous, tomentellous, to 5 mm long, very ear-
ly caducous. Leaves chartaceous to thinly coria-
ceous, oblong, 4- 1 1 by 1 .8-4.2 cm, glabrous above,
Fig. 11. Distribution of Parinari nonda F.v.M. ex
with stomatal crypts filled with lanate pubescence
Benth.
beneath, rounded to acute (or rarely bluntly acumi-
nate) at apex, subcuneate at base; midrib plane or
prominulous, sparsely tomentellous when young
above, prominent beneath; primary veins 10-17 curved and anastomosing at margins; secondary
pairs, curved at margins; secondary nerves reticulate nerves reticulate, slightly flattened, with a series of
slightly flattened, with a series of marginal glands at marginal glands at vein endings on lower portion;
veins on lower portion; petioles 5-10 mm long, petioles 2-8 mm long, tomentellous when young,
tomentellous, terete, with 2-4 prominent, conspic- terete or slightly canaliculate, with 2 conspicuous or
uous glands near mid point. Inflorescence of spread- sometimes obscure glands. Inflorescence of terminal
ing terminal and subterminal panicles, 5-11 cm and subterminal panicles, 2-6 cm long, the rachis
long, the rachis and branches rather sparsely grey- and branches densely tomentose or tomentellous;
brown tomentellous; bracts and bracteoles large, bracts and bracteoles large, ovate, 2-2.5 mm long,
ovate, 2.5-3 mm long, tomentose, caducous. Recep- tomentose on exterior; pedicels 0.2-1.5 mm long.
tacle campanulate, 2-3 mm long, tomentose on ex- Calyx lobes triangular, acute, c. 1 mm long, tomen-
terior; pedicels 0.5-1 mm long. Calyx lobes triangu- tose on exterior, tomentellous within. Petals 5,

lar, acute, mm long, tomentose on exterior,

c. 1 white, acute. Fertile stamens 7-8, with tooth-like
tomentellous within. Petals 5, white, acute. Fertile staminodes opposite, lanate around base. Ovary
stamens 7-9, with tooth-like staminodes opposite, villous, style villous on lower portion, glabrous
lanate around base. Ovary villous. Style villous above; stigma capitate. Fruit ovoid, 4-6 cm long;
lanate on lower portion, glabrous above; stigma epicarp sparingly to densely lenticellate; mesocarp
capitate. Fruit ovoid, epicarp sparingly lenticellate. thin, fleshy; endocarp hard, thick, marbled, lanate
D i s t r . Australia (Queensland and Northern Ter- within.
ritory) and in Malesia: southern extreme of Papua
New Guinea and Irian Jaya. Fig. 11. KEY TO THE SUBSPECIES
Ecol. Savanna, open forest, forest on rocky
areas in lowlands. 1. Leaves coriaceous, 4-11 by 1.9-5 cm. Mature
Vern. Papua New Guinea: warrem. fruit c. 6 cm long when dry. Montane
a. ssp. papuana
6. Parinari papuana C.T.White, J. Am. Arb. 31 1. Leaves chartaceous, 7-18 by 2.5-7 cm. Lowland.
(1950) 86. - Parinari nonda auct. non Benth.: Kos- 2. Fruit sparsely lenticellate, small, c. 4 cm long.
term. Reinwardtia 7 (1965) 170, p.p. Leaf base subcuneate b. ssp. salomonense
Large trees to 40 m tall, buttressed or unbut- 2. Fruit densely lenticellate, large, c. 6.5 cm long.
tressed, the young branches puberulous, soon gla- Leaf base usually rounded c. ssp. whitei
brous, with clusters of large prominent Ienticels with
centralslit. Stipules lanceolate, very early caducous. a. ssp. papuana
Leaves thickly coriaceous to chartaceous, oblong, Unbuttressed tree. Leaves thickly coriaceous,
4-18 by 1.5-6.5 cm, glabrous above, with stomatal 4- 1 1 by 1.9-5 cm, subcuneate at base. Mature fruit
crypts filled with lanate pubescence beneath, acu- c. 6 cm long when dry; densely lenticellate on ex-
minate at apex, the tip 3-10 mm long, rounded to terior.
subcuneate at base; midrib plane or slightly impress- Distr. Malesia: Northern, Central and Eastern
ed and sparsely tomentellous when young above, Papua New Guinea. Fig. 12.
prominent beneath; primary veins 16-22 pairs, Ecol. Mountains, 500-2000 m altitude. Fig. 12.
1989] Chrysobalanaceae (Prance) 659

Fig. 12. Distribution of Parinari papuana C.T. White ssp. papuana (dots), ssp. salomonensis (C.T. White)
Prance (triangles), and ssp. whitei Prance (stars).

Vern. Korafe, morni, Aiyura, puwirini, Was- brescent, conspicuously prominently lenticellate.

kuk. Tor, Anona. Stipules ovate to lanceolate, acute, 3-5 mm, pilose
on Leaves coriaceous, ellip-
exterior, early caducous.
tic to oblong, 14-23 by 4-9 cm, glabrous above,
b. ssp.salomonense (C.T.White) Prance, Brittonia
- Parinari salomonense C.T.White, with stomatal cavities filled with grey lanate pubes-
39 (1987) 369.
cence beneath, shortly acuminate at apex, the tip
J. Arn. Arb. 31 (1950) 87.

Buttressed tree. Leaves chartaceous, 7-12 by 3-7

3-13 mm long, rounded to subcordate at base;
midrib plane above, glabrous when mature except at
cm, subcuneate at base. Mature fruit c. 4 cm long,
base, prominent, glabresceiit beneath; primary veins
sparsely lenticellate on exterior.
23-35 pairs, erect, plane above, flattened and prom-
Distr. Solomon Islands. Fig. 12.
inent beneath; secondary veins prominulous and
Ecol Lowland forest, hillsides and ridges to 300
.

m altitude.
parallel ± ladder-like beneath; petioles 9-17 mm
Vern. Malmone, Kwara'ae, nakisi, one one, long, thick, tomentellous, when young, glabrescent,
eglandular or glandular. Inflorescences of large,
sauialu, susui.
spreading terminal panicles, 10-21 cm long by 7-12
cm broad, the rachis and branches yellow-grey to-
c. ssp. Prance, Brittonia 39 (1987) 369.
whitei
mentellous; bracts and bracteoles ovate, 3 mm long,
Unbuttressedtree. Leaves chartaceous, 7-18 by
early caducous. Receptacle campanulate, slightly
2.5-6.5 cm, rounded at base. Mature fruit c. 6.5 cm mm long, densely grey tomentose on ex-
gibbous, 3
long, densely lenticellate on exterior. terior; pedicels 1-3 mm long; calyx lobes ovate,
Distr. Malesia: West Irian and Papua New Gui- acute, 1.5-2 mm long, unequal, grey tomentose.
nea along northern coast from extreme west to east. Petals white to bluish, lanceolate to spathulate, nar-
Fig. 12.
rowed towards base, c. 2 mm long, glabrous. Sta-
Vern. Lowka, Manikiong, ogelel, Mooi. mens 8-10, with tooth-like staminodes opposite.
Ovary pilose; style glabrous; stigma truncate. Fruit
7. Parinari oblont>ifolia Hook. /. Fl. Brit. India 2 ellipsoid, 5-9 by 3-4 cm, epicarp den.sely len-
(1878) 309; King, J. As. Soc. Beng. 66 (1897) 279; ticellate; mesocarp 1.5-2 mm thick; endocarp hard,

RiDLKY. Agr. Bull. Str. & Fed. Mai. St. 1 (1902) 144; thick, marbled, 7-13 mm thick, fibrous, densely
Fl. Mai. Pen. I (1922) 668; Foxw. Mai. For. Rec. 3 lanate within.
(1927) Corner, Wayside Trees (1940) 527;
175; Distr. Malesia: Malay Peninsula (S. Kelantan to
Kosterm. Rcinwardtia 7 (1965) 165,/. 8; Prance & Johorc), Sumatra, Borneo (Sabah, Kalimantan).
Whitm. Tree Fl. Malaya 2 (1973) 335. - Ferolia Fig. 13.
ohionfii/olia (HfXJK./.) O. Kt/c, Rev. (icn. PI. 1 Ecol . Lowland rain-forest and beside rivers or in

(1891) 216. - Partnanum hurneense Mkrr. Univ. valleys extending lo 450 m altitude.
Calif. Publ. Bot. 15 (1929) 93. Vern. Malay Peninsula: hedara hulan, kemalau,
1 recs to 40 m tall, trunk low thick buttrcs.scd lo 2 nwntelor, merhalu; dunfiun hukit, Malay; Borneo:
m. the young branches minutely tomentellous, gla- mankudar, mengkudu, Kalimantan.
660 Flora Malesiana [ser. I, vol. 10^

at base; midrib plane above, prominent beneath;

primary veins 10-15 pairs, prominulous to plane
above, prominent beneath, erect, curved only at
margin; petioles 14-20 mm long, glabrescent, with
inconspicuous glands near to lamina base, puberu-
lous, glabrescent. Inflorescence of axillary little-

branched panicles, 4-10 cm long, the rachis and

branches densely brown tomentellous; bracts and
bracteoles ovate, early caducous. Receptacle cam-
panulate, slightly gibbous, 2-3 mm long, ferrugin-
eous pubescent on exterior; pedicels 0.5 mm long;
calyx lobes lanceolate, acute, 1 mm long, tomentel-
lous. Petals lanceolate, glabrous. Stamens c. 8 with
short tooth-like staminodes opposite. Ovary densely
Fig. 13. Distribution of Parinari oblongifolia pilose. Style glabrous, equalling stamens; stigma
Hook./, (dots), P. gigantea Kosterm. (stars), and P. truncate. Fruit not seen.
metallica Kosterm. (triangles). Distr. Known only from Brunei, Sabah, and
Sarawak. Fig. 13.
Ecol. Forests on well-drained soil, hillsides,
8. Parinari gigantea Kosterm. Reinwardtia 7 (1965) 50-300 m altitude.

182, f. 19.
Large trees to 40 m tall, trunk fluted at base, the 10. Parinari prancei Kosterm. Reinwardtia 10(1985)
young branches densely lanate pubescent, glabres- 124.
cent, with conspicuous small lenticels. Stipules lan- Trees to 25 m tall, the young branches densely
ceolate, acute, to 25 mm long, caducous, membra- brown lanate and pilose, lenticellate. Stipules
neous, densely appressed tomentellous on exterior, caducous (not seen). Leaves rigidly coriaceous, ellip-
elliptic, 9-17 by
glabrous within. Leaver coriaceous, tic, 9-21 by 6.5-12 cm, glabrous and shiny above

5-8 cm, glabrous above, with dense conspicuous when mature, lanate when young, with conspicuous
stomatal crypts beneath, bluntly acuminate at apex, stomatal crypts filled by lanate pubescence beneath,
the tip 3-6 mm long, rounded at base; midrib plane broadly apiculate at apex, rounded to broadly sub-
above, prominent and pilose, glabrescent beneath; cuneate at base; midrib ± plane above, prominent
primary veins 20-28 pairs, slightly impressed on up- beneath; primary veins 14-16 pairs, plane or slightly
per portion, prominulous on lower portion of upper impressed above, prominent beneath, arcuate near
surface, straight, erect, parallel; secondary veins ± margins, secondary venation parallel and forming a
parallel; petiole thick, 4-7 mm long, tomentellous ladder-like reticulum; petioles 10-12 mm long,
when young, with 2 small round glands on mid point densely ferrugineous lanuginose when young, eglan-
above. Flowers not seen. Infructescence axillary, dular. Inflorescences of axillary little branched small
3-5 mm long. Fruit irregularly ellipsoid, 6.5 cm panicles or racemes, to 3 cm long, the rachis and
long, 4 cm broad; epicarp densely lenticellate; meso- branches densely appressed tomentellous; bracts and
carp fleshy; endocarp hard, bony, irregularly ribbed, bracteoles caducous. Receptacle campanulate-cupu-
lanate within. liform, 3-4 mm long, appressed tomentellous on ex-
D i s t r . Malesia: Borneo (W. Kalimantan, Sabah). terior; pedicels 1.5 cm long; calyx lobes triangular,
Fig. 13. acute, 1.5 mm long, tomentellous. F/-U/7 ellipsoid, c.
Ecol. Lowland forest. 4 by 6 cm diam.; epicarp densely lenticellate; meso-
Vern. Lempong, Kalimantan. carp fleshy, 2mm thick; endocarp woody, very hard
and thick, marbled, densely lanate within.
9. Parinari metallica Kosterm. Reinwardtia 7 (1965) Distr. Malesia: E. Papua New Guinea (Milne
49, f. 3; 160, f. 3. Bay Prov., Northern Prov.). Fig. 14.
Trees to 16 m
tall, unbuttressed, the young bran- Ecol. Lowland rain-forest to 400 m altitude.
ches appressed strigose, glabrescent, conspicuously
lenticellate. Stipules ovate-lanceolate, acute, 8-15 11. Parinari rigida Kosterm. Reinwardtia 7 (1965)
mm long, densely brown tomentose, membraneous, 53, f. 6a, b; 163. - Parinari ashtonii Kosterm. Rein-
early caducous. Leaves thickly coriaceous, elliptic, wardtia 7 (1965) 53, f. 7; 164.
8-17 by 4-9 cm, glabrous and shiny with metallic Trees to 30 m tall, unbuttressed, the young branch-
sheen above when dry, with dense stomatal crypts es tomentellous, glabrescent, inconspicuously len-
filled with hairs, apex rounded to shortly blunt acu- ticellate. S//pu/e5 caducous (not seen). Leaves rigidly
minate, the tip 0-3 mm long, rounded or subcuneate coriaceous, elliptic to oblong ovate, 7.5-23 by 3-8
1989] Chrysobalanaceae (Prance) 661

Fig. 14. Distribution of Pahnari prancei Kosterm. (diamonds), P. rigida Kosterm. (stars), and P. sumatrana
(Jack) Benth. (triangles).

cm, those near to inflorescence much smaller than 12. Parinari sumatrana (Jack) Benth. in Hook.,
others, broadest below mid point, glabrous and Niger Fl. (1849) 335; Blume, Mus. Bot.Lugd.-Bat. 2
shiny above, sometimes slightly bullate, the lower (1852) 97; MiQ. Fl. Ind. Bat. 1, 1 (1855) 353; ibid.
surface with stomatal crypts filled with pubescence, (1858) 1084; Suppl. Sumatra (1860) 115; ibid. (1861)
with 2 glandular areas at junction of midrib and 306; C.Muell. in Walp., Ann. 4 (1857) 644; Flora 41
petiole below, shortly and broadly acuminate at (1858) 255; Hook. /. Fl. Brit. India 2 (1878) 309;
apex, the tip 3-17 mm long, rounded or subcordate Miers, J. Linn. Soc. Bot. 17 (1879) 336; K. & V.
at base; midrib plane or impressed for upper portion Bijdr. 5 (19(X)) 340, p.p. excl. P. costatum auct. non
above, prominent and appressed pilose beneath Blume; Merr. J. Arn. Arb. 33 (1952) 239; Backer
when young; primary veins 13-20 pairs, slightly im- & Bakh./. Fl. Java 1 (1964) 522, p.p. excl. P.
pressed above, prominent beneath, slightly curved at costatum; Kosterm. Reinwardtia 7 (1965) 176. -
margins only; secondary venation flattened or Petrocarya sumatrana Jack, Mai. Misc. 2 (7) (1822)
rounded, parallel; petioles thick, 3-10 mm long, 67 (repr. Calc. J. Nat. Hist. 4 (1843) 165). - Lepido-
grey-pilose pubescent, rugose, with 2 small glands on carpa ovalis (KoRTH.) Blume ex Miq. Fl. Ind. Bat. 1,

mid point of upper side. Inflorescences of narrow 1 (1855) 353. - Ferolia sumatrana (Jack) O. Ktze,
terminal panicles to 13 cm long, the rachis and Rev. Gen. PI.1 (1891) 216. - Parinarium auct. non

branches tomentose; bracts and bracteoles lanceo- Blume: Backer, Schoolfl. Java (1911) 445, p.p. - I

late, to 2 mm long, early caducous. Receptacle cam- Fig. 15.

panulate, slightly gibbous, 5 mm long, densely vil- Trees to 30 m tall, without buttresses or small ones
lous-tomentose on exterior; pedicels c. 1 mm long; to 50 cm; the young branches densely tomentellous,
calyx lobes elongate triangular, 2-2.5 mm long. glabrescent, lenticellate. Stipules oblong, to oblong-
Petals spathulate. Stamens 6-8. Ovary densely vil- ovate, 5 - 2 1 mm
long by 3 - 5 mm
wide at base, mem-
lous. Style equalling stamens; stigma capitate. Fruit braneous, early caducous, pilose on exterior. Leaves
irregularly ellipsoid, 5 cm long, to 4 cm diam., chartaccous to subcoriaccous, elliptic to oblong ellip-

tapered towards base almost into a stipe; epicarp tic, 7 14( 21) by 3-7.5
cm, obtuse to shortly broad
densely Icnticcllate; mcsocarp thin fleshy; cndocarp acuminate at apex, the tip up to 3 mm long, rounded
thick,woody, marbled, lanatc within. to subcordate at base; glabrous and shiny above,
Distr. Malesia: S. Malay Peninsula, Sumatra, with deep-set stomatal crypts beneath obscured by
Borneo (Sarawak, L. Kalimantan). Fig. 14. dense caducous lanatc pubescence when young;
Ecol. Heath and swamp forests, lowland forest; midrib plane to slightly impressed above, pilose
0-1400 m altitude. towards base, prominent beneath; primary veins
662 Flora Malesiana [ser. I, vol. 10^

Fig. 15. Parinari sumatrana (Sack) BEt^TH. A Habit; B. leaf undersurface with pubescence removed
. in small
area to show stomatal cavities; C. flower; D. flower section; E. petal; F. ovary and style; G. ovary section;
H. young fruit (Kostermans 21859).
1989] Chrysobalanaceae (Prance) 663

9-14 pairs, arcuate, prominulous above, prominent a. ssp. costata. - Lepidocarpa coslata Korth. Ned.
beneath; petioles 4-8 mm long, with 2 conspicuous Kruidk. Arch. 3 (1855) 387; MiQ. Fl. Ind. Bat. 1, 1

glands near middle, lightly canaliculate, glabrescent. (1855) 354, in syn., sphalm. Lepidocarya costata. -
Inflorescence of short axillary panicles 2-6 cm long, Ferolia costata (Korth.) O. Ktze, Rev. Gen. PI. 1

the rachisand branches brown tomentose; bracts and (1891)216.

bracteoles membraneous, tomentellous on exterior, Tree to 60 m tall, trunk buttressed up to 2 m, the
puberulous within, caducous. Receptacle conical- young branches densely appressed tomentellous,
campanulate, 3 mm long, densely pilose on exterior, glabrescent, with small conspicuous lenticels. Stip-
almost sessile. Petals spathulate, bluish. Calyx lobes ules lanceolate, membranaceous, 3-7 mm long,
elongate-triangular, 2 mm long, acute, pilose on pilose on exterior, early caducous. Leaver coriaceous
both surfaces. Fertile stamens 8, unequal. Ovary or rigidly chartaceous, elliptic, subovate-elliptic to

densely pilose. S/v/e glabrous, equalling stamens, the oblong (leaf index 1.7-2.7), 5-10.5 by 1.8-4 cm,
stigma truncate. Fruit ellipsoid, 4 by 2.5 cm, epicarp glabrous above when mature but with sparse lanate
densely lenticellate; mesocarp 3-4 mm thick; en- covering when very young, with stomatal cavities
docarp marbled in cross section, hard, 5 mm thick, filled with grey lanate pubescence beneath, acumi-
densely lanate within. nate at apex, the tip 3-5 mm long, round to sub-
Distr. Malesia: Sumatra, W. Java. Fig. 14. cuneate at base; midrib prominulous above, tomen-
Vern. Java: kanjere badak. tellous towards base, prominent beneath; primary
Note. This species is distinct from others by the veins 10-16 pairs, arcuate, prominulous above,
predominantly axillary inflorescences. The material prominent beneath; secondary veins rounded or only
described as Lepidocarpa ovalis has much larger, slightly flattened; petioles 5-9 mm long, slender,
more pointed leaves than most of the collections, but tomentellous when young, soon glabrous, usually
KosTERMANS is correct in placing that name in eglandular or with 2 inconspicuous median glands.
synonymy under P. sumatrana. Inflorescences of predominantly axillary or terminal
few-flowered lax panicles to 8 cm long, the rachis and
13. Parinari costala (Korth.) Blume, Melang. Bot. branches appressed grey to brown appressed tomen-
2 (1855) 10; Miq. FI. Ind. Bat. 1, 1 (1855) 354; ibid. tellous; bracts and bracteoles lanceolate, c. 2 mm
(1858) 1084; Suppl. Sumatra (1860) 115; C.Muell. long, caducous. Receptacle campanulate, slightly
in Walp., Ann. 4 (1857) 644; Flora 41 (1858) 255; gibbous, grey-brown pubescent on exterior, 3-3.5
Hook. /. Fl. Brit. India 2 (1878) 309; King, J. As. mm long; pedicels 0.5-1 mm long; calyx lobes ovate,
Soc. Beng. 66 (1897) 277; Ridley, Agr. Bull. Str. & acute, 1.5-2 mm long, grey tomentellous on exte-
Fed. Mai. St. 1 (1902) 145; Brandis, Indian Trees rior. Petals white, spathulate, 1.5-2 mm long, cadu-

(1906) 278; Burk. J. Str. Br. Roy. As. Soc. n. 73 cous, glabrous. Stamens 7-8, with small tooth-like
(1916) 200; Merr. J. Str. Br. Roy. As. Soc. n. 76 staminodes opposite, slightly unequal; style gla-

(1917) 81; Enum. Born. PI. (1921) 290; Ridley, Fl. brous; stigma capitate. Fruit ellipsoid, to 3.5 by 4.5
Mal.Pen. (1922) 666; Merr. Enum. Philip. Fl. PI.
I cm; epicarp usually sparsely vcrrucose; mesocarp 2
2(1923)236; Burk. Diet. (1935) 1667; Heyne, Nutt. mm, fleshy; endocarp hard, marbled, 3-5 mm thick,
PI. Ned. Ind. ed. 3 (1950) 697; Kosterm. Rein- fibrous, densely lanate within.
wardtia 7 (1965) 179, f. 17a, b; Prance & Whitm. Distr. Malesia: Malay Peninsula, Sumatra,
Tree Malaya 2(1973) 333.
Fl. Borneo, Philippines (Mindanao, Culion, Samar).
For further synonyms, see under the subspecies. Fig. 16.
Ecol. Lowland forest, hillsides, ridges; altitude
key to the subspecies up to 300 m.
Vern. Malay Peninsula: kemalau, mambatu,
I. Inflorescence and flowers densely ferrugineous merbatu; Borneo: augok, Piak, bugan, Iban.
villous pubescent. Often at high altitudes
b. ssp. rubJKinosa b. ssp. rubii(inosa (Ridley) Prance, Brittonia 39
I . Inflorescence and flowers sparsely to densely grey (1987) 368. Parinarium helferi Hook./. Fl. Brit.
or brown appresscd pubescent. Lowlands. India 2 (1878) 311, excl. syn. Parinarium .sumatra-
2. Primary leaf veins 16-26 pairs; mature leaves num sensu Kurz; Brandis, Indian Trees (1906) 278;
9- 15.7 cm long, oblong (index 2.3-3.65). Fruit Kostkrm. Rcinwardtia 7 (1965) 175. - Parinari
cxocarp usually densely vcrrucose ruhiginosa Ridley, J. Sir. Br. Roy. As. Soc. n. 75
c. ssp. polyneura (1917) 29; Fl. Mai. Pen. I (1922) 668; Foxw. Mai.
2. Primary leaf veins 10-16 pairs; mature leaves For. Rcc. 3 (1927) 175; Burk. Diet. (1935) 1667;

5 10.5 cm long, elliptic (index 1.7 2.7), rarely KosTiJRM. Rcinwardtia 7 (1965) 168. f. 10; Prance &
oblong. Fruit cxocarp u.sually sparsely vcrrucose Whitm. Tree II. Malaya 2 (1973) 336. Parinarium
a. ssp. coslata custatum Ulumi. var. rubiginusum Ridley, J. led.
664 Flora Malesiana [ser. I, vol. 10"*

Fig. 16. Distribution of Parinari costata (Kosterm.) Blume (diamonds), P. costcta ssp. rubiginosa (Ridley)
Prance (triangles), and ssp. polyneura (Miq.) Prance (inverted triangles).

Mai. St. Mus. 6(1915) 143. - Parinari bicolorMERR. c. ssp. polyneura (Miq.) Prance, Brittonia 39 (1987)
Philip. J. Sc. 10 (1915) Bot. 309; Enum. Philip. Fl. 368. - Parinarium polyneurum Miq. Fl. Ind. Bat.,

PI. 2 (1923) 235; Kosterm. Reinwardtia 7 (1965) 172, Suppl. Sumatra (1860) 115; ibid. (1861) 306; Hook.
/. 12. /. Fl. Brit. India 2 (1878) 309; King, J. As. Soc. Beng.
Leaves 4-11.5 by 1.6-4.3 cm, oblong elliptic to 60 (1897) 278; K. & V. Bijdr. 3 (1901) 340; Kosterm.
oblong lanceolate; primary veins 11-19 pairs; pet- Reinwardtia 7 (1965) 167, f. 9a, b; Prance & Whitm.
ioles 4-8 mm long, thickly tomentose. Inflorescence Tree Fl. Malaya 2 (1973) 336. - Ferolia polyneura
dense to lax, ferrugineous villous pubescent. Fruit (Miq.) O. Ktze, Rev. Gen. PI. 1 (1891) 216.
exocarp sparingly lenticellate. Leaves 9-15.7 by 3.7-6.3 cm, oblong (index 2.3
Distr. Burma; Malesia: Malay Peninsula, Bor- -3.65); primary veins 16-26 pairs; petioles 3-7 mm
neo (Sabah, Sarawak, Kalimantan), Philippines long, thick, tomentose. Inflorescence lax, inflores-
(Mindanao, Bucas Grande I.). Fig. 16. cence and flowers with grey appressed tomentellous
Ecol . In lower montane forests of Malay Penin- pubescence. Fruit exocarp usually densely verrucose.
sula and Borneo (750-1500 m) and lowland forests Distr. Malesia: Malay Peninsula (Kelantan,
of the Phihppines. Perak, Pahang, Malacca), Singapore, Sumatra, Bor-
Vern. Merbatu, Malay (= Malesian standard neo. Fig. 16.
timber name for various genera); Borneo: mengku- Ecol. Lowland forest and occasionally in hills
dur, Balikpapan. and seasonal swamps.
1989] Chrysobalanaceae (Prance) 665

Excluded species 7520] ex Hook. /. Fl. Brit. India 2 (1878) 311;

KosTERM. Reinwardlia 7 (1965) 178, /. 16. =
Parinari wallichiana R.Br, [in Wall., Cat. (1832) Dipterocarpus cornutus Dyer (Dipterocarpaceae).

6. ATUNA

Rafin. Sylva Tellur. (1838) 153; Kosterm. Reinwardtia 7 (1969) 421; Prance
& Whitm. Tree Fl. Malaya 2 (1973) 323; Smith, Fl. Vit. Nova 3 (1985) 47. -
Atunus RuMPH. Herb. Amb. 1 (1741) 171, t. 66; Lamk, Encycl. Meth. 1 (1783)
329, non Atunus Rumph. (1743); Panigrahi & Purohit, Taxon 32 (1983) 122.
- Cyclandrophora Hassk. Flora 25^ Beibl. 1 (1842) 47; Steen. Bull. Jard. Hot.
Btzg III, Kosterm. Candollea 20 (1965) 118. - Moquilea sect.
17 (1948) 461;
Cyclandrophora (Hassk.) Endl. Gen. PI. Suppl. 3 (1843) 103. - Parinarium
subg. Cyclandrophora (Hassk.) Blume, Melang. Bot. 2 (1855) 10; repr. Flora
N.R. 16 (1858) 255. - Parinarium subg. Macrocarya Miq. Fl. Ind. Bat. 1, 1
(1855) 354. - Parinarium sect. Cyclandrophora (Hassk.) C.Muell. in Walp.,
Ann. (1857) 644. - Entosiphon Bedd. Madr. J. Lit. Sci. ser. 3, 1 (1864) 44. -
Parinarium subg. Ill Hook. /. Fl. Brit. India 2 (1878) 308, p.p. - Petrocarya
auct. non Schreb.: Jack, Mai. Misc. 2 (7) (1822) 68 [repr. Hook. Comp. Bot.
Mag. 1 (1836)220; Calc. J. Nat. Hist. 4(1843) 164]. - Parinari auct. non Aubl.
(Parinarium auct. non Juss.): Benth. in Hook., Niger Fl. (1849) 333, p.p.;
Blume, Mus. Bot. Lugd.-Bat. 2 (1852) 94; Benth. in Benth. & Hook./., Gen.
PI. (1865) 607; Boerl. Handl. Fl. Ned. Ind.
1 (1890) 431, 424; Focke in E.
1

& P. Nat. Pn. Fam. 3, 3 (1891) 60; Koord. Exk. Fl. Java 2 (1912) 338; Ridley,
Fl. Mai. Pen. 1 (1922) 666. - Fig. 19.
Small to large trees, ultimate shoots with complicated system of divaricate
branching. Stipules large, prominently keeled, lateral, persistent or subpersis-
tent. Leaves almost glabrous on both surfaces, often with minute papillae on
venation giving beaded appearance, without stomatal crypts, with a pair of
glands on midrib at or near base of lower surface. Petioles eglandular. In-
florescence a raceme, or sparsely branched, contracted panicle. Bracts and
bracteoles persistent, eglandular, not enclosing groups of flower buds. Flowers
hermaphrodite. Receptacle obconical to cylindrical, as long as or exceeding
calyx lobes, hollow, hairy inside throughout, throat blocked by retrorse hairs.
Calyx lobes broadly ovate to lanceolate, tomentellous on both surfaces.
5,

Petals 5, glabrous, exceeding calyx lobes. Stamens 10-20, posterior, inserted

unilaterally on margin of disk; filaments free, exserted; staminodes forming a
barely visible denticulate margin to throat. Ovary inserted at mouth of recep-
tacle tube, pilose on exterior; carpel bilocular with 1 ovule in each loculus. Fruit
large; epicarp glabrous, densely verrucose-crustaceous; mesocarp transversely
fibrous; endocarp hard, thick, shortly and sparsely hairy inside, breaking up ir-
regularly at germination. Cotyledons large and strongly ruminate. Germination
cryptocotylar, eophylls alternate.
666 Flora Malesiana [ser. I, vol. 104

Distr. About 11 species in Southern India, Thailand, E. to Fiji and Samoa in the Pacific; in Malesia 5

species in the Malay Peninsula throughout Indonesia, and New Guinea.

Vern. Merbatu, Malay = Malesian standard timber name for various genera.

KEY TO THE SPECIES

(including species of India and the Pacific)

1 . Leaf apex rounded; primary veins 6-8 pairs. Fiji A. elliptica (Kosterm.) Kosterm.
1. Leaf apex acuminate or acute; primary veins usually more than 10 pairs. India. Malesia, or Pacific: Fiji,
only A. racemosa.
2. Receptacle tube cylindrical and narrow.

3. Leaves broadly elliptic, 8- 10 cm broad; rounded at base; apex shortly acuminate, the acumen 2-3 mm
long 1. A. latifrons
3. Leaves oblong, 2.5-6 cm broad; subcuneate to rounded at base; apex with long thin acumen 4-22 mm
long.
4. Receptacle 8-13 mm long. Leaf apex long acuminate; base rounded 2. A. nannodes
4. Receptacle 5-7 mm long. Leaf apex short acuminate, the acumen 3-10 mm long; base cuneate
3. A. penangiana
2. Receptacle tube funnel-shaped to campanulate.
5. Leaves broadly ovate, thickly coriaceous, cordate at base, 4.5-12 cm long 4. A. cordata
5. Leaves usually elliptic, chartaceous to thinly coriaceous, usually rounded at base (if cordate then ex-
ceeding 10 cm in length).

6. Fertile stamens 12-14. Inflorescence sericeous or sparsely pilose. India.

1 Inflorescence sparsely pilose. Leaves elliptic-lanceolate, with 8-10 pairs of primary veins. India
.

A. indica (Bedd.) Kosterm.

7. Inflorescence densely sericeous. Leaves lanceolate, with 12-16 pairs of primary veins. India
A. travancorica (Bedd.) Kosterm.
6. Fertile stamens 15-20. Inflorescence tomentellous. Not in India 5. A. racemosa

1. Atuna latifrons (Kosterm.) Prance & White, sericeous on exterior, sessile; calyx lobes lanceolate
Phil. Trans. Roy. Soc. Lond. 320 (1987) 132. - to oblong-ovate, 5-10 mm long, unequal, densely
Parinarium latifolium Hend. Card. Bull. Str. Settl. sericeous on exterior, tomentellous within. Petals
7 (1933) 102, nom. illeg., non latifolium Exell. - obovate narrowed to base, 10-11 mm long. Stamens
Parinari latifrons Kosterm. Reinwardtia 7 (1965) 54. c. 20, inserted on faucal annulus 2 mm high with

- Cyclandrophora latifolia (Hend.) Prance in tooth-like staminodes opposite, the filaments 10-12
Kosterm., Candollea 20 (1965) 121. ~ Atuna latifolia mm long. Ovary densely strigose. Style slender, gla-
(Hend.) Kosterm. Reinwardtia 7 (1969) 421. brous; stigma truncate. Fruit unknown.
Small tree to m
young branches densely
5 tall, the Distr. Known only from Malay Peninsula on
lanate-tomentellous becoming glabrous, obscurely Kedah-Perak border. Fig. 17.
lenticellate. Stipules lanceolate, to 11 mm long,
acute, keeled, sparsely appressed pubescent. Leaves
chartaceous, broadly elliptic, 11-13 by 8-10 cm,
glabrous and shiny above, slightly bullate, glabrous
beneath except for sparsely pilose venation, apex
very shortly abrupt acuminate, the acumen 2-3 mm
long, rounded at base with base contracted into
petiole; midrib prominent on both surfaces, slightly
pilose towards base above, pilose beneath; primary
veins 12-14 pairs, prominulous inset in a groove
above, prominent and pilose beneath, venation
prominulous; petioles thick, 5-7 mm long, terete,
densely brown lanate when young. Inflorescences of
axillary little-branched panicles or spikes, to 5 cm
long, densely brown sericeous; bracts and bracteoles
to 15 mm
long, ovate-lanceolate, acute, densely seri- Fig. 17. Distribution of Atuna latifrons (Kosterm.)
ceous on exterior, appressed puberulous within. Re- Prance & White (star) and A. cordata Cockburn
ceptacle tube narrowly cylindrical, 7-11 long, mm ex Prance (triangles).
1989] Chrysobalanaceae (Prance) 667

2. Atuna nannodes (Kosterm.) Kosterm. Rein- 3. Atuna penangiana (Kosterm.) Kosterm. Rein-
wardtia 7 (1969) 422; Prance & Whitm. Tree Fl. wardtia 7 (1969) 422; Prance & Whitm. Tree Fl.
Malaya 2 (1973) 325. - Parinari nannodes Kosterm. Malaya 2 (1973) 326. - Cyclandrophora penangiana
Reinwardtia 7 (1965) 50, f. 4. - Cyclandrophora Kosterm. & Prance, Candollea 20 (1965) 124. -
nannodes (Kosterm.) Kosterm. & Prance, Can- Parinari asperula auct. non MiQ.: King, J. As. Soc.
dollea 20 (1965) 122. Beng. 66 (1897) 2U, p.p.
Trees to 20 m, usually smaller, unbuttressed; the Trees to 20 m tall, unbuttressed, the young branch-
young branches sparsely appressed hirsutulous- lets glabrescent, obscurely lenticellate. Stipules lan-
strigose, soon glabrous, obscurely lenticellate. Stip- ceolate, acute, to 7 mm long, glabrous, stiff, subper-
ules narrowly lanceolate, acute, 6-12 long, mm sistent. Leaves thinly subcoriaceous, oblong to
strigose to glabrous, subpersistent. Leaves thinly co- oblong lanceolate, 3.7-13 by 2-5.5 cm, glabrous on
riaceous, oblong-lanceolate, 6.7-19 by 2.5-5.5 cm, both surfaces, acuminate at apex, the acumen 3-10
glabrous on both surfaces, sometimes slightly bullate mm, cuneate at base; midrib flattened prominulous
above, long slender acuminate at apex, the acumen above, prominent beneath; primary veins 10-13
7-22 mm rounded at base; midrib promin-
long, pairs, arcuate, prominulous on both surfaces; pet-
ulous above, prominent beneath; primary veins ioles 3-5 mm long, eglandular, glabrescent, smooth,
10-12 pairs, arcuate, prominulous on both surfaces not swollen or curved. Inflorescences axillary ra-
or sometimes prominent beneath; petioles 2-4 mm cemes 3-7 cm long, the rachis densely appressed
long, glabrescent, eglandular, the lower part swollen, pilose; bracts and bracteoles sericeous to 10 mm
usually curved. Inflorescences axillary racemes 3-7 long, persistent. Receptacle cylindrical, 5-7 mm
cm long, the rachis densely sericeous-tomentellous; long, sericeous pubescent on exterior; calyx lobes
bracts and bracteoles lanceolate, 3-7(-13)mm long, acute, mm long, slightly unequal. Stamens 20,
4-5 c.

persistent, sericeous. Receptacle cylindrical, 8-13 the filaments to 8 mm long with tooth-like stami-
mm long, densely sericeous on exterior, sessile; calyx nodes opposite. Style to 10 mm long, stigma capitate.
lobes to 6 mm long, unequal, acute, sericeous on ex- Ovary pilose. Fruit (immature) ellipsoid, epicarp
terior. Petals white, spathulate to ovate, 8-12 mm crustaceous, verrucose.
long, narrowed to base. Stamens 18-20, black to Distr. Malesia: Malay Peninsula (Penang,
purple, the filaments 10-15 mm long, slightly unilat- Perak, Johore, Kelantan and Trengganu). Fig. 18.
eral with tooth-like staminodes opposite. Style to 15 Ecol. Well drained forests to 500 m altitude.
mm long, glabrous; stigma capitate. Ovary pilose. Vern. Membatu, Malay.
Fruit ellipsoid, 3-4 by
cm, slightly tapered to
1.5 Note. The two species Atuna nannodes and A.
base, crustaceous verrucose on exterior; mesocarp penangiana are hard to separate. The larger flowers
2-2.5 mm, fibrous, hard, endocarp thin. of A. nannodes seem consistent and the species gen-
D s t r Malesia: Malay Peninsula (Trengganu and
i . erally has leaves with a much longer apex. These may
Pahang southward), Borneo (Sabah, Sarawak). Fig. be one variable species.
18.

Ecol. Well drained forests to 500 m altitude. 4. Atuna cordata Cockburn ex Prance, Brittonia
Vern. Merbatu, Malay. 39 (1987) 364. - Atuna cordata Cockbvrh, Trees of
Sabah 2 (1980) 82, nom. inval.
Tree to 40 m tall, the trunk often with thick but-
tresses; young branches glabrescent, inconspicuously
lenticellate. Stipules to 1 .7 cm long, very early cadu-
cous. Leaves coriaceous, broadly ovate, 4.5-12 cm
long, 3-9.5 cm wide, abruptly acuminate at apex,
the acumen 1-3 mm long, cordate at base, glabrous
and shiny above, glabrous beneath; midrib promin-
ulous above, prominent beneath; primary veins 9- 12
pairs, lightly prominulous above, prominulous and
glabrous beneath; petioles 1-3 mm long, short and
thick, glabrous. Inflorescences of terminal and
subterminal racemes 4-8 cm long, borne in single or
more often in paired branches, densely tomcntellous
on exterior, pubcrulous within; bracts and bracteoles
ovate, tomcntellous, early caducous. Receptacle 5-7
Fig. 18. Distribution of>4/i/na/ion/io^e5 (Kosterm.) mm long, conical to campanulatc, tomcntellous on
Kostfrm. (iriangles), A. penanffiana (Kosterm.) exterior, sessile; calyx lobes slightly unequal, tomcn-
KosTtRM. (dots). tellous on both surfaces. Petals c. 7 mm long,
 Flora Malesiana [ser. I, vol. 10^
668

Fig. 19. Aluna racemosa Rafin. ssp. excelsa (Jack) Prance. A. Habit; B. leaf undersurface; C. flower bud;
D. flower section; E. ovary section; F. base of stamens; G. petal; H. fruit {A-G Whitmore 3542, // Agama
4222).
1989] Chrysobalanaceae (Prance) 669

)bovate, glabrous. Stamens c. 10, inserted on one Kanehira. Bot. Mag. Tokyo 45 (1931)282. - Petro-
ide of ring, the filaments 10-12 mm long. Ovary carya glaberrima (Hassk.) Miers. J. Linn. Soc. Bot.
lensely pilose. Style slender, hirsutulous on lower 17 (1879) 336. - Ferolia glaberrima (Hassk.) O.
)ortion. Fruit 6 cm long, 5 cm wide, ovoid; epicarp Ktze. Rev. Gen. PI. 1 - Ferolia scabra
(1891) 216.
Tustaceous verrucose, mesocarp 5 mm thick, fi- (Hassk.) O. Ktze. I.e. - Petrocarya scabra
216.
)rous, hard, endocarp thin. (Hassk.) Miers, J. Linn. Soc. Bot. 17 (1897) 336. -
Distr. Malesia: Borneo (Sabah). Fig. 17. Parinarium elatum King, J. As. Soc. Beng. 66 (1897)
Ecol. Hill forests on ultrabasic rock. 280; Ridley, Fl. Mai. Pen. 1 (1922) 669. - Parina-
rium hahlii Warb. Tropenpfl. 6 (1902) 370. - Pari-
>..\tuna racemosa Rafin. Sylva Tellur. (1838) 153; narium mindanaense Perk. Fragm. Fl. Philip.
^ERR. Index Rafin. (1949) 136; Kosterm. Reinward- (1904) 119. - Parinarium curranii Merr. Philip. J.
ia7 (1969)422. - Fig. 19. Sc. 4 (1909) Bot. 264. - Parinarium warburgii Perk.
For further synonyms, see under the subspecies. ex Merr. J. Str. Br. Roy. As. Soc. n. 76 (1917) 82.
- Cydandrophora elata (King) Kosterm. Candollea
KEY TO THE SUBSPECIES 20 (1965) 122. - Cydandrophora scabra (Hassk.)
Kosterm. I.e. 126. - Cydandrophora laurina
1. Leaves 10-25(-35) cm long, usually elliptic, (Gray) Kosterm. I.e. 135. - A. elata (King)
oblong or lanceolate but sometimes ovate, char- Kosterm. Reinwardtia 7 (1969) 421; Prance &
taceous or thickly coriaceous, the apex long finely Whitm. Tree Fl. Malaya 2 (1973) 324. - Atuna
acuminate, 6-25 mm
long; petioles thick. Flowers scabra (Hassk.) Kosterm. Reinwardtia 7 (1969) 422.
10-17 mm
long. Medium to large trees often with Trees to 45 m tall, usually smaller, the bole often
fluted bole a. ssp. racemosa fluted, young branches glabrous or appressed strig-

I. Leaves 4.5-12 cm long, usually ovate or oblong- ose. Stipules lanceolate, stiff, to 20 mm long, acute,
ovate, subcoriaceous or coriaceous, the apex glabrous to strigose. subpersistent. Leaves usually
bluntly acuminate, 3-10 mm long; petioles thin. chartaceous. more rarely stiffly coriaceous, broadly
Flowers 8-11 mm long. Large trees with cylin- ovate, elliptic, oblong or even lanceolate, 10-25
drical bole b. ssp. excelsa (-35) by 3.5- cm, acuminate at apex, the acumen
1 1

6-25 mm long, rounded to subcordate at base,

I. ssp. racemosa. - Atunus alba Rumph. Herb. glabrous on both surfaces when mature, sometimes
\mb. 1 (1741) 171, t. 66, nan Atunus litorea Rumph. sparsely strigose beneath on lower portion when
4erb. Amb. 3 (1743) 96, t. 63. - Cydandrophora young; midrib prominent on both surfaces; primary
\laberrima Hassk. Flora 25 (2), Beibl. 1 (1842) 47; veins 10-13 pairs, prominulous above, prominent
bid. 11 (1844) 583; Cat. Hort. Bog. (1844) 269. - beneath, straight or arcuate; the venation conspicu-
°arinari glaberrimum (Hassk.) Hassk. Tijd. Nat. ously papillose and often giving leaf a scabrous ap-
3es. Phys. 10(1843) 147; C.Muell. in Walp., Rep. pearance; petioles thick, 3-7 mm long, glabrous or
i(1845/46) 647; in Walp., Ann. 4 (1857) 645; Blume, pilose glabrescent . Inflorescences of axillary racemes
Vlus. Bot. Lugd.-Bat. 2 (1852) 98; Miq. F1. Ind. Bat. or branched with up to 3 racemose branches on
little

I. 1 (1855) 355; K. & V. Bijdr. (1900) 338, incl. var. short main peduncle, 5-15 cm long, the rachis
anceolatum (Teijsm. & Binn.) K. & V., p.p. quoad tomentellous to sericeous; bracts and bracteoles
;pec. Java; Burk. Diet. (1935) 1696; Backer & ovate, acute, to 8 mm long, caducous. Receptacle
3akh./. FI. Java 1 (1964) 522. - Parinarium sca- turbinate-campanulate, 5-10 mm long, tomentose
irum Hassk. Tijd. Nat. Ges. Phys. 10 (1843) 147, to sericeous on exterior; pedicels 0.5-1 mm long,
lomen; Cat. Hort. Bog. (1844) 269, nomen; Flora 27 calyx lobes 4-7 mm long, ovate to ovate-oblong,
1844) 585; C.MuELL. in Walp., Rep. 5 (1845/46) densely tomentellous on both surfaces. Petals ovate-
>47; in Walp., Ann. 4(1857)645; Blume. Mus. Bot. oblong, to 10 mm long, blue or white. Stamens
Lugd.-Bat. 2 (1852) 95, p.p.; Mio. Fl. Ind. Bat. 1, 1 15-20, pale blue, to 15 mm long with tooth-like
;i855) 354, t. 5; K. & V. Bijdr. 5 (1900) 337. p.p.; staminodes opposite. Ovary densely villous. Style
Backer, School fl. Java (191 1)445; Ridley, Fl. Mai. equalling filaments, stigma small. Fruit ellipsoid to
Pen. (1922)669. ~ Parinarium lanceolalumJtusM.
1 subglobose. to 7.5 cm diam.; epicarp crustaceous
k Bins. Cat. Hor(. Bog. (1854) 253. 255. nomen. - verrucose; mesocarp to 1 1 mm thick, endocarp thin,
Parmarium amhomense Teijsm. & Binn. I.e. 254, 1-3 mm, dcn.scly pilose within.
nomen. - Parmarium margarata A.Gray. Bot. Distr. A wide range from Thailand to the
kVilkcs U.S. Expl. Expcd. (1854) 489. I. 55; 1 and Solomon Islands,
Pacific: Admiralty, Caroline,
C.Muell. in Walp., Ann. 4 (1857) 646. - Parina- Fiji. Tonga. Samoa; in Malesia: Malay Peninsula

num laurinum A.Gray. Bot. Wilkes U.S. Expl. Ex- (Pcrak). Singapore. Sumatra, Borneo (Sarawak,
Xd. (1854) 490. I. 55; C.Muell. in Walp.. Ann. 4
1 Brunei), Sulawesi. Philippines. Ambon. Ternate.
[1857)646; Merr. Philip. J. Sc. 10(1915) Hot. 210; Ccram. New Ciuinca, New Hrilain. lig. 20.
670 Flora Malesiana [ser. I, vol. 10

Fig. 20. Distribution of A tuna racemosa Rafin. racemosa (dots) and ssp. excelsa (Jack) Pranc
ssp.
(triangles). Atuna racemosa ssp. racemosa also occurs in Tonga, Fiji,and Samoa outside the area shown i

the map.

Eco 1 . Usually occurring in well-drained lowland Note. KosTERMANS included Cyclandrophoi

or hill forest, up to 600 m altitude, but also found on glaberrima as a synonym of Atuna excelsa rath(
riverbanks, freshwater or brackish swamps and even than where it is placed here. There seems little doul
in mangrove. based on the original description and herbariui
Uses. The fruit (cotyledon) is grated and made material at Leiden bearing Hasskarl's writing thi

into a putty for caulking canoes, widely used in C. glaberrima is equal to Atuna racemosa ssp. rac
Pacific islands. An oil is extracted from the seeds mosa as defined here. The original description of <
used variously in different areas, e.g. to scent glaberrima indicates leaves that are far too large f(

coconut oil and for hairdressing. The leaves are used ssp. excelsa. Atuna was distinguished t
excelsa
to thatch the outside walls of houses in Fiji. The KosTERMANS by its coriaceous leaves and short pe
wood is used locally for posts and poles, but is not of ioles. However, many sheets which he determined {

good quality. A. racemosa have equally short petioles and there

Vern Jangong, membatu, Malay; kisokka, Jav.;
. much variation in leaf texture. Therefore it is n<
Borneo: belibu, senumpol, Iban, kukut, Sarawak, possible to maintain A. elata. Similarly the distin(
merampangi, tatambu, Sabah, torog, Orang Sungei, tion of A. scabra was the scabrous texture of tl

K'tangan; Sulawesi: lomo, Makassar; Philippines: leaves and their more lanceolate shape. Many colIe(
aluma, Ceb., botabon, butabul, getabon, Tagb., tions of /4. racemosa are equally scabrous {e.g. LA
bolga, Bik., pantog-usa, Kuy., pinae, tabontaba, 52392 from New Guinea) and there is so much varii
takoutaban. Bis., tabong. Bag., tabon-tabon, C. tion in leaf shape that it would be quite impossible t

Bis., Bik., Mbo., samake, Bug.; New Guinea: separate /I. scabra on that feature. This was alread
asikua,asista, Saki, bata-bata, koewao, Kwerba, placed under Parinari glaberrimum by Backer an
dela,Mooi, kan, Oriomo, low tukwa, lowtukwa, Bakhuizen van den Brink (I.e. 1964).
Manikiong, mangosowai, Japen; New Britain: latita,
tita; New Georgia: y//", tavai, tita, Uso; Caroline Is.: b. ssp. excelsa (Jack) Prance, stat. nov. - Petroci
agaratim, ais, eis, eritem, grihing, Palau, adidi. Yap, rya excelsa Jack, Mai. Misc. 2 (7) (1822) 68 [rep:
Solomon Is.: do-omu, oso, saia, tij, Kwara'ae; Fiji: Hook. Comp. Bot. Mag. 1 (1836) 220; Calc. J. Na
makita; Tonga: hea, seea; Samoa: ifi-ifi. Hist. 4 (1843) 164]; Walp. Rep. 2 (1843) 7. - Parint
1989] Chrysobalanaceae (Prance) 671

rium jackianum Benth. in Hook., Niger Fl. (1849) teoles oblong, e. 3 mm long, persistent. Reeeptacle

335; MiQ. Fl. Ind. Bat. 1, 1 (1855) 356; C.Muell. in turbinate-campanulate, 4-7 long, sericeous on ex-

Walp., Ann. 4 (1857) 644; Hook./. Fl. Brit. India 2 terior; calyx lobes ovate, equal, to 4 mm, sericeous

(1878) 312. - Parinarium aspentlum MiQ. Fl. Ind. on exterior, tomentellous within. Petals white to

Bat., Suppl. Sumatra (1860) \\5,nomen; ibid. (1861) bluish white, oblong, to 5 mm long, caducous. Sta-

307, descr.; Hook. /. Fl. Brit. India 2 (1878) 310; mens 13-18, to 8 mm long with tooth-like stamin-
King, As. Soc. Beng. 66 (1897) 281; K. & V. Bijdr.
J. odes opposite. Ovary pilose. Style glabrous, equal-
(1900) 337, p./?.; Ridley, Fl. Mai. Pen. 1 (1922)670. ling filaments, glabrous above, stigma small. Fruit
- Ferolia asperula ^iQ.) O. Ktze, Rev. Gen. PI. 1 subglobose to slightly pyriform, 5-7 cm diam. or
(1891) 216.- Ferolia jackiana /Benth.) O. Ktze, I.e. 5-7 by 3.5-4.5 cm; epicarp crustaceous, verrucose;
- Parinarium spicatum King, J. As. Soc. Beng. 66 mesocarp fibrous, 5-8 mm thick, endocarp thin,
(1897) 279; Ridley, Fl. Mai. Pen. (1922) 669. -
1 densely pilose within.
Parinarium maingayi King, J. As. Soc. Beng. 66 Distr. Malesia: Malay Peninsula (Kedah and
(1897) 280; Ridley, Fl. Mai. Pen. 1 (1922) 669.
- Trengganu southward), Sumatra, Java, Borneo, N.
Parinarium villamilii Merr. Philip. J. Sc. 10 (1915) Sulawesi. Fig. 20.
Bot. 308; Enum. Philip. Fl. Pi. 2 (1923) 236. - Ecol. Lowland on well drained soils ex-
forests

Cyclandrophora villamilii ^err.) Prance ex tending to 750 mon ridges and hillsides.
altitude

Kosterm. Candollea 20 (1965) 126. - Cyclan- Vern. Malay Peninsula: kemalau utat, merbatu;
drophora excelsa (Jack) Kosterm. I.e. 128. - Sumatra: kemiling utan, klappa soepai, pelee kamb-
Cyelandrophora asperula (Miq.) Prance ex Kos- ing, salak; Borneo: membatu, Sabah, mahadiu, Ban-

term. I.e. 130. - Atuna villamilii (Merr.) Kosterm. djar, temalang.

Reinwardtia 7 (1969) 422. - Aluna exeelsa (Jack) Notes . Kostermans is probably correct in inter-

Kosterm. I.e. All; Prance & Whitm. Tree Fl. preting Petrocarya exeelsa Jack as the species

Malaya 2 (1973) 324. - Fig. 19. described here. The original description is quite
Tree to 45 m tall, the trunk buttressed up to 2 m, detailed and fits this taxon better than any other
not fluted, the young branches sparsely strigose, Atuna.
glabrescent, obscurely lenticellate. Stipules lanceo- Kostermans treated these two subspecies as
late, 8-15 mm long, acute, sparsely strigose, subper- separate species. They were differentiated by small
sistent. Leaves rigidly chartaceous to coriaceous, characteristics of leaf shape, the acumen and the
ovate to oblong-ovate or less frequently oblong, base. While there do seem to be two elements involv-
4.5-12 by 2-5 cm, acuminate at apex, the acumen ed in this complex, there is a complete graduation of

3-10 mm long, subcordate, rounded or subcuneate any single character such as leaf length, apex length,
at base, glabrous on both surfaces; midrib prominent petiole thickness, leaf shape or fiower size. Ssp. ex-
on both surfaces; primary veins 9-13 pairs, arcuate, eelsa is much commoner in Sundaland and ssp.
prominulous above, prominent beneath, the vena- raeemosa in the Sahul shelf and Pacific islands, but
tion papillose giving a beaded appearance; petioles the two subspecies have considerable geographical
slender, 3-6 mm long, puberulous, glabrescent or overlap with ssp. raeemosa occurring sporadically on
glabrous. Infloreseenees of axillary racemes to 7.5 the Malay Peninsula. Since all characters merge and
cm long, or little branched with 2 or more racemose are only weakly correlated, these two species are
branches on short main peduncle, the rachis and reduced to subspecies, a rank more in accord with
branches densely short sericeous; bracts and brac- their variational and geographical patterns.

7. MARANTHES

Blume, Bijdr. (1825) 89; Kosterm. Candollea 20 (1965) 196; Prance, Bol. Soc.
Brot. s^r. 2, 40 (1966) 183; Brittonia 20 (1968) 203; Fl. Neotrop. 9 (1972) 201;
Prance & Whitm. Tree Fl. Malaya 2 (1973) 329; White, Bull. Jard. Bot. Nat.
Belg. 46 (1976) 294; Distr. PI. Afr. 10 (1976) 313; Fl. Zamb. 4 (1978) 41;
Letouzey & White, Fl. Gab. 24 (1978) 29. - Exitelia Blume,
Cameroun 20, Fl.
Fl. Jav. 1, Praef. (1828) vii, nom. illeg. - Grymania Presl, Epim. Bot. (1851)

193, p.p. quoad G. salicifolia tantum. - Parinari sect. Sarcostegia Benth. in

Hook., Niger Fl. (1849) 335, excl. P. jackiana (Petrocarya excelsa). - Parinari
Mihg. Sarcostegia (Besth.) MiQ. Fl. Ind. Bat. 1, 1 (1855) 355, ^jrcV. P. Jackiana;
672 Flora Malesiana [ser. I, vol. lO"

Fig. 21. Maranthes corymbosa Blume. A. Habit; B. leaf base and glands; C. flower and bud; D. flower se(

tion; E. petal; F. anthers; G. ovary section; H. fruit (A-G Sulit 19, H Sinclair 10687).
1989] Chrysobalanaceae (Prance) 673

Hauman, Bull. Jard. Bot. Brux. 21 (1951) 185. - Pahnari subg. Exitelia Bluue,
Melang. Bot. 2 (1855) 10; Hassk. Flora 16 (1858) 255. - Pahnari sect. Exitelia
(Blume) C.Muell. in Walp., Ann. 4 (1857) 645. - Fig. 21.
Medium-sized to large trees. Stipules deltate, intrapetiolar, stiff, caducous.
Leaves glabrous on both surfaces when mature (or lanate in African species),
with dense caducous cobweb-like indumentum when young, without stomatal
crypts; with paired glands at junction of lamina and petiole. Petioles eglandular.
Inflorescence a many-flowered corymbose panicle. Bracts and bracteoles eglan-
dular, caducous, not enclosing flower buds in small groups. Flowers herma-
phrodite. Receptacle 6bcomcdi\, narrowed into pedicel, solid, almost completely
filled with nectariferous tissue, short tomentose to glabrous on exterior, gla-

brous within, calyx lobes suborbicular, deeply concave, unequal. Petals 5, not
clawed. Stamens 25-40, inserted on margin of disk, unilateral with tooth-like
staminodes opposite to almost in a complete circle; filaments far exserted
beyond calyx lobes, in a tangled mass. Ovary inserted laterally at mouth of
receptacle; carpel bilocular with 1 ovule in each loculus. Style pubescent at base
only, curved upwards, exserted. Fruit a large fleshy drupe; epicarp smooth,
glabrous, not lenticellate; mesocarp fleshy; endocarp very hard, fibrous with a
rough exterior, densely tomentose within, with 2 lateral plates which break away
on germination. Germination phanerocotylar. Cotyledons fleshy, pale green;
cataphylls absent; first 2 eophylls opposite, the others alternate or opposite.

Distr . In tropical Africa 10 species, one native to Central America and one widespread species in Malesia,
NE. Australia and W. Pacific.

1. Maranlhes corymbosa Blume, Bijdr. (1825) 89; Craib, F1. Siam. Enum. 1 (1931) 563. - Grymania
KosTERM. Candollea 20 (1965) 107; Prance & salicifolia Presl, Epim. Bot. (1849) 193; Walp.
Whitm. Tree Fl. Malaya 2 (1973) 330, excl. syn. Ann. 3 (1853) 854. - Parinarium griffithianum
Couepia panamensis. - Exitelia corymbosa (Blvme) Be.nth. in Hook., Niger Fl. (1849) 334; Fl. Austr. 2
Blume, Fl. Java 1, Praef. (1828) vii. - Maranlhes (1864) 426; Walp. Ann. 2 (1851/52) 463; Blume,
multiflora Korth. Verh. Nat. Ges. Ned. Overz. Mus. Bot. Lugd.-Bat. 2 (1852) 98; Melang. Bot. 2
Bezitt., Bot. (1839/42) 259; Ned. Kruidk. Arch. 3 (1855) 10; Miq. Fl. Ind. Bat. 1. (1855) 356; ibid. 1

(1855) 281; Teijsm. & Bins. Cat. Hort. Bog. (1866) (1858) 1084; Hook. /. Fl. Brit. India 2 (1878) 310;
253. - Exitelia multiflora (Korth.) Walp. Rep. 5 .Miers, J. Linn. Soc. Bot. 17 (1879) 336; Vidal,
(1845/46) 115; Miers, J. Linn. Soc. Bot. 17(1879) Sinopsis Atlas (1883) 25; Maingay. Kew Bull. (1890)

336, sub Exileles. - Parinarium griffithianum 122; King, J. As. Soc. Beng. 66 (1897) 283; Bailey,
Benth. in Hook.. Niger Fl. (1849)334; Fl. Austr. 2 Queensl. Fl. 2(1900) 524; K. & V. Bijdr. 5 (1900) 334;

(1864) 426; Walp. Ann. 2 (1851/52) 463; Blume, K.Sch. &

Laut. Fl. Deut. Schutzgeb. Siidsee (1901)
Mus. Bot. Lugd.-Bat. 2 (1852) 98; Melang. Bot. 2 341; Perk. Fragm. Fl. Philip. (1904) 118; Brandis,
(1855) 10; Mio. Fl. Ind. Bat. I, 1 (1855) 356; ibid. Indian Trees (1906) 278; Foxw. Philip. J. Sc. 2

(1858) 1084; Hocjk. /. Fl. Brit. India 2 (1878) 310; (1907) Bot. 386; Backer, Schoolfi. Java (191 1) 446;
Miers, J. Linn. Soc. Bot. 17 (1879) 336; Vidal, Ridley, Fl. Mai. Pen. 1 (1922)670; Disp. (1930)400;
Sinopsis Atlas (1883) 25; Maingay, Kew Bull. (1890) Craib, Fl. Siam. Enum. 1 (1931) 563. - Parinarium
122; King. J. As. Soc. Beng. 66(1897) 283; Bailey, maranlhes BiVMt., Mus. Bot. Lugd.-Bat. 2(1852)99;
Queensl. FI.2(I900)524;K.&V. Bijdr. 5(1900)334; Melang. Bot. 2 (1855) 10. - Parinarium corym-
K.Sc-H. & Laut. Fl. Deut. Schutzgeb. Sudsec (1901) hosum (Blume) Miq. Fl. Ind. Bat. 1, (1855) 356; 1

341; Pf.rk. Fragm. Fl. Philip. (1904) 118; Brandis. ibid. (1858) 1084; Ann. Mus. Bot. Lugd.-Bat. 3

Indian Trees (1906) 278; Foxw. Philip. J. Sc. 2 (1867) 237; Walp. Ann. 4 (1857)645; Vidal, Cat. PI.
(1907) Boi, 386; BArKFR. SchoolH. Java (191 1) 446; I.cn. Silv. Cull. Manila (1880) 29; Merr. Philip. J.

RirjLF.Y,Fl. Mai, Pen. I (1922)670; Disp. (1930)400; Sc. 10 (1915) Bot. 309; Spec. Blanc. (1918) 162;
674 Flora Malesiana [ser. I, vol. 10^

>..
••^

V-
Fig. 22. Distribution of Maranthes corymbosa Blume.

Enum. Born. (1921) 290; Enum. Philip. Fl. PI. 2 tressed or slightly enlarged at base. Stipules intrapet-
(1923) 235; Craib, Fl. Siam. Enum. 1 (1931) 563; iolar, lanceolate, acute, 5-10 mm long, sparsely
BuRK. Diet. (1935) 1695; Corner, Wayside Trees pilose on exterior, glabrous within, early deciduous.
(1940) 527; Backer & Bakh./. Fl. Java 1 (1964) 522. Leaves coriaceous, usually oblong-lanceolate to
- Parinarium mulliflorum (Korth.) Miq. Fl. Ind. oblong-elliptic, 6.5-14 by 2.5-8 cm, acuminate at
Bat. 1, 1 (1855) 356; ibid. (1858) 1084; Suppl. apex, the acumen 8-20(-30) mm long, cuneate at
Sumatra (1860) 115; ibid. (1861) 307; C.Muell. in base, glabrous when mature but often sparsely cadu-
Walp., Ann. 4 (1857) 646. - Parinarium salicifolium cous arachnoid-lanate when young, usually with 2
(Presl) Miq. Fl. Ind. Bat. (1855) 357; C.Muell.
1 , 1 conspicuous prominent glands at junction of petiole
in Walp., Ann. 4 (1857) 646. - Maranthes speciosa and decurrent lower surface; primary veins 7-10
KoRTH. ex Miq. Fl. Ind. Bat. 1, 1 (1855) 357. - pairs, arcuate, prominulous on both surfaces; midrib
Chrysobalanus ciliatus Korth. ex Miq. I.e. 357. - plane above, prominulous beneath; petioles 4-9 mm
Petrocarya griffithiana (Benth.) Miers, J. Linn. long, glabrous when mature, flattened above. In-
Soc. Bot. 17 (1879) 336. - Parinarium racemosum florescences of flattened many-flowered corymbose
ViDAL, Cat. PI. Len. Silv. Cult. Manila (1880) 29. - panicles, rachis and branches sparsely pilose,
Ferolia griffithiana (Benth.) O. Ktze, Rev. Gen. PI. glabrescent. Bracts and bracteoles ovate to lanceo-
1 (1891)216. - Ferolia corymbosa (BtvuE) O. Ktze, late, sparsely pubescent, caducous. Receptacle tur-
I.e. 216. - Ferolia salicifolia (Presl) O. Ktze, I.e. binate, tapering into pedicels 2-4 mm long, grey
216. - Parinarium nitidum auct. non Benth.: tomentose to glabrous on exterior, glabrous within,
KooRD. Meded. Lands Planten Tuin Btzg 19 (1898) calyx lobes fleshy, ovate to elliptic, obtuse, 2.5-4
448. - Polyalthia pulchrinervia Boerl. Cat. PI. mm long, unequal. Petals white tinged pink, gla-
Hort. Bog. (1899) 20; Icon. Bog. (1899) 106. - 1 brous, 3-6 mm long, caducous. Stamens 25-35 in-
Parinarium palauense Kanehira, Bot. Mag. Tokyo serted in several rows on one side of throat, with
45 (1931) 282;~ Fl. Micrones. (1933) 129; J. Dept. tooth-like staminodes opposite. Ovary bilocular,
Agr. Kyushu Imp. Univ. Fukuoka 4 (1934) 325. - densely lanate and villous. Style glabrous except at
Fig. 21. base; stigma truncate. Fruit ellipsoid, 3-4 mm long,
Small to large tree up to 40 m, sometimes flower- 1 .5-2 cm broad, tapered towards base; epicarp thin,
ing when only a few metres high, trunk not but- glabrous on exterior when mature, sometimes lanate
1989] Chrysobalanaceae (Prance) 675

when young; endocarp hard, 5 mm thick, rough on kawang, bonsissian, Malay; bansisian, Sabah,
exterior; densely lanate within; bilocular usually with Tengara; nyalin Sarawak; buenza, kajebabu,
laat,
seed in one locule only. Cotyledons plane-convex. kajoe kambang, kambang, potang, Kalimantan;
Disir. S. Thailand extending east to Solomon Sulawesi: kolaka; Tidore: latan, Aru Is.; Philip-
and Caroline Islands and Australia (Queensland, pines: almag, delebaybai, kaphangan, kolaka, kola-
Northern Territory); in Malesia: Malay Peninsula, sa, kulingan, malapiga, malapuyan, sampinit, tak-
Sumatra, Java, Borneo, Lesser Sunda Islands, dangan. Tag., aningat, binggas, caratacat,
Sulawesi, Philippines, Moluccas, New Guinea, New kagemkena, karatakat. Ilk., arangan, Tagb., daka-
Britain and Admiralty Islands. Fig. 22. yau. Pang., bakoyan, tapas, P. Bis., bongog, dau,
Ecol. Common in coastal areas on rocky and mata-mata, sarangun, S.-L.Bis., dumaga, Kuy., ka-
sandy hills and extremely inland up to 600 m altitude. gangan, kalakangon, ogat. Bag., kamuli tingan,
Also in gallery forest and in Australia on sand dunes Pamp., lank angan, Lan., langog, Buk., lumaluas,
behind mangrove swamp. In Kalimantan the fruit is sigaadan, Mag., maluktik, Sul., salipungan, salutui,
eaten by many bird species, including hornbills and Neg., bareraga, baril, Bik., C.Bis., laiusin, Bik.,
fruit pigeons, which probably disperse the seed. The S.C.Bis., liusin, Sbl., Tag., Bik., sabongkaag. Ilk.,
seed is also scatter-hoarded by the squirrel Sun- Ting., tadiang manok. Ting., Tag.; New Guinea:
dasciurus hippurus. African species of Maranthes badigal, Wagu, djuramun, Kemtuk, jambuan,
are bat-pollinated. Kaigorin, kaupen, Jal, kawol, kowot, Muyu, kwanu,
Uses. Wood used for house-building and for Maprik, lakan, luikoko. Bush Mekeo, marigag,
posts. Fruit edible. Sinai, mehlue, Bembi, morolee, mun, Dagu, naas,
Vern. Thailand: chi-kai-pen, chi-ot-pen, Korat; ningua, njali, Nemo, njiwa, niwa, Sidei, paguh,
Malay Peninsula: chana, lejin, merbatu, m. layang, Timbunke, phu, Wasuk, watu, Karopa; Solomon
mujagon, sau hulan, sunko rimau; Sumatra: damor Is.: asikisiki, giza, mon warlu, morigag, now-wa-ru,

lilis, kajiebatu, kaju baiu, Banka, kalek kureseng, k. santalan; Bougainville: mon-warku, Kugumaru,
parada; Java: gesing, kituwat, solo, sulo, triwulan, marigai, Siwai, Bouin; Palau Is.: apgau.
Huloh, Jav., taritik, l. monjet, Sund.; Borneo: bang-

8. KOSTERMANTHUS

Prance [Tree FI. Malaya 2 (1973) 327, unpublished], Brittonia 31 (1979) 91;
Prance & White, Phil. Trans. Roy. Soc. Lond. 320 (1988) 149, f. 40, 41. -
Parinari auct. non Aubl.: quoad P. heteropetala Scortech. ex King et P.
myriandra Merr., tan turn. - Acioa auct. non Aubl.: Kosterm. Reinwardtia
7 (1965) 9. - Fig. 23.
Large trees, ultimate shoots not divaricate. Stipules to 7 mm long, foliaceous,
persistent, lanceolate to ovate. Leaves glabrous on both surfaces with minute
papillae on veins giving a beaded appearance. Petioles eglandular. Inflorescence
an unbranched or little-branched terminal or axillary raceme with shortly
stalked congested cymules proximally and singly inserted flowers distally.
Bracts and bracteoles small, suborbicular, persistent, eglandular, not enclosing
groups of flower buds. Flowers hermaphrodite, strongly zygomorphic. Recep-
tacle broadly obconic-campanulate, shorter than calyx lobes, asymmetric,
hollow, hairy on both surfaces, but throat not blocked by retrorse hairs; calyx
lobes 5, markedly unequal, suborbicular to lingulate, strongly imbricate. Petals
5, unequal in size and shape, the 2 posterior larger than the others, markedly
ungulate and enclosing stamens in bud. Stamens 8-30, inserted unilaterally on
margin of disk; filaments united for half to three quarters of length into a strap;
sta.ninodcs 5 8, inserted opposite stamens. Ovary inserted laterally at mouth
of receptacle; unilocular with 2 ovules. Fruit large, hard; epicarp glabrous,
 [ser. I, vol. 10^
676
Flora Malesiana

f D
K.TincL

Habit; ^
Fig 23. Kostermanthus heteropetalus (Scortech. ex King) Prance. A. ^•^"°^^^^^f^'°"' f„f
C. ovary section (>»-£, G Ogata KEP 105153, FMeijer SAN 34279).
petals; £. stamen; F. fruit;
 1989] Chrysobalanaceae (Prance) 677

crustaceous-verrucose; endocarp hard, thick, glabrous within, breaking ir-

regularly on germination. Cotyledons slightly ruminate.

Distr. Malesia: Malay Peninsula, Sumatra, Borneo, Sulawesi, Philippines (Mindanao); 2 species.

KEY TO THE SPECIES

1. Leaves coriaceous; petioles 6-12 mm long; calyx tube 2-3 mm long, broadly campanulate
1. K. heteropetalus
1 . Leaves chartaceous; petioles 2-3 mm long; calyx tube 5 mm long, slender 2. K. malayanus

1. Kostermantbus heteropetalus (Scortech. ex

King) Prance, Brittonia 31 (1979) 91; Prance &
White, Phil. Trans. Roy. Soc. Lond. 320 (1988) 152
- Parinarium heteropetalum Scortech. ex King, J.
As. Soc. Beng. 66 (1897) 283; Ridley, F1. Mai. Pen.
1 (1922) 670; Nayaranaswami, J. As. Soc. Beng.
n.s. 27 (1931) 368. - Parinarium kunstleri King, J.
As. Soc. Beng. 66 (1897) 282; Ridley, F1. Mai. Pen.
1 (1922) 670. - Parinarium myriandrum Merr.
Univ. Cal. Publ. Bot. 15 (1929) 93. - Acioa hetero-
peiala (Scortech. ex King) Koster.m. Reinwardtia 7
(1%5) 11. - Fig. 23.
Tree to 35 m tall, older trees buttressed to 1 m up
trunk; young branches glabrous, lenticellate. Stip-
ules 6-7 mm long, partly intrapetiolar, carinate,
Fig. 24. Distribution of Kostermanthus heterope-
ovate, foliaceous, acute to acuminate, persistent to
talus (Scortech. ex King) Prance (dots) and K.
subpersistent.Leaves coriaceous, usually elliptic-
malayanus (Kosterm.) Prance (star).
subovate to rarely lanceolate, 5-20 by 2.5-6 cm,
bluntly acuminate at apex, cuneate to rounded at
base, glabrous on both surfaces, minutely papillose 2. Kostermanthus malayanus (Kosterm.) Prance,
on venation of both surfaces giving a bead-like ap- Brittonia 31 (1979) 94; Prance & White, Phil.
pearance; midrib prominulous above, prominent Trans. Roy. Soc. Lond. 320(1988) 152. - Acioa ma-
beneath; primary veins 6-10 pairs, arcuate, slender, layana Kosterm. Reinwardtia 7 (1965) 13.
prominent beneath; petioles 6-12 mm long, some- Small tree to 10 m; young branches glabrous, len-
times lightly alate from decurrent leaf margins, ticellate. Stipules lanceolate, acute, glabrous, sub-
slightly flattened above, eglandular. Inflorescences persistent, c. 5 mm long. Leaves chartaceous, ellip-
little-branched, to 10 cm long, the rachis and branch- tic,14-20 by 6.5-8.5 cm, acuminate at apex, the
es lightly tomentellous; bracts and bracteoles ovate, acumen 4-10 mm long, cuneate at base, glabrous on
acute, to 3 mm long, caducous. Receptacle broadly both surfaces; midrib slightly prominulous to plane
campanulate, 2-3 mm long, tomentose on both sur- above, prominent beneath, with a pair of round
faces; calyx lobes fleshy, unequal, acute, to 7 mm glands at base; primary veins 10-13 pairs, pro-

long, pilose on both surfaces, reflexed in open flow- minulous above, prominent beneath; petioles 2-3
ers. Petals white tinged pink, fleshy, ellliptic, con- mm long, glabrous, slightly alate with decurrent leaf
cave, largest up to 15 mm long, tomentellous on ex- margins. Inflorescence of subterminal racemes or lit-

terior, enveloping staminal ligule, the others much tle branched, the rachis brown pilose pubescent;
smaller to 6 mm long. Stamens 25-30 united into a bracts and bracteoles ovate, acute, to 3 mm long,
unilateral ligule for 2/3 length, to 12 mm long, caducous. /?ecfp/flc/e slender cylindrical, 5 mm long,
glabrous; anthers pubescent. Ovary densely pilose. sessile tomentose on exterior, densely tomentose
.S7>'/^ densely appressed pilose, stigma truncate. Fruit within; calyx lobes ovate, acute, 4-5 mm long,
ovoid, unilocular 4 by 3 cm; epicarp glabrous, crus- densely tomentose on exterior, glabrous within ex-
taceous; endocarp hard, thick. Cotyledons slightly cept at apex. Petals spalhulate, 6 mm long, clawed.
ruminate, 1.5 by 3 cm. Stamens S- 10, united into a unilateral ligule for half
Distr. Malesia: Malay Peninsula, Sumatra, of length. Ovary densely pilose. Style pilose for most
Borneo, Sulawesi, Philippines (Mindanao). Fig. of length. Fruit unknown.
24. Distr. Malesia: Malay Peninsula (Penang).
Ecol . From sea level up to 500 m altitude. Known only from the type collection.
 10^
678 Flora Malesiana [ser. I, vol.

Insufficiently known

Acioa percohacea Kosterm. Reinwardtia 7 (1965)

14.
andj j- . •• u ^ from
distmgu.shed
«•

sterile collection from the Malay Penmsula,

This species was described from a single
branches and caducous pubescent leaf undersurfaces.
Kostermanthus heteropetalus Prance by its pubescent
collected, but almost certamly belongs withm K.
It is impossible to evaluate until further material is

heteropetalus.
 SABIACEAE (C.F. van Beusekom & Th.P.M. van de Water, Leiden)*

Trees, scandent shrubs or woody climbers. Leaves alternate or spirally ar-

ranged, penninerved, simple or imparipinnate, the leaflets in the latter case op-
posite on often somewhat swollen nodes of the rachis; exstipulate. Flowers
small, bisexual, rarely polygamo-dioecious, in terminal or axillary racemose
panicles, or cymose: paniculately arranged cymes, or these reduced to solitary
axillary flowers. Sepals (3-)5, imbricate, free or ± connate at the base, equal
or unequal. Petals mostly opposite the sepals (rarely alternate:
(4-)5,
Ophiocaryon spp. South America). Stamens (including staminodes) 5, opposite
,

the petals, all polliniferous (Sabia) or only 2 inner ones opposite the reduced
petals polliniferous and the other 3 staminodial. Disk small, annular, surround-
ing the base of the ovary. Ovary of 2(-3) carpels united to form a compound
superior ovary, carpels very rarely free in the apical part, in that case tapering
to 3 short styles with a capitate stigma; otherwise normally a short, cylindric or
conical style; cells 2(-3), each with 1 or 2 pendulous or horizontal, axile hemi-
tropous, unitegmic, crassinucellar ovules. Fruit either 1 -celled or 2-coccous,
drupaceous or dry, indehiscent; endocarp often wrinkled. Endosperm scanty or
wanting. Embryo with a curved radicle and 2 folded or coiled cotyledons.

Distribution. Three genera: Sabia Indo-Malesian, from the S. Deccan and Kashmir to S.
Japan, throughout Malesia as far as the Solomons; Meliosma with a similar range but also occur-
ring in tropical America; Ophiocaryon in the Neotropics. The family is absent in Australia and
Africa.
Fossils of both Malesian genera are found onwards of the Oligocene and Eocene in Asia and
Europe. See under the genera.
Ecology. Tropical forests, mostly below 2000 m altitude.
Taxonomy & Delimitation. There is no concensus of opinion on the affinity, hence the
systematic position of Sabiaceae. Some even doubt whether Sabia and Meliosma are correctly
placed in one family.
After the description of Sabia by Colebrooke (1818), Blume (1851) accommodated it in a new
monogeneric family, Sabiaceae, suggesting its affinity with Menispermaceae. Shortly afterwards
MiERS (see Lindley, 1853), while working on Menispermaceae, placed Sabia between that family
and Lardizabalaceae Hooker/. & Thomson (1855) considered the genus intermediate between
.

Menispermaceae and Schisandraceae.

The scandent habit and the resemblance of the drupelets of Sabia with those of Menispermaceae
undoubtedly were a major argument for supposed affinity.
Subsequently Bentham & Hooker (1862) extended the then monogeneric family Sabiaceae to
include Meliosmaceae Endl., adding the genera Meliosma Blume and Ophiocaryon Schomb.;
both are trees, the first Asian-American, the latter tropical American. They removed the family
in its new concept from the Menispermaceous affinity and accommodated Sabiaceae near Sapin-

daceae and Anacardiaceae. This position has been stable for a century and was adhered to by
many Warburg (1895), von Wettstein (1911), Hutchinson (1926, 1973),
leading botanists:
Melchior (1964). Takhtajan (1969), Dahloren (1975, 1983), and Thorne (1976, 1983). Some
of these authors showed some doubt about the position and some made suggestions, e.g. War-
burg {I.e. 370), who believed one could possibly derive the flower of Meliosma from the Meni-

( I ) Accommodated from he monographs of both authors

t in Biumea volumes 9 and
1 26, and provided with
an introduction.

(679)
 Flora Malesiana [ser. I, vol. 10^
680

an affinity with Sapin-

spermaceous scheme and mentioned that Radlkofer was not in favour of
daceae or Anacardiaceae.
In recent years there is a tendency to return to Blume's
opinion towards affinity with Menisper-
maceae. Pollen morphology (Erdtman, 1952) and embryology (Mauritzon, 1936) have been in-
terpreted in favour of a relationship with Menispermaceae. Airy Shaw (1973) remarked that the
of calyx, corolla and stamens is a most unusual feature, but can
probably be derived
opposition
from the Menispermaceous type of fiower. In his recent classification Cronquist (1981)
tentative-

ly placed Sabiaceae near Menispermaceae in the Ranunculales. Also Forman, in his treatment of

the Menispermaceae (Fl. Males. I, 10^ 1986, 157-253), shares this opinion.
Another matter is whether Sabia and Meliosma/Ophiocaryon should be accommodated in one
family; hitherto they are represented by two tribes in Sabiaceae
(Warburg, 1890), differing in
habit (climbers versus trees), the leaves, and in the androecium. Moreover,
Cronquist (1981)
mentioned in his discussion that, according to Wolfe, the leaf venation of Sabia is highly com-
Menispermaceae, but that of Meliosma more similar with some
patible with a position near
members of may be more arguments to accommodate Meliosma in a separate
the Rosidae. There
family Meliosmaceae Endl., apart from Sabiaceae sensu stricto. This opinion was held by Airy
Shaw (1973).
References: Airy Shaw in Willis, Diet. ed. 8 (1973) 1017; Bentham & Hooker, Genera Plan-
tarum (1862)413; Blume, Mus. Bot. Lugd. -Bat. 1 (1851) 369; Cronquist, An integrated system
1

of classification of flowering plants (1981) 140; Dahlgren, Bot. Notis. 128 (1975) 126; Nordic
J. Bot. 3 (1983) 144; Erdtman, Pollen morphology and plant taxonomy (1952) 380;
Hooker/.
& Thomson, Flora Indica 1 (1855) 208; Hutchinson, Families of flowering plants 1 (1926) 254;
ed. 3 (1973) 449; Lindley, Vegetable kingdom ed. 3 (1853) 467; Mauritzon, Acta Hort. Goth.
1 1 Melchior, Engler's Syllabus 2 (1964) 285; Takhtajan, Flowering plants: origin and
(1936) 18;
dispersal (1969) 226; Thorne, Evol. Biol. 9 (1976) 61 Nordic J. Bot. 3 (1983) 106; Ware, in E. &
;

P., Nat. Pfl. Fam. 3, 5 (1895) 367; Wettstein, Handb. Syst. Bot. ed. 2 (1911) 633.

Vegetative Anatomy. - Leaf anatomy. Hairs unicellular in Saft/o; uniseriate nonglandular

and capitate glandular in Meliosma. Stomata confined to the lower leaf surface, anomocytic or
paracytic. Mesophyll dorsiventral, with arm palisade cells in Meliosma. Veins embedded in
mesophyll and sheathed by sclerenchyma. Petiole in distal end with a closed vascular cylinder.
Crystalliferous cells containing clusterscommon near the veins.
Young stems. Cork Cortex with stone cells in some species oi Meliosma. Pericyclic
superficial.
sclerenchyma forming a composite, closed ring in Sabia, and composed of isolated fibre groups
in Meliosma. Phloem with broad lignified rays in Sabia, and with non-lignified, dilatating (tri-
angular) rays in Meliosma. Vessels with mixed simple and scalariform perforations in first formed
xylem. Cluster crystals common in cortex, phloem, and pith. Secretory cells with unidentified con-
tents noted in parenchyma of several Meliosma species.
Wood anatomy. Vessels exclusively solitary in Sabia, solitaryand in radial multiples or small
clusters in Meliosma; vessel perforations typically simple in Sabia; mixed simple and scalariform
or exclusively scalariform to reticulate in Meliosma. Intervessel pits alternate. Vessel— ray and
vessel-parenchyma pits simple, and often large. Fibres, usually thin-walled, with minutely
bordered to simple pits, and mainly confined to the radial walls in Meliosma (libriform fibres);
with distinctly bordered pits common in both the radial and tangential walls in Sabia; occasionally
septate. Parenchyma scanty paratracheal to vasicentric with occasional lateral extensions in
Meliosma, very sparse to almost absent in Sabia, usually in 8-celled strands. Rays sometimes of
two different sizes, the broad ones 4-8(- 15) cells wide in Meliosma, up to 20 cells wide in Sabia,
usually over 2 mm high, heterogeneous (Kribs type II), often with sheath cells.
Taxonomic note based on vegetative anatomy. The above description is mainly based on early
studies of a very limited number of species, so that the information is far too limited to serve in
the discussion of infrageneric classification and delimitation. The two genera are anatomically
quite distinct in their leaf and wood anatomy. Partly this is related to general anatomical dif-
1989] Sabiaceae (van Beusekom & van de Water) 681

ferences between climbers (Sabia) and erect shrubs or trees (Meliosma). Thus, the anatomical
evidence can be interpreted both in favour of the separation of Meliosma and Sabia into two
same family. Anatomically Sabia is
families, or alternatively to retain their tribal position in the
quite distinctfrom the Menispermaceae to which it has been compared (see above, under tax-
onomy); affinity oi Meliosma and Sabia with families of the Sapindales, especially A nacardiaceae
seem to find more support in vegetative anatomy.
References: Carlqvist, Aliso 11 (1985) 139-157; Desch, Manualof Malayan Timbers 2 (1954)
522-523; Metcalfe & Chalk, Anatomy of the Dicotyledons 1 (1950) 448-452; Moll &
Janssonius, Mikrographie des Holzes 2 (1922) 424-437; Solereder, Systematische Anatomic
der Dicotyledonen (1899) 276-278; & Erganzungsband (1908) 108-109. - P. Baas.
Palynology. Pollen grains in Sabiaceae are prolate spheroidal to prolate. Size ranges from
20 to 33 |im. The apertural system is always tricolporate. Ectoapertures are long colpi, endoaper-
tures are lalongate pori or short colpi. The shape of the endoapertures is oblong to elliptic,
sometimes appro.ximately rectangular or meridionally constricted. Exine stratification is easily to
observe in the light microscope. Each layer is about uniformly thick throughout. The tectum is
equally thick or up to twice as thick as the nexine. It is mostly more than twice as thick as the
columellate layer. Total exine thickness is 1-2.5 |im. The ornamentation is usually finely to
coarsely reticulate; sometimes it is finely or indistinctly perforate.
Meliosma and Sabia show only little infrageneric variation. Moreover, the ranges in both
genera are rather similar. Only minor differences exist: Sabia mostly has a thinner exine with a
finer reticulate ornamentation than Meliosma. Pollen morphology does not support accom-
modating the genera in separate families (Mondal & Mitra, 1982).
As taxonomists, pollen morphologists are ambiguous with respect to the position of the
Sabiaceae. Erdtman (1952) reported pollen similar to that of Sabiaceae to occur in several other
families. However, heactually mentioned only the A/e/?/5/?ermcrceae. PoUen of A nacardiaceae and
Sapindaceae was considered less similar or different. According to Mondal & Mitra (I.e.)
Sabiaceae pollen differs from that of Aceraceae, Hippocastanaceae, Lardizabalaceae, Melian-
thaceae, Menispermaceae, Sapindaceae, and Schizandraceae. On the basis of grain shape and
size, P/E ratio, exine structure and aperture characters they suggested to classify the Sabiaceae

nearest to the Anacardiaceae. It must be stressed, however, that it is extremely difficult to infer
relationships from resemblances between rather simple pollen types. Obviously unrelated taxa
may show very similar pollen, whereas closely related taxa sometimes have completely different
pollen.
References: Erdtman, Pollen morphology and plant taxonomy, Angiosperms (1952) 390;
Mondal & Mitra, Geophytology 12 (1982) 166-180. - R.W.J.M. van der Ham.
Phytochemistry. The only observations worth to be reported here are the presence of pen-
and the absence of starch in seeds. The 3-acetates of
tacyclic triterpenoids of the oleanene series
oleanolic acid and oleanolic aldehyde were isolated from bark of Meliosma simplicifolia. Seeds
of Meliosma myriantha Sieb. & Zvcc. (continental SE. Asia) were reported to give positive reac-
tions for alkaloids and to contain 8% of protein and 10<^o of fatty oil but no starch.
References: Desai c.s., Indian J. Chem. 15B (1977) 291; Hegnauer, Chemotaxonomie der
Pnanzen 6 (1973) 240. - R. Hegnauer.
Note. Though the genera are extremely clearly defined, specific delimitation has in both
genera been difficult, as it seems that racial segregation is common in both. Van de Water has
in Sabia employed a finer specific distinction than van Beusekom did in Meliosma.

KEY TO the genera

I . Climbers or scandent shrubs. Flowers with 5 equal, fertile stamens, in usually rather few-flowered thyrses
or cymes, sometimes reduced to a single axillary flower. Leaves simple, entire or subcntirc, alternate
I. Sabia
682 Flora Malesiana [ser. I, vol. 10"^

1. Trees. Flowers in usually large, racemose, terminal or axillary panicles. Fertile

stamens only 2, the other
3 abortive and reduced to scales or nectary-like bodies.
Leaves uneven pinnate, leaflets opposite on ± nod-
ed rachis, rarely simple, entire or toothed, spirally arranged 2. Meliosma

1. SABIA

CoLEBROOKE, Trans. Linn. Soc. Lond. 12 (1818) 355, t. 14; Wall, in Roxb., Fl.
Ind. 2 (1824) 308; Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 368; Ware, in E. &
P., Nat. Pn. Fam. 3, 5 (1895) 367, f. 183A, 184A-H; Chen, Sargentia 3 (1943)
1; VAN DE Water, Blumea 26 (1980) 1.
- Meniscosta Blume, Bijdr. (1825) 28;
DiETR. Syn. PI. 2 (1840) 923 ('Menicosta'). - Fig. 2-4.
Evergreen or deciduous, woody climbers or more or less scandent shrubs
(rarely recorded as small trees). Twigs terete, striate (see note), with ± promi-
nent leaf cushions, unarmed, mainly in deciduous species with some cataphylls

at their base, spirally arranged. Buds either ± globular and obtuse to rounded,
or ovoid and acute; scales glabrous to pubescent, ciliolate or not, persistent at
the base of the twigs. Leaves simple, ovate or elliptic to lanceolate, 2-25 by
1-10 cm, herbaceous to coriaceous, petioled, entire or very rarely subentire;
nerves 3-12 ascending to patent, curved to straight. Flowers bisexual, 5-
pairs,
merous, actinomorphic, up to c. 15 mm
diam., green to white, yellow, or pur-
ple, axillary, either solitary, or arranged in a few- to many-flowered cyme, ap-
pearing before or with the new leaves. Cymes axillary, either solitary, or, when
the subtending leaves are shed or are bract-like, arranged in racemose to thyr-
soid or sometimes corymbose inflorescence, pedicel ± thickened upwards in
fruit; bracts ovate to lanceolate, up to 6 mm, bracteoles as bracts but usually

smaller, or sepal-like, or minute and then often situated near calyx. Sepals 5(-7,
see bracteoles), equal to very unequal mutually, mostly ± confluent at the base,
variable in size and shape but often suborbicular or broad-ovate to ovate, persis-
tent. Petals 5, rarely 6 or 7, episepalous, imbricate, suborbicular to lanceolate,
glabrous, sometimes (sub)ciliolate, persistent or not; nerves ± parallel, branch-
ing or not, sometimes conspicuous when dark-coloured. Stamens 5, epipeta-
lous, ± equal, persistent or not; filaments more or less flattened, adherent to
the base of the subtending petals; anthers globular to ellipsoid, introrse, upright
or inflexed. Disk in most species ± crown-shaped, sometimes short-cylindrical
{S. sumatrana), truncated conical, or ± cushion-shaped; lobes and ribs, if pres-
ent, alternating with the stamens. Pistil: style conical to cylindrical, rarely ab-
sent, persistent. Ovary superior, 2-celled, (sub)globose to subreniform, usually
laterally somewhat compressed, very rarely subapocarpous. Ovules 2 per cell,
more or superimposed, attached to the septum, hemi-anatropous. 'Drupe-
less
lets' 1-seeded or very rarely with 2 seeds, (sub)globose, obovoid, oblong-
obovoid (or pyriform), or subreniform, laterally ± compressed, green or white
to red or deep blue when fresh; mesocarp rather thin, pulpy, sometimes with
many dark 'granules', endocarp crustaceous, very often with ± prominent ribs
1989] Sabiaceae (van Beusekom & van de Water) 683

Fig. 1 . The Southeast Asian and Malesian distribution area of Sabia Colebrooke. The numbers refer to the
number of species in that area.

forming a fine to coarse reticulate pattern, margin sometimes distinctly keeled.

Seed conform to the drupelet; testa usually conspicuously dark-dotted, inside
often lined with a very thin layer of endosperm. Embryo with two flat, smooth,
somewhat undulated, or sometimes strongly folded cotyledons and a cylindrical
rootlet curving to the hilum.

Distr. Indo-Malesia, along the Himalayas (1 species disjunct, also in the S. Deccan) through Burma and
China to S. Japan; throughout Malesia (not yet known from the Lesser Sunda Islands), as far as New Guinea,
the Louisiades and Solomon Islands. In all 19 species, of which 7 in Malesia. Fig. 1.
Ecol. Inconspicuous climbers (rarely reported as small trees), except two continental Asian species all
evergreen, found in forests and thickets, from the lowland up to c. 1000-1200 m altitude, 5. javanica up to
1500 m and 5. pauciflora to 2000 m; S. racemosa ssp. kinabaluensis is mainly montane, at 800-1500 m.
Flowering occurs mostly throughout the year.

KEY TO THE SPECIES

I . Flowers solitary, sometimes 2 or 3 together, or arranged in a thyrsus; ovary glabrous; style in flower 3-6
mm long, conspicuous in fruit and about half as long as the adjacent side(s) of the drupelet(s)
7. S. sumatrana
1 . Flowers in cymes, these solitary, axillary, up to 30(-40)-nowered; ovary densely pubescent; style in flower
2.25-2.5 mm long; drupelets not known 1. S. erratica
I Flowers in few- to many-flowered cymes; cymes either solitary, axillary, or arranged in an up to 15 cm long
racemose to thyrsoid inflorescence, (I - )2-25-nowercd; ovary glabrous; style in flower up to 1.5(-1.75)
mm long, inconspicuous in fruit and much shorter than the adjacent side(s) of the drupelct(s).
2. Leaves oblong to lanceolate, 3- I2(- 15) by 1-5 cm, beneath usually distinctly paler than above; nerves

(5-)6-9(-IO) pairs, patent, straight; cymes solitary, axillary, (4-)7-25-nowered; style either absent or
obscurely or normally developed, (0.75-) - .5(- .75) mm long
I 1 1 4. S. parviflora
2. Leaves elliptic-oblong to sublanccolatc, 5-25 by 2- 10 cm, beneath usually somewhat paler than above

but not conspicuously so; nerves 4 8(-9) pairs, ± patent, straight to curved; cymes often arranged in
an up to 15 cm long racemose to thyrsoid inflorescence, (l-)2- I0( l2)-nowercd, sometimes solitary, ax-
illary, up to 4( 6)-nowcred; style normal-developed, 0.2- mm long. I

3. Cymes up to 2 cm, 4( 6)-nowcrcd; petals suborbicular to elliptic, 1.75 2.5 by 1.25-2 mm, obtuse
1
684 Flora Malesiana [ser. I, vol. 10^

to rounded; stamens nearly as long as petals; drupelets globular to obovoid, very compressed, 1 1 - 14 by
10-13 mm; reticulate pattern usually faint or absent 3. S. limoniacea
3. Cymes up to 1 cm, l-4(-7)-nowered; petals either oblong-ovate to ovate-lanceolate, acute, acuminate
or not, or elliptic-oblong to oblong, obtuse, 3.5-6.5 by (1 .25-)l .5-2.5 mm; stamens distinctly shorter

than petals; drupelets obovoid, ± compressed, c. 10-12by(7-)8- 10 mm; reticulate pattern rather faint

but usually visible, often limited to the margin 6. S. racemosa

3. Cymes up cm, (l-)2-10(-12)-flowered; petals oblong, 2.5-4(-4.5) by c. 1-1.5 mm, obtuse;

to 2(-3. 5)
stamens distinctly shorter than petals; drupelets obovoid or ± globular, ± compressed, 7.5-11 by
8-10(-ll) mm; reticulate pattern usually clearly visible, sometimes obscure, limited to the margin or
not.
4. Leaves oblong to sublanceolate, 5- 14(- 8) by 2-6(-8) cm; nerves (5-)6-8(-9) pairs; cymes either ar-
1

ranged in a racemose to thyrsoid inflorescence, or solitary, axillary, 1-4-flowered; style 0.6-1 mm;

drupelets ± globular, sometimes somewhat obovoid, compressed, 7.5-11 by 8-10(-ll) mm

5. S. pauciflora

4. Leaves elliptic-oblong to oblong, sometimes sublanceolate, 6-19 by 2-8(-10) cm; nerves 4-7(-8)
pairs;cymes usually arranged in a thyrsoid inflorescence, sometimes subtended by small leaves,
3- 10(- 12)- flowered; style 0.2-0.5 mm; drupelets obovoid, sometimes globular, somewhat com-
pressed, 9-11 by c. 9-10 mm 2. S. javanica

1. Sabia erratica van de Water, Blumea 26 (1980) 2. Backer ex Chen, Sargen-

Sabia javanica (Blume)
35. tia 3 Backer & Bakh./. F1. Java 2 (1965)
(1943) 59;
Evergreen, woody. Twigs glabrous to somewhat 144; VAN DE Water, Blumea 26 (1980) 39. -
pubescent; flowering twigs up to 2.5 mm diam., ± Meniscosta javanica Blume, Bijdr. (1825) 29. -
lax-pubescent. Buds ovoid, acute; scales ± pubes- Meniscosta scandens Blume ex Spreng. Syst. Veg. 4,
Leaver oblong, 5-8 by 2.5-3 cm, in-
cent, ciliolate. 2 (1827) 114, nom. illeg.; Dietr. Syn. PI. 2 (1840)
dex 2-2.7, pergamentaceous, above glabrous or still 923.- Sabia meniscosta Blume, Mus. Bot. Lugd.-
sparsely pubescent especially at the base and on Bat. (1851) 369, f. 44, nom. illeg., incl. var. firma
1

midrib, beneath laxly pubescent especially on midrib Blume, var. latifolia Blume et var. glabriuscula
and nerves; base acute, apex acute or short-acumi- Blume; Miq. F1. Ind. Bat. 1, 2 (1859) 6 1 8 ( 'me«/co5-
nate; nerves 6-7 pairs, patent, ± .straight to some- tay, Fl. Arch. Ind. (1870) 71; //j/c^. (1871) pi. 31, /nc/.

what curved; petiole up to 1 .5 cm, glabrous to pubes- var. elliptica Miq.; Hook./. Fl. Brit. India 2 (1876)
cent. Cymes solitary, axillary up to 4.5 cm, up to 3 ('menescortay. Backer, Schoolfl. Java (1911) 273;
40-flowered, ± lax-pubescent; pedicels up to 4 mm; KooRD. Exk. Fl. Java 2 (1912) 544. - Sabia elliptica
bracteoles oblong to oblong-ovate, up to 0.8 mm, (Miq.) Miq. Sum. (1861) 203, 521. - Sabia javanica
pubescent, ciliolate. Sepals ovate to somewhat ellip- ^lume) Chen var. glabriuscula (Blume) Chen,
tic, 0.8-1 by 0.5-0.75 mm, obtuse to acute, ± Sargentia 3 (1943)61.
pubescent, ciliolate. Petals oblong or oblong-ovate Evergreen woody climber or scandent shrub, up to
to sublanceolate or ovate-lanceolate, 3.75-4 by 10 m. Twigs glabrous; flowering twigs up to 5 mm
1-1.5 mm, acute to narrow-obtuse, subciliolate, diam., glabrous or ± pubescent. Buds ovoid, up to
nerves up to 6, dark-coloured. Stamens 2.3-3 mm; 2 mm, acute; scales glabrous or with few hairs, ±
filament flattened, 1 .8-2.6 by 0.25-0.4 mm; anther Leaves elliptic-oblong to sublanceolate,
ciliolate.
ellipsoid to oblong-ellipsoid, c. 0.4-0.6 mm, 6-19 by 2-8(-10) cm, index 2-3(-4), pergamen-
upright. Disk crown-shaped; lobes very short or ab- taceous to pergamentaceous-coriaceous, above and
sent; ribs ± prominent. Pistil 2.75-3 mm; style beneath glabrous or with some hairs on midrib; base
narrowly-conical to cylindrical, 2.25-2.5 mm, with acute to rounded, apex acute, acuminate; nerves
some hairs at the base; ovary somewhat globular to 4-7(-8) pairs, patent, curved to straight; petiole up
subreniform, 0.5-0.6 by 0.6-0.8 mm, densely to 2.5 cm, glabrous to sparsely pubescent, ± (fine-)
pubescent. Drupelets not available. wrinkled. Cymes arranged in an axillary, up to 12 cm
Distr Malesia: Singapore (Bt. Timah Res.), only
. long, glabrous to pubescent, thryrsoid inflorescence,
known from the type, collected in 1940. subtended by bracts or sometimes by small leaves
Notes. In habit somewhat resembling S. par- and then inflorescence up to 17 cm long; cymes up to
viflora but readily distinguished by floral characters. 3 cm, forming a lax to dense cluster of 3-10(-12)
On the label noted as a 'tree, 100 ft', but this is flowers, subglabrous to pubescent. Bracts ovate to
suspected to be a wrong annotation or field observa- sublanceolate, up to 5 mm, subglabrous to more or
tion or a wrong label. less pubescent, ± ciliolate; bracteoles as bracts but
1989] Sabiaceae (van Beusekom & van de Water) 68S

and then
smaller, or bracteoles minute or sepal-like 3. Sabia limoniacea Wall. [Cat. (1829) n. 1000,
situated near calyx; pedicel up
Flowers to 4 mm. nom. nud.] ex Hook./. & Th. Fl. Ind. 1 (1855) 210;
green to yellow or white. Sepals sometimes 6 (see Walp. Ann. 4 (1857) 139; Benth. Fl. Hongk. (1861)
bracteoles), ± ovate or broad-ovate, 0.75- 1{- 1.25) 70; Hook./. Fl. Brit. India 2 (1876) 3; Kurz, J. As.
by 0.5-0. 8(-l) mm, acute to obtuse, ± pubescent, Soc. Beng. 45, ii (1876) 204, excl. syn. Sabia sp.
ciliolate. Petals oblong, 2.5-3.5(-4) by 1-1.5 mm, Griffith (= S. parvijlora ssp. parviflora); For. Fl.
obtuse, nerves up to 5, often dark-coloured and then Burma 1 (1877) 300 {'timonaceay, Forbes &
conspicuous. Stamens (1-)1.25-1.5 mm; filament Hemsley, J. Linn. Soc. Bot. 23 (1886) 144; King, J.

± flattened, (0.75-)l-1.25 mm long, 0.25-0.5 mm As. Soc. Beng. 65, ii (1896) 454; Prain, Beng. PI. I

wide; anther globular to ellipsoid, 0.2-0.3 mm, in- (1903) 246; Brandis, Indian Trees (1906) 194; Dunn
flexed. Disk crown-shaped; ribs sometimes faint or & Tutcher, Kew Bull. Add. Ser. 10 (1912) 68;
absent. P/5/// 0.8-1.2 mm; style ± conical, 0.2-0.5 Ridley, Fl. Mai. Pen. 1 (1922) 513; Merr. Lingnan
mm, much shoner than the adjacent side(s) of the Sc. J. 5 (1927) 19; Kanjilal c.s. Fl. Assam 1,2 (1936)
drupelet(s); ovary globular to subreniform, 0.5-0.6 326; Chen, Sargentia 3 (1943) 56, f. 7; Biswas, PI.
by 0.5-0.7 mm, glabrous. Drupelets obovoid or Darj. Sikkim Himal. van de Water,
1 (1966) 261;
sometimes globular, ± compressed, 9-11 by 9-10 Blumea 26 (1980) 44, / 6b, 8. - Androglossum
mm, without persistent petals and stamens at the reticulatum Champ, ex Benth. Hook. J. Bot. Kew
base; reticulate pattern often coarse and limited to Gard. Misc. 4 (1852) 42; Benth. Fl. Hongk. (1861)
the margin. Embryo with somewhat undulated or 70; Chen, Sargentia 3 (1943) 58, non S. reticulata
faintly folded cotyledons. Elmer (1909). - Sabia celastrinea Muell. in Walp.,
Distr. Malesia: Sumatra (East Coast Res., In- Ann. 6 (1865) 1269. - Sabia malabarica Bedd. Ic.
dragiri,Lampongs), W. Java. In all c. 30 collections. PI. Ind. Or. 1 (1874) 39. t. 177; HooK./ Fl. Brit. In-
Ecol. Forests, at (20-)200-1500 m. Ft. fr. dia 2 (1876) 2; Brandis, Indian Trees (1906) 194;
Jan. -Dec. Gamble, Fl. Pres. Madras 1 (1918) 254; Chen,
Vern. Java: areuj bebentjojan, a. kahawatang, Sargentia 3 (1943) 48. - Fig. 2, 3.
a. katjapi, S. Evergreen woody climber, up to 10 m. Twigs
Notes. Sabia javanica strongly resembles S. glabrous or sometimes sparsely pubescent; flowering
pauciflora from the Philippines, the Moluccas, New twigs up to 5 mm diam., glabrous to lax-pubescent.
Guinea, and the Solomon Islands. It can be Buds broad-ovoid to ovoid, up to 2.5 mm, acute;
distinguished from that species by its often more- Leaves oblong-
scales (sub)glabrous, often ciliolate.
flowered cymes, its shorter style, and some other ovate to lanceolate, 4-18 by 1.5-6.5(-8) cm, index
slight differences. Since both species are geographi- 2-4(-4.5), ± pergamentaceous-coriaceous, above
cally separated, it was also possible to combine them and beneath glabrous or with some hairs especially
into one species and give them the rank of subspecies. on midrib; base acute to rounded, apex acute,
Although the differences are rather small, 1 believe sometimes obtuse, acuminate or not; nerves 5-9
that S. javanica and 5. pauciflora represent two dif- pairs, ± patent, sometimes somewhat ascending,
ferent, well-delimited, but very closely related spe- curved to straight; petiole up to 2.5 cm, glabrous to
cies.Moreover, a reduction of both species to a single lax-pubescent. Cymes either solitary, axillary,
one would increase the variability of several taxo- subtended by small and often herbaceous leaves, or
nomic important characters, in consequence of when either the leaves are fallen or the cymes are
which the delimitation with some other related subtended by bracts arranged in an up to 1 5 cm long,
species, like S. parvijlora and 5. racemosa, and glabrous to ± lax-pubescent or tomentellous, race-
possibly also S. limoniacea, would become less mose to thyrsoid inflorescence, cymes up to 2 cm,
distinct. Finally, this might result into a far-going l-4(-6)-nowered; pedicels up to 7 mm; bracts
lumping and a reduction of all these species to, say, oblong, up to 4 mm, glabrous to pubescent, ciliolate;
subspecies. Contrary to the situation in the extra- bracteoles ovate to oblong, up to .75 mm, glabrous 1

Malesian species .S. campanuiata Wall., however, in to pubescent, ciliolate, often situated near calyx.
this case believe that the differences between these
I Flowers green to yellow or white. Sepals sometimes
taxa have reached a higher level already, resulting in 6 or 7 (see bracteoles), broad-ovate to elliptic,

the distinction of mutually closely related but ± 0.7- .2(-

1 1 .5) by 0.6- I mm, acute to rounded, gla-
well-delimited species, each with its own specific brous to ± pubescent, ciliolate. Petals suborbicular
combination of characters. to elliptic or ± obovate, 1.75-2.5 by 1.25-2 mm,
In vegetative characters and in drupelets S. obtuse to rounded, sometimes broad-acute, nerves 5,
javanica resembles .S. racemosa from Borneo. It can, usually obscure. Stamens 1.5-2 mm; filament
however, easily be distinguished from that species by somewhat flaticncd, 1 .25- 1 .75 by 0.3-0.4 mm; an-
its morc-flowcred inflorescences and its floral ther ellipsoid, 0.25-0.35 mm, inflexed. Disk crown-
characters, es{>ccially its petals. shaped, thin; ribs often faint or absent. Pistil
686 Flora Malesiana [ser. I, vol. lO^

revel
1977

Fig. 2. Sabia limoniacea Hook./. & Thoms. a. Habit, x2/3; b. ditto, with axillary cymes, x2/3; c. open
flower, x4; d. petal and the opposed stamen, x 8; e. disk and pistil, x8 (a & c-e C.W. Wang 79409; b
Wallich 1000).
1989] Sabiaceae (van Beusekom & van de Water) 687

up to 4 mm
diam., glabrous to pubescent. Buds
broad-ovoid to ovoid, up to 2 mm, acute; scales
glabrous to short-pubescent, ciliolate. Leaves oblong
to (sub)lanceolate, 3-12(-15) by 1-5 cm, index 2-4
(-4.5), ± pergamentaceous, above glabrous to
subglabrous or sometimes sparsely pubescent espe-
cially when young, beneath glabrous to lax-pubes-
cent especially on midrib; base acute to rounded, at-
Fig. 3. Sabia limoniacea Hook./. & Thoms. a. fruit; tenuate or not; apex acute, acuminate; nerves (5-)
a', embryo, both xl.5 (a Poilane 24769; a' 6-9(-10) pairs, patent, straight or sometimes ±
POILANE 18918). curved; petiole up to 1.5 cm, glabrous to mainly
above lax-pubescent. Cymes solilary, axillary, 1.5-8
(-10) cm long, 4-25-flowered, sometimes widely
0.7-1.2mm; style conical to cylindrical, 0.2-0.6 spreading, lax, and with up to 35 or more flowers,
mm, much shorter than the adjacent side(s) of the glabrous to sparsely pubescent; pedicels up to 1 cm;
drupelet(s); ovary globular to subreniform, 0.5-0.6 bracts ovate to lanceolate, up to 2 or, when sub- mm
by 0.5-0.8 mm, glabrous. Drupelets globular to tending a cyme up to 6 mm, subglabrous to pubes-
obovoid, strongly compressed, 11-14 by 10-13 mm, cent, ciliolate; bracteoles as bracts. Flowers green to
red to blue or black when fresh, without persistent yellow or white. Sepals broad-ovate to ovate,
petals and stamens at the base; reticulate pattern 0.7- 1 .5 by 0.5- 1 mm, acute to rounded, glabrous to
usually faint or absent, sometimes more prominent pubescent, ciliolate. Petals elliptic-oblong to lanceo-
at the margin. Embryo with somewhat undulated late or sometimes oblong-ovate, 2-4(-4.5) by
cotyledons. 0.7-1.3 mm, acute to obtuse; nerves up to 7, dark-
Distr. Continental SE. Asia (throughout India, coloured or sometimes obscure. Stamens 1.2-2.25
Burma, Bangladesh, Thailand and Indochina to (-2.5) mm; filament flattened, 0.9-2(-2.25) by
China); in Malesia: Malay Peninsula (incl. also P. 0.25-0.5 mm; anther ellipsoid to ovoid, 0.25-0.4
Penang), Central Sumatra and Borneo (Sarawak), in mm, often ± inflexed. Disk crown-shaped, usually
all 7 collections. and narrow,
thin; lobes often distinct, relatively long
Ecol . Thickets and forest, 300-1200 m altitude. sometimes short or margin of disk irregular; ribs
FL Sept. -Jan.,//-. Dec. -April. often faint or absent. Pistil l-2(-2.5) mm; style
either absent or obscure, or conical, (0.75-)l-1.5
4. Sabia parvinora Wall,
in Roxb., Fl. Ind. 2 (1824) (-1.75) mm, much shorter than the adjacent side(s)
310; G.Do.M, Gen. Hist. 2 (1832) 69; Walp. Rep. 1 of the drupelet(s); ovary globular to subreniform,
(1842) 557; Hook./ & Th. Fl. Ind. (1855) 210; 1 0.4-0.7 by 0.5-0.75 mm, glabrous. Drupelets glob-
Walp. Ann. 4 (1857) 139; Hook./ Fl. Brit. India 2 ular to somewhat obovoid, ± compressed, 7-9 by
(1876) 2; Stapf, Trans. Linn. Soc. Lond. 4, 2 (1894) 6-8 mm, green to red or blue when fresh, without
142; Bra.ndis, Indian Trees (1906) 194; Lecomte, Fl. persistent petals and stamens at the base; reticulate
Gen. I.-C. 2 (1908) 2, incl. var. harmandiana Le- pattern rather fine, but often inconspicuous or ob-
comte, Bull. Soc. Bot. Fr. 54 (1907) 674; Kanjilal scure. Embryo with faintly wrinkled cotyledons.
Assam 1, 2 (1936) 325; Chun, Sunyatsenia 4
c.s. Fl. Distr. Widely ranging in SE. Asia from Nepal to
(1940) 242; Merr. Brittonia 4 (1941) 112; Chen, China; in Malesia: N. Borneo (Sabah) and the Philip-
Sargentia 3 (1943) 64; Gagnep. & Vidal, Fl. Camb. pines (Luzon).
Laos, Vietnam 1 (1960) 16; Biswas, PI. Darj. Sikkim
Himal. 1 (1966) 261; Sen Gupta, Bull. Bot. Soc. KEY TO the subspecies
Bcng.22, ii(1968) 196; Hara. Fl. E. Himal. 2 (1971)
74; Sen Gupta, Rec. Bot. Surv. India 20, 2 (1973)65; I. Style normally developed, distinctly conical,
Hah A & Williams, Enum. Fl. PI. Nepal 2(1970) 100; (0.75-)l-1.5(-1.75) mm long
VAs de Water. Blumea 26 (1980) 48, f. 3c, 9. - a. ssp. parviflora
\al)ia harmandiana Pierre, Fl. For. Coch. 5 (1897) 1. Style usually absent or obscure, the upper part of
pl 360B; Craib, Fl. Slam. Enum. (1926) 340. - I the pistil carpel-like, sometimes normally devel-
Sahia phitippinensis Robins. Bull. Torrcy Bot. Club oped and then up to 0.75 mm
'
I9(J8) 70; .Merr. Enum. Philip. 2 (1923) 516; b. ssp. philippinensis
•.. Sargentia 3 (1943) 67.
" a complete synonymy, see van de Water a. .w/7. parviflora - Sabia parviflora Wall. - Sabia
'M harmandiana Pii.rri. .

crgrccn climber or scandcnt shrub, up to 6 m. leaves oblong, somclimcs oblong-ovatc to (sub)

'.'."> glabrous to laxly pubescent; flowering (wigs lanceolate, 3~12(-I5) by 1-5 cm. C>mfi 2-8(-I0)
688 Flora Malesiana [ser. I, vol. 104

cm long, 7-25-flowered, sometimes widely spread- especially conspicuous. In ssp. philippinensis the
and with up to more than 35 flowers. Petals
ing, lax, pale margins and undersides of the leaves provide a
oblong to lanceolate, sometimes oblong-ovate, useful character to distinguish vegetative specimens
2.25-4(-4.5) by 0.7-1.25 mm. Style distinctly con- from those of S. pauciflora, another Philippine
ical, (0.75-)l-1.5(-1.75) mm long. species
Distr. SE. Asia; in Malesia: Borneo (Sabah), 9
collections. 5. Sabia pauciflora Blume, Mus. Bot. Lugd.-Bat. 1

Ecol. Roadsides, in thickets, and in forests, (1851) 370; MiQ. Fl. Ind. Bat. 1, 2 (1859) 619; Fl.

mainly 600-2000 m altitude. Fl. Jr. probably Arch. Ind. (1870) 72; ibid. (1871) pi. 32; Chen,

throughout the year. Sargentia 3 (1943) 61; van de Water, Blumea 26

(1980) 51. - Sabia papuana Warb. in K.Sch. &
b. ssp. philippinensis (Robins.) van de Water, Laut., Fl. Deut. Schutzgeb. Sudsee (1900) 425. -
Blumea 26 (1980) 50. - Sabia philippinensis Robins. Sabia reticulata Elmer, Leafl. Philip. Bot. 2 (1909)
- Fig. 4. 579; Merr. Enum. Philip. Fl. PI. 2 (1923) 516;
Leaves oblong or oblong-ovate to lanceolate, Chen. Sargentia 3 (1943) 62.
3-11 by 1-3.5 cm. Cymes 1.5-4.5 cm, 4-20-flow- Evergreen woody climber or scandent shrub, up to
ered. Petals elliptic-oblong to sublanceolate, 2-3.5 20 m. Twigs glabrous; flowering twigs up to 5 mm
by 1-1.25 mm. and often
Style absent or obscure diam., glabrous or sparsely pubescent. Buds ovoid,
carpel-like, sometimes normal-developed and then up to 2.5 mm, acute; scales glabrous to pubescent,
up to 0.75 mm. Fruits not seen. (sub)ciliolate. Leaver oblong to sublanceolate, 5-14
(-18) by 2-6(-8) cm, index (2-)2.5-3.5(-4), above
and beneath glabrous or with very few hairs on
midrib, pergamentaceous; base acute to rounded,
apex acute, acuminate; nerves (5-)6-8(-9) pairs,
patent, straight to curved; petiole up to 2 cm, gla-
brous to sparsely pubescent. Cymes either arranged
in an axillary, up to 12 cm long, glabrous to sparsely
pubescent, racemose to thyrsoid inflorescence, sub-
tended by bracts, or solitary, axillary, often subtend-
ed by small leaves, up to 3.5 cm, 1-4-flowered, gla-
Fig. 4. Sabia parviflora ssp. philippinensis (Robin-
brous to sparsely pubescent; bracts oblong to
son) VAN DE Water, a. & b. disk and pistil showing
lanceolate, up to 3.5 mm, subglabrous to somewhat
the absence of a style; c. a feebly developed one; all
pubescent, (sub)ciliolate; bracteoles as bracts but
X 12 (fl Ramos 26973; b Jacobs 7402; c Merrill
smaller, or minute, or sepal-like and then often
7708).
situated near calyx; pedicels up to c. 1 cm. Flowers
green to yellow or white. Sepals sometimes 6 (see
Distr. Malesia: Philippines (Luzon: Benguet bracteoles), ovate to broad-ovate, 0.75-1.25 by
Prov.), 11 collections. 0.7-1 mm, acute to obtuse, glabrous to somewhat
Ecol. Forests, 71000-2100 m. Fl. mainly pubescent, (sub)ciliolate. Petals oblong, sometimes
Febr. -April. somewhat oblong-ovate, 2.5-4(-4.5) by (0.75-)
Vern. Baybayok, kopdas, uakal, udok, Ig. 1-1.3(-1.5) mm, (narrow-)obtuse, sometimes sub-
Notes. Ssp. philippinensis can be distinguished ciliolate, nerves up to
5, sometimes dark-coloured
rather easily from by the absence of
ssp. parviflora and then conspicuous. Stamens (l-)1.25-1.75(-2)
a normally developed style. In all the specimens I mm; filament flattened, (0.75-)l-1.5(-1.75) by
have seen (except one) the upper parts of the two 0.25-0.5 mm; anther globular to ellipsoid, 0.2-0.3
carpels of each flower are not connate with each mm, Disk crown-shaped; lobes often short
inflexed.
other and differentiated into a style as usual, but re- or irregular; ribs sometimes faint or absent. Pistil
main free and carpel-like, although the tip of each 1.3-1.7 mm; style conical, 0.6- 1 mm, much shorter
carpel is sometimes slightly stigmatic. Moreover, the than the adjacent side(s) of the drupelet(s); ovary
margins of the upper part of a carpel are not fused, globular to subreniform, 0.5-0.7 by 0.5-0.8 mm,
so that the upper half of each carpel remains open. glabrous. Drupelets ± globular, sometimes some-
Although this phenomenon is unique within the what obovoid, compressed, 7.5-11 by 8-10(-ll)
genus, I have reduced S. philippinensis to a subspe- mm, white to red or dark-blue when fresh, without
cies of S. parviflora because it agrees very well with persistent petals and stamens, reticulate pattern fine
that species in all other main characters. to rather coarse, sometimes indistinct, limited to the
Like Sabia species the leaves are dark above,
in all margin or not. Embryo with somewhat undulated or
paler beneath, but in the present one the contrast is faintly folded cotyledons.
1989] Sabiaceae (van Beusekom & van de Water) 689

Distr. Malesia: Moluccas (Buru, Halmaheira, reticulate pattern faint to rather coarse, often limited
Batjan), Philippines (Luzon, Negros, Mindanao), to the margin. Embryo with somewhat to very
New Guinea; Solomon Islands. wrinkled or folded cotyledons.
Ecol. Forests, from sea-level up to 2300 m. Fl.fr. Distr. Malesia: Borneo.
throughout the year. Note. In vegetative characters and somewhat in

Uses. Fresh leaves eaten against wound fever in the fruit this species resembles S. javanica. It differs,

New Guinea. however, from that species in its inflorescence (few-

Vern. Philippines: bungoi, dadabu. Bag.; New flowered cymes) and in its floral characters, especial-
Guinea: hambui, Poio, Enga lang., kubiakan, ly the petals.
Hagen-Chimbu, Yoowi dial., mongotya ka. North- Since the fruiting collections oi ssp. racemosa bear
ern Prov., pehkuma, Mumuni, Orokaiva lang, pipi, only immature or damaged fruit, the description of
E. Highlands, pukhabu, S. Highlands. the drupelets has mainly been based on the fruit of
Note. This species is closely related to S. javanica ssp. kinabaluensis.
from Java and Sumatra, but can be distinguished The two subspecies can easily be distinguished
from that species by its always few-flowered cymes, from each other by the difference in the shape of
its longer style, and its often ± globular drupelets (S. their petals. Since they can be distinguished from
javanica often obovoid). each other only when flowers are available, the iden-
tification of most of the vegetative and fruiting
6. Sabia racemosa Chen, Sargentia 3 (1943) 36, f. 2; specimens has mainly been based on the locality from
VAN DE Water, Blumea 26 (1980) 54. where they have been collected.
Evergreen woody climber or scandent shrub, up to
6 m. Twigs glabrous; flowering twigs up to 4 mm KEY TO THE SUBSPECIES
diam., glabrous or somewhat short-pubescent. Buds
ovoid, up to 1.5 mm acute; scales (sub)ciliolate or 1. Petals oblong-ovate to ovate-lanceolate, acute,
not. Leaves ob\ong or somewhat oblong-ovate, 6-25 somewhat acuminate or tapering to the apex
by 2-10 cm, index 2-3(-3.5), pergamentaceous, a. ssp. racemosa
glabrous or with some hairs on midrib, rarely 1. Petals elliptic-oblong to oblong, acute to obtuse
beneath all over sparsely short-pubescent; base acute b. ssp. kinabaluensis
to rounded, apex acute, acuminate; nerves 4-8 (or 9)
pairs, ± patent, curved to straight; petiole up to 2.5 a. ssp.racemosa - Sabia racemosa Chen.
cm, glabrous or with some very short hairs. Cymes Sepals 0.6-
1.1 by 0.5- mm. Petals oblong-ovate
1

arranged in an axillary, up to 8 cm long, glabrous to to ovate-lanceolate, (3.5-)4.5-6.5 by (1.25-)1.5-

puberulous or short-tomentellous, racemose to thyr- 2.5 mm, acute, somewhat acuminate or tapering to
soid inflorescence, subtended by bracts but often the apex. Pistil 1-1.2 mm; style 0.5-0.7 mm long.
bracts fallen or sometimes leaf-like, cymes up to 1 Distr. Malesia: Borneo (Kalimantan), 7 collec-
cm, -4(-7)-nowered, glabrous to somewhat puber-
I tions.
ulous or short-tomentellous; bracts ovate to oblong, Ecol. Low altitudes, up to 100 m. Fl. fr.
up to 3 mm, glabrous to somewhat pubescent, throughout the year.
(sub)ciliolate; bracteoles as bracts but usually smal-
ler, and then often situated near calyx;
or minute b. ssp. kinabaluensis van de Water, Blumea 26
pedicelsup to 4 mm. Flowers (pale-)green to yellow. (1980) 55.
Sepals ± ovate to broad-ovate, 0.6-1.3 by 0.5-1 Sepals 0.9-1.3 by 0.6-1 mm. Petals elliptic-
mm, acute to obtuse, glabrous to somewhat pubes- oblong to oblong, 3.5-5 by 1.5-2.5 mm, acute to
cent, (sub)ciliolate. Petals elliptic-oblong to ovate- obtuse. Pistil 1.2-1.5 mm high; style 0.6-0.9 mm
lanceolate, 3.5-6.5 by(1.25-)l.5-2.5 mm, acute to long.
obtuse, or acuminate, or gradually narrowed,
± Distr. Malesia: Borneo (Sabah: Mt Kinabalu),
nerves up to 7, thin but distinct. Stamens 1.2-2.2 15 collections.

mm; filament flattened, i -2 by 0.2-0.5 mm; anther Ecol. Forests, mainly al 800- 500 1 m altitude. Fl.
globular to ellipsoid, 0.2-0.3 mm, inflexed. Disk fr. throughout the year.
crown-shaped; lobes sometimes very short or in-

distinct; ribs sometimes faint or absent. Pistil i - 1.5 7. Sabia sumalruna Biumi., Mus. Bot. Lugd.-Bat. I

mm; style ± conical, 0.5-0.9 mm, much shorter (1851) 370; Miy. II. Ind. Bat. 1, 2 (1859) 619; Fl.
than the adjacent side(s) of the drupclct(s); ovary Arch. Ind.(1870)72;;7>/ry. (187l)pl. 33; Kino, J. As.
globular to subrcniform, 0.5-0.6 by 0.5-0.7 mm, Soc. Beng. 65, ii (1896) 454; Ridley, Fl. Mal. Pen.

glabrous. Drupe/e/5 obovoid, ± compressed, 10-12 1(1922) 513; Chen, Sargentia 3 (1943) 39; van de
by (7- )8- 10 mm, while to pink or red when fresh, Water. Blumea 26 (1980) 56.
without persistent petals and stamens al the base; Evergreen woody climber, up to c. 3.5 m. Twigs
690 Flora Malesiana [ser. I, vol. 10"^

glabrous; flowering twigs up to 4 mm diam., globular to subreniform, 0.5-0.8 by 0.7-1 mm,

glabrous. Leaves elliptic to oblong, sometimes (sub) glabrous. Drupelets obovoid, somewhat compress-
lanceolate, (5-)7- 15(- 18) by (1 .5-)2.5-7(- 10) cm, ed, 11-13 by 8-9 mm, white to blue when fresh,

index 2-3(-4), pergamentaceous, above and be- without persistent petals and stamens, reticulate pat-
neath glabrous; base acute, apex acuminate to sub- tern absent, often more or less rugged on the outside.

cuspidate; nerves 5-7 pairs, patent, curved to Distr. Malesia: Sumatra (W. Coast Res., Palem-
straight; petiole up cm, glabrous. Flowers yel-
to 2 bang), 7 collections.
lowish-green to white, either solitary, sometimes 2 or Ecol. Forests, 60-1000 m altitude. Fl. May-
3 together, axillary, or arranged in a thyrsoid, ax- Aug.,//-. July-Sept., Febr.
illary, up to 6.5 cm long, glabrous inflorescence; Note. Only a few collections are available. For
pedicels up to 2.5 cm, glabrous, with few small that reason no buds and embryos could be described,
budscales at the base when flowers solitary; bracts ± whereas the description of the flowers has partly
oblong-ovate, up to 1 .5 mm long, glabrous, ciliolate; been based on rather young ones.
bracteoles as bracts. Sepals broad-ovate to ovate,
1.25-1.75(-2) by (0.75-)l-1.75 mm, acute to ob- Excluded
tuse, glabrous, (sub)ciliolate or not. Petals oblong or
ovate-lanceolate, c. 6- 10 by .5-2.5 mm,
1 sometimes Sabia densiflora Miq. Sum. (1861) 203, 520 =
the upper part somewhat channeled, tapering to the Meliosma angulata Blume: K. & V. Bijdr. 9 (1903)
apex, acute to narrow-obtuse, nerves obscure. 131 = Meliosma simplicifolia (Roxb.) Walp. ssp.
Stamens 2.5-1.5 mm; filament ± flattened, 3-7 by simplicifolia: van Beusekom, Blumea 19 (1971) 476;
0.4-0.75 mm; anther ellipsoid, 0.5-0.7 mm, Fl. Males. 10" (1989) 698 (this issue).

upright. Disk short-cylindrical, small, the upper part

not enclosing the base of the ovary and without Sabia floribunda Miq. Sum. (1861) 203, 521 =
lobes; ribs ± prominent. Pistil 3.5-c. 7 mm; style Meliosma angulata Blume: K. & V. Bijdr. 9 (1903)
narrow-conical, 3-6 mm, ± half as long as the adja- 131 = Meliosma simplicifolia (Roxb.) Walp. ssp.
cent side(s) of the drupelet(s); ovary somewhat simplicifolia : /. c.

2. MELIOSMA

Blume, Cat. (1823) 32; Rumphia 3 (1849) 196; Miq. Fl. Ind. Bat. 1 2 (1859) 612 ,

Benth. & Hook./. Gen. PI. 1 (1862) 414; Hook./. Fl. Brit. India 2 (1876) 3
BoERL. Handl. Fl. Ned. Ind. 1 (1890) 290; Warb. in E. & P., Nat. Pfl. Fam
3, 5 (1895) 371; van Beusekom, Blumea 19(1971)355. - Millingtonia Roxb
[Hort. Beng. (1814) 3, nomen] PI. Corom. 3 (1820) 50, t. 254, non Linn./
(1781), nee Donn (1807). - Kingsboroughia Liebm. Vid. Medd. Nat. For
Kjobenhavn 2 (1850) 67; Walp. Ann. 2 (1852) 834. - Fig. 5-8, 10, 12.
For a complete synonymy, see van Beusekom (1971).
Evergreen or sometimes deciduous shrubs or trees, up to 42 m, 1 m diam.,
sometimes buttressed. Twigs more or less lenticellate, often with conspicuous
leaf-scars. Buds densely pubescent. Leaves simple or imparipinnate with
(sub)opposite leaflets, ending in 3 or 1 leaflet(s), in the latter case its petiolule
articulated with the rachis; leaves or leaflets entire or dentate, with or without
hairy domatia beneath; rachis and petioles, usually also petiolules, with a usual-
ly shallow and narrow, more or less conspicuous longitudinal groove above,
usually with swollen base, articulately attached. Inflorescence terminal, some-
times axillary, a pyramidal panicle, poor to usually profuse, up to 4 times
ramified, with alternate, articulately attached, often lenticellate axes. Bracts
small, those of lower order usually soon caducous; cataphylls often present.
Bracteoles absent, but sometimes one (or two) bracteole-hke sepals present.
1989] Sabiaceae (van Beusekom & van de Water) 691

Flower of Meliosma. A. Semi-diagrammatical sketch of flower (subg. Meliosma) with opened outer
Fig. 5.
stamens still in bud position. B. Semi-diagrammatical length section of bud (subg. Meliosma). C.
petals, but
Diagram {subg. Kingsboroughia and subg. Meliosma). Names of the flower parts: a. sepals; b. outer petals;
c. inner petals; d. fertile stamens; e. staminodes; /. disk; g. style.

lowered on the pedicel. Flowers numerous, sessile or short-pedicelled, small,

bisexual. Sepals 5, by reduction sometimes 4, rarely 3, sometimes by addition
of empty bracts seemingly more, up to c. 13, and together forming a kind of in-
volucre, usually unequal and then mostly 3 about equal. Petals 5, episepalous,
3 outer ones more or less unequal, alternisepalous, mostly suborbicular and con-
vex, rarely the largest one much wider than long and more or less reniform, the
smaller ones irregularly shaped; 2 inner ones equal, much smaller, reduced, op-
posite the fertile stamens and more or less adherent to the base of the filaments,
entire to bifid. Disk generally present, sometimes very reduced or absent, often
irregularly shaped, as a rule with 5 more or less developed teeth, 4 of which
paired, 1 unpaired, each pair opposite a fertile stamen. Stamens 5, epipetalous,
2 fertile, filament short, strap-shaped, flat, incurved at the top, abruptly ter-
minating in a wide, varyingly shaped cup which bears two globose to elliptic
transversely dehiscent anther-cells which are ripe in bud, springing back elasti-
cally when the flower opens; 3 staminodial, opposite the larger petals and more
or less adherent to the base of these, deformed, broad, irregularly shaped, with
1 or 2 holes near the top in which fit the anther-cells of the fertile stamens, often
coherent and forming a cup over the pistil. Ovary globose to ovoid or conical,

2-, very rarely 3-locular, apically contracted in a rather short, simple or 2-

parlible, cylindric or subulate to conical, rarelyminute style, with simple or
somewhat minute stigma. Ovules 2 (or 1) in each cell, more or less
bifid,
superimposed, attached to the partition, hemi-anatropous. Fruit a drupe,
subglobose to pyriform, small, glabrous, with one stone; rarely two ovules in-
stead of one per ovary develop, resulting in a didymous fruit; mcsocarp pulpy,
mostly thin; endocarp globose, pyriform, or semiglobose, 1-cellcd, stony to
crustaceous, splitting in two valves, inside with a basilar rounded projection
over which the seed is curved. Vascular bundle connecting pedicel and seed
either running outside the endocarp wall (free in the pulpy mcsocarp or in a
groove on the ventral endocarp wall), or through a canal inside the endocarp
692 Flora Malesiana [ser. I, vol. 10^

Fig. 6. Diagrammatical length sections of three types of fruit in Meliosma. A. Subg. Kingsboroughia sect.
Hendersonia: vascular bundle running freely in the mesocarp. B. Subg. Kingsboroughia sect. Kings-
boroughia: vascular bundle running in a groove of the endocarp, entering the wall through the ventral pore.
C. Subg. Meliosma sect. Meliosma: similar to B, but the marginal canal lengthened through the endocarp.
All x3.
 1989] Sabiaceae (van Beusekom & van de Water) 693

wall.Seed sub- to semiglobose, more or less concave at the ventral side, with
membranous testa, without endosperm. Embryo with rather long, 2-3 times
folded radicle and more or less folded cotyledons.

Distr. About 20-25 species, 15 of which in SE. Asia, and not more than c. 10 in Central and South
America. In Malesia: 8 species.
The New World species belong to Meliosma subg. Meliosma sect. Lorenzanea, a section restricted to the
New World; besides, there is one species oi subg. Kingsboroughia which is widely spread in China but also
occurs in Mexico (A/, alba Walp.).
Correctly namedfrom the Tertiary are found widely distributed on the northern hemisphere, in
fossils
Europe, Asia, and North America; see van Beusekom, I.e. 384-424, fig. 16-18 (maps). The oldest known
fossils, of both subgenera, date from the Eocene. All localities lie south of the 60° parallel of latitude and
almost all beyond the present range of the genus. It is remarkable that still in the Pliocene the genus occurred
in Europe, S. Russia, but no longer in North America. Only in southern Japan Pliocene fossils and recent
species are found together.
Ecol. In primary and secondary forests, especially on hills and mountains up to c. 3300 m, but also in
lowlands. All or almost all species prefer everwet to moist, tropical to subtropical conditions. Some are hardy
in mild temperate climates; these are deciduous and grow flush-wise.
Morph. Trees, mostly small, sometimes shrubs, rarely mentioned to be subscandent, but A/, pinnata ssp.
ferruginea and ssp. macrophylla are recorded to reach 42 m height and M. lanceolata to reach 30 m by 1 m
diam.
The margin of may be entire or dentate and is often variable. In saplings, watershoots and
leaf or leaflet
seedlings the margin mostly dentate. In species with pinnate leaves the size of the leaflets mostly increases
is

apically and their greatest width tends to shift towards the upper half. The leaves, when pinnate, have I or
3 top-leaflets; in the first case the petiolule of the top-leaflet has an articulation with the rachis.
The inflorescence consists of a racemosely arranged, rich-fiowered panicle.
Van Beusekom {I.e. 361-364, fig. 2 & 3) amply discussed the peculiar fiower structure. Although Baillon
assumed the fiower to be basically 3-merous, he agrees with the majority of authors that it is 5-merous. The
3 outer petals are differently shaped from the 2 inner ones; the latter may be of the lanceolate or bifid type,
and taxonomically their shape is important.
The structure of endocarp and seed(/.c. 364-369, fig. 4)isof great importance. The ovary contains 4 ovules
but only one develops into a seed (exceptionally 2, resulting in an anomalous didymous fruit). The fruit is
a drupe with rather thin, pulpy mesocarp and a stony to crustaceous endocarp, more or less globular to
pyriform, smooth or often with a reticulate surface. When dehiscent, it splits into two valves, the plane of
dehiscence usually marked by a ± prominent keel running all around the endocarp. At the ventral side there
is a usually narrow pore through which the seed is connected with the vascular bundle towards the pedicel.

There are two main types: 1) endocarps which only enclose the seed, whereas the vascular bundle connecting
pedicel and seed is running outside the endocarp wall; 2) endocarps which enclose both seed and vascular bun-
dle, the latter being situated in a marginal canal inside.
Taxon The . subdivision of the genus Meliosma is as follows:

1. Leaves simple or pinnate; when pinnate rachis terminating in 3 leaflets (anomalously 2 or 1). Sepals mostly
5. Outer petals narrowly imbricate, subrotund to broad-elliptic, all regularly shaped. Vascular bundle con-

necting pedicel and seed situated in a long or short marginal canal inside the endocarp. About 12 species
in SE. Asia. Spp. 1-7 Subg. Meliosmii
2. Leaves simple or pinnate. Ovary glabrous or pubescent. Endocarp wall relatively thin, not drawn out
around the ventral perforation; endocarp mostly (sub)globosc, sometimes semiglobose, or ellipsoid to
obovoid. About 12 spp. in SE. Asia. Spp. 1-7 Sect. Meliosma
3. Leaves simple. Spp. I A 2 Subsecl. Simplices (Warb.) Beus.
4. Deciduous shrubs or small trees. Nerves all or almost all straight or almost straight, ("oniincnial Asia

Ser. Krclinervia Beus.

4. Evergreen shrubs or trees. Nerves all or almost all distinctly ascending. Spp. I A 2
Ser. Curvinervia Beus.
3. Leaves pinnate. Spp. 3-7 Subsect Pinnalae (Warb.) Beus.
2. Leaves simple. Ovary always glabrous. Endocarp wall relatively thick, more or less drawn out around the
694 Flora Malesiana [ser. I, vol. 10^

ventral perforation which often gives the mostly (sub)globose endocarp a somewhat pyriform shape.
About 10 species in Central and tropical South AmericaSect. Lorenzanea (Liebm.) Beus.

1 . Leaves pinnate, petiolule of terminal leaflet articulate with the rachis. Sepals mostly 4. Outer petals widely
irregular shape ±
imbricate, the largest one widely reniform, much wider than long, the smaller ones of
not wider than long. Vascular bundle connecting pedicel and seed situated outside the endocarp, either run-
ning in a groove at the ventral side or freely in the pulpy mesocarp
Subg. Kingsboroughia (Liebm.) Beus.
5. Deciduous trees. Vascular bundle connecting pedicel and seed running in a ventral groove of the endocarp
wall. Two SE. Asia and S. Mexico
species. Sect. Kingsboroughia

5. Evergreen Vascular bundle connecting pedicel and seed running freely in the mesocarp. One species.
trees.

Malay Peninsula, N. Borneo. Sp. 8 Sect. Hendersonia Beus.

KEY TO THE SPECIES

1 . Leaves simple. Subg. Meliosma sect. Meliosma subsect. Simplices.

2. Petioles 1/5-1/3 the length of the lamina. Panicles mostly axillary or sometimes ramiflorous, sometimes
terminal. Inner petals bifid or entire. Endocarps ellipsoid to obovoid, with reticulate to smooth surface,
and then smooth, 6-14 cm diam
rarely (sub)globose 1. M. lepidota

2. Petioles 1/20-1/5 the length of the lamina. Panicles always terminal. Inner petals always bifid. Endo-
carps always (sub)globose, always with reticulate surface, 3.5-8 mm diam 2. M. simplicifolia

1 . Leaves pinnate. Subg. Meliosma and subg. Kingsboroughia.

3. Leaf-rachis terminating in 3 (sometimes 2, rarely 1) leaflets. Outer petals widely ovate to orbicular, entire.

Endocarps inside with a marginal canal in which runs the vascular bundle connecting pedicels and seed.
Sect. Meliosma subsect. Pinnatae.
4. Leaves 2-5(-6)-jugate; leaflets usually glabrous, sometimes (in Bornean specimens) pubescent beneath,
above always with a prominent midrib. Inner petals (1.2-)1.5-2(-3) mm, entire to retuse or slightly
bifid at apex. Ovary glabrous. Endocarps 0.7-2 cm diam 3. M. sumatrana

4. Leaves 2-23-jugate; leaflets glabrous or pubescent, midrib usually flat to sulcate above. Inner petals

(0.3-)0.5-l(-1.5) mm, always distinctly and rather deeply bifid. Ovary glabrous or pubescent. En-
docarps 0.2-1 cm diam.
5. Leaves (3-)6-18(-23)-jugate, with (10-)20-100 cm long rachis. Leaflets only very rarely with slight

pubescence on midrib and nerves above. Panicles large and lax, 0.5-1.5 m, pendulous, usually suddenly
bent down at the base, with up to 90 cm long primary side-axes which are never subtended by (small)
leaves.
6. Leaflets in middle and upper part of the leaf elliptic to lanceolate, very rarely a few linear-lanceolate,
index (0.5-)2-5(-7). Petiolules 2-30 mm 4. M. lanceolata
6. Leaflets in middle and upper part of the leaf linear-lanceolate, index 5-10. Lateral petiolules absent
or very short, to c. 1 mm 5. M. hirsuta
5. Leaves 2-7(-9)-jugate, rachis up to c. 40(-60) cm. Leaflets usually more or less pubescent on midrib
and nerves above, sometimes glabrous. Panicles lax to dense but not very large, 10-50(-70) cm, usually
erect, sometimes ± pendulous, but almost never suddenly bent dov/n at the base, with up to 35(-60)
cm long side-axes which may be subtended by decrescent leaves.
7. Sepals pubescent. Outer petals glabrous. Leaves 2-3(-4)-jugate. Endocarps 7-8 mm diam. Small

trees 7. M. sarawakensis
7. Sepals glabrous (rarely with a few hairs), sometimes pubescent but then also outer petals pubescent.
Leaves 2-7(-9)-jugate. Endocarps 3-9(-10) mm diam. Small to large trees.
8. Plants from Malesia except Sumatra and Java 6. M. pinnata
8. Plants from Sumatra and Java.
9. Leaflets pubescent at least on midrib and nerves, usually entire. Panicles often with decrescent
leaves. Lowland and mountains up to 2500 m 6. M. pinnata
9. Leaflets (sub)glabrous, usually (obscurely) dentate. Panicles without decrescent leaves. Mountains,
1300-2900 m 4. M. lanceolata/. nervosa
3. Leaf-rachis terminating in of which the short petiolule is always well markedly articulated
1 (or 2) leaflets
with the rachis. Outer petals mostly widely reniform, or of irregular shape, mostly wider than long, with
± irregular margin and often emarginate. Endocarps inside without a marginal canal in which runs the
vascular bundle, large, wider than 1 cm. Inner petals hardly or not incised at apex. Subg. Kingsboroughia
sect. Hendersonia 8. M. rufo-pilosa
 1989] Sabiaceae (van Beusekom & van de Water) 695

1. Meliosma lepidota Blume, Rumphia 3 (1849) 199; key to the subspecies

Walp. Ann. 2 (1852) 224; MiQ. Fl. Ind. Bat. 1, 2
(1859) 614; Sum. (1861) 203; van Beusekom, Blumea 1. Inner petals distinctly bifid.
19(1971)451,/. 25. - Fig. 7. 2. Leaves 1 .5-2(-2.5) times as long as wide; petiole
For further synonyms, see under the subspecies; 1/4-2/3 as long as the blade. Panicles 5-15 cm.
for a complete synonymy, see van Beusekom (1971). Mature buds 2-2.5 mm diam. Endocarps ellip-
Evergreen shrub or tree, up to c. 15(-22) m. Flow- soid to obovoid d. ssp. kinabaluensis
ering twigs pubescent when young, glabrescent. 2. Leaves (1.6-)2-3 times as long as wide; petiole
Leaves elliptic or obovate to lanceolate, 2-32 by (l/6-)l/5-l/3 (-1/2) as long as the blade.
0.7-12(-18)cm, inde.\(1.2-)1.5-3(-4), atthebase Panicles 3-30 cm. Mature buds 1 .5-2(-2.2) mm
acute, at apex acute to caudate, rarely obtuse, usual- diam. Endocarps long- to short -ellipsoid (always
sometimes remotely spinously dentate to-
ly entire, distinctly higher than wide) ... .a. ssp. lepidota
wards the apex, beneath sometimes pubescent on 1. Inner petals entire, usually lanceolate.
midrib and nerves, without domatia; nerves 7-15 3. Inner petals 2.5 mm. Panicles distinctly axillary
pairs, usually strongly ascending, petioles usually or ramiflorous. Mature buds 2.5-3 mm diam.
rather long, 1-10 cm, 1/5-1/3 as long as the blade. b. ssp. dolichomischa
Panicles usually axillary and erect, widely to usually 3. Inner petals 1-1.5 mm. Panicles terminal or
narrowly pyramidal, 3-30(-200) cm, usually dense- crowded end of the twigs, rarely distinctly
at the
ly pubescent, bearing numerous solitary to crowded axillary. Mature buds 2-2.5 mm diam.
flowers which are sometimes spicately arranged; c. ssp. vulcanica
side-axes usually many, usually short, up to c. 15
(-40) cm, sometimes subtended by normal to small a. ssp. lepidota. - Meliosma lepidota Blume, Rum-
leaves; bracts ovate to linear-lanceolate, up to c. phia 3 (1849) 199; Walp. Ann. 2 (1852) 224; MiQ. Fl.
2(-6) mm, usually densely pubescent. Pedicels ab- Ind. Bat. 2 (1859) 614; Sum. (1861) 203; Illustr.
1,

sent or present, up to c. 3(-5) mm. Mature buds (1871) 73. - Meliosma pedicellata K. & V. Bijdr. 9
1.5-3 mm diam. Sepals (4) 5, (round-)ovate, sub- (1903) 134; Koord. Exk. Fl. Java 2 (1912) 545; Atlas
equal, 1 -2 mm, or the outer 1 or 2 smaller, often one 2 (1914) t. 379; Backer & Bakh./. Fl. Java 2 (1965)
lowered on the pedicel, all entire, ciliolate. Outer 144.
petals glabrous. Inner petals ± lanceolate and entire, Leaves oblong, sometimes somewhat ovate-
or bifid, (0.6-)0.8-2.5 mm, glabrous or somewhat oblong, rarely elliptic, 5-26 by (1.5-)2-12 cm, en-
ciliolate at margin or tip, when bifid never with a cen- tire, base acute, apex acute to caudate, glabrous
tral lobule. Filaments 0.7-1.5 mm. Ovary 0.5-1 when mature; nerves 8-12(-14) pairs; petiole 1.5-6
mm, very exceptionally pubescent. Fruit (sub)glo- cm. Panicles axiWary, rarely terminal or ramiflorous,
bose, sometimes elliptic, when ripe 5-10 mm diam.; often several together near the end of a branch, 3-30
endocarp globose to ellipsoid, 6-8(-9) mm diam., cm, rather poor and lax, ramified up to the 2nd
usually with a slightly elevated rather fine reticulum; order; primary (essentially secondary!) side-axes
median keel distinct, more or less prominent; ventral short, up to c. 6(-10) cm. Mature buds 1.5-2 mm
pore whether or not sunken but never spouted. diam. Inner petals about halfway bifid, 0.7-1 mm;
Distr. SE. & E. Asia; in Malesia (with 4 lobes rather narrow. Endocarp obovoid to ellipsoid,
subspecies): Sumatra, Malay Peninsula, W. Java, N. (8-)9-14 mm long, 5.5-8 mm diam., with or
Borneo (Sabah), and the Philippines (Luzon, Min- without rather wide and feeble reticulum; median
doro). keel distinct, more or less prominent, blunt to rather
Ecol . In evergreen forests under tropical or sub- sharp, at one or both ends running out into a ventral,
tropical conditions, at medium to high altitudes; for often somewhat beak-like processus; ventral pore
details, see under the subspecies. rather wide, somewhat sunken.
Notes Meliosma . lepidota displays a rather wide Distr. Malesia: Sumatra (not uncommon in
variation, especially in the ramification of its pan- Aceh, Tapanuli, and West Coast), W. Java.
icles which covers almost the whole range of possibil- Ecol Primary montane rain-forest; 900-26(X) m
.

ities found throughout Meliosma. altitude in Sumatra, 1050-1600 m in Java.

Within M. lepidota seven subspecies are recog- Field notes Outer bark dark brown, finely cor-
.

nized, four of which in Malesia. The differences be- ky, 0.5 mm; inner bark turning rcdbrown, 0.5 cm;
tween them arc on the same level as in other sub- wood ochre with reddish stripes.
species in Meliosma. Transitional forms between Vern Sumatra: antuang, hontuang, Batak
. lang.,
these subspecies, however, occur in only a few cases, Toba, kalompang bag^h, CJn. Talamau.
which is logical since there is perfect geographical Note. .Ssp. lepidota is similar and probably most
isolation between most of them. Sec further the notes closely related to the adjacent s.<!p. dolichomi.\cha
under the subspecies. and ,v.v/>. kinabaluensis. However, ssp. lepidota also
696 Flora Malesiana [ser. I, vol. 104

Fig. 7. Meliosma lepidota Blume ssp. dolichomischa (Vidal) Beus. a. Flowering twig; xO.5; b. half-opened
flower, x4.5; c. outer petal with adhering staminode; d. stamen with adhering inner petal, adaxial view; e.

stamen, abaxial view;/, pistil with surrounding disk; g. ovary, length section, all x9; h. fruit, x 1.5 {a-g
Henderson SF 23488; h Henderson SF 23492).

shows a close resemblance to certain forms of ssp. 514; Vidal, Not. Syst. 16 (1960) 306. - Fig. 7.
longipes (Merr.) Beus. from Vietnam, from which Leaves elliptic to oblong, 4-22 by 2-10 cm, en-
it can sometimes only be distinguished by the shape tire, base mostly attenuate, apex usually cuspidate,
of the endocarp. glabrous or subglabrous, nerves 7-13 pairs, petiole
(l-)3-10 cm. Panicles axillary or ramiflorous,
b. ssp. dolichomischa (Vidal) Beus. Blumea 19 solitary or afew together, 6-25 cm, rather poor and
(1971) 458, f. 25. - Meliosma dolichomischa Vidal, lax, ramified up to the 2nd or 3rd order; primary (es-
Not. Syst. 16 (1960) 304. - Meliosma monophylla sentially secondary!) side-axes up to c. 10 cm.
Ridley, J. Str. Br. Roy. As. Soc. n. 54 (1910) 40, Mature buds 2.5-3 mm
diam. Inner petals lanceo-
nom. illeg.,non Merr. (1909); Fl. Mai. Pen. 1 (1922) late, c. 2.5 mm, entire, hooding over the stamens,
 1989] Sabiaceae (van Beusekom & van de Water) 697

d
Fig. 8. Various types of inner petals in Meliosma simplicifolia Walp. a. ssp. pungens (Walp.) Beus., b. ssp.
rigida (Sieb. & Zucc.) Reus., c. ssp. fruticosa ^lume) Beus., d. ssp. simplicifolia; all x 18.

glabrous. Endocarp as in ssp. lepidota. d. ssp. kinabaluensisBeus. Blumea 19 (1971) 455. -

Distr. Malesia: Malay Peninsula
(Pahang: Meliosma pedicellata {nan K. & V.) Merr. & Perry,
Eraser's Hill, Cameron Highlands; Perak: Hermi- J. Arn. Arb. 20 (1939) 356.
tage Hill, once). leaver elliptic, rarely oblong, 3-15 by 1.5-9 cm,
Ecol. Primary montane rain-forest, c. 1200- usually entire, base acute to rounded and somewhat
1500 m altitude. attenuate, cuspidate, above glabrous or ± pubescent
Field notes. Bark thick, red. Wood first white on the midrib, subglabrous beneath, usually with a
when cut, darkening to orange-brown. Leaves white waxy layer beneath, which gives a glaucous ap-
glaucous below. pearance; nerves 8-14 pairs; petiole 1-7.5 cm.
Panicles terminal or axillary, solitary or a few togeth-
c. ssp. vulcanica (Merr.) Beus. Blumea 19 (1971) er,5-15 cm, rather poor and lax, ramified up to the
460. - Meliosma vulcanica Merr. Philip. J. Sc. 11 2nd (3rd) order; primary (essentially secondary!)
(1916) Hot. 15; Enum. Philip. Fl. PI. 2 (1923) 518. - side-axes up to c. 8 cm. Mature buds 2-2.5 mm
Machilus nervosa Merr. Philip. J. Sc. 4 (1909) Bot. diam. Inner petals halfway or somewhat less bifid,
262; Enum. Philip. Fl. PI. 2 (1923) 189; Salvore & 1-1.2 mm; lobes rather narrow. Endocarp ±
Lagrimas, Philip. J. For. 4 (1941) 309; c/. Kosterm. obovoid, c. 8 mm
long, c. 6 mm
diam., with rather
Reinwardtia 5 (1960) 377; Bibl. Laur. 1 (1964) 919. wide and feeble reticulum; median keel only slightly
- .Meliosma boniocensis Merr. Philip. J. Sc. 20 elevated, blunt, at one end running out into a minute
(1922) 403; Enum. Philip. Fl. PI. 2 (1923) 517; ventral processus; ventral pore wide, not sunken.
Kosterm. Reinwardtia 5 (1960) 377. Distr. Malesia: Borneo (Mt Kinabalu).
Leaves obovate-oblong or oblong, 5-16 by 2-6 Ecol. Montane forest, 1700-2700 m altitude.
cm, base acute, apex acute to acuminate or Field notes. The lower surface of the leaves is

sometimes rounded, glabrous or subglabrous; nerves often said to be white to light grey; in herbarium
8-11 pairs; petiole 2-4 cm. Panicles terminal, specimens indeed a whitish waxy layer can be ob-
sometimes axillary, 3-20 cm, rather profuse to poor, served often.The general colour of the leaves is re-
ramified up to the 3rd or 4th order; primary side-axes ported to be glaucous.
(mostly essentially primary!) up to c. 15 cm, usually Note. Ssp. kinabaluensis has a very low degree of
subtended by normal to reduced leaves. Mature buds variability, a characteristic found in
which is also
2-2.5 mm diam. Inner petals lanceolate, 1-1.5 mm, some other subspecies of M. is most
lepidota. It

entire, sometimes frayed at the tip. Endocarp subglo- similar to ssp. dolichomischa from the Malay Penin-
bosc, rather oblique, 6-7 mm diam., apart from a sula and to ssp. lepidota from Sumatra, with which
few ribs smooth, median keel distinct, rather promi- it shares, amongst others, the more or less ellipsoid

nent, at one end running out into a minute ventral endocarp; all other subspecies have (sub)globosc en-
processus; ventral pore somewhat sunken. docarps.
Distr. Malesia: Philippines (Luzon, Mindoro).
Ecol. Primary rain-forest, low altitude up to c. 2. Meliosma simplicifolia (Roxb.) Waip. Rep. 1

2000 m. (1842) 103; HA.ssK.Cat. Hon. Bog. (1844) 226; Miy.

Note . Ssp. vulcanica is the only subspecies of M. Fl. Ind. Bat. 1, 2 (1859) 613; Sum. (1861) 203; van
lepidota in which normal terminal panicles have been BhUSEKOM, Blumea 19(1971)462, f. 28. Milling-
found. In general habit it is more similar to certain tonia simplicifolia Roxb. [Horl. Bcng. (1814) 3,

forms of ssp. longipes from Vietnam than to .ssp. nomen] PI. Corom. 3 (1820) 50, t. 254. FIr. 8. 10.
squamulata (Han* e) Bius. from Taiwan or to ssp. For further synonyms, see under the subspecies;
kinabatuensis from Borneo, to which i( is obviously for a complete synonymy, see van Beusekum 97 ). ( 1
1

less closely related. Evergreen shrub or tree, up to 20 m. Leaves elliptic

698 Flora Malesiana [ser. I, vol. 104

or obovate to lanceolate. 3-50 by 1-18 cm, base brous to densely pubescent, rarely tomentose,
cuneate, apex acute to acuminate, rarely caudate or with or without domatia. Style about as long as

rounded, entire to spinously dentate, sometimes with the ovary or shorter.

hairy domatia; nerves 7-25 ± ascending,

pairs, 3. Leaves with or without domatia; midrib on the
sometimes looped; petiole 0.5-6(-7) cm, 1/20-1/3 upper side of the full-grown leaf glabrous or
as long as the blade. Panicles terminal, very rarely nearly so, more or less prominent, rarely flat.

axillary, erect, lax to rather dense, widely to narrow- Inner petals with entire lobes, which are

lypyramidal, (4-)10-50(-60) cm, usually profusely sometimes slightly fimbriate or ciliolate at the

branched up to the 2nd-4th order, bearing numer- very tips. Endocarps 3.5-5(-7) mm diam.
ous solitary to crowded or glomerulate flowers which a. ssp. simplicifolia

are usually spicately arranged; primary side-axes 3. Leaves with or without domatia; midrib on the
usually many, up to c. 25 cm, often subtended by upper side of the full-grown leaf more or less
leaves; bracts ovate to linear-lanceolate, up to c. 8 but distinctly pubescent, ± impressed to flat.
mm. Pedicels sometimes present, up to c. 3 mm. Inner petals usually with fimbriate, rarely entire
Mature buds (1-) 1.5-3 mm
diam. Sepals (4) 5, lobes which are rarely minutely ciliolate at the
sometimes by addition of empty bracts seemingly very tips. Endocarps (4.5-)5.5-8 mm diam.
more, up to 11(-13), (round-)ovate, equal or usually c. ssp. fruticosa

more or less unequal, the inner ones 0.7-2 mm, the 1. Sepals (8-)9-ll(-13). Leaves usually with do-
outer one(s) smaller, often minute. Inner petals more matia. Endocarps 3.5-5.5 mm diam.
or less deeply bifid, 0.5-1.5 mm, with glabrous, d. ssp. pungens
sometimes fimbriate or ciliolate lobes, never with a
central lobule. Filaments 0.5-1.5 mm. Ovary a. ssp. simplicifolia. - Millingtonia simplicifolia
0.5-0.7(-l) mm. Mature fruit (sub)globose, 4-10 RoxB. [Hort. Beng. (1814) 3, nomen] PI. Corom. 3

mm diam.; endocarp globose to subglobose, often (1820) 50, t. 254; Fl. Ind. 1 (1820) 103; Nees, Flora
depressed or oblique, 3-9 mm diam., with very 8 (1825) 106; Griff. Not. PI. As. (1854) 162; Ic. PI.

vague to very strong and prominent reticulum; me- As. (1854) Anon. Ic. Roxb. 4 (1970) 40, t. 20;
t. 442;
dian keel more or less prominent; ventral pore VAN Beusekom, Blumea 19 (1971) 476. - Meliosma
somewhat sunken to somewhat spouted. simplicifolia Walp. Rep. (1842) 103; Hassk. Cat.
1

Distr. Continental SE. Asia (from Ceylon to Hort. Bog. (1844) 226; Thw. Enum. PI. Zeyl. (1858)
China, Taiwan and S. Japan); in Malesia: Sumatra, 59; MiQ. Fl. Ind. Bat. 1, 2 (1859) 613; Sum. (1861)
Malay Peninsula, Borneo, Java, and Lesser Sunda 203; Illustr. (1871) 73; Bedd. Fl. Sylv. 3 (1871) 77;
Islands. Fig. 9. Brandis, For. Fl. (1874) 116; Hook./. Fl. Brit. India
Ecol. Subtropical to tropical forests, under 2 (1876) 5; KuRZ, J. As. Soc. Beng. 45, ii (1876) 204;
various conditions, usually in mountains up to c. Fl. Burma 1 (1877) 301; Trim. Fl. Ceyl. I (1893) 315;
3000 m, but also at sea-level. For details see under the Prain, Bengal PI. 1 (1903) 246; Brandis, Indian
subspecies. Trees (1906) 194; Merr. Contr. Arn. Arb. 8 (1934)
Note. Meliosma simplicifolia is a very variable 95; Brittonia 4 (1941) 110; Vidal, Not. Syst. 16
species, covering an enormous area in which it is (1960) 307. - Meliosma angulata Blume, Rumphia 3
adapted to many different habitats. It can be divided (1849) 197; Walp. Ann. 2 (1852) 224; K. & V. Bijdr.
into eight well-marked subspecies, five of which cen- 9 (1903) 131; Koord. Exk. Fl. Java 2 (1912) 545;
tre in SW. Yunnan, and diverge over different parts Atlas 2 (1914) t. 378; Baker/, in Rendle, J. Bot. 62
of the area. (1924) Suppl. 30; Vidal, Not. Syst. 16 (1960) 304. -
Sabia densiflora Miq. Sum. (1861) 203, 520. - Sabia
KEY TO THE SUBSPECIES floribunda Miq. I.e. 203, 521; Kurz, J. As. Soc.
Beng. 39, ii (1870) 74. - Fig. 8d.
1. Sepals (4-)5. Leaves obovate-oblong to -lanceolate, up to c. 50
2. Panicles branched up to the 2nd (3rd) order, by 18 cm, base cuneate, apex acute to short-cuspi-
nearly always (very) densely tomentose; primary date, beneath often with domatia; nerves 8-23 pairs.
side-axes subtended by leaves. Leaves
rarely Panicles rather lax, 10-45 cm, branched up to the
sparsely to densely but always distinctly pubes- 3rd or 4th order; axes sparsely to densely pubescent
cent to tomentose, at least on midrib and nerves; but never tomentose, the lower primary ones sub-
without domatia. Style c. 1.5-2 times as long as tended by leaves. Flowers more or less crowded to
the ovary b. ssp. rigida solitary, (sub)sessile; mature buds 1.5-2 diam. mm
2. Panicles branched up to the (2nd) 3rd or 4th Sepals 5 (4). Inner petals 0.6-0.8 mm, usually over
order, sparsely pubescent to moderately tomen- halfway bifid, lobes more or less divergent, narrow,
tose; lower
primary side-axes often subtended by glabrous, sometimes slightly fimbriate or ciliolate at
normal to small or reduced leaves. Leaves gla- the very tips. Style about as long as ovary or shorter.
1989] Sabiaceae (van Beusekom & van de Water) 699

M. simplici folia
^^m ssp.fordii
• •M ssp. rigida
_^_ ssp. thomsonii
^^= ssp. laui
ssp. frulicosa
ssp. pungens
._- ssp.yunnanensis
ssp. simplicifolia

Fig. 9. Generalized areas of the subspecies of Meliosma simplicifolia Walp.

Endocarps subglobose, usually rather oblique, near- MiQ., nomen; Maxim. Bot. Jahrb. 6 (1884) 60;
ly triangular at ventral view, 3.5-5(-7) mm diam., Forbes&Hemsley, J. Linn.Soc. Bot. 23 (1886) 145,
with more or less prominent, rather coarse reticu- p.p., excl. M. pungens; Dunn, J. Linn. Soc. Bot. 38
lum; median keel usually very prominent, at one end (1908) 358; Hayata, Ic. PI. Formos. (1911) 161 1

sometimes running out into a minute ventral pro- Dunn & Tutch. Kew Bull. Add. Ser. 10 (1912) 68
cessus; ventral pore somewhat or not sunken, not Chun, Sunyatsenia 1 (1933) 180; Hand.-Mazz
spouted. Beih. Bot. Centralbl. 52(1934) 166; Kaneh. Formos
Distr. Widely distributed in continental SE. Trees ed. 2 (1936) 416, f. 372; Cufod. Oest. Bot. Z
Asia; in Matesia: northern half of Sumatra, W. Java 88 (1939) 267, incl. var. patens; Hara, Enum
(not found since Blume's time). Fig. 9. Sperm. Japon. 3 (1954) 121; Making, III. Fl. Jap
Ecol. Primary and secondary evergreen forest, (1954) 348, f. 1044; Walker, Imp. Trees Ryukyu Is
from sea-level up to c. 1200<- 1500) m altitude. It is (1954)200, f. 121; How, Acta Phytotax. Sin. 3 (1955)
often reported to occur along watercourses. 444; Gagnep. & Vidal, Fl. Camb. Laos & Vietnam

Vern. Sumatra: medanf; sungu, M, simulingga, 1 (1960) 47, in obs.; Liu, Taiwan 2
III. Lign. PI.
sumpa mana belawah, Karo, kayu gadis. West (1962) 925, f. 762; Li, Woody
Taiwan (1963) 503;
Fl.

Coast. Ohwi, Fl. Japan (1965) 613. - Quercus jama-buwa

Note. A rather uniform, well recognizable sub- SiEB. in sched. ex Miq. Ann. Mus. Bot. Lugd.-Bat.
species all over its area. 3 (1867) 93, nom. inval. - Meliosma pungens auct.
non (W. & A.) Walp.: Hook./. Fl. Brit. India 2
(Sieb.&Zucc.)Beus. Blumca 19(1971)
b. ssp. rigida (1876) 4, p.p., quoad pi. Japon. - Melio.sma patens
473. Meliosma rigida Sieb. & Zurr. Abh. K. Hkmsley ex Forbes &. Hemsley, J. Linn. Soc. Bot.
(Bayer.) Ak. Wiss. M. Ph. Kl. Munchcn 4, 2 (1845) 23 (1886) 145. Meliosma harmandiana Pierre,
153; Mio. Ann. Mus. Hot. Lugd.-Bat. 3 (1867) 93; II. For. Cochinch. 5 (1897) I. 360. Meliosma -

Cat. Mus. Bot. (1870) 23, incl. var. angusli/ulia

I gtomerulata Rehu. & Wils. in Sarg., PI. Wils. 2
700 Flora Malesiana [ser. I, vol. 10^

Fig. 10. Meliosma simplicifolia Walp. ssp. fruticosa (Blume) Beus. a. Fruiting twig, xO.5; b. detail of leaf
undersurface, x2.5; c. endocarp, in different positions, x2.5 (a-c Kadim & Noor 395).
1989] Sabiaceae (van Beusekom & van de Water) 701

(1914) 203. - Meliosma

loheri Merr. Philip. J. Sc. Lugd.-Bat. 1 (1850) 306, was included in the synony-
10(1915) Bot. 38; Enum. Philip. Fl. PI. 2 (1923) 517. my of M. rigida by Hara, I.e., but I found it to
- Meliosma pannosa Hand.-Mazz. Anz. Ak. Wiss. belong to Quercus gilva Blume.
Wien M.-N. Kl. 58 (1921) 179; How, Acta Phytotax.
Sin. 3 (1955) 442; Gagnep. & ViDAt, Fl. Camb. Laos c. ssp. fruticosa (Blume) Beus. Blumea 19 (1971)
& Vietnam 1 (1960) 50, in obs., p.p. - Meliosma 477, f. 28. - Meliosma fruticosa Blume, Rumphia 3
costata CuFOD. Oest. Bot. Z. 88 (1939) 266; How, (1849) 198; Walp. Ann. 2 (1852) 224; MiQ. Fl. Ind.
Acta Phuotax. Sin. 3 (1955)444; Gagnep. & Vidai, Bat. 1,2 (1859) 614; Illustr. (1871) 73; K. & V. Bijdr.
Fl. Camb. Laos & Vietnam 1 (1960) 45; Vidax, Not. 9 (1903) 133; Koord. Exk. Fl. Java 2 (1912) 545. -
Syst. 16 (1960) 304. - Meliosma evrardii Gagnep. Meliosma elliptica Hook./. Fl. Brit. India 2 (1876) 5,
Not. Syst. 14(1952) 212, p.p. p.p., excl. Sabia floribunda Miq.; King, J. As. Soc.
Leaves usually obovate-oblong to obovate-lanceo- Beng. 65, ii (1896) 456; Ridley, J. Str. Br. Roy. As.
sometimes oblong to lanceolate, 4-25(-32) by
late, Soc. n. 33 (1900) 67; Fl. Mai. Pen. 1 (1922) 514;
1.5-8(-ll) cm, base long-cuneate to acute, apex BuRK. & Hend. Card. Bull. S. S. 3 (1925) 364. -
acute to cuspidate, without domatia; nerves 7-19 Meliosma lancifolia Hook./. Fl. Brit. India 2 (1876)
pairs. Panicles lax to rather dense, 10-30 cm, 5; King, J. As. Soc. Beng. 65, ii (1896) 456; Ridley,
branched up to the 2nd (3rd) order; axes more or less Fl. Mai. Pen. 1 (1922) 514. - Meliosma monophylla
tomentose, sometimes woolly-pubescent, the pri- Merr. Philip. J. Sc. 4 (1909) Bot. 286; Enum. Philip.
mary ones only exceptionally subtended by small Fl. PI. 2 (1923) 517; Vidal, Not. Syst. 16 (1960) 306.
leaves. Flowers more or less crowded, usually in - Fig. 8c, 10.
dense glomerules, sessile; mature buds 1.7-2.2 mm Leaves usually oblong to lanceolate, 5-40(-45) by
diam. Sepals 5 (4). Inner petals 0.6-O.S mm, usually 2-15 cm, base acute, apex acute to acuminate,
less than halfway bifid, lobes hardly or not densely pubescent on midrib and sometimes on
divergent, rather narrow, fimbriate or ciliolate at the nerves and lamina, beneath glabrous to tomentose,
tips. Style about (1 .5-)2 times as long as ovary. En- sometimes with domatia; nerves 7-25 pairs. Panicles
docarps (sub)globose, not or not much oblique, usually lax, sometimes more dense, 10-50 cm,
(3.5-)4-5 mm
diam., with fine reticulum; median branched up to the 3rd (4th) order; eixes pubescent to
keel blunt to rather sharp, hardly to distinctly promi- short-tomentose, the lower primary ones subtended
nent, often at one end running out into a minute ven- by small leaves or not. Flowers more or less crowded
tral processus or tubercle; ventral pore not sunken, to solitary, (sub)sessile; mature buds 1.5-2 mm
often somewhat spouted. diam. Sepals 5 (4). Inner petals c. 0.7 mm, about
Distr. Widely distributed in continental SE. halfway or somewhat less bifid; lobes divergent or
Asia, incl. China, Laos, S. Vietnam (only at Hue), not, usually rather narrow, more or less fimbriate,
Taiwan (incl. Pescadores), Ryu Kyu Islands, Japan; sometimes entire. Style about as long as ovary or
in Malesia: Philippines (Luzon: Mountain Provi- shorter. Endocarps globose or ± ellipsoid,
nce). Fig. 9. (4.5-)5.5-8 mm diam., with rather wide, coarse
Eco I . In evergreen broad-leaved or laurophyllous reticulum; median keel prominent, at one end often
forests, on different soils; in dry as well as in wet running out into a minute ventral processus; ventral
100-1000 m, but in Luzon
places; altitude usually pore not or not much sunken, not spouted.
reported from 1200- 16(X) m. Distr. S. Peninsular Thailand (Sural) and
Field notes. Bark grey, smooth. Branches Taiwan; in Malesia: common in the Malay Peninsula
brown. Leaves lustrous green above, sometimes and Sumatra; W. Java, Lesser Sunda Islands (Sum-
glaucous beneath. Fruit blue-purple to purplish bawa, Flores), Borneo (Central Sarawak, Kinabalu,
black. W. Kutai), and the Philippines (Luzon). Fig. 9.
Vern. Philippines: gahatan. If., Luzon; lasuit, Ecol. Primary rain-forest, on various soil types,
Bondoc dial. reported to occur on limestone, sand, volcanic loam,
Notes. Ssp. rigida is variable in quite some and andesite; altitude from sea-level up to 24(X) m.
characters in its area outside Luzon, for instance in Field notes. Bark smooth, grey to brown, len-
the degree of pubescence, leaf shape, and dentation. ticellatc, paperthin. Inner bark pale brown to dark
In continental SE. Asia the area of ssp. rigida bor- brownrcd. Wood reddish to rcdbrown. Fruit yellow
ders on or overlaps the areas of five or six other to pale red when ripening, dark red to brown when
subspecies of M. which substantially
simplici/olia, ripe.
adds to the chance of confusing them, several Vern. once noted). Malay Peninsula: bua
(all

specimens being hybrids, it is probable that these palu, Sclangor, medang kerkulu, mengading, Malac-
subspecies are ecologically isolated to a large extent ca; Sumatra: lelagan, Gajo lang., Aceh, kaju djarap,
and thus contact between them is prevented. k. gasir, k. si raga, Asahan, kaju ardong ardong,
Quercus gilva var. prucera Blumf:, Mus. Bot. Toba, kabung kabung, Tapanuli, masadih pajo.
702 Flora Malesiana [ser. I, vol. 10^

Simalur, kendung, Palembang, redjang, Djambi; 520; Illustr. (1871) 74; King, J. As. Soc. Beng. 65, ii

Java: ki (iwu, Preanger; Flores: kaju sar, Philip- (1896) 457; K. & V. Bijdr. 9 (1903) 117; Koord. Exk.
pines: malaligas. Tag. Fl. Java 2 (1912) 546, f. 81; Atlas 2 (1914) 377;
Ridley, Fl. Mai. Pen. 1 (1922) 515; Baker/, in Ren-

d. spp. pungens (Wall, ex W. & A.) Beus. Blumea dle, J. Bot. 62 (1924) Suppl. 30; Burk. & Hend.
19 (1971) 466. - Millingtonia pungens Wall, ex Card. Bull. Str. Settl. 3 (1925) 364; Heyne, Nutt. PI.

W. & A. Edinb. New Phil. J. 15 (1833) 178; Prod. 1 (1927) 1002; Merr. &
Perry, J. Arn. Arb. 20 (1939)
(1834) 115; Wight, Ic. 3 (1845) t. 964/3. - Meliosma 357; Backer & Bakh./. Fl. Java 2 (1965) 145. - Irina

pungens (Wall, ex W. & A.) Walp. Rep. (1842) 1 integerrima Blume, Bijdr. (1825) 231, non Hassk.
423; Ann. (1848) 135; Thw. Enum. PI. Zeyl. (1858)
1
PI. Jav. Rar. (1848) 284 ('Iriney, Walp. Rep. 1

59; Bedd. F1. Sylv. 3 (1871) 77; ibid. t. 160; Merr. (1849) 416; Blume, Rumphia 3 (1849) 202, in syn.
Contr. Am. Arb. 8 (1934) 94; Vidal, Not. Syst. 16 sub M. nitida. - Millingtonia nitida Schult. &
( 1 960) 306. - Meliosma wightii Planch, ex Brandis, ScHULT. Syst. Veg. Mant. 3, add. 2 (1827) 250;
For. Fl. (1874) 116;HooK./. Fl. Brit. India 2 (1876) DiETR. Syn. PI. 1 (1839) 103. - Meliosma confusa
4. - Fig. 8a. Blume, Rumphia 3 (1849) 200; Walp. Ann. 2 (1852)
Leaves elliptic to oblong, sometimes lanceolate, 225; Miq. Fl. Ind. Bat. 1, 2 (1859) 616;Sum. (1861)
5-20(-30) by 2-8(-10) cm, without or with some 203, 520; Illustr. (1871) 74. - Meliosma cuspidata
distant teeth, acute to rounded at the base, acute to Blume, Rumphia 3 (1849) 202; Miq. FI. Ind. Bat. 1,

acuminate at the apex, usually distinctly pubescent 2 (1859) 617; Illustr. (1871) 74; Hall./ Meded.
on midrib and sometimes on nerves above, sparsely Rijksherb. (1910) 2; Merr. Enum. Born. (1921)
1

to moderately pubescent beneath especially on mid- 362. - Meliosma pinnata (non Walp.) Koord.
rib and nerves, usually with domatia; nerves 7-18 Minah. (1898)408. - Meliosma diepenhorstii Valet.
pairs. Panicles lax to dense, (5-) 10-55 cm, branched Ic. Bog. 2 (1904) 195, t. 150. - Meliosma elmeri

up to the 2nd (3rd) order; axes rather coarse, densely Merr. PI. Elm. Born. (1929) 177. - Meliosma
short-tomentose, the lower primary ones almost philippinensis Merr. & Perry, J. Arn. Arb. 20
always subtended by small leaves. Flowers crowded (1939) 357.
in dense glomerules, mature buds 2-2.5 mm
sessile; Evergreen tree, up15-20(-25) m. Leaves
to

diam. Sepals {8-)9-ll(-\i). Inner petals c. 1 mm, 2-5(-6)-jugate; rachis terete, 6-50 cm, including
slightly bifid; lobes divergent, wide, glabrous. Style the up to c. 25(-30) cm long petiole, up to c. 10(- 15)
about as long as ovary. Endocarps (sub)globose, mm across, rarely slightly pubescent, usually with
often rather irregular, 3.5-5.5 diam., with mm distinctly swollen base; leaflets usually elliptic to
usually lax reticulum; median keel distinct but not lanceolate, (3-)5-35(-50) by (1.5-)2.5-I5(-20)
very prominent, not running out into a ventral pro- cm, base cuneate to rounded, shortly narrowed into
cessus; ventral pore hardly or not sunken, not spout- the petiole, apex acuminate to caudate, usually en-
ed. tire, beneath rarely more or less pubescent, without

Distr. Sri Lanka and Deccan Peninsula; in domatia; midrib slightly prominent above; nerves
Malesia: N. Sumatra (Gajo Lands, Takengon), one (5-)7-13(-19) pairs, ascending, nearly always
collection. Fig. 9. looped and joined; petiolules very short or up to c.
Ecol. Mountain forest, 1500-2000 m altitude. 6 cm, usually distinctly swollen at the base especially
in older leaves. Panicles usually terminal, usually
3. Meliosma sumatrana (Jack) Walp. Ann. 1 (1848) narrowly, sometimes widely pyramidal, 7-50(-75)
135; MiQ. Fl. Ind. Bat. 1, 2 (1859) 617; Sum. (1861) cm, usually profusely branched up to the 4th order,
203; lUustr. (1871) 75; Hook./. Fl. Brit. India 2 rather stiff and coarse, puberulous, bearing numer-
(1876) 6; KooRD. Minah. (1898) 408; Suppl. Cel. 2 ous crowded flowers; primary side-axes usually
(1922) 7, t. 56; ibid. 2 (1922) 28; Merr. Enum. Born. rather short, up to c. 30 cm, the lower ones excep-
(1921) 363; Enum. Philip. Fl. PI. 2(1923) 518; Contr. tionally subtended by small to reduced leaves; bracts
& Perry, J. Am. Arb.
Arn. Arb. 8 (1934) 95; Merr. ovate to narrowly triangular, up to c. 6 mm, ±
20(1939) 357; van Beusekom, Blumea 19 (1971) 485. puberulous. Pedicels absent or short, up to c. 2 mm.
- Millingtonia sumatrana Jack, Mai. Misc. 2 (7) Mature buds (1 .5-)2-3(-3.5) mm diam. Sepals 5 or
(1822) 30;Hook. J. Bot. 1 (1834) 378; Merr. J. Arn. 4, ovate, unequal, the inner 3 or 4 c. 1-2 mm, the
Arb. 33 (1952) 236. - Meliosma nitida Blume, Cat. outer 1 or 2 usually smaller, often minute, sometimes
(1823) 32; Nees, Flora 8 (1825) 106; Hassk. Tijd. lowered on the pedicel, sometimes puberulous out-
Nat. Gesch. Phys. 10 (1843) 139; Cat. Hort. Bog. side, especially the outer ones, with entire or 2- or 3-
(1844) 226; Blume, Rumphia 3 (1849) 202, t. 169, lobed, often ciliolate margin. Outer petals glabrous.
incl. var. Blume, var. cerasiformis Blume et
tridenta Inner petals elliptic to lanceolate or strap-shaped
var. splendens Blume; Walp. Ann. 2 (1852) 225; with wide-truncate tip, (1.2-)1.5-2(-3) mm, acute
Miq. Fl. Ind. Bat. 1, 2 (1859) 617; Sum. (1861) 203, to slightly bifid or retuse and frayed at the tip. Ovary
1989] Sabiaceae (van Beusekom & van de Water) 703

Fig. 11. Generalized areas of Meliosma sumatrana (Jack) Walp. (thick line) and M. lanceolata Blume (thin
line); the small ova! areas indicate the localities of M. lanceolata var. polyptera (Miq.) Beus. The distribution
of M. hirsuta Blume is indicated by a dot, that of M. rufo-pilosa Hend. by squares.

0.5-1 mm, glabrous. Fruit globose to short-ellip- often with shallow fissures, sometimes said to be
soid, when 1-3 cm diam., with rather thick
ripe dimpled, patchy or scaly. Inner bark 0.5- 1 cm thick,
spongy to pulpy mesocarp; endocarp ellipsoid, some- soft, fibrous, light yellow or dirty white, soon turn-
times nearly globose, 0.7-2 cm diam., with almost ing pink, brownish, reddish, or rusty after exposure.
smooth to somewhat lumpy surface, often with a few Sapwood said to be whitish, yellowish, creamy
faint to sharply prominent ribs; median keel distinct, orange, or brownish. Sap without special smell or
slightly elevated to sharply prominent, at one end taste.Leaves bright green on both sides. Flower col-
often running out into a more or less prominent curv- our varying from white, cream, or greenish, to partly
ing, at the other end sometimes into a minute tuber- or entirely pinkish to red. Fruit first yellow, then
cle; ventral pore mostly rather wide, usually some- yellow with red to red when ripe; pulp white, turning
what sunken. quickly blood-red on exposure, finally becoming
Distr. Malesia: Sumatra (incl. Nias, Batu & black, sweetish to tasteless.
Sipora Is., Banka), Malay Peninsula (incl. Penang Uses. The species was proposed by Koorders &
I.), Anambas Is., W. half of Java, throughout Bor- Valeton, I.e., for reforestation purposes. In Min-
neo, Sulawesi, and the Philippines (Mindanao, danao the triturated bark and leaves are several times
Palawan). Common. Fig. II. reported to be in use as a medicine applied for
Lcol . Primary and secondary lowland and mon- wounds, to soothe itchy skin or - charred and put in
tane rain-forest, up to c. 2200 m altitude. Found on water - against tympanites. It was also .said to be
various soils, fertile as well as infertile, in dry to wet used in agricultural rituals. The fruits are many times
localities, in dense to open forests, by streams as well reported to be edible.
as on hilltops and ridges. Vern. Malay Peninsula: pa-on^, Saki name, and
Field notes. Often a crooked tree, irregularly mennadinfi hesar, both from Pahang, huah mata
branched. Trunk sometimes with small buttresses. ikan, Icmuan, Sclangor, pokoh haran, Negri Scm-
Bark surface grey to brown, smooth, with lenticels. bilan, kaju kahwa kanlu, membuloh, pokoh gra6
704 Flora Malesiana [ser. I, vol. 10^

jantan, p.mala gajah, p. paigigi, p. pinang plandog, 2 (1912) 546; Merr. Enum. Born. (1921) 363;

p. ravoa antoo, pudding utan, Malacca, pelanlu; Ridley, Mai. Pen. 1 (1922) 516, f. 51; Baker/.
Fl.

Sumatra: laon, si paturut, sringkut, Karo country; in Rendle, J. Bot. 62 (1924) Suppl. 30; Craib, Fl.

kaju durung durung, k. ining ining, Tapianuli, tarn- Siam. Enum. 1 (1926) 340; Ridley, Kew Bull. (1926)
pa bussie, Priaman, marazat, Mt Kerinci; Java: ki 63; Merr. PI. Elm. Born. (1929) 176; Hochr. Can-

liwu (landuk) (bodas), S, ki huut, Udjong Kulon; dollea 6 (1936) 467, incl. var. genuina Hochr.;

Borneo, Sarawak: buliliap, Kenyah dial., maiak, Backer & Bakh./ Fl. Java 2 (1965) 145; van
Kayan dial., buiu manuk, Iban, bilonok, Dyak; Beusekom, Blumea 19 (1971) 489. - Millingtonia
Sabah: bung lai, Sungai, gapas gapas, kapas kapas, lanceolata Schult. & Schult. Syst. Veg. Mant. 3,
limpangol, tunjang, Murut; add. 2 (1827) 250; Dietr. Syn. PI. (1839) 103. -
keriyan, Dusun, illulal, 1

SE. Borneo: tambalilin, tandao, Dyak, Tidung dial.; Meliosma polyptera Miq. Sum. (1861) 203, 520;
djangkanggunung, Bandjar lang., Riom dial.; lllustr. (1871)73. - Meliosma levis KwG, J. As. Soc.

Sulawesi: see Koord. Minah. (1898) 408; eng- Beng. 65, ii (1896) 457; Ridley, Fl. Mai. Pen. 1

golokia, W. Toradja dial., pulu putu, situi, Tobela (1922) 515. - Meliosma nervosa K. & V. Bijdr. 9
pobumengo, Gorontalo; dama, Torai
lang., Malili, (1903) 129; Koord. Exk. Fl. Java 2 (1912) 546; Atlas
Menado; Philippines: carabo-rabo, daborabo,
dial., 2 (1914) t. 376; Fl. Tjibodas 2 (1923) 158; Merr. &

kadabudabo, karabu-rabu, magobaylung, mahag- Perry, J. Arn. Arb. 20 (1939) 359, in obs.; Backer
yagabogan, Mbo, Buk., bentinguasay, gepulu,
kol, & Bakh./ Fl. Java 2 (1965) 145.
Zamboanga, waat, Cebuano, Mt Apo, salalab, Evergreen tree, up to c. 25(-30) m. Twigs often
Moro dial., garong, gimbingimbing. Sub., sumaga- with conspicuous leaf-scars. Leaves (3-)7-18(-25)-
sa, Bag. jugate; rachis terete, (10-)30-100 cm, including the
Notes. Meliosma sumatrana is very constant in 5-30 cm long petiole, up to c. 8 mm diam., usually
its discriminative characters (especially the promi- with distinctly swollen base, usually ± lenticellate;

nent midrib and entire inner petals), but there is leaflets usually oblong to lanceolate, hardly or not
nevertheless some geographical variation, especially asymmetrical, 5-20 by 2-7 cm, not or only slightly
in the northern part of Borneo (Sarawak, Sabah). As increasing in size towards the top of the leaf, often
general tendencies may be noticed that towards the the lowermost pairs much smaller, base usually acute
centre of the area leaflets and fruits increase in size to rounded, apex acuminate to cuspidate, glabrous
and dentate leaflets become more common. More- to moderately pubescent, always without domatia;
over, the number of when the
leaflet pairs decreases midrib usually deeply impressed above; nerves 5-16
leaflets are larger. pairs, ascending, looped. Panicles terminal, nearly
Sterile hybrids between M. sumatrana and M. pin- always pendulous and lax, rarely erect (then also

nata ssp. ridieyi are rarely found (Sabah). small), pyramidal, usually large, (15-)50-150 cm
DocTERS VAN Leeuwen (Zoocecidia Neth. East and profusely branched up to the 3rd order, ± pu-
Indies, 1926, 339, f. 612) described a leaf-gall on a bescent, bearing numerous glomerulate or crowded
specimen from Sulawesi. This type of galls (usually flowers which are usually spicately arranged, the
ball-shaped, c. 4 mm, ending in a short mucro, and glomerules often with regular space; main axis terete,
surrounded by a calyx-like circumvallation) is rather often bent down abruptly at the base; primary side-
commonly met with in this species, not only in axes many, usually long, up to c. 90 cm, never
specimens from Sulawesi, but also from Borneo, subtended by leaves; bracts ovate to narrowly
Sumatra, and the Malay Peninsula. The galls do not triangular, up to c. 5 mm, ± pubescent. Pedicels ab-
only occur on the lower surface of the leaflets, but sent, up to c. 1 mm. Mature buds 1.5-2 mm diam.
occur also on the upper surface, and on rachis and Sepals 5 (4), ovate, more or less unequal, the inner 3

petiolules, often very many crowded together. or 4 c. 1 mm, the outer 2 or 1 usually much smaller,
often minute and sometimes slightly keeled,
sometimes somewhat lowered on the pedicel, all
4. Meliosma lanceolata Blume, Cat. (1823) 32; Nees, glabrous, and with an entire margin. Outer petals
Flora 8 (1825) 106; Hassk. Cat. Hort. Bog. (1844) 1.5-2 mm. Inner petals about halfway bifid, c. 0.6
226; Blume, Rumphia 3 (1849) 200, t. \6i,p.p., incl. mm, with ciliolate, rarely glabrous lobes, usually
var. pendula Blume, membranacea Blume, var.
var. with a minute central lobule. Filaments c. 1 mm.
chart acea Blume et var. obliqua Blume; Walp. Ann. Ovary (0.5-)0.7(-l) mm, usually densely, some-
2 (1852) 224; Miq. F1. Ind. Bat. 2 (1859) 614; Sum.
1 , times sparsely pubescent, rarely glabrous. Fruit
(1861) 203, 520; lllustr. (1871) 74, p.p.; Hook./. F1. (sub)globose, when ripe 7-10 mm diam.; endocarp
Brit. India 2 (1876) 7; King, J. As. Soc. Beng. 65, ii subglobose, often somewhat depressed to applanate
(1896) 458; Ridley, J. Str. Br. Roy. As. Soc. n. 33 at the ventral side, usually strongly obUque, (5-)6-9
(1900) 67; K. & V. Bijdr. 9 (1903) 125; Hall./. Med. mm diam., with usually distinct, rather coarse, most-
Rijksherb. 1 (1910) 2, in obs.; Koord. Exk. Fl. Java ly sharply prominent reticulum; median keel sharp
1989] Sabiaceae (van Beusekom & van de Water) 705

and prominent, at one end often running out into a (2-)2.5-7(-10) cm, index (l.5-)2-5(-6), without
small to minute ventral processus or tubercle; ventral or with teeth, glabrous or pubescent.
pore not or not much sunken. Notes. In the lowland parts of its area var.
D i s t r . Nicobar Is. , extreme South of Peninsular lanceolata is nearly always very constant in the main
Thailand; in Sfalesia: Sumatra (incl. Simalur, Batu, characters. Mainly at higher elevations, however,
and Banka Is.), W. Java, Borneo (northern halO- forms occur which deviate considerably, often to
Not uncommon, scarce in Borneo. Fig. 11. such an extent that it is very difficult to separate them
Ecol. Primary and often secondary forests, at from less typical forms of the otherwise well distinct
low and medium altitudes, occasionally ascending to M. pinnata ssp. ferruginea and ssp. ridleyi; in a few
1500 m,/. nervosa to 2900 m, on various soil types. cases, especially when the material is incomplete, this
Field notes. Outer bark grey to brown, rather can only be done by a specialist who is thoroughly ac-
smooth, later with longitudinal cracks, thin, often quainted with habitus and variability of both species.
lenticellate. Inner bark 0.5-1 cm, several times said For instance, a form with erect, unusually short
to be (light) red, orange brown, or redbrown, also panicles (sometimes only 15 cm long) and other de-
dirty white and then turning rusty after e.xposure. viating characters may be met with. It occurs mainly
Wood soft, white or pale yellow to light yellow in the montane zone; transitional forms are found
brown. Crown low, irregular and lax, with few usual- lower, and these show a more or less gradual fading
ly crooked branches. The conspicuous large leaves of typical lanceolata characters. Specimens of this
are rather crowded at the end of the twigs. Leaflets mountain form have been described from Java by
when young red-brownish. Flowers white or yellow- KooRDERS & Valeton, I.c, as M. nervosa. In my
ish to pink or red (sometimes different colours in the opinion this species should be reduced to the rank of
same panicle). Fruits, first dirty red, then bluish black a form only; see below.
when ripe.
Vern. Malay Peninsula: medang siri, Malacca; forma lanceolata.
Sumatra: kabung kabung (blumut), Batak lang., Leaves (3-)6-18-jugate, with elliptic to lanceo-
Simelungun dial., bulung manuk, Batak lang., Karo late, glabrous to pubescent leaflets. Panicles pen-
dial., sondang, sonlang, Timor on N. Sumatra, kaju dulous, usually much longer than 50 cm. Inner petals
buluk hudjan, Lampong, angke foluh pajo, silaora, ciliolate. Ovary pubescent.
sarin silo bulung, tulun surin or seulang (pajo), t.

lungke ali, Simalur I.; W. Java: ki tiwu, S, often /ormo nervosa (K.&V.)Beus. Blumea 19(1971)493.
used as well for M. pinnata and M. sumatrana (also - M. nervosa K. & V., vide supra.
with the addition lalaki, mindi bodas or persawon), Leaves not more than 8(-10)-jugate, with usually
suren ieuweung, S. See also under var. lanceolata f. elliptic glabrous leaflets. Panicles erect, shorter than
nervosa and var. polyptera. c. 50cm, minimum length c. 15 cm. Inner petal most-
Notes. Meliosma lanceolata is generally very ly glabrous. Ovary pubescent to glabrous.
well characterized by its large pendulous panicles and Distr. Malesia: Sumatra (G. Leuser, G. Talak-
its long leaves with many usually lanceolate leaflets. mau), W. Java.
Nevertheless shows a wide variation especially in
it Ecol. Mountain forest, 1300-2900 m altitude.
number but also in shape and size of the leaflets and The tree can reach a height of 30 m by Im diam.
the panicles. On the islands west of Sumatra Vern. Java: ki Ijermeh badak, ki tjermeh beu-
(Simalur, Nias, Batu) specimens are found with nor- reum, S.
mal inflorescences but only 3-5-pinnate leaves, and
elliptic, sometimes subrotund, large leaflets. Transi- b. var. polyptera (Miq.) Beus. Blumea 19 (1971) 492.
tions to this extreme are common. There is another - M. polyptera Miq., vide supra.
deviating form, however, which takes a separate Leaves 12-25-jugate, with most 50 cm long
at
position. It has many small, mostly lanceolate leaf- rachis (including the oblong to
petiole); leaflets
lets which otherwise do not differ from those of M. linear-lanceolate, small, 4-11 by 1-2 cm, entire,
lanceolata. Also the panicles agree with that species. glabrous.
In view of the wide variability in the leaves of M. Distr. Malesia: Sumatra (Asahan, W. Coast).
lanceolata, I have
prefer to include it here and I Fig. II.
reduced it to a variety. The varieties and forms can Ecol. At low altitudes.
be distmguished as follows: Vern. Sumatra: badar badar, Lubuk Alung, tan-
dikat batu, Priaman, simarpapahu, Mula Padang.
a. var. lanceolata.

f.cavcs(3 )6 18 jugate, with up IOC. lOOcm long 5.Meliosma hirsuta Blumh, Rumphia 3 (1849) 200;
rachis (including the petiole); leaflets elliptic to Waip. Ann. 2 (1852) 225; Mio. Fl. Ind. Bat. I, 2

lanceolate, mcdium-si/cd to large, 5 20( 25) by (1859) 616; Sum. (1961) 203; lllustr. (1871) 74;
706 Flora Malesiana [ser. I, vol. 10^
 1989] Sabiaceae (van Beusekom & van de Water) 707

Merr. Enum. Born. (1921) 363; van Beusekom, 1.5-25 by l-IO cm, usually increasing in size to-
Blumea 19 (1971) 493. wards the top of the leaf, base usually acute to
Evergreen small tree, c. 5 m. Leaves 15-20-(or rounded, rarely slightly emarginate, apex acuminate
probably more-)jugate; rachis 50-100 cm including to cuspidate, entire or dentate, usually slightly to
the10-20 cm long petiole, up to c. 6 mm across, densely pubescent, often with domatia; midrib flat to
more or less hirsute, usually with distinctly swollen impressed above; nerves 3-15 pairs, ascending,
base, sometimes sparsely lenticellate; leaflets (sub) looped; petiolules up to 5 cm, terminal one usually
sessile, those in medium and upper part of the leaf longest, not or not much swollen at the base. Panicles
linear-lanceolate, 10-20(-25) by (1.5-)2-3 cm, in- terminal, sometimes somewhat pendulous,
erect,
dex 5-10, the lower ones (ovate-)lanceolate to ovate, dense to lax, widely to narrowly pyramidal,
gradually decreasing in length towards the base of 10-55(-70) cm, usually profusely branched up to
the leaf, up to only c. 3 cm, base rounded to acute, the 4th order, bearing numerous solitary to usually
sometimes slightly oblique, apex acuminate to crowded flowers; primary side-axes usually many, up
caudate, with entire to remotely spinously dentate to 35(-60) cm, lower ones sometimes subtended by
margin, thin-chartaceous, above glabrous except for small to reduced leaves; bracts ovate to narrowly
some pubescence on the midrib, beneath moderately triangular, up to c. 5(-10) mm, more or less pubes-
to sparsely hirsute especially on midrib and nerves, cent. Pedicels absent or up to 3(-4) mm. Mature
without domatia; midrib above flat to slightly im- buds (1.5-)2(-3) mm diam. Sepals 5 or 4, ovate,
pressed; nerves widely apart, (5-)8-12 pairs, as- unequal, the 3 or 4 inner ones 1-1.5 mm, the outer
cendmg, looped and joined into a distinct marginal 1 or 2 usually smaller, often minute, sometimes
nerve situated at 2-4 mm from the margin; venation lowered on the pedicel, sometimes slightly keeled,
distinct, wide, reticulate; petioles absent or up to c. glabrous or pubescent outside, all entire, usually
1 mm, terminal one often longer, up to c. 8 mm, ciliolate. Outer petals usually glabrous. Inner petals
densely hirsute, not swollen at the base. Panicles and more or less deeply bifid, (0.3-)0.6(-l) mm, gla-
flowers as in typical M. lanceolala, but sepals up to brous, ciliolate or fimbriate at the tips, often with a
c. 1.5 mm. Fruit as in M. lanceolala. minute central lobule, often frayed at the tips.

Distr. Malesia: Sumatra (West Coast: G. Malin- Filaments c. 1 mm. Fruit (sub)globose to obovoid,
tang), only one collection. Fig. 11. when ripe (3-)4-10(-ll) diam., with thin mm
Notes. This species was by Blume erroneously mesocarp; endocarp (sub)globose, oblique or not,
recorded to occur in S. Borneo. (2.5-)3.5-9(-10) mm
diam., with more or less pro-
Meliosma hirsuta is doubtless very closely related minently reticulate surface; median keel usually
to M. lanceolala, but very well distinct by its leaf distinct and more or less prominent, at one end
characters. sometimes running out into a small to minute pro-
See also Pimela angustifolia under the dubious cessus or tubercle, and sometimes curving outwards
species. at the other end; ventral pore usually rather narrow,
whether or not sunken.
6.Meliosma pinnata (Roxb.) Maxim. Bull. Ac. Imp. Distr. Throughout SE. Asia, from Sri Lanka and
Sc. St. Petersb. 12 (1867) 64; Melanges 6: 263. - China to Japan; throughout Malesia as far as New
Millingtonia pinnata Roxb. F1. Ind. (1820) 103. -
1 Guinea (incl. New Britain). Fig. 13.
Fig. 12. E c o Forests under moist tropical to subtropical,
1 .

For further synonyms, see under the subspecies; sometimes warm-temperate conditions, on various
for a complete synonymy, see van Beusekom (1 97 1 soils, from sea-level up to c. 3000 m altitude.
494). Notes. Meliosma pinnata covers a very large area
Evergreen, sometimes deciduous tree, small to up in which it has developed a complex and wide varia-
to c. 42 m. Twigs often with conspicuous leaf-scars. tion pattern. It can be divided up into nine well-
Leaves 2- II -jugate; rachis terete, (2-)5-40(-60) marked subspecies. Four of these are widely distrib-
cm, including the up to c. 15(-25) cm long petiole; uted, whereas five have a limited distribution. The
obovate to ovate-
leaflets usually ovate, elliptic, or first group, the subspecies arnottiana, ridleyi,

oblong, sometimes lanceolate, often asymmetric. macrophylla aind ferruginea , are considered primary

Fig. 12. Meliosma pinnata (Roxb.) Walp. s.sp. macrophylla (Murr.) Beus. a. Flowering twig, xO.33; b. half-
opened flower, y 5; c. outer petal with adhering staminodc; d. flower with outer petals removed and stamens
snapped backward; e g. stamen with adhering inner petal, in different positions; h. pistil with surrounding
disk; (. ovary, length section, all x IO;y. ripe fruit, x 3; *-/. endocarp in different positions, x 3 (a Sulit /

F'NH 32941 j I Kostermans 691 ).

. 1
708 Flora Malesiana [ser. I, vol. 10^

Fig. 13. Generalized areas of the subspecies of Meliosma pinnata (Roxb.) Walp., and distribution of M.
sarawakensis Ridley.

subspecies; they centre in W. Malesia. The sub- quired is still wanting. In this respect the picture is

species of the second group occur scattered at the not so complete as it is in M. simplicifolia.
periphery of the area of M. pinnata; I consider them The type subspecies does not occur in Malesia.
secondary offsplits from the primary subspecies, viz.
KEY TO THE SUBSPECIES
ssp. pinnata and ssp. angustifolia (Merr.) Reus.
from ssp. arnottiana, and ssp. pendula, ssp. sylva- 1. Ovary glabrous or only with a few hairs. Sepals
tica, and ssp. humilis from ssp. macrophylla. and petals always glabrous.
The areas of the secondary subspecies fall partly or 2. Leaves 3-5-jugate; leaflets dentate (sometimes
entirely within the area of the primary subspecies only a few teeth), with domatia in the axils of the
from which they are derived, but they are ecological- nerves beneath which are sometimes obscured by
from these, usually by preference for dif-
ly isolated very dense tomentum of the leaf-blade
forms
ferent altitudinal zones; transitional or hybrid g. ssp. humilis
are sometimes found. The areas of the four primary 2. Leaves (3-)4-6(-7)-jugate; leaflets dentate or
subspecies, on the other hand, all touch or only not, without domatia, never with very dense
slightly overlap mutually, but generally they are per- tomentum.
fectly replacing, and usually there is also different 3. Leaflets entire, index (l-)1.5-3, mostly round-
ecological preference. Due to the scarcity of collec- ed or obtuse to truncate or emarginate at the
tions from critical regions, especially Sumatra, base. Medium-sized to large trees
Borneo and Sulawesi, it is mostly not clear how the d. ssp. macrophylla
relation is in contact zones. There is some evidence 3. Leaflets dentate (sometimes very sparsely), in-
that one or two mutually may behave as good dex (l-)1.5-4(-5), acute or rounded, obtuse,
species, where one or two others may be connected truncate or emarginate at the base. Small to
by transitional forms, but in general the evidence re- medium-sized trees, rarely shrubs.
 1989] Sabiaceae (van Beusekom & van de Water) 709

4. Leaflets moderately to rather densely villous- Meliosma floribunda Blume, Rumphia 3 (1849) 200;
pubescent (often more or less glabrescent when Walp. Ann. 2 (1852) 225; MiQ. FI. Ind. Bat. 1, 2
older), mostly (especially lower ones) rounded (1859) 615; lllustr. (1871) 74; K. & V. Bijdr. 9 (1903)
to truncate at the base, index 1.5-3(-4). En- 137; Hall./ Meded. Rijksherb. 1 (1910) 2; Koord.
docarps 6-7 mm diam., without ventral pro- Exk. FI. Java 2 (1912) 546. - Meliosma sambucina
cessus. Above c. 1800 m alt. e. ssp. pendula MiQ. lllustr. (1871) 74; K. & V. Bijdr. 9 (1903) 137,
4. Leaflets sparsely to densely short-pubescent, in obs. - Meliosma luzonensis Merr. Publ. Govt.
rarely subglabrous, mostly with acute base, in- Lab. Philip. 29 (1905) 24; Elmer, Leafl. Philip. Bot.
dex I.5-4(-5). Endocarps 5-7.5 diam., mm 2 (1908) 492, in obs. ('luzonica'); Merr. Enum.
mostly with a small but distinct ventral pro- Philip. FI. PI. 2 (1923) 517. - Meliosma multiflora
cessus. Below c. 1000 m altitude. Merr. Publ. Govt. Lab. Philip. 29 (1905) 25; Enum.
5. Leaves (3-)4-6(-7)-jugate; leaflets acute at Philip. FI. PI. 2 (1923) 517. - Melliosma ferruginea
the base f . ssp. sylvatica (non Blume) Koord. Gedenkb. Jungh. (1910) 177. -
5. Leaves 3-5(-6)-jugate; lateral leaflets Meliosma apoensis Elmer, Leafl. Philip. Bot. 10
rounded to truncate at the base (1939) 3784, descr. angl. - Meliosma cannarioides
d. ssp. macrophylla (Celebes form) Elmer, Leafl. Philip. Bot. 10 (1939) 3785, descr.
1. Ovary entirely, rarely partly, but always densely angl. - Meliosma ferruginea (non Blume) Backer &
pubescent. Sepals and petals glabrous or pubes- Bakh./ FI. Java 2 (1965) \45, p.p., quoad M. glauca
cent. et floribunda.
6. Sepals and usually also outer petals moderately Small to medium-sized, rarely big tree, up to c.

to densely pubescenton the outside. Leaflets en- 20(-30) m. Leaves (2-)3-7(-8)-jugate; leaflets
tire, index (l-)1.5-3 c. ssp. ferruginea ovate to ovate-oblong, elliptic, or lanceolate, small
6. Sepals and petals glabrous or rarely a few hairs to up to c. 25 by 10 cm, index (l-)1.5-4(-5), acute
on the outer sepals only. Leaflets entire or den- to truncate at base, entire or dentate, chartaceous to
tate, index (1-)1 .5-4(-5). coriaceous, often with domatia. Panicles erect,
7. Endocarps 4.5-9(-10) mm diam., usually with spreading, lax to dense, lower primary side-axes
more or less sunken ventral pore. Inner petals usually subtended by small or reduced leaves. Sepals
with fimbriate or ciliolate, rarely glabrous glabrous or the outer ones rarely with a few hairs.
lobes. Leaflets never with domatia Petals glabrous, inner ones sometimes a bit ciliolate

b. ssp. ridleyi or frayed at the tips of the lobes. Ovary densely

7. Endocarps (2.5-)3-4.5 mm diam., not with pubescent, very rarely subglabrous. Endocarps (sub)
sunken ventral pore. Inner petals with usually globose, not or not much depressed, hardly or not
glabrous, sometimes at the tips ciliolate or oblique, (2.5-)3-4.5 mm diam., with distinct, more
frayed lobes. Leaflets with or without domatia or less prominent, fine reticulum, with slightly to
a. ssp. arnottiana rather strongly prominent, blunt to rather sharp me-
dian keel which does not run out into a ventral pro-
a. ssp. arnottiana (Wight) Beus. Blumea 19 (1971) cessus or tubercle; ventral pore not sunken, some-
499. - Sapindus ? microcarpus W. & A. Prod. 1 times a bit elevated.
(1834) 112, non R. & P. (1804); Wight,
nom. illeg., Dist r . Sri Lanka throughout SE. Asia to China,
III. Ind. Bot. (1840) 142; Walp. Rep.
1 (1842)416, 1 S. Korea, Japan and Taiwan; in Malesia: N. Sumatra
423. - Millingtonia arnottiana Wight, III. Ind. Bot. (Karo), Malay Peninsula, W. & Central Java, Philip-
1 (1840) 144, t. 53. - iVellinglonia arnottiana Meiss . pines (Batan Is., Luzon, Mindanao). Rare in W.
PI. Vase. Gen. (Comm.) 2 (1840) 207, in nolo. - Malesia. Fig. 13.
Millingtonia samhucina Jungh. Tijd. Nat. Gesch Ecol Primary or secondary montane rain-forest,
.

Phys. 8 (1841) 365. - Meliosma arnottiana Walp 600-2500 m altitude, on loamy or volcanic soils, also
Rep. I (1842) 423; Thw. Enum. PI. Zcyl. (1858) 59 on limestone if the climate is wet enough. At higher
Bedd. FI. Sylv. 3 (1871)77; ibid. t. 160; Hook./. FI altitudes the subspecies is deciduous. In Malesia but-
Brit. India 2 (1876) 6; Trim. FI. Ceyl. 1 (1893) 315 tresses are sometimes developed, up to 1.5 m high.
Bra.sdis, Indian Trees (1906) 195; Gamble, FI. Pres Field notes Bark dark to light grey, smooth, in
.

.Madras (1918) 256. - Meliosma glauca Blume

I old trees sometimes distantly shallowly fissured. In-
Rumphia 3 (1849) 200. I. I68B, nom. illeg. \ Walp ner bark soft, fibrous, with 'fingers' tapering out-
Ann. 2 (1852) 225; Hassk. Hon. Bog. I (1858) 140 wards into granular tissue, pale pinkish brown to dull
MiO. M. Ind. Bat. I, 2 (1859) 615; K. & V. Bijdr. 9 red or redbrown, also said to be white, and turning
(1903) 135. ind. var. floribunda (Biijmi.) K. & V. salmon red on exposure. Wood light and soft,
•Hall./. Mcdcd. Rijkshcrb. I (1910) 2; KrK)Rn. Fxk fibrous, easily split, white, with large pores and
n. Java 2 (1912) 546; M. Tjibodas 2 (1923) 157 beautiful grain, with prominent rays, hcarlwood in
Baker/ in Rcndlc. J. Bot. 62 (1924) Suppl. 30. older trees striped reddish and white. Leaflets be-
710 Flora Malesiana [ser. I, vol. \0^

neath pale green, often glaucous. Fruits said to be Inner petals with fimbriate or ciiiolate tips, rarely

reddish, green brown, or black when ripe. glabrous. Ovary densely pubescent. Endocarps
Vern. Sumatra: kabung sillang bulung, Batak subglobose to very depressed and oblique, 4.5-9
lang.; Java: dangdur bulu, kawayang, ki sum, ki (- 10) mm
diam., with vague to distinct, more or less

tiwu lalaki, S; Philippines: adope. adupong, prominent, rather wide reticulum, with slightly to
aropong. bantinan, kamug, Ig., bae, If. strongly prominent, blunt to very sharp median keel
Notes. Attention should be given to the relation which often one end runs out into a minute ventral
at

between ssp. arnottiana and ssp. pendula in the processus, the curving at the other end sometimes far

Philippines (for the relation to ssp. macrophylla, see drawn out into a blunt beak; ventral pore hardly to
under that subspecies). In the mountains of Luzon rather deeply sunken.

both subspecies have been collected, ssp. pendula Distr. Malesia: Central Sumatra, Malay Penin-
above 1800 m altitude and ssp. arnottiana from c. sula,Borneo (Sarawak, Sabah, W. Kutai), Philip-
800-900 m up to c. 2400 m. Locally, e.g. on Mt San- pines (Mindoro). Fig. 13.

to Thomas, they have been found together, but Ecol Primary and secondary rain-forest, both in
.

doubtless intermediate specimens are not observed. mixed dipterocarp and in heath forest, on various
It is possible that in such localities these subspecies
soil types, from sea-level up to 1400 m altitude.

mutually behave as species; population studies in the Field notes. Bark mostly smooth, sometimes
field might yield more evidence with regard to this. somewhat scaly or slightly fissured, grey to brown.
The same problem arises in W. Malesia, where 55/?. Inner bark fibrous, pinkish to red or redbrown,
arnottiana has been collected (rarely). In Sumatra turning brown after exposure. Young branches,
and in Java its relation to ssp. ferruginea is in- inflorescence-axes, and leaf-rachises are sometimes
teresting since there is an altitudinal zone of overlap (Singapore) covered with a dense layer of soft dark
between both, though ssp. ferruginea generally oc- reddish brown hairs. Sepals sometimes said to be
curs lower than ssp. arnottiana. In Java the situation purple. Fruit often ± hairy ( 'trichocarpa'), once said
is as follows: ssp. ferruginea is by far the most com- to be bright purple.
mon of both, ssp. arnottiana having only been col- Vern . Sumatra: kaju rokkam, k. rube gala, k. si
lectedon a few mountains. Of these it is only G. Sa- hasur, k. si(mardjuhut) (nij manuk, Asahan, mo-
lak and G. Gedeh where both subspecies have been dang halimponan, Tapanuli.
found. Only of G. Gedeh more detailed ecological Notes . Ssp. ridleyi is rather variable when com-
evidence is available: Koorders (FI. Tjibodas 2, pared to the other subspecies of A/, pinnata, especial-
1923, 157) stated that ssp. ferruginea occurs at c. ly in number and dentation of leaflets, in the degree

1400 m and that ssp. arnottiana ('M. glau-

altitude of pubescence, and in shape and size of the endo-
ca') occupies a zone between 1800 and 2400 m, being carps. In theMalay Peninsula, for instance, a form
especially abundant at c. 2200 m. This does suggest with few subglabrous and somewhat dentate leaflets
the existence of ecological differentiation, but since has been found ('M. elegans'), as well as a beautiful,
ssp. arnottiana on other mountains also grows at densely rufous-pubescent form with distinctly more
lower altitudes, the situation remains unclear. and entire leaflets ('M. ridleyi'). It is not astonishing
that such different plants have been described as
b. spp. ridleyi (King) Beus. Blumea 19 (1971) 505. - separate species; only by studying material from
Meliosma ridleyiKwG, J. As. Soc. Beng. 65, ii (1896) Borneo it becomes clear that these extremes are con-
458; Ridley, J. Str. Br. Roy. As. Soc. n. 33 (1900) nected by a range of transitions. Another form from
67; FI. Mai. Pen. 1 (1922) 516. - Meliosma elegans Dallas (Kinabalu), which has rather condensed
Ridley, J. Str. Br. Roy. As. Soc. n. 54 (1910) 40; FI. panicles, has been described as M. confertiflora.
Mai. Pen. I (1922) 515. - Meliosma paucinervia This again is merely a local form without any
Merr. Philip. J. Sc. 10(1915) Bot. 39; Enum. Philip. systematical significance, as is M. paucinervia, with
FI. PI. 2 (1923) 518. - Meliosma trichocarpa Merr. very lax panicles, from Mindanao. Yet, in spite of
Pap. Mich. Ac. Sc. 24 (1938) 80, nom. non
illeg., this variation, it is obvious that ssp. ridleyi is a natu-
Hand.-Mazz. (1934). - Meliosma Merr.
bartlettii ral unit, probably most closely related to the adjacent
& Perry, J. Am. Arb. 20 (1939) 356. - Meliosma ssp. arnottiana from which it differs least of all
confertiflora Merr. & Perry, I.e. 359. subspecies, mainly in shape and size of the en-
Shrub or tree, up to c. 20 m. Leaves 3-7-jugate; docarps, but also in some
important characters; less
leaflets oblong to lanceolate, small to usually me- an especially close resemblance has been observed
dium-sized, up to c. 20 by 6 cm, base acute, rarely between ssp. ridleyi and some deviating specimens
rounded, usually entire, densely villous to glabrous, from South Vietnam which have been tentatively in-
without domatia. Panicles erect, usually rather lax cluded in ssp. arnottiana. Furthermore, the area of
and slender; lower primary side-axes mostly subtend- ssp. ridleyi borders on or somewhat overlaps the
ed by small leaves. Sepals and outer petals glabrous. areas of ssp. ferruginea and macrophylla. The rela-
1989] Sabiaceae (van Beusekom & van de Water) 711

tion between ssp. ridleyi and these subspecies has pale greyish to glaucous green beneath. Fruits
been discussed under ssp. macrophylla. brownred to black when ripe.
Finally, it should be noted that the area of ssp. Uses Advocated for reafforestation purposes by
.

ridleyi fully overlaps that of M. sarawakensis; this is KOORDERS.

not accidental, since the latter probably is a Vern. Sumatra: sekapong, Takengon, sontang,
derivative of ssp. ridleyi (see the note under M. Simelungun, sihubung, Kerinci; Java: ki tiwu, ki
sarawakensis). tjerme badak, S, gempong, gijubuk, gompong, J;
Lesser Sunda Is.: gempong, sambuk, Bali, mladja,
c. ssp. Blumea 19 (1971)
ferruginea (Blume) Beus. tanggo, tawu, Flores, Endeh lang., lohot, raok,
507. - Meliosma ferruginea Blume, Cat. (1823) 32, Flores, kaju mangkok, W. Sumbawa.
non SiEB. & Zucc. e.x Hook./. (1876), nee Kurz ex Note. Ssp. ferruginea is usually well recognizable
King (1896); Nees, Flora 8 (1825) 106; Hassk. Cat. by its outside pubescent sepals and petals. However,
Hort. Bog. (1844) 226; Blume, Rumphia 3 (1849) in N.Sumatra and the Lesser Sunda Islands
200; Walp. Ann. 2 (1852) 225; MiQ. Fl. Ind. Bat. 1, specimens occur in which these characters are im-
2 (1859) 616; Illustr. (1871) 74; K. & V. Bijdr. 9 perfectly or not developed, and they may also lack
(1903) 121; KooRD. Exk. Fl. Java 2 (1912) 546; Atlas the pubescence on the ovary and may have almost
2 (1914) t. 375; Fl. Tjibodas 2 (1923) 157; Backer & glabrous leaves. They are not easily identifiable and
Bakh./. Java 2 (1965) 145, p.p., exct. M. glauca
Fl. may be confused with A/, lanceolata var. lanceolata
et Jloribunda. - Millingtonia ferruginea Schult. & f.nervosa or with the closely related M. pinnata ssp.
ScHULT. Syst. Veg. Mant. 3, add. 2 (1827) 250; macrophylla and ssp. ridleyi.
DiETR. Syn. PI. 1 (1839) 103.
Medium-sized to big tree, up to c. 42 m. Leaves A. ssp. macrophylla (Merr.) Beus. Blumea 19(1971)
2-6(-7)-jugate; leaflets elliptic to oblong, basal ones 510. - Meliosma macrophylla Merr. Philip. J. So.
sometimes a bit ovate, upper ones sometimes ± 7 (1912) Bot. 294; Enum. Philip. Fl. PI. 2 (1923) 517.
obovate, usually rather large, up to 25(-38) by - Meliosma lanceolata var. obliqua {non Blume)
10(-18) cm, base rounded to truncate, sometimes KooRD. Minah. (1898) 408; Suppl. 2 (1922) 7, t. 55;
acute, entire, firmly coriaceous, pubescent, rarely ibid. 3 (1922) 28. - Meliosma wallichii (non Planch.

subglabrous, rarely with domatia. Panicles erect, ex Hook./.) Koord. Minah. (1898) 408. - Meliosma
spreading, lax to rather dense; lower primary side- tongcalingii Elmer, Leafl. Philip. Bot. 8 (1915)
axes usually subtended by small leaves. Sepals usual- 2815. - Meliosma megalobotrys Mekr. Philip. J. So.
ly densely pubescent, rarely on the outside sparsely 11(1916) Bot. 16; Enum. Philip. Fl. PI. 2 (1923) 517.
so to subglabrous. Outer petals pubescent outside, - Meliosma macrocarpa Elmer, Leafl. Philip. Bot.
rarely glabrous. Inner petals with fimbriate or 10 (1939) 3786, descr. angl. - Meliosma ferruginea
ciliolate tips.Ovary partly or entirely but almost (non Blume) Merr. & Perry, J. Arn. Arb. 20(1939)
always distinctly and densely pubescent, very rarely 356. - Fig. 12.
nearly glabrous. Endocarps subglobose, often some- Medium-sized to large tree, up toc. 42 m. Leaves

what depressed and oblique, 3.5-5.5(-8) mm diam., (3-)5-9-jugate; leaflets elliptic to oblong to ovate-
with rather vague to distinct, ± prominent reticulum, oblong, medium-sized to rather large, up to c. 20 by
with usually very prominent, rather sharp median 9 cm, base rounded or obtuse to truncate, entire,
keelwhich does not run out into a ventral processus or rarely with a few teeth (Sulawesi), chartaceous to
tubercle; ventral pore not or not much sunken. firmly coriaceous, very sparsely to densely pubes-
Distr. Malesia: N. & Central Sumatra, through- cent, always without domatia. Panicles erect and
out Java, and the Lesser Sunda Islands (Bali, Sum- spreading, lax and slender to rather dense; lower
bawa, Flores, Timor), locally common, especially in primary side-axes usually subtended by small leaves.
Java. Fig. 13. Sepals and petals glabrous. Ovary glabrous, rarely
Ecol. Rain-forest, preferably on fertile, often with a few scattered hairs. Endocarps subglobose,
volcanic soils, 250-1600 m altitude. sometimes more obovoid or depressed, more or less
Field notes. Bole cylindrical, straight, some- oblique, 3.5-5mm diam., exceptionally 5-7.5 mm
times crooked, at the base up to c. 2.5 m diam. Bark diam. (Sulawesi), with vague to distinct and promi-
on the surface grey to brown, smooth, sometimes a nent reticulum, with rather sharp and prominent me-
bit peeling or shallowly fissured to (deeply) cracked, dian keel which at one end mostly runs out into a
about 0.7-1.5 cm thick, easily detachable. Inner small but distinct ventral processus or tubercle; ven-
bark pale brown to brownrcd or orange, with tral pore .somewhat sunken.
streaks, also said to be dirty while and turning orange Distr. Malesia: E. Borneo (E. Sandakan, Bcrao,
brown when exposed to the air as a result of the W. & E. Kutai, Tandjung), Sulawesi (Minahasa,
discolouring of the initially colourless watery exuda- Malili), Moluccas (Halmahcra, Scram). Philippines
tion. Wood soft, yellowish to pinkish white. Leaflets (Luzon, Lcytc, Mindanao. Palawan), throughout
712 Flora Malesiana [ser. I, vol. lO-*

New Guinea (incl. New Britain). Fairly common in seems to be restricted to lowland forests below c. 100
most parts of the area. Fig. 13. m altitude!). To the SW the area o^ ssp. macrophylla
Ecol. Usually in primary, rarely in secondary borders on that oi ssp. ferruginea which inhabits the
rain-forest, at low to medium altitudes; in Borneo Lesser Sunda Islands. The latter two subspecies are
only collected below 100 m. in the other parts of the huge trees, very similar in general habit, and some-
area also higher, up to c. 100-1200 m, in W. New1 times they have been confused. Nevertheless, they
Guinea once at 1800 m. in Borneo usually found in are usually well distinct, mainly by flower characters,
lowland dipterocarp forests. Generally reported to though in both subspecies there is a tendency to lose
occur on clayish, loamy, or sandy clayish soils, also some of these characters.

on red earth, on volcanic soil, and on loamsoil on In the Philippines ssp. macrophylla is sympatric
limestone. It found in occasionally sub-
is rarely with ssp. arnottiana, but they prefer different

merged areas. Once (New Guinea) said to occur on altitudinal zones, the first being a lowland subspecies
peaty soil, and there developing stiltroots. not exceeding c. 900 m altitude, the latter being a

Field notes. Bole mostly straight, cylindrical, montane subspecies occurring from c. 800 up to c.

up to at least 1 m diam. at the base, usually develop- 2400 m (once recorded from c. 600 m).
ing 1.5-2.5 m high buttresses, sometimes without
buttresses,once observed stiltrooted. Bark grey to Blumea 19(1971)512.
e.spp. pendula (Merr.) Beus.
brown, or patchy brown-white-grey, smooth, some- - Meliosma pendula Merr. Publ. Govt. Lab.
times with shallow vertical cracks, not or little peel- Philip. 29 (1905) 25;Enum. Philip. Fl. PI. 2 (1923)
ing, with vertical rows of lenticels. Inner bark c. 1 cm 518. - Meliosma reticulata Merr. Philip. J. Sc. 5
thick, soft, light brown or pink to brownred, inside (1910) Bot. 195; Enum. Philip. Fl. PI. 2 (1923) 518.
paler,sometimes said to be streaked with cream, with - Meliosma macgregorii Merr. Philip. J. Sc. 10
some colourless sticky exudate (which is also said to (1915) Bot. 37; Enum. Philip. Fl. PI. 2 (1923) 517.
be redbrown!); it is said to be rapidly darkening upon Small to medium-sized tree, up to c. 20 m. Leaves
exposure or 'a bright orange-brown stain quickly ap- (3-)4-6-jugate; leaflets elliptic to oblong, rarely lan-
pears between bark and sapwood.' Wood very light ceolate, up by 7(-ll) cm, the lower ones
to 18(-20)
and soft; sapwood white to pale pink or brown, when at the base nearly always rounded or (sub)truncate to

fresh with bright brown sap streaks,heartwood ab- obtuse, the upper ones more or less acute, nearly
sent or present, darker than the sapwood. The fruit always distinctly dentate, villous-pubescent, ± gla-
is brown to black.
said to be brescent, without domatia. Panicles erect and
Vern. Borneo: surian, E. Kutai; Sulawesi: kaju- spreading to somewhat pendulous and rather flaccid,
saul-rintek, Tooelooe lang., papako, Tontemboan slender and rather lax; lower primary side-axes most-
lang., mumping, Tonsea lang., Hasan, Ratahan ly subtended by small leaves. Sepals and petals
lang.; Moluccas: bais, Seram; Philippines: arocong, glabrous. Ovary glabrous. Endocarps subglobose,
Ig., agosos, balilang-uak (a corruption of barilan ng slightly oblique, 6-7 mm diam., with rather vague,
uak). Tag., morau, S.L.Bis., mungapong, Bik., ma- slightly elevated reticulum, with hardly to moderate-
gasorod. Bag.; New Guinea: sebotebuk, tubuk, ly prominent, blunt median keel, the latter not run-
Mooi, serajema, Manikiong, marwaskeipi, Japen, ning out into a distinct ventral processus or at most
bagare, Kapauku lang., biedewon, iediewat, Muju, into a very minute tubercle; ventral pore hardly or
morrotuno, waito, Wapi, frikipa, Orne lang., not sunken.
tapuha, Managalase, kufi, Kutubu, uliga, Madang, Distr. Malesia: Philippines (Luzon: Mountain
kombowase, Waskuk, wagebi, Wagu. Prov.). Fig. 13.
Notes. Ssp. macrophylla is the most common Ecol. Montane rain-forest, 1800-2500 m alti-
and widespread of the East Malesian subspecies tude. In mossy forest, in ravines as well as on expos-
group, characterized by a glabrous ovary by which it ed ridges.
is readily distinguished from the West Malesian Field notes. Bark thick, checked. Wood soft,
subspecies. Within its large area a few other sub- soon assuming an orange-brown colour.
said to be
species occur, viz. ssp. pendula, sylvatica, and Uses. The leaves are once said to be used for
humilis, which have much more limited areas and smoking by the Igorots.
probably represent offsplits from it. These three Vern. Anitap, Ig.
subspecies are ecologically well isolated from ssp. Note. Ssp. pendula replaces the lowland and
macrophylla. lower macrophylla at high elevations (in this
hill ssp.
In Borneo the area oissp. macrophylla is, as far as respect being comparable to ssp. humilis from New
can be judged from the available evidence, sharply Guinea).
delimited against that of the West Malesian ssp.
ridleyi, which, moreover, appears to prefer a higher f.ssp. sylvatica (Elmer) Beus. Blumea 19(1971)513.
altitudinal zone (only in Borneo ssp. macrophylla - Meliosma sylvaticaElmer, Leafl. Philip. Bot.
1989] Sabiaceae (van Beusekom & van de Water) 713

2 (1908) 492; Merr. Enum. Philip. Fl. PI. 2 (1923) and with slender axes; lower primary side-axes often
518. - Meliosma acuminatissima Merr. Philip. J. subtended by small leaves. Sepals and petals gla-
Sc. 10 (1915) Bot. 36; Enum. Philip. Fl. PI. 2 (1923) brous. Ovary glabrous. Endocarps subglobose,
517. - Meliosma brachyboirys Merr. Philip. J. Sc. somewhat depressed and rather oblique, 5.5-7.5
12 (1917) Bot. 275; Enum. Philip. Fl. PI. 2 (1923) mm diam., with more or less vague, slightly elevated
517. reticulum, with very prominent, rather sharp median
Slender shrub or treelet, up to c. 5 m. Leaves keel which at one end runs out into a small but
(3-)4-6(-7)-jugate; leaflets elliptic to usually ob- distinct ventral processus; ventral pore somewhat
long or lanceolate, usually medium-sized, up to c. 18 sunken.
by 6 cm, acute at the base, sparsely to rather closely, Distr. Malesia: Papua New Guinea (Highlands
always distinctly dentate, very sparsely to moderately Provinces, Madang, Morobe, Milne Bay). Common.
pubescent, without domatia. Panicles erect, spread- Fig. 13.
ing, usually slender and rather la,\; primary side-axes Ecol. Montane rain-forests, 1000-30(X) m
(mostly?) not subtended by small leaves. Sepals and altitude.Observed as an understorey tree in dense
petals glabrous. Ovary glabrous. Endocarp subglo- Castanopsis-Nothofagus forest, on ridges as well as
bose, oftensomewhat ellipsoid, more or less oblique, on streambanks, but also often reported from several
5-6 mm diam., with distinct, more or less prominent kinds of disturbed forest, such as bamboo regrowth,
reticulum, with rather sharp and prominent median old garden land, transition between coniferous forest
keel which at one end runs out into a small but and treefern grassland, and even from open grass-
distmct ventral processus; ventral pore somewhat land. Once reported from limestone ridge.
sunken. Field notes. Bark greybrown, smooth, with big
Distr. Malesia: Sulawesi (Minahasa, Latimod- lenticels. Inner bark straw-coloured to pink, red, or
jong Mis), Philippines (Luzon, Negros). Fig. 13. reddish brown (due to discolouring, as in other sub-
Ecol Lowland rain-forest, usually not above 750
. species?), exuding 'resin'. Wood white to light

m altitude, growing in the shrub layer. brown, with conspicuous rays and clear growth
Field notes. Slender, suberect or bent shrub or rings, said to be of moderate weight and hardness.
treeletof a sparsely branched habit. Bark smooth, Petioles, peduncle, and pedicels purplish to red-
grey and brown mottled. Wood white, soft, easily brown; buds reddish. Fruits dark red to black when
breakable. Leaves once said to be light bluish green ripe.

beneath. Uses. Once said to be used as housing timber,

Note. Ssp. sylvatica is closely related to ssp. free from borers.
macrophylla. The most striking difference between Vern. Mansalong, Finschhafen, kokopong,
them found in their physiognomy, the former be-
is Nako lang., E. Madang Prov., kass, Maring lang.,
ing a small undergrowth treelet, the latter a large mappam, Enga lang., W. Highl. Prov.
forest tree. The main systematical differences are Notes. Ssp. humilis is closely allied to ssp. ma-
found in the dentation and the shape of the base of crophylla, mainly differing by its dentate leaflets
the leaflets. These fit nicely in the spectre of with domatia. In Papua New Guinea it replaces ssp.
character combinations present in the subspecies of macrophylla mainly found below 1000 m altitude (cf.
M. pinnata, and it seems justified to consider ssp. ssp. pendula from Luzon). It is remarkable that ssp.
sylvatica a subspecies of that species, instead of a humilis has as yet been collected, even rather abun-
separate species. dantly, only in Papua New Guinea and not in W.
New Guinea.
R. spp. humilis (Merr. &Perry) Beus. Blumea 19
(1971) 514. - Meliosma humilis Merr. & Perry, J. 7. Meliosma sarawakensis Ridley, Kew Bull. (1933)
Arn. Arb.20(1939)358;/*/Vy. 22 (1941) 263, //j 065. 193; Merr. & Perry, J. Arn. Arb. 20 (1939) 359. -
- Meliosma schlechteri Merr. & Perry, J. Arn. Meliosma grandifolia Lecomte, Bull. Soc. Bot. Fr.
Arb. 22(1941) 262. 54(1909)676, now. ///e/?.. non Urban (1895); Merr.
Small to medium-sized tree, up to c. 20 m. Leaves Enum. Born. (1921) 362; van Beusekom. Blumea 19
3-5 jugate; leaflets elliptic to oblong, rarely short- ( 971 5 5 - Meliosma confusa var. laxior Baker /.
1 ) 1 .

lanceolate, usually ralhcr small, up to c. 15(~24) by in Rendle, J. Bot. 62 (1924) Suppl. 30. - Meliosma

6<-9) cm, the base acute, lower leaflets sometimes lattfolia RitUEY. Kew Bull. (1933) 193; Merr. &
rounded at the base, sparsely to rather closely den- Perry, J. Arn. Arb. 20 (1939) 359. m obs.
tate, beneath subglabrous to ralhcr densely pubes- Evergreen, small tree, up to c. 10 m. Leaves
cent, sometimes densely villous-tomcntose, always 2-3(-4)-jugate; rachis terete, 12-30 cm, including
with more or
less distinct domatia in the axils of the the 6-15 cm long petiole, up to c. 5 mm diam..
nerves beneath (obscure in densely lomentose densely short-tomcntosc. later ± glabresccni; leaf-
leaflets). Panicles erect, spreading, mostly rather lax lets usually elliptic to oblong, the lower ones often
714 Flora Malesiana [ser. I, vol. 10^

more or less ovate to ovate-oblong, the upper ones ly pubescent sepals. These characters indeed are also
often more or less obovate to obovate-oblong, found in M. pinnata ssp. ferruginea, but this subspe-
sometimes ± asymmetrical, (2-)5-22 by (1.5-) cies is quite different from M. sarawakensis in
3-12 cm, mostly distinctly increasing in size towards various other aspects. An additional argument to the
the top of the leaf, base acute to rounded, apex more specific status of M. sarawakensis
is found in the fact

or less acuminate, sometimes subacute or cuspidate, that found together with M. pinnata ssp. ridleyi
it is

with entire to remotely spinously dentate margin, in the same area and at the same altitudes in Sarawak

chartaceous, moderately to rather densely pubescent (near Kuching) and in Sumatra (Asahan), without
especially beneathand on midrib and nerves, often any sign of hybridization.
when older, never with domatia;
partly giabrescent
midrib more or less impressed above; nerves 6-12 8. Meliosma rufo-pilosa Hend. Card. Bull. Str.
pairs, ascending, usually looped; venation distinct, (1933) 96, t. 18; Merr. & Perry, J. Arn.
Settl. 7

reticulate; petiolules up to c. 1.5(-3) cm, terminal Arb. 20 (1939) 360; van Beusekom, Blumea 19
one usually longest, tomentose. Panicles terminal, (1971) 517.
usually more or less pendulous, flaccid, lax, narrow- Evergreen rather large tree up to c. 30 m. Flower-
ly pyramidal, (20-)25-55 cm, not profusely ing twigs terete, 5- 10 mm diam., stout, abruptly ter-
branched up to the 2nd or 3rd order, branches minating in a tuft of leaves and inflorescences, gla-
spreading, ± flaccid, usually slender, densely brous, often with many
large conspicuous leaf-scars.
tomentose, bearing numerous flowers crowded in Leaves (6-)7-9-jugate; rachis terete, (13-)25-50
dense spikes; primary side-axes few to rather many, (-65) cm, including the (3.5-)6- 16 cm long petiole,
up to c. 25(-35) cm, the lower ones usually subtend- pubescent, not swollen at the base, hardly or not len-
ed by reduced leaves; bracts ovate to usually narrow- ticellate; leaflets elliptic to oblong, sometimes ovate
ly triangular or linear-lanceolate, up to c. 4 mm, (-oblong), 3- 15 by 2-6 cm, base obtuse to truncate,
densely pubescent. Pedicels (almost) absent. Mature apex acuminate, entire, glabrous or ± puberulous
buds c. 2 mm diam. Sepals 5 (4), ovate to ovate- on nerves above, (sub)glabrous beneath, pubescent
lanceolate, the 3 or 4 inner ones 1-1.5 mm, the outer on nerves, always without domatia; midrib im-
1 or 2 usually much smaller, often minute, densely pressed above; nerves 7-18 pairs, ascending, looped;
pubescent on the outside, with entire margin. Outer venation fine, very distinct, reticulate; petiolules 1-6
petals glabrous. Inner petals about halfway or some- mm, densely pubescent. Panicles terminal, one or a
what less bifid, 0.5-0.7 mm, glabrous, sometimes few crowded together at the end of a twig, erect,
with a minute central lobule. Filaments c. 1 mm. rather dense to lax, pyramidal, 30-50 cm, including
Ovary 0.5-0.7 mm, densely pubescent. Fruit the0-20 cm long peduncle, profusely branched up to
(sub)globose, when ripe 0.7-1 cm diam.; endocarp ± flaccid, densely
the 4th order, branches spreading,
depressed-globose, applanate at the ventral side, pubescent, bearing numerous solitary flowers;
strongly oblique, 6-7(-8) mm diam., with usually primary side-axes well-spaced, c. 8-15, up to c. 30
distinct, more or less sharply prominent reticulum; cm, not lenticellate, the lower ones never subtended
median keel sharp and very prominent, not at one by small or reduced leaves; bracts narrowly triangu-
end running out into a ventral processus or tubercle, lar to lanceolate, up to c. 4 mm, densely pubescent.
at the other end rather far curving outwards; ventral Pedicels 1-3 mm, densely pubescent. Sepals (3) 4,
pore rather sunken. ovate, (sub)equal, c. 1.5-2 mm, the outer one often
Distr Malesia: Sumatra (Asahan to Palembang)
. much smaller, rarely minute, usually lowered on the
and NW. Borneo (Sarawak, around Kuching and pedicel, glabrous or somewhat pubescent outside;
Pontianak; common). Fig. 13. margin flimsy, more or less ciliolate, entire or some-
Ecol. Lowland rain-forest, up to c. 800 m times with some coarse irregular teeth. Outer petals
altitude. c. 1.5 by 1.5-2.5 mm. Inner petals ligular, usually
Field notes. The redbrown to purple.
sepals are somewhat widened towards the top, 0.7-1 mm; top
Vern. Sumatra: kaju rube boras, Asahan; Bor- entire or with a shallow incision, blunt, minutely
neo: bulu manok, Iban name, Kuching. ciliolate. Filaments 0.7-1 mm. Ovary 0.5-0.7 mm,
Note. The closest affinity of M. sarawakensis is glabrous. Fruit globose, when ripe 1.5-2 cm diam.,
doubtless with M. pinnata ssp. ridleyi to which it is with moderately thick, fleshy mesocarp; endocarp
very similar in all characters (they even share the red semiglobose, broad-ovate to subcordate at ventral
sepals).Only after some hesitation M. sarawakensis view, 11-13 mm
long and wide, 7-8 high, with mm
ismaintained as a separate species and not made a relatively thin wall, with slightly lumpy surface,
subspecies of A/, pinnata. It would fit rather well into especially lumpy and somewhat furrowed at the ven-
that species but is distinguished from it by a wider tral curving of the wall; median keel faint, hardly
range of characters. The most important of its elevated but at one end drawn out into a con-
characters are the 2- or 3-jugate leaves and the dense- spicuous, laterally flattened, downwards-curved.
1989] Sabiaceae (van Beusekom & van de Water) 715

blunt beak; ventral side rather deeply concave with a crown, once reported with c. 2 m high buttresses,
smooth, wide-ovate to suborbicular central part Bark smooth, grey to brown, with lenticels in vertical
from the centre of which protrudes the ± conical rows {'scarred', 'dippled'). Inner bark soft, fibrous,
hilum of the seed. orange to reddish outside, pale fawn to white to-
Distr. Malesia: Malay Peninsula (Pahang), wards the cambium. Sapwood pale brown. Twigs
Borneo (Sarawak, Sabah, Kinabalu complex). Fig. pale brown, rough, lenticellate, with darker leaf-
1 1 scars. Leaves pale green. Fruit yellow to orange when
Ecol. Montane rain-forest, 1350-1700 m alti- ripe,

tude. Vern. Malay Peninsula: sengkuang, Genting

Field notes. Large tree with deep, rounded Highlands.

Excluded and dubious

Meliosma celebica Warb. ^.v Dihm, Beih. Bot. Centralbl. 21, 1 (1907) 125, nomen. - I have not seen the
type specimen (Warburg 15416, Sulawesi, Bojong), which probably got lost during World War \\.

Meliosma laurina Bllme, Rumphia 3 (1849) 198; Walp. Ann. 2 (1852) 224; Miq. F1. Ind. Bat. 1, 2 (1859)
Merr. Enum. Born. (1921) 363; Hall./. Beih. Bot. Centralbl. 39, 2 (1921) 161;
614; lllustr. (1871) 73;
KosTER.M. Bibl. Laur. (1964) 951. - As was noted by Hallier/., I.e., the type specimens (S. Muller^.a;.,
1

Borneo, G. Sakumbang) consist of a mixture, viz. inflorescences of M. sumatrana and leaves of Cryptocarya
reticulata Blume (Lauraceae).

Meliosma petiolaris Miq. Sum. (1861) 519, 203; lllustr. (1871) 73, in obs. - This species was later referred
by MiQUEL himself to Xylosma leprosipes Clos which is now known as Bennettiodendron leprosipes (Clos)
.Merr. (Flacourtiaceae).

Meliosma timorensis Blume ex Blenk, Flora 67 (1884) 370, nomen. - This name was cited in an enumera-
tion oi Meliosma species having leaves with pellucid dots; it was probably copied from a label on a sheet. The
specimen could not be traced.

Pimela angustifolia BivMt, Mus. Bot. Lugd.-Bat. (1850)226. - Canariopsis angustifolia Blvme e.x Miq.
1

Fi. Ind. Bat. 1, 2 (1859) 653. - Canarium angustifolium Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 117;
H.J. Lam, Bull. Jard. Bot. Btzg III, 12(1932) 179. t. 11 f. l\d,subC. rigidum Zipp.; Leenh. F1. Males. 1,

5 (1956)296.
The material under this name was excluded from the Burseraceae by Leenhouts, I.e., and tentatively
assigned to Meliosma. This may be correct, and should then be placed close to M. laneeolata and M. hirsuta.
it

At first sight it isvery similar to the latter species, but there are important differences in nervation and
pubescence. If it belongs to Meliosma it would certainly be a new species, but I refrain from including it
because I am not sure about its identity. Unfortunately, the specimens consist of young leaves only, with many
characteristic narrow leafiets, but in absence of woody parts it cannot be identified with certainty. Moreover,

on the original labels (Zippel^./;.) the place of origin is mentioned as being 'Nova Guinea', in Blume's hand-
writing. However, this would not fit in the distribution pattern of Meliosma, since species of this kind only
occur in western Malesia; if New Guinea indeed is the correct locality, 'Canarium angustifolium' can hardly
belong to Meliosma. Its identity will probably remain uncertain until more satisfactory material has been
found.
 ADDENDA, CORRIGENDA ET EMENDANDA
C. G.G.J. VAN SteenisI, W.J.J.O. de Wilde, c.s.

As was done in it seemed useful to correct some errors which have crept into the text
the preceding volumes,
of volumes 4- 1 add some additional data, new records and references to new species which came
as well as to
to our knowledge and are worth recording.
Volume and page number are separated by a colon. Page numbers provided with either a ot b denote the
left and right columns of a page respectively.

Araliaceae Cyperaceae

9: 47b Osmoxylon sessiliflorum (Laut.) Philip- 7: 653a Cyperus diaphanus Schrader ex R. & S.
SON. var. latespicatus (Boeck.) Kern.

Add to Distr.: Moluccas (Halmaheira), 2 Add to Distr.: Lesser Sunda Islands

coll. (Flores: Schmutz 5767).
Add to Vern.: saha-saha, Ternate lang., 9: 173a Carex bilateralis Hayata.
tele, Sahu lang. Add to Distr.: SE. to E. Asia.

Bignoniaceae Dipterocarpaceae

8: 142b Dolichandrone spathacea (L.f.) K. Sch. 9: 436b Hopea gracilis Miq. (under Excluded).
Add to Distr.: A.G. Wells, in H.J. Teas It was unfortunately overlooked that the
(ed.). Biology and ecology of mangroves type at L (W. Central Sumatra, Padang,
(1983) 61, map, records this species for the Teijsmann HB 424) was already in 1968
first time from Australia: the northern tip referred by Ashton to Meiogyne and later
of Cape York Peninsula (2 localities). in the same year identified by F.H. Hilde-
BRAND as Meiogyne virgata (Bl.) Miq. {An-
Cardiopteridaceae nonaceae).

7: 93 Cardiopteris [Wall, ex] Royle. Ericaceae

Add The genus extends to the
to Distr.:
Solomon Islands (Bougainville) and N. 6: 856b Vaccinium angulatum J.J.S. 1914, non
Queensland; cf. Bailey, Queensl. Fl. (Griff.) Theobald, 1883 = Vaccinium
(1899) 251; Compreh. Cat. Queensl. Pi. commutatum Mabberley & Sleumer,
(1912) 93, f. 76. Taxon 34 (1985) 155, nom. nov.

Celastraceae Flacourtiaceae

6: 257b Glyptopetalum loheri Merr. 5: 24b Hydnocarpus nana King.

Distr.: Philippines: add Palawan (Rids- Add to Distr.: NE. Sumatra (Besitang R.,
DALE 683). Sikundur For. Res., de Wilde & de Wilde-
6: 266a Lophopetalum beccarianum Pierre. Duyfjes 19540).
Add to Distr.: Central Sumatra (Djambi: 5: 35a Scaphocalyx spathacea Ridl.
Vreeken-Buys 69). Add the synonym: S. parviflora RrDL. etc.
6: 291 b Perrottetia alpestris (Bl.) Loes. ssp. philip- 5: 35b Reduce Scaphocalyx parviflora Ridl. to S.
pinensis (Vidal) Ding Hou. spathacea Ridl.
Add to Distr. (and map): Lesser Sunda 5: 56a Homalium dasyanthum.
Islands (Flores: Loeters 650, 1796). Change authorship: (Turcz.) Theob. in
Mason, Burma ed. Theob. 2 (1883) 451;
Convolvulaceae Warb. etc. (see Mabberley, Taxon 34,
1985, 155).
4: 434b Jacquemontia brown iana Ooststr. 5: 98b Reduce Casearia pallida Craib. to C
Add the synonym: J. pannosa (R. Br.) flavovirens Bl.
Mabberley, Bot. Macar. 6 (1980, '1978') 5: l(X)b Casearia flavovirens ^i..
63. Add the synonym: C. pallida Craib, etc.

(716)
 1989] Addenda, corrigenda et emendanda 717

Add to Distr.: Thailand; in Malesia: Suma- Iridaceae

tra, Java, Lesser Sunda Islands (Bali).
8: 83 The correct name for Gladiolus natalensis
Haemodoraceae (Ecklon) Reinw. ex Hook, should be
Gladiolus daienii Geel, Sen. Bot. fasc. 28
5; 113a Haemodorum corymbosum Vahl should (1829). See Milliard & Burtt, Notes Roy.
be called Haemodorum coccineum R.Br., Bot. Card. Edinb. 37 (1979) 297.
according to T.D. Macfarlane, Perth,
Australia. Olacaceae

Hamamelidaceae 10: 7a The correct name

for Olax scandens Roxb.
should Olax psittacorum (Willd.)
be
5: 366 Maingaya malayana Oliver. Vahl; see Almeida, J. Bomb. Nat. Hist.
A marvellous colour photograph of this Soc. 81 (1985) 742.
very rare species (and genus), endemic in 10: 9b E.\cluded, add:
Malaya and Penang, was given by Francis Olax baliculin Blanco, F1. Filip. ed. 2
Nc in Nature Malaysiana 7, no 4 (1982) 8; (1854) 589; ed. 3 (1877) 38; Kosterm. Bibl.
the species is thought to be ± extinct and is Laur. (1964) 1153 = Litsea baliculin
now grown at the Forest Research Institute, (Blanco) Kosterm. Bull. Bot. Surv. India
Kepong. 10(1968) 268 (Lauraceae).
Merrill (Enum. Philip. Fl. PI. 2, 1923,
Hydrocharitaceae 195) recognized it as a Litsea and said that
it might prove to belong to L. leytensis
5: 390b. Blyxa aiiheriii Rich. Merr. Since no material is known to exist,
391 Add: var. echinosperma (Clarke) Cook & the matter will remain speculative.
LiJOND, Aquat. Bot. 15 (1983) 14.

Add the synonym: B. echinosperma Passifloraceae

(Clarke) Hook./, elc.
5: 392a Blyxa leiosperma Koidz. 7: 416a Passiflora aurantia Forst. /.
Reduced to B. japonica (Miq.) Maxim, ex Add to Distr.: W. Central Celebes, near
AscHERS. & Gl'rke var. japonica. Palu, at 200 m, Meijer 10172.
5: 392b Blyxa aliernifolia (Miq.) Hartog.
Reduced to B. japonica (Miq.) Maxim, ex Pittosporaceae
Aschers. & GuRKE var. alternifolia (Miq.)
Cook & Luond, Aquat. Bot. 15 (1983) 25. 5: 356b Pittosporum moluccanum (Lamk) Miq.
5: 393a fl/vx^crvopoAj/co (Miq.) Maxim, ex Aschers. Add the synonym: Coffea angustifolia
& Gurke. RoxB.;c/. FoRMAN,Kew Bull. 38(1983)64.
Add: var. japonica and var. aliernifolia
(Miq.) Cook. & Luond, Aquat. Bot. 15 Podostemaceae
(1983) 22 and 25 respectively.
Add the synonyms: B. leiosperma Koidz. 6: 66b Cladopus nymani H. Moll.
and B. alternifolia (Miq.) Hartog. Add to Distr.: Central Borneo, en route
from Sinar Baru to Ryan Ruwan, N of
Hypericaceae Long Bawan, Krayan, 5° 45 E, 4° 5 N,
1 1 ' '

on rock in a rapid stream (Pa Raya), sub-

8: 18b Replace Hypericum uralum Bif h.-Ham. merged and sterile, 1150 m alt., M.
ex D.Don by Hypericum henryi Lev. & Okamoto C.S., s.n. (L, Osaka). The
Vaniot ssp. hancockii Robson, Bull. Br. material matches the figure on p. 66 of Fl.
Mus. Nat. Hist. Bot. 12(1985)261, map 22. Males. I, 4 almost exactly.
Distr. of var.: Continental SE. Asia (S.
China, Vietnam, Burma, Thailand). Rhizophoraceae

Icacinaceae 5: 461b Bruguiera gymnorrhiza (L.) Saviony.

Distr. It is worthy of record that according
:

7: 6a Cilronelta suaveolens (Hi.) Howard. to an internal report by A.N. Gillkson

Add to Distr.: Lesser Sunda Islands (C'SIRO, Canberra) this mangrove species

(Flores: Schmutz 5819). is found on the lower limestone terrace on

I
718 Flora Malesiana [ser. I, vol. 10"^

Christmas I. (Indian Ocean), in a healthy long. Male flowers: filaments up to 6 or

and self-maintaining community near the more mm long; anthers (0.8-)l -2(-2.2)
treshwaler Hosnie's Springs al 40 m alt., mm long. Fruits fusiform, sometimes con-
and not tar from it a stand of a remnant ht- around the middle, 5-6 mm long,
stricted

toral forest of Heriliera littoralis Dryand. lightbrown, dull, tapering below into a
in Ait., another mangrove species. 1.5-3 mm long pedicel. Endocarp c. 2 mm
5 : 471a Ceriops decandra. wide.
The authorship must be (Griff.) Distr. E. India to Japan; in Malesia:
Theobald; see Taxon 34 (1985) 154. Sumatra, Papua New Guinea (up to 2000 m
alt.).

Simaroubaceae
Sparganium subglobosum Morong, Bull.
Torrey Bot. Club 15 (1888) 81, t. 79, f. 1;
6: 218b Ailanthus inlegrifolia LAMk.
Cook & Nicholls, Bot. Helv. 97 (1987) 4,
Add to Distr. (and map): N. Queensland
t. 13b, 14b, map 15.
(several localities) (L. Pedley, in litt.).
Basal leaves usually exceeding the in-
florescence, (l-)2-4(-9) mm wide. Low-
Sparganiaceae ermost bract up to 15 cm long, 0.5-1 (rare-
ly more) times as long as the inflorescence.
4: 233 The correct name for Sparganium simplex Female heads solitary, axillary or on lateral
HuDS./. simplex as used by Backer should branches, occasionally one head supra-
be: axillary, sessile, rarely with an up to 5 mm
Sparganium fallax Graebn. in E. & P. Pfl. long peduncle. Male heads solitary. Female
R. 2 (1900) 15, t. 3H; Cook & Nicholls, flowers: perianth segments free, in fruit

Bot. Helv. 96 (1986) 253; ibid. 97 (1987) 3, 0.3-0.5 times as long as the fruit; pedicel

t. 13a, 14a, map 14. - S. simplex auct. non O-I mm long. Male flowers: filaments
HuDS./. simplex: Backer, FI. Males. I, 4 2.5-3. 2(-3. 8) mm long, anthers 0.5-0.9
(1951) 233, fig. (-1) mm long. Fruits obovoid to almost
Basal leaves usually exceeding the stem, globose, yellowish to pale brown, shiny,
(4-)5-10(-15) mm wide. Lowermost bract subsessile or with an up to mm long, not
1

20-35(-60) cm long, -several times as 1

persistent pedicel. Endocarp 1.6-1.8 mm
long as the inflorescence. Female heads wide.
3-4(-6), usually supra-axillary, sometimes Distr. India (once, in Khasia), Yunnan,
extending beyond the next internode; ped- Vietnam to Manchuria and to Okinawa,
uncle up to 3 cm long. Male heads 5-8 or Australia, New Zealand; in Malesia: Papua
more. Female flowers: perianth segments at New Guinea.
least connate below, in fruit more than 0.5 Note. The identity of a collection from
times as long as the fruit; pedicel 1-3 mm Arfak is not resolved by Cook & Nicholls.

Symplocaceae

8: 217 Symplocos J acq. subg. Hopea Clarke.

Add in key couplet la as follows:

1. Leaves (pseudo-)verticillate.
la. Leaves obovate 55. S. verticillifolia
la. Leaves elliptic.

2. Upper side of leaves hairy. Twigs tomentose 30. S. herzogii

2. Upper side of leaves glabrous. Twigs appressedly to patently short-hairy S. rayae
1 . Leaves not verticillate, etc.

8: 218 Add couplet 32a as follows:

32. Leaves longer than 5 cm (mean length).
32a. Leaves 14-36 by 5-11 cm. Twigs tomentose S. riangensis
32a. Leaves 3- 13(- 18) by 1.5-4.5(-6)cm. If leaves longer than 13 cm or broader than 4.5 cm then
twigs not tomentose.
33. Flowers etc.
1989] Addenda, corrigenda et emendanda 719

8: 227 Add couplets 5a, 5b, 15a, and 16b as follows:

4. Twigs hairy.
5. Leaves distichous.
5a. Leaves 14-36 cm long S. riangensis

5a. Leaves at most 12 cm long 33. S. laeteviridis

5. Leaves spirally arranged or pseudo-verticillate.
5b. Leaves pseudo-verticillate or at least 3-5 close together at the end of the flushes . S. rayae
5b. Leaves spirally arranged.
6. Leaves etc.

15. Leaves longer than 5 cm.

15a. Inflorescence a fascicle S. iliaspaiensis

15a. Inflorescence a (short) spike, raceme, or panicle.

16a. Inflorescence a (compound) spike S. cochinchinensis ssp. laurina

16a. Inflorescence a short raceme or a panicle 42 (3). S. ophirensis ssp. cumingiana

14. Bracts eic.

8: 230 Add couplet 25a as follows:

25. Inflorescence a raceme but pedicels sometimes very short. Bracts caducous.
25a. Leaves 5-22 cm long
1 36. S. maliliensis

25a. Leaves 3-6 cm long S. ambangensis

25. Inflorescence etc.

8: 235 Add the couplets 10a, 22a, and 22b as follows:

10. Inflorescence not a spike.
10a. Inflorescence a fascicle S. iliaspaiensis

10a. Inflorescence a raceme or panicle 42. S. ophirensis

10. Inflorescence etc.

22. Leaves distichous.

22a. Leaves 14-36 cm long S. riangensis

22a. Leaves at most 12 cm long.

23. Underside etc.
22. Leaves spirally arranged or pseudo-verticillate.
22b. Leaves pseudo-verticillate or at least 3-5 close together at the end of the flushes . S. rayae

22b. Leaves spirally arranged.

29. Leaves eic.

8: 237 Add couplet 17a in between couplet 17:

17a. Leaves 15-22 cm long 36. maliliensis
17a. Leaves 3-6 cm long S. ambangensis
or next to S. cochinchinensis as couplet 20a:
20. Stone different.
20a. Leaves more than 6 cm long 16. S. cochinchinensis

20a. Leaves shorter than 6 cm S. ambangensis

f&mWy Symplocaceae Symplocos costalifrucia Nom. Blumea 31 (1986)

277, f. 1.

Add the following species: Dislr. Malesia: Borneo (Sarawak, Brunei,

Sabah).
S>mplocos rayae Nrxn. Blumea .30 (1984) 73.
Symplocos iliaspaiensis Noot. Blumea 30(1984) 279.
D.str, Malesia: Borneo (Kalimantan Tcngah.
Di si r Mfl/rs/a: Borneo (Sarawak and possibly E.
.

Bukit Raya, only known from the type collection). L-,.i;.,r....,i..r,»

Null in till lull /.

s\mpl<Kos riangensis N^kh. Blumea 30 (1984) 74. Symphuos a

Disir. Malesia: Borneo (Kalimantan Tcngah, 263,1. 1.

I', it 11 Raya). Oislr. Malesia: North Sulawesi (Mcnado, Poso).

720 Flora Malesiana [ser. I, vol. 10^

Triuridaceae

10: 109 According to Ms. T. Rubsamen (Bochum, ferentiated, bitegmic, only in ripe seed the

West Germany; in litt.) the following inner integument wholly or largely sup-
amendments should be made: pressed.

10: 110 Family distribution map: extend the central Violaceae

area to the west to include part of East
Africa; see Kew Bull. 36 (1982) 733. 7: 198 Add under Viola the following species:
Sciaphila Blume. Viola rheophilaOkamoto, Bull. Osaka
Add/change description: Anthers some- Mus. Nat. Hist. no. 37 (1984) 4-15, 3 fig.
times 4-celled (America). Endosperm pres- Distr. Malesia: Borneo (Kalimantan
ent; embryo small, orthotropous, undif- Timur)
 INDEX TO SCIENTIFIC PLANT NAMES
compiled by

E.E. VAN NiEUWKOOP

Families and higher taxa have been entered under their name.
Names of families which have been revised in volumes 4-10 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
indication of their rank (subfamilies, tribes, etc.).
Infrageneric epithets have been entered immediately under the generic name to which they belong, pre-
ceeded by the indication of their rank (subgenera, sections, series, etc.).
Infraspecific epithets have been entered under the specific name to which they belong preceded by the
indication of their rank (subspecies, variety, forma, etc.).
Epithets of new names and new combinations have been printed in bold type, synonyms in italics.
Page numbers bold type denote main treatment; an asterisk behind a page number denotes the
in
presence of a figure of the concerned texon; '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.

Abies dammara Desf. 433 (Agathis ceiebica) (Agathis)

Aceraceae 4: 3-4, 592; 6: ssp. flavescens (Ridl.) robusia (Moore) Bailey 442
915; 7: 820; 8: 549 Veldk. & Whitm. ex map
Acioa Aubl. 639 Veldk. & de Laub. 438 ssp. nesophila Whitm.
heieropeiala (ScorL ex dammara (Lamb.) Richard 436
King) Kosierm. 677 433, 434 var. nesophila auct. 442
malayana Kosterm. 677 ssp. dammara aucl. 435, var. robusia Whitm. 442
pcrcoriacea Kosterm. 678 437 spathulata dc Laub. 433,
Acioa aucl. 675 ssp. dammara VJhilm. 433 435, 436 map
Acmopyle 343, 352, 354 ssp. flavescens (Ridl.) Agonandra 31-34
Actinidiaceae s.str.4: 37-39 Whitm. 438 raccmosa 33
Adenia 166 dammara auct. 435,437 Ailanthus intcgrifolia Larnk 718
Afrocarpus 354 endcrtii Mcijer Drees 439, Aizoaceae 4: 267-275; 6:

Afrolicania Mildbr. 645 440* map 915

Agathis Salisb. 339,341-343, flavescens Ridl. 340, 341, Alberlisia Becc. 160-162, 164,
347, 354,420,421, 429, 433, 438 167-169, 171, 179
430', 431 map hamii Meijer Drccs 435 crassaForman 179, 181
sect. Agathis 431 kinabalucnsis de Laub. 433, mccistophylla Forman 181
sect Macrobracteaiae Mcijer 438 map, 439 mcgacarpa Diels ex Forman
Drees 431 labiilardieri Warb. 436 map, 159, 179, 181
sect. Microbracleatae Mcijer 440, 441* papuana Becc. 166, 179,
Drccs431 latifotia Mcijer Drees 433 180*, 181
sect. Prismatobractcatae Icniicula de Laub. 433, Alc.imandra Dandy 565, 566.
Meijer Drees 431, 440 436*, 437 map 567, 569
sect. Rostrata dc Laub. 431, loranthifolia Salisb. 433 cathcariii (Hk.f. & Th.)
439 loranthifolia auct. 435 Diindy 569
alba Foxw. 433 macrostachys Warb. 433 Alismataceae 5: 317-334;
fl/6flauct. 435,437 motleyi (Pari.) Diimmcr 6: 915, 9: 553
beccarii Warb. 433 393 Alismauilcs 109
beckingii Mcijer 435 orbicuia de Laub. 433, 437* Allantospcrmum Forman 621,
bornccnsis Warb. 340, 34 1 map 622
430*. 432*. 433, 434* palmerstonii (F.v.M.) Bailey Alnus 338
map 442 Alpam Rhccde 65
cclcbica (Koord.) Warb. 433. Warb. 433,
philippincn.sis Alscuosmia 335
434. 435 map 437. 438 map AlMCUosmiaccac 335-336
ssp. ceiebica Vcldk. & regia Warb. 437 Aisoinitra macrcxarj^a (Bi.)
Whitm. 435 rhomboidales Warb. 433 Rocm. 29
(721)
722 Flora Malesiana [ser. I, vol. 104

(Andruris) (Araucaria)
Aliingiacxccl.sa341
Amaranthaceae 69-98,
4: viiiensis (A.C.Smith) Giesen sect. Eutacta Endl. 422, 423,

593; 5: 554; 6: 915; 118 425

7: 820; 8: 549 wariana Schltr 1 17 sect. Euiassa (Salisb.) Benth.
Ambora ficus Tul. 326 Angelesia Korth. 645 & Hook. 425
Amborclla 261 papuana Baker f. 65 sect. Intermedia White 423
Amborcllaccae261 splendens Korth. 646 subg. Colymbea Antoine 423
Amorpha pedalis Blanco 488 Anisadcnia 609 subg. Eutacta (Link) Antoine
Anacardiaceae 8: 395-548; Anisophyllaea gaudichaudiana 425
9: 553; 10: 679-681 Baill. 253 beccarii Warb. 427
Anaoirdium occidcntale 352 Annonaceae 63, 64, 261, 605, bidwiiiii 420, 423
Anacolosa Bl. 1,3-6, 23 716 cunninghamii Ait. ex D. Don
arborea K. & V. 25 Anthobembix Perkins 306, 308 340, 347, 425
cauliflora Sleum. 25 brassii A.C. Smith 315 var. papuana Laut. 426*,
celebica Valet. 25 dentatus \ a\Q.iox\ 310 427 map, 428*
frutcscens (Bl.) Bl. 2, 24*, hospilans (Becc.) Perkins 311 cunninghamii auct. non Ait.

25, 26* map ledermannii Perkins 319 427

heptandra Maing. ex Mast. 25 moszkowskii Perkins 3 1 hunsteinii K.Sch. 339, 347,

luzoniensis Mcrr. 25 myrtifolia A.C. Smith 313 420, 423, 424 map
maingayi Mast. 22 oligantha Perkins 315 klinkii Laut. 423, 424
papuana Schelle.nb. 25, 26 parvifolia Perkins 315 schumanniana Warb. 424
Zollinger I Baill. 25 Antidesma ghacsembilla Gaertn. Araucariaceae 338, 339, 343,
Anamirta Colcbr. 157, 158, 52 347, 354, 419-442
160-162, 166-168, 170, pcntandrum (Blco) Merr. 52 Araucariacites 338
171, 211 Ami taxis Miers 172 Arcangelisia Becc. 158, 160,
cocculus (L.) Wight & Am. calocarpa Kurz 174 161, 166-171, 209
159, 165. 166, 212*, cauliflora (Miers) Dicls 174 flava(L.) Mcrr. 160, 210,
213, 214* fasciculata Miers 174 211
flavescens (Lamk) Miq. 210 longifolius (Decne ex Miq.) inclyta Becc. 210
jucunda Micrs 213 Miers 174 lemniscata (Miers) Becc. 210
lemniscata Miers 210 Apama Lamk 62, 63, 65 loureiri (Pierre) Dicls 210
210
loureiriPicTTC affinisWeisse 79 tympanopoda (Laut. &
luctuosa Miers 210 brevipes Weisse 79 K. Sch.) Diels 210, 211
Anaxagorea A. St. Hil. 605 corymbosa (Griff.) Willd. ex Archichlamydeae 127
Ancistrocladaceae 4: 8-10; Soler. 78 Ardisia glabra (Thunb.) DC. 134
5: 553 macranlha Weisse 81 Aristolochia L. 53-55, 57-59,
Ancistrocladus pentagynus tomeniosa Engl, ex Soler. 61-64,83, 166
Warb. 612 55,79 acuminata Lamk 94
Androglossum reliculaium var. lanuginosa (Hk.f.) barbata 55
Champ, ex Benth. 685 & G. 79
K. brasiliensis Mart. & Zucc.
AndrurisSc\\\\s\09, 110 iAp/zy/Ze/a Champion 110 55,64
andajensis (Becc.) Schltr 118 erubescensCUdsn^). 114 clematitis 59
anisophylla Cicscn 112, 118 Aponogetonaceae 4: 11—12; coadunata Backer 57-59, 84,
australasica (Hemsl.) Giesen 5:553; 7: 213 86, 95, 96*
118 Aptandra 3, 4 var. bosschai Backer 95
bur uensis J. J. Smith 118 Aptandropsis 4 var. coadunata 95

celebica Schltr 118 Aquifoliaceae 145 crassinervia Schmidt 58, 82*,

clemensae (Hemsl.) Giesen Aquilegia 165 104*
85, 87,
112, 118 Ardbidopsis 543 King 84, 86, 89
curtisii
clemensae (Hemsl.) Giesen Arabis 54 1 decandra Ding Hou 57-59,
var. borneensis Giesen 118 Araliaceae 9: 1-105, 553; 64, 83, 85, 87, 100,
crinila (Becc.) Schltr 1 17 10: 716 101*, 102*
elegans G'lescn 118 Araucaria Juss. 337, 339, 421, deltantha F.v.M. 97
gracillima Giesen 1 17 423 map dictyophlebia Merr. & Perry
javanica Giesen 118 sect. Araucaria 422, 423 97
loheri Giesen 117 sect. Bunya Wilde & Eames dielsiana Schmidt 57, 61, 64,
nana Giesen 117
(Bl.) 423 84, 85, 88, 105
palawensis Tuyama 118 sect. Columbea All elegans Mart. & Zucc. 55,
/ene//a Schltr 118 sect. Colymbea Endl. 422 58,64
1989] Index to scientific plant names 723

(Aristolochia) (Aristolochia) Ascarina J.R. & G. Forst. 123-

engleriana Schmidt 85, 88, philippinensis Warb. 60*, 129, 137, 140 map
"106 66, 84, 87, 102 sect- Ascarina 137, 139, 140
foveolata Merr. 55, 58, 85- piihecurus Ridl. 97 map
87, 91 map ramosii Merr. 92 sect. Madascarina Leroy &
gaudichaudii Duchartre 56*, ringens Vahl 55, 64 Jeremie 123, 137, 140
57, 84, 88, 107 roxburghiana Klotzsch 94 map
gigantea Mart. & Zucc. 64 6 angustifolia Duchartre diffusa A.C. Smith 139,
glaucifolia Ridl. 55, 85, 86, 94 141
93 ssp.kankauensis (Sasaki) lanceolata (non Hk.f.) auct.
gracilifoUa Schmidt 97 Kitamura 92 141
grandiflora Sw. 57, 59, 62, roxburghiana (non Klotzsch) lucida 124, 139
64 Warb. 107 maheshwarii Swamy 139,
Duchartre 59, 91
griffiihii rumphii KosteleL?Jcy 85, 86, 140, 141
hastata Jack 88 97, 101* philippinensis C.B. Rob.
hastata (non Jack) Klotzsch samarensis Merr. 85, 87, 138*, 139, 141
92 102 reticulata Merr. 139
humilis Merr. 55, 85, 87, schlechteri Laut. 58, 59, 85, rubricaulis 127
103 88, 97, 105, 106, 107 subsessilis Verdcourt 139,
imbricaia Mast. 103 sericeaBlco55, 57, 58, 61, 141
indica L. 99 66, 85, 87, 103 Ascarinopsis 123, 128
indica (non L.) BIco 94 singalangensis Korth. ex Ascarinopsis Humb. & Capuron
indica (non L.) L. 97 Ding. Hou 61, 84, 86, 123, 128, 137, 140 map
jackii Steud. 55, 58, 59, 84, 89, 90* As hto n s Aromadendron Mcijcr
86, 87, 88, 89* tagala Chamisso 54, 55, 56*, 579
japonica Miq. 94 57, 63, 85-88, 94 Asiphonia Griff. 65
kaempferi WiUd. 92 tagala (non Chamisso) Halus. piperiformis Griff. 78
kankauensis Sasaki 92 92 sp. Griff. 78
kaoi Liu & Lai 91 var. hankaoensis 92 Aspidocarya 162
klossh Ridl. 85, 86, 93 var. kankauensis (Sasaki) dissitiflora Laut. & K. Sch.
kwangsiensis Liang 91 Yamazaki 92 197
lautcrbachiana Schmidt 108 timorensis Dccne 94 hirsuta^cjcc. 193
ledermannii Schmidt 106, Lranstillifera Ding Hou 60*, kelidophylla Laut. &
107 map 84, 87, 99 K. Sch. 253
Icuconeura 62 iripariiia Backer 88 uvifcra Hk. f. & Th. 189*
leytensis Merr. 84, 85, 87, ungulifolia Mast. 88 Aihcrospcrmataceae 261
104 zollingeriana Miq. 57, 85, Athcrospermoxylon 256
linnemannii Warb. 55, 85, 86, 92 Athrotaxis 339
88, 107 Aristolochiaceae 53-108, Au-iaecarpum 158
longifolia Roxb. 94 126, 166 Atroxima 456
macgregorii Merr. 55, 56*, Aristolochiales 567 Atuna Rafm. 636-642, 665
57, 85, 87, 103 Aromadendron Bl. 567, 568, cordata Cockburn ex Prance
maurorum 58 576 636, 666 map, 667
megalophylla K.Sch. 94 ashionii Dandy ex Cockbum data (King) Kostcrm. 669
memhranacea Merr. 103 579 elliplica Ko.stcrm. 666
mindanaensis Warb. 94 borneensis Dandy ex Cock- excelsa (Jack) Kostcrm. 636,
minuiiflora Ridl. ex Gamble bum 579 640,641,671
55. 85-87, 93 elegans Bi. 577 indica (Bald.) Kostcrm. 666
var. dolahrata Gamble 93 var. glauc.a (Korth.) latifolia (Hcnd.) Kostcrm.
moluccana Duchartre 94 Dandy 577 666
momandul K. Sch. 58. 59, glaucum Korth. 577 lalifrons (Kostcmi.) Prance &
82*, 85, 86, 88, 97 nutans Dandy 577 While 666 map
naviculilimba Ding Hou 85, oreadum (Diels) Kanch. & nannodes (Kostcrm.)
87. 100* Hatus. 583 Koslerm. 666. 667 map
novogumcCnsis Schmidt 108 Arorrtadendrum 568 pcnangiana (Kostcnn.)
ornilhrjcephala 55 Aromoracia rusticana G.M. & Ko.stcrm. 666. 667 map
papillifolia Dmg Hou 58, 84. Schcrb. 543 raccmo.sa Rafin. 666. 669
87. 98*. 99 Asarum canadcn.sc 57 ssp. cxccl.sa (Jack) I'rancc
pcliata 59 virginitum 62 668*. 669. 670 map
724 Flora Malesiana [ser. I,vol. 104

Betulaceae 5: 207-208; 6: Bruguiera gymnorrhiza (L.)

(Atuna racemosa)
ssp. racemosa 669, 670 917 Savigny 717
map Bignoniaceae 8: 114-186; Bubbia 123
scabra (Hassk.) Koslerm. 9: 554; 10: 716 Burmanniaceae 4: 13-26,
641,669 Bixaceae s.str. 4: 239-241 592; 5: 553; 7: 820;
Bladhia glabra Thunb. 34 9: 554
travancorica (Bedd.) Kosterm. 1

666 Blumia Nees 569 Burseraceae 5: 209-296,

villamilii (Merr.) Kosterm. candollii (Bl.) Necs 582 567; 6: 917; 7: 820;
671 Blyxa alternifolia (Miq.) Hanog 9: 555; 10: 715

Aiunus Rumph. 665 717 Butomaceae 5: 118-120,

alba Rumph. 669 aubenii Rich. 717 566
Austrocedrus Florin & Boutelje var. echinosperma (Clarke) Byblidaceae 7: 135-137
443 Cook &Luond 717
Austrotaxus 339, 348 echinosperma (Clarke) Hk.f.
717 Callitrichaceae 4: 251-252
japonica (Miq.) Maxim, ex Callitris sp.453
Badiera (non DC.) Hassk. 459, Aschers. & Gurke 717 Callitroideae 444
467 var. alternifolia (Miq.) Calocedrus 444
pulchra (Hassk.) Hassk. 468 Cook&Luond^n Camelina 541
venenosa (Poir.) Hassk. 468 var. japonica 717 Campanulaceae 6: 107—141,
Bafodeya Prance 642 leiosperma Koidz. 717 928; 8: 549; 9: 556
Balanophoraceae 7: 783—805; Boraginaceae 29 Canariopsis anguslifolia Bl. ex
8: 549; 9: 554 Borneo Aromadendron Meijer Miq. 715
Balantium Desv. ex Buch.-Ham. 579 Canarium angustifolium Miq.
654 Bracea King 29 715
Balsaminaceae 539 Brachynema 3, Candjera 5
Bania Becc. 183 Braganlia Lour. 65 Cannabinaceae 4: 222-223
thyrsiflora Becc. 183 affinis Planch, ex Rolfe 79 Cansiera 50, 52
Banisterodes O.K. 493 blumii Lindl. 79 zyziphifolia 50
affine (Miq.) O.K. 503 brevipes Merr. 79 Cansjera Juss. 31-35, 48, 49
ellipilcum (Korth. ex Miq.) corymbosa (non Griff.) map
O.K. 530 F.-Vill. 103 grossularioides Blco 52
excelsum (Bl.) O.K. 501 corymbosa Griff. 78 lanceolala Bth. 50
glaucum (Wall, ex Hassk.) macranlha Boerl. 81 leplostachya Bth. 31, 33, 34,
O.K. 527 melastomaefolia Duchartre 48*, 49 map, 51*
griffiihii (Hk.f. ex Benn.) 78 leplostachya Koord. 35
O.K. 513 tomentosa Bl. 79 malabarica Lamk 50
insigne (Benn.) O.K. 537 var. lanuginosa Hk. f. 79 var. 6 50
/on^//b//a(Bl.)O.K. 514 Brassica L. 541, 543, 545 manillana Bl. 36
maingayi (Benn.) O.K. 524 besseriana Andrz. ex Trautv. monostachya (Willd.)
obscurum (Benn.) O.K. 537 546 M. Roemer 50
rufum (Benn.) O.K. 505, campestris L. 543 parvifolia Kurz 48*, 49 map
507 chinensis L. 543 penlandra Blco 52
stipitatum (Benn.) O.K. 535 integrifolia (West) Rupr.546 polystachya (Willd.)
vitellinum (Bl) O.K. 515 juncea (L.) Czem. 544*, 545 M. Roemer 50
Bannisierioides 493 napus L. 543 rheedii J.F.Gmelin 31-33,
Barbarea vulgaris R. Br. 545 oleracea L. 543 48*, 49 map, 50*
Bamharlia Gleason 493 rapa L. 543 rheedii Blco 52
Basellaceae 5: 300-304 rugosa Prain 543 scandens Roxb. 9, 50
Baiania Ualus. 172 Bredemeyera 456, 457, 465 timorensis Decne 5
insignis Hatus. 174 sect. Melchiora Steen. 465 zizyphifolia Griff. 50, 51
Baterium Miers 183, 184 floribunda 457 Cappar(id)aceae 6: 61-105;
validum Miers 1 84 papuana Steen. 465 7: 822
Bat(id)aceae 5: 414-415; 6: Brewsiera M.J. Roemer 622, Caprifoliaceae 4: 175-194;
917 625 6:928; 9: 556; 10: 335
Benneiiiodendron leprosipes crenaia M.J. Roemer 625 Capsella Medicus 541, 543,
(Clos) Merr. 715 Brongniartia Bl. 287 545, 549
Bcrberidaceae 163 coriacea Bl. 298 bursa-pastoris (L.) Medicus
Berbcridales 163 Brownetera L.C. Rich. 356 550
1989] Index to scientific plant names 725

Cardamine L. 541, 543, 545, (Chamaebuxus) (Chloranthus)

550 arillaia (D. Don) Hassk. 469 hainanensis Pei 1 36
africana L. 541, 551 pulchra (Hassk.) Hassk. 468 henryi Hemsley 130, 133
ssp. borbonica (Pers.) venenosa (Poir.) Hassk. 468 inconspicuus Svj2iT\z 133
Schulz sub var. elliptica Miq. 468 inconspicuus (non Swartz)
var. papuana Laul. 55 subvar. obovata Miq. 468 Blco 131
alligena O.E. Schulz 551, var. apiera Miq. 468 indicus Wight 133
553 var. gracilis Miq. 468 japonicus 127, 130
borbonica Pers. 551 var.minor Miq. 468 monander R. Br. ex Sims
decurrens (Bl.) Z. & M. 554 var. robusla Miq. 468 134
flexuosa With. 551, 554 Champaca Adans. 598 multisiachys Pei 128
flexuosa auct. 554 lurbinata Nor. 583 obtusifalius Miq. 133
hirsuta L. 551, 554 Champereia Griff. 31-34, 35 officinalis Bl. 124, 127,
javanica (Bl.) Miq. 552 cumingiana (Baill.) Merr. 36 128, 131, 132
keysseri O.E. Schulz 551, gaudichaudiana (Baill.) Tiegh. oldhamii Solms-Laub. 133
553 36 oldhamii (non Solms-Laub.)
papuana (Laut.) O.E. Schulz griffithii Planch, ex Kurz 36 Merr. & Quis. 133
551, 552* lanceolata Merr. 36 philippinensis Merr. 133
re ge liana Miq. 554 manillana (Bl.) Merr. 31, 34, salicifolius Presl 131
sublyrata Miq. 559 35, 36*, 37*, 38 map, 40* serratus 127
Cardiopteridaceae 7: 93—96; oblongifolia Merr. 36 spicalus (Thunb.) Makino
10: 166, 253, 716 perrolteiiana Baill. 52 127, 128, 130, 132
Cardioptcris Royle 166, 716 platyphylla Merr. 36 veriicillatusMen. 133
moluccana Bl. 19, 253 Champereya 35 Chrysobalanaceae 635-678
Carex bilateralis Hayata 716 gnetocarpa Kurz 36 tribe Chrysobalaneae 635
Caricaceae 61 griffiihiana Planch, ex Kurz tribe Couepieae 636
Carpolobia 456, 457 36 tribe Hinelleae 636
Carronia F.v.M. 158, 160, 162, Chandlera 158 tribe Parinarieae635
167, 168, 170, 172, 183 Chasmanihera 164 Chrysobalanus L. 635-637,
thyrsiflora (Becc.) Dicls Chaunochiton 3, 4 639, 640, 642, 643
182*, 183 Cheirolepidaceae 338 ciliatus Korth. ex Miq. 674
Carson's Magnolia Meijer 571, Chenopodiaceae 4: 99—106, icaco L. 643, 644* map
573 594; 6: 932; 8: 549; 9: splendens Korth. ex Miq.
Cascaria flavovirens Bl. 716 557 646
pallida Craib 716 Chlaenandra Miq. 158, 161, Cissampelos L. 157, 158. 160-
Casuarinaccae 453 162, 167, 168, 170, 171, 162. 165. 167. 169-171,
Cathedra 3, 187 234
Cayloniales 338 ovaia Miq. 159. 186*, 187, convolvulacea Willd. 236
Caytonipollcniies 338 189* var. hirsuta (DC.) Hassk.
Celastraceae 6: 227-291, Chloranthaceae 123-144 236
389-421,930; 10: Chloranihales 127 cumingiana Turcz. 236
539, 716 Chloranlhus Swariz 123-128, discolor DC. 236
Cclaslrales 145, 629 129. 139. 143, 144 var. cardiophylla A. Gray
Celasiranac 151 brachysiachys Bl. 236
Ccliis paniculaia (Endl.) Planch. var. melanocarpus Ridl. hernandifolia 245
29 136 hernandiifotia Willd. 245
Ccntrolepidaceae 5: 421-427 brachysiachys scnsu Benlh. insularis Makino 240
Ccphaiotaxaccac 343, 347 134 ovala Poir. 217
Ccphaloiaxus 354 chincnsis 124 pareiraL. 165
celebica Warb. 349 denliculalus Cordemoy 34 1 p236
mannii (non Hk.f.) F*rilzcl ex elalior R.Br, ex Sims 131 var. hirsuia (Buch. ex
Dicls 349 creclus (Buch.-Ham.) Verde. DC.) Forman 235*.
sumatrana Miq. 349 124. 127. 128. 130*. 236
Ceramium Bl. 65 131. 132*. 133*. 136 var. orbiculaia (DC.) Miq.
lomenlosum fil. 79 ereclusSwccl 130 236
Ceratophyilaceae 4: 41-42 fortunci 127 236
var. peltata Schcff.
CV.ri<jps dctandra 7 1 glabcr scnsu Backer & var. typica Diels236
ChamaehuxiLS (DC.) Spach 459, Bakh.f. 136 parcira (non L.) Ridl. 229
467 glaber (Thunb.) Makino \?A psilophylla Prcsl 25
726 Flora Malesiana [ser. I, vol. 104

Citronella suaveolens (Bl.) (Cocculus) Crassulaceae 4: 197-202;

Howard 717 rimosus Bl. 207 9: 558

Cladopus nymani H. M611. 717 sarmenlosus Dicls 233 Cri.spiloba Stccn. 335
Classopollis 338 var. stenophyllus Merr. Croialaria duboisii Lev. 469
Clethraceae 7: 139-150 233 Cruciferae 541-560
Clypea BI. 243 tr iandr us CoXchx. 185 tribe Arabideae 543
acuminalissima Bl. 252 /r//?o/'U5DC. 219, 233 tribe Brassiceae543
capitata Bl. 252 /n7(3feu5 (Thunb.) DC. 159, tribe Heliophilcae543
corymbosa Bl. 249 163, 165,231,233,234 tribe Lepidicae 543

discolor BI. 245 velutinus Wall. 223 tribe Stcnopctaleae 543

glaucescens Decne 245 Cochlearia officinalis L. 543 Cryphaea Buch.-Ham. 129

tomentosa Bl. 229 Cochlospermaceae 4: 61—63 erecta Buch.-Ham. 131, 132
venosa Bl. 248 Coffea anguslifolia Roxb. 717 Crypteroniaceae 8: 187-204
Coccomelia Ridl. 645 Columbea All, Al^ Cryplocarya reticulata BI. 715
CocculusDC. 157, 158, 161, Colymbea Spreng. 422 Ctenolophon Oliver 29, 629
162, 165, 169, 231 Combretaceae 533-589; 4: englerianus Mildbr. 630, 631
angustifolius Hassk. 234 5: 564; 932; 7: 823
6: grandifolius Oliver 631
bantamensis Bl. 199 Comesperma 456, 461, 465 parvifolius Oliver 630, 631,
blumei Boerl. 223 sect. Prosthemosperma 632*, 633* map
brachysiachyus DC. 217 F.v.M. 465 philippinensis Hall.f. 630,
celebicus Boer). 175 Coniferales 337-453 631
cinereus Zoll. & Mor. 229 Connaraceae 5: 495-541; Ctenolophonaceae 607, 609,
coriaceus Bl. 199 6: 933; 7: 823; 8: 549; 621,629-634
corymbosus Bl. 229 557
9: Cucurbitaceae 29, 166, 253
crispus [non (L.) DC] Hassk. Convolvulaceae 4: 388- Cupressaceae 338, 339, 343,
199 512,599; 5: 558; 6: 347,442-447
cuspidatus V^ all. 2\9 936; 7: 823; 9: 558; Curupira 3, 4
cynanchoides PresI 233 10: 29,716 Cyclandrophora Hassk. 642, 665
elegans (Ridl.) Ridl. 233 Cordai tales 126 asperula (Miq.) Prance ex
ferrandianus Gaudich. 233 Cornaceae 8: 85-97 Kosterm. 671
flavescens (Lamk) DC. 210 Coronopus 541 elata (King) Kosterm. 669
flavicans Wall. 253 Coryneliales 353 excelsa (Jack) Kosterm. 671
forsteri DC. 245 Corynocarpaceae 4: 262— glaberrima Hassk. 669
glaucescens BI. 230 264; 5: 557; 6: 941 lalifolia (Hend.) Prance 666
^/aucwi (Lamk) DC. 229 Corytholobium Mart, ex Benth. laurina (Gray) Kosterm. 669
incanus Colebr. 229 483 nannodes (Kosterm.)
japonicus (Thunb.) DC. 245 Coscinium Colebr. 160-162, Kosterm. & Prance 667
var. limoriensis DC. 245 166-168, 170, 171, 215 penangiana Kosterm. &
kunslleriKing 231 blumeanum Micrs ex Hk.f. & Prance 667
lanuginosus Bl. 229 Th. 212*, 215, 216, scabra (Hassk.) Kosterm. 669
laurifolius DC. 160, 166, 217 Prance ex
villamilii (Merr.)
169, 171,231,232*, blumeanum (non Miers ex Kosterm. 671
233, 234 Hk.f. &Th.) Miq. 216 Cyclea Am. ex Wight 160-162,
var. angustifolius (Hassk.) var. epel latum Boerl. 216 165, 167, 169, 170, 172,
Boerl. 233 fenestratum (Gaertn.) Colebr. 237
var. triplinervis Boerl. 233 160, 165, 212*, 215, acuminatissima Merr. 239
leptostachyus DC. 217 217 apoensis Yamamoto 24
limacia DC. 223 var. macrophyllum atjehensis Forman 237, 238*,
longifolius Decne ex Miq. Yamamoto215 239
174 var. ovalifolium barbata Miers 159, 136, 139,
lucidus Teijsm. & Binn. 174 Yamamoto215 242, 243, 253
macrocarpus W. & A. 230 maingayi Pierre 216 barbata Craib 239
mollis Hk.f. & Th. 233 mangayi 216 caudata Merr. 239
orbiculalus (L.) DC. 159, miosepalum Diels 216 cauliilora Merr. 237, 241
160, 162, 170, 172, peltatum Merr. 216 Craib 242
ciliata
231, 232* wallichianum Miers 215 elegans King 187, 237, 239
ovalifolius (Vahl ex Pers.) Couepia 640 insularis (Makino) Hatus.
DC. 233 Coula 3, 4 237, 240
populifolius DC. 213 Crambe541,543 var. luxurians Hatus. 240
1989] Index to scientific plant names 727

(Cyclea) (Dacrycarpus imbncatus) (Dacrydium)

kinabaluensis Forman 159, var. robustus de Laub. xanihandrum Piiger 363,
237, 239 377, 379 map 369*, 370* map
var. hispida Forman 239 kawaii (Hayata) Gaussen Dactyladenia 639
korihalsii Diels ex Norman 379 Dahuronia Scop. 645
241,242 kinabaluensis (Wasscher) de Dammara Link 429
Miers 237, 239,
laxiilora Laub. 376, 381 map alba Rumph. ex Hassk. 435
241, 242, 243 leptophylla (Wasscher) var. alba Hassk. 435

merrillii Diels 237, 240 Gaussen 371 var. celebica Hassk. 435

peliata [non (Lamk) Hk. f. & steupii (Wasscher) de Laub. celebica Koord. 435
Th.] Becc. 241 340, 376, 380 loranthifolia Link 433
var. arnouiiMiers 241 steupii (non de Laub.) de motleyi Pari. 393
peliata [non (Lamk) Hk. f. & Laub. 379 orienialis Lamb. 433
Th.] Miq. 163, 242 Dacrydium Soland. ex Forsl. f. var. alba Knight ex Henkel
percgrina Miers 237, 239, 337, 338, 342, 343, 347, & Hochst. 435
241. 242 352-355, 360, 362 map var. orienialis Carr. 433
robusia Becc. 237, 240 balansae 365 var. pallens Carr. 435
scyphigera Suesseng. beccarii Pari. 362, 366, 367 palmerstonii F.v.M. 442
Heine 239 map robusia Moore ex F.v.M.
forma anguslifolia var. rudens de Laub. 368 442
Suesseng. & Heine 239 var. subelatum Comer 363 rumpfii auct. 437
tomcntosa 243 comosum Comer 363, 370, rumphii Presl 435
tonkinensis (non Gagnep.) 371 map Daphnandra Perkins 256, 261,
Yamamoto 239 cornwalliana de Laub. 340, 263, 265
wallichii Diels 242, 243 362, 364 map, 366* aromadca 261
Cyclodiscus tomentosus cupressinum 365 novoguineensis Perkins 266
Klolzsch 79 datum (Roxb.) Wall, ex Hk. perkinsiae Gilg & Diels 266
Cyperaceae 7: 435—753, 341, 362, 363. 364 map Daphne monosiachya Willd. 50
823; 9: 107-187,560; ericoides de Laub. 340, polysiachya Willd. 50
10: 716 341, 363, 371 map Datiscaceae 4: 382-387; 7:

C>perus diaphanus Schrader ex falciforme (Pari.) Pilgcr 372 823

R. & S. falciforme [non (Pari.) Pilgcr] Davisicarpum 158
var. latespicalus (Boeck.) Foxw. ex Mcrr. 373 Decussocarpus de Laub. 389
Kern 716 falciforme [non (Pari.) Pilgcr] sect. Dammaroides (Bennett)
CyrillopsisKuhlm. 621,622 Laut. 373 de Laub. 390
gibbsiae Siapf 363. 369 .sect. Decussocarpus de Laub.

gracilis de Laub. 363, 367 394

Dacrycarpus (Endl.) dc Laub. map maximus de Laub. 394
337, 338,343, 347, 351, junghuhnii Miq. 363 motleyi (Pari.) de Laub. 393
353-355. 374, 376 map leptophyllum (Wa.sschcr) viiiensis (Seem.) dc Laub.

cincius (Piiger) de Laub. 376, dcLaub. 363,371 395

383*, 384 map magnum de Laub. 363, 368 wallichianus (Presl) dc Laub.
compaclus (Was.scher) de map 393
Laub. 376, 382* map, medium dc Laub. 339-341, Dcgcncriaceac 562
383*. 384 363, 368 map Dichapetalaceae 5: 305-316.

cumingii (Pari.) de Laub. micropedunculatum 340 567; 6: 941; 7: 823

376, 381 map nidulum de Laub. 362. 365 Diclidanthcra456,457,493
dacrydiifoUa (Was-schcr) map Diemenia Korth. 648
Gausscn 383 var. araucarioides dc Laub. racerrwsa (Korlh.) Miq. 649
cxpansus dc Laub. 342. 376, 366 Dillcnia 123
381, 382 map novo-guincensc Gibbs 362, Dilleniaceae 4: 141-174;

imbncatas (Bl.) dc Laub. 364 map 5:557; 7: 824

340, 341. 376 pctlmatum de Laub. 340. Dimocarpus longan 159
var. turvulus (Miq.) dc .34 362, 364 map
1 . Diogoa 3, 4
Laub. 340, 341. 377. var. robusium dc Laub. Dioscorca 166
379 map, 380* 364 aculcata (non L.) Zoll. 199
var. imbncatus 377 map pterrii Hickcl 363 spiculala Bl. 199
var. palulas dc l^ub. 377 spathoidcs dc I-aub. 340, 363, slcnomcridora Pram ik Hurk.
map. 378*. 379 367 map. 368 253
728 Flora Malesiana [ser. I, vol. 104

Dioscoreaceae 4: 293-335; {Durandea) (Elaeagnus triflora)

5: 557; 10: 253 robinsonii (Merr.) Hall.f. 612 ssp. leiragonia Serv. 153

Dioscorcophyllum cumminsii rolundaia K. Sch. 612 ssp. leiragonia (non Serv.)

165 Merr. & Perry 155

Diploclisia Miers 158, 160, var. brcvilimbata 't Han
169, 170, 172, 229 Elaeagnaceae 151-156 154*, 155
glaucescens (Bl.) Diels 230 Elaeagnales 1 5 var. brevilimhalus 155
glaucescens (non (King) Elaeagnus Toum. ex L. 151, var. brevipes Serv. 153
Diels) sensu Forman 23 155* var. longipes Serv. 153

kunstleri (King) Diels 228*, seel. Elaeagnus 152 var. triflora 154*
230 sect. Sempervirentes Serv. Zollinger i Serv. 154
macrocarpa (W. & A.) Miers 152 Eiaeocarpus 123
230 anguslifolia (non L.) Blco Elatinaceae 4: 203-206
Dipsacaceae 4: 290-292; 154 Elmerrillia Dandy 562, 564-
5: 557 arborea Roxb. 567, 593, 595
Dipterocarpaceae 9: 237—552; var. dendroidea sect. P seudoaromadendron
561; 10: 665, 716 Schlechtend. 153. Dandy 595
Dipierocarpus comutus Dyer 665 conferta Roxb. 152, 153, celebica (Koord.) Dandy 596
Discogyne Schitr 622 154* mollis Dandy 596, 598
papuana Schitr 627 ssp. dendroidea ovalis (Miq.) Dandy 595,
Dolichandrone spalhacea (L.f.) (Schlechtend.) Serv. 596
K.Sch. 716 153 papuana (Schitr) Dandy 595,
Dombeya Lamk. 422 ssp. euconferia Serv. 153 596
Dor>'phora 261 ssp. javanica (Bl.) Serv. var. adpressa Dandy 596

aromatica 261 153 var. glaberrima Dandy

Drebbelia Zoll. 6 var. calcullensis Serv. 153 596, 598
subarborescens Zoll. 7 var. malaccensis Serv. 153 platyphylla (Merr.) Noot.
Droseraceae 4: 377—381; var. pallescens Serv. 156 595, 596
557; 6: 943; 9: 562
5: var. sepientrionalis Serv. pubescens (Merr.) Dandy 595,
Dryadodaphne S. Moore 255- 153 596
257, 262, 263 var. silheiensis Serv. 153 sericea C.T.White 598
celastroides S. Moore 266 cumingii Schlechtend. 154 tsiampacca (L.) Dandy 595,
crassa Schodde 264*, 265, ssp. perrollelii Serv. 154 596
266 ssp. philippinensis Serv. ssp. mollis (Dandy) Noot.
novoguineensis (Perkins) 154 567, 598, 599*
A.C. Smith 261, 265, dendroidea Sc\\\tz\\\sx\± 153 ssp. tsiampacca 598
266, 267 ferruginea A. Rich. 153 var. glaberrima (Dandy)
ssp. novoguineensis 266, ssp. sumairana Serv. 153 Noot. 595, 598
267 var. airovirens Serv. 153 var. tsiampacca 597*,
var. macra Schodde var. richardia Serv. 153 598
266 fruticosa (Lour.) Cheval. vriesearuj (Miq.) Dandy 596
var. novoguineensis 153 Elsoia Adans. 483
266, 267 gaudichaudiana Schlechtend. bracieala (Benn.) O.K. 484
ssp. occidentalis 153 corymbosa (Turcz.) O.K.
Schodde 265, 267 javanica Bl. 153 484
novoguineensis (Perkins) latifoliaL. 152 lavoyana (Benn.) O.K. 484
A.C. Smith p.p. 265 laiifolia (non L.) A. Rich. Embelia urophylla Wall, ex
Drymis-leaved Magnolia Meijer 153 A. DC. 649
571, 573 var. iriflora Schlechtend. Emmenanihus Hk.f. & Am.
Dugoriia Scop. 654 153 622, 626
Dulacia 4 /7erro»e/H Schlechtend. 154 chinensis Hk.f. & Am. 626
Durandea Planch. 608, 609 philippinensis Perrotlet 153 Entosiphon Bedd. 665
jenkinsii (F.v.M.) Siapf 612 rigida Bl. 153 Epacridaceae 6: 422-444,
magnifolia Stapf 615 rosiraia Serv. 154 943; 10: 335
pallida K.Sch.eW iriilora Roxb. 152, 153 Ephedra 338
pentagyna (Warb.) K. Sch. ssp. obsoleia Serv. 153 Ephippiandra 258
612 ssp. polymorpha Serv. Epicryanlhes 489
var. rolundaia (K. Sch.) 153 Epinetrum 164
Laul. 612 ssp. rigida Serv. 153 Epirhizanlhes 489
1989] Index to scientific plant names 729

Epirixanthes Bl. 455, 456, 459, Falcalifolium de Laub.343, 347, Ganua palembanica (Miq.) v.d.
486, 488 352, 354, 355, 371 Assem & Kosterm. 453
aphylla (Griff.) Merr. 490 angustum de Laub. 372, 373 sp. 453
cylindricaBl. 457,490, 491 map, 374 Geobalanus Small 645
elongata Bl. 489, 490* falciforme (Pari.) de Laub. Geraniaceae 6: 445-449;
kinabaluensis Wendt 490, 372, 373* map 9: 565; 10: 607, 639
491 gruezoi de Laub. 372, 373 Geraniales 609, 639
linearis Bl. 490 map Gesneriaceae 335
pallida Wendt 490, 492 papuanum de Laub. 372, 373 Gigantopteris 338
papuana J.J. Sm. 489, 490, map Gjellerupia Laut. 31-35, 45
491 Fawceltia F.v.M. 164, 188 papuana Laut. 45*, 46
tenella Hk. f. 490 me rril liana (Diels) Yamamoto Gladiolus dalenii Geel 717
Epirixanihus 489 193 natalensis (Ecklon) Reinw. ex
Epirizanthes 489 Ferolia O.K. 655 Hk. 717
Epirrhizanihe 489 asperula (Miq.) O.K. 671 Glossocalyx 258
Epirrhizanlhes 489 corymbosa (Bl.) O.K. 674 Glyptopetalum loheri Merr. 716
Eriandra Royen & Sleen. 455, cosiata (Konh.) O.K. 663 Gnetaceae 4: 336-347; 6:
457, 459, 492 glaberrima (Hassk.) O.K. 944
fragrans Royen & Sieen. 493 669 Goodeniaceae 5: 335-344;
Ericaceae 6: 469-914; 943; griffithiana (Benth.) O.K. 567; 6: 949; 7: 827; 9:
7: 827; 8: 549; 9: 562; 674 566
10: 335,716 jackiana (Benth.) O.K. 671 Gordonia decandra Roxb. 626
Ericybe Roxb. 29 /jmda(Hk.f.)Ridl. 646 Govantesia Llanos 35
Eryihropalla 17 nonda (F.v.M. ex Benth.) malulucban Llanos 36
Erythropalum Bl. 1-6, 17, 166 O.K. 658 Grossulariaceae 335
grandifoliwn Elmer 17 oblongifolia (Hk.f.)O.K. Groutia Guill. & Perr. 46
scandens Bl. 17, 18* 659 cellidifolia Guill. & Perr. 47
var. abbrevialum Hochr. 17 polyneura (Miq.) O.K. 664 Grymania Presl 67
iriandrum Quis. Merr. 29& salicifolia (Presl) O.K. 674 salicifolia Presl 671, 673
vagum (Griff.) Mast. 17 scabra (Hassk.) O.K. 669 Guatteria incerta Bl. 515
Erythroropalum 17 sumatrana (Jack) O.K. 661 Guttiferae 145
Erythroxylaceae 5:543-552; Fibraurea Lour. 157, 160-162, Gymnostoma sp. 453
8: 549; 10: 607, 609, 622 166-168, 170, 171, 207
Escalloniaceae 335 chloroleuca Miers 160,207
Euglypha 53, 61,63 chloroleuca (non Miers) Merr. Haematocarpus Miers 160, 161,
Euphorbiaceae 29, 59 184 167-169, 171, 183
Euiacta cunninghamii (Ait.) elliplica Yamamoto 184 complus Miers 1 84
Link 426 fasciculata Miers 209 incusus Miers 1 84
Eutassa Salisb. 425 haematocarpus Hk.f. & Th. subpeltatus Merr. 182*. 184
cunninghamii Spach 426 184 ihomsonii Miers 184
Evodia 123 laxa Miers 209 validus (Miers) Bakh.f. ex
Exellodendron Prance 642 recisa 207 Forman 184
coriacca 64 lincloria Lour. 160, 165, Haemodoraceae 5: 111-113;
Exiielea Bl. 642 207, 208* 717
10:
Exiteles 673 Fibraureopsis Yamamoto 83 1 Haemodorum coccineum R. Br.
ExiteliaBl61\ smilacifolia Yamamoto 84 1 717
corymbosa (Bl.) Bl. 673 Ficus/7u/cAra Wall. 323 corymbosum Vahl 7 1
muliiflora (Konh.) Walp. Fissipelalum Merr. 29 Halocarpus 354
673 Flacourlia camptoceras Miq. Haloragaceae 7: 239-263,
Eystathes Lour. 493, 507 618 828
Flacourtiaceae 5: 1-106.565; Ilamamelidaceae 5: 363-
6: 943; 7: 827; 9: 563; 379; 6:952; 10: 341,717
Fagaceae 7: 265-403; 8: 10: 333. 539, 715, 716 Harmandia Pierre ex Baill. 1-6.
549; 9: 563 Flagellariaceae 4: 245-250; 9. 10 map
Faika Philipson 255. 262. 263. 5:557; 9: 564 kunstleriKing 9
284 map Folium lunalum minus Rumph. mckongensis Pierre ex Bail!.

villosa (Kanch. & Hatus.) 229 9, 10*

Philip.son 284. 285*. Frcnclopsis 338 Hcbcpctalum 608
286* Funis felleus Rumph. 194 Ucckelia nymanii K.Sch. 253
730 Flora Malesiana [ser. I, vol. 104

Hedycarya 255, 259, 260 Hyalisma 112 {Idenburgia)

arborea 256 Hydnocarpus nana King 716 elaeocarpoides Gilg & Schllr
salomoncnsis Hemsl. 311 Hydrocaryaceae 4: 43-44 147
Hedycrea Schrcb. 645 Hydrocharitaceae 5: 381— novoguineensis Gibbs 147
Hcdyosmum Swarlz 123-129, 413,569; 6: 952; 7: pachyphylla Gilg & Schltr
143. 144 828; 9: 566; 10: 112, 147
arboresccns 125, 127 717 pauciflora A.C. Smith 147
brasiliense 143 HydrophyUaceae 4: 207-209 Indorouchera HaU.f. 607-609,
mexicanum 125 Hypericaceae 8: 1-29; 10: 615
nutans (non Sw.) Merr. 144 717 contestiana (Pierre) Hall. f.

orientale Merr. & Chun Hypericinea macrocarpa Wall. 616*, 619 map
142*, 144 map 626 griffithiana (Planch.) Hall.f.
144
sp. Sieen. Hypericum henryi L6v. & Vaniot 616*, 617*, 618* map
sumatranum 144 ssp. hancockii Robson rhamnifoUaWaXU. 6\9
Hcisieria 3,4 717 Iridaceae 8: 77-84; 10: 717
micranlha 3 uralum Buch.-Ham. ex Irina integer rima Bl. 702
Hcnnecartia 257 D. Don 717 Irvingiaceae 621 , 622
Ilernandia kunslleri 9 Hyperixanthes 489 Isomerocarpa A.C. Smith 263
Ueierosamara birmanica (O.K.) Hypserpa Miers 157, 161, 167- novoguineensis (Perkins)
Chodat 463 169, 172, 222, 218 A.C. Smith p.p. 265, 266
Heterotropa 53, 63 borneensis (Miq.) Becc. Ixionanthes 622
Hexastylis 53, 63 219 Ixonanthaceae 607, 609, 621-
Hibbenia 123 cu5p/Jafa(Hk.f. &Th.) 627, 629
Himantandraceae 562 Miers 219 Ixonanthes Jack 621, 622, 623*
Hippophae L. 151 var.imcrophylla (Miq.) sect. Brewstera (M.J.
Ilolopeira Miers 231 Boerl. 219 Roemer) Hall.f. 621, 625
australis Miers 234 heteromera Miers 219 sect. Emmenanthes Hall.f.
laurifolia (DC.) Miers 234 jagorii'D'itXs 2\9 626
Holostylis 53,61,63 latifolia Miq. ex Diels 221 sect. Ixonanthes 621, 625,
Homalium dasyanthum 716 laurina (F.v.M.) Diels 218, 626
Hopca gracilis Miq. 716 219 beccariiWalU.en
Horsfieldia iryaghedi (Gaertn.) monilifera (Burk.) Diels chinensis (Hk. f. & Am.)
Warb. 605 221 Champ. 626
Hortonia 258, 259 nandinifolia Yamamoto cochinchinensis Pierre 626
Hortoniaceae 261 219 crassifolia Hall.f. 627
Hugonia L. 608, 609 nitida Miers 219, 220* cuneata Miq. 625
sect. Durandea (Planch.) parvifolia Kaneh. & Hatus. dodecandra Griff. 625
BaUIon609, 611 219 grandiflora Hochr. 626
sect. Hugonia 611 polyandra Becc. 219, 221 grandifolia Ridl. 627
costata Miq. 610*, 611, 612 var. tomentosa Forman hancei Pierre 626
map 221 icosandra Jack 624*, 625
jenkinsii F.v.M. 611, 612 praevaricaia Miers 219 var.cuneata Miq. 625
map, 613* propensa Miers 219 var.obovata Ridl. 625
pentagyna (Warb.) K.Sch. raapii Diels 221 khasiana Hk. f. 626
612 selebica Becc. 221 longipedunculata Merr.
robinsonii Merr. 612 selwynii F.v.M. 219 627
sumatrana Miq. 618 iriflora [non (DC.) Miers] lucida Bl. 625
Hugoniaceae 607, 608, 629 Miers 219 multiflora Stapf ex Ridl.
Humiria 629 Hypsipodes Miq. 188 626
Humiriaceae 607, 609, 629, 630 obovata mi.i. 615
Hunga Pancher ex Prance 635- papuana (Schltr) Hub.Winkler
637, 642, 643, 650 Iberis amara L. 543 627
fusicarpa Kosterm. 647 umbellaia L. 543 petiolaris Bl. 623, 625, 626,
longifolia Prance 651, 653* Icacinaceae 7: 1-87, 828; 9: 627*
map 566; 10: 29, 33, 91, 145, petiolaris (non Bl.) Hall.f.
novoguineensis Prance 651, 166, 629, 717 627
652*, 653 map Idenburgia Gibbs 145, 146, philippinensis Elmer 626
papuana (Baker f.) Prance 326, 333 reticulata Jack 625, 626
651, 652*. 653 map arfakensis Gibbs 149 subdodecandra 625
1989] Index to scientific plant names 731

Jackia longifolia 514 (Kibara) (Kibara)

vitellina514 fugax Philipson 290, 300, sleumeri Philipson 290, 303
Jacquemontia browniana Oosistr. 301, 302* stapfiana Perkins 298
716 grandifolia Merr. 298 streimannii Philipson 288,
pannosa (R.Br.) Mabberley hartleyi Philipson 289, 295, 289, 296
716 296 sudestensis Philipson 290,
Jakkia Bl. 493, 508 hirsutaV^aih. 315 303
excelsam.5Q\ hospitans Becc. 311 symplocoides Perkins 289,
longifolia Bl 514 inamoena Perkins 298 290, 303
vitellina Bl. 514 karengana Philipson 289, leijsmanniana Perkins 296
Juglandaceae 6: 143-154, 293 timorensis Boerl. 305
953 katikii Philipson 290, 298 tomeniosa Perkins 298
Juncaceae 210-215; 5:
4: kosiermansii Philipson 289, irichantha Perkins 298
557; 953; 9: 566
6: 294 versieeghii Philipson 289,
Juncaginaceae 4: 57; 5: 554 lalifolia Philipson 257, 289, 294
Juniperus 345 291 vidalii Perkins 298
chinensis 453 laurifolia A.C. Smilh 290. vrieseana Perkins 296
elata Roxb. 363 300 warburgii Perkins 298
Juppia borneensis Merr. 253 leachii Philipson 289, 294 warenensis Kaneh. & Hatus.
ledermannii Perkins 296 289, 295
longipes Perkins 298 Kibaropsis 260
Kairoa Philipson 255, 256, 258, macrantha Philipson 288*. Kingsboroughia Liebm. 690
259, 262, 263, 284 map, 289, 297, 300 Kmeria 566
305 macrocarpa Perkins 298 Korihalsella 353
suberosa Philipson 304*, macrophylla Perkins 298 dacrydii 353
305, 306* merrilliana Perkins 298 Kosiermanthus I*rance 636-
Kibara Endl. 255, 257, 259, microphylla Perkins 289, 639, 641, 642, 675
260, 262, 263, 287, 308 296 heteropetalus (Scort. ex King)
298
angusiifolia Perkins mollis Merr. 298 Prance 676*. 677 map,
archboldiana A.C. Smith moluccana Perkins 288*, 678
257, 289, 291, 293 289, 295, 296, 303 malayanus (Kosterm.) Prance
arucnsis Becc. 305 monticolia Perkins 290, 303 677 map
blumei Sleud. 298 motleyi Perkins 298 Krameriaceae 457, 458
borneensis Boerl. 305 myrtoidea Perkins 289, 291
buergersiana Perkins 305 ncriifolia Perkins 305
bullata Philipson 289, nitens Philipson 290, 299 Labiatae 8: 301-394; 9:

293 novobrilanica Philipson 290, 566

carrii Philipson 257, 290, 301 Lagarosirohus Quinn 360, 361
291, 297 oblongata Philipson 288*. Lardizabalaceae 163, 679
chariacea Bl. 298 289, 295 Larix 343
chimbuensis Philipson 290, oblusa Bl. 290, 300 Lauraceac539,715,717
301 oligocarpclla (Kaneh. & Lauralcs 127,327,330
clemensiae Perkins 298 Hatus.) Philipson 256, Lautcrbachia Perkins 263. 284
coriacea (Bl.) Tulasne 262, 289, 291 map, 326
290, 298 olivaeformis Becc. 305 novoguineensis Perkins
cuspidaia Bl. 298 papuana A.C. Smith 290. 325*. 326
depauperata Merr. 3(X) 299 Lavallea Baill. 19

dichasialis Suesscng. & pcrkinsiae K.Sch. & Laul. ceylanica (Gardn.) Baill.
Heme 298 305 22
ellipsoidea Merr. 298 A.C. Smilh 256.
rigidifolia philippinensis Baill. 22
clmeri Perkins 305 258, 288, 290, 302 Leeaceae 7: 755-782
clongata A.C. Smilh 290. rocmeri (Perkins) Philip.son Legnephora Micrs 158. 165,
297 289, 290 169, 170, 172,225
fcrox Philipson 257. 258. rossclcnsis Philipson 289. acuta Forman 159. 225.
289. 291. 292* 295 226*
flagclliformis Philipson 290. roycnii Philipson 289, 295 microcarpa Forman 225.
301 serrulula Pcrkms 298 226*. 227
form icar urn Becc. 305 shungolcnsis Philipson 289. minutidora (K.Sch.) Dicis
fragrans Philipson 290. 300 296 225. 226*. 227. 228*
732 Flora Malesiana [ser. I, vol. 104

(Lcgnephwa) (Libocedrus) Linum 609

subg. Eulibocedrus Pilger Liriodendron 562, 563, 565-
moorei (F.v.M.) Miers 225,
226* 443 567, 605

nyctericarpa Diels 227 446

arfakensis Gibbs liliiferum L. 582
philippinensis Forman 225, papuana F.v.M. 342, 444, tulipifera 567
226* 445* Liriopsis Spach 598

Lemnaceae 7: 219-237 var. arfakensis (Gibbs) fuscata (Andr.) Spach 598

Lentibulariaceae 8: 275-300 de Laub. 446, 447 map Litsea baticulin (Blco) Kosterm.

Lepidium L. 541,543,545,547 var. papuana 445, 446*, 717

lacteviride (P. Royen) 447 map insignis (Bl.) Boerl. 539

Hewson 548, 549* torricellensis Schltr ex Laut. leytensis Merr. 717

maccowagei Hews. 547, 444 Lobbia Planch. 65

549 Licania Aubl. 635-640, 642, dependens Planch. 75
minuliflorum (Ridl.) Hews. 643, 645 Lobularia maritima (L.) Desv.
548 angelesia Bl. 646 545
sativum L. 547, 548 diemenia Bl. 649 Loganiaceae 6: 293-387,
virginicum L. 547, 548 elaeosperma 640 953,960; 7: 828; 9:567
Lepidocarpa Korlh. 655 fusicarpa (Kosterm.) Prance Longpistillate Manglieiia Meijer

costata Korth. 663 646, 647, 650 map 591

ovalis (Korth.) Bl. ex Miq. macrophylla 639 Lophopetalum beccarianum
661 palawanensis Prance 646, Pierre 716

Lepidocarya costata 663 647 map Lophopyxidaceae 7: 89-91

Lepidothamnus Phil. 360 splendens (Korth.) Prance Lophostylis Hochst. 483
Lepionurus Bl. 31-35, 42, 43 640, 646, 647 map javanica Miq. 484
javanicus G. Don 43 Liliaceae 9: 189-235; 10: Loranthaceae 2, 3, 34, 35
oblongifolius (Griff.) Mast. 109, 112, 166
44 Limacia Lour. 158, 160, 161,
var. angustifoUus Roxb. 167, 168, 170, 172, 221, Macadamia 123
44 222 Macrococculus Becc. 157, 158,
pubescens Ridl. 52 blumei (Boerl.) Diels 222, 160-162, 167-169, 171,
sylvestris Bl. 31, 33, 34, 43, 223 178
44* map, 46 borneensis Miq. 219 pomiferus Becc. 159, 178,
var. lanceolata Val. 43 cerasifera Becc. 224 179, 180*
Leptonium Griff. 43 cerasifolia 224 tympanopodus Laut. &
oblongifolium Griff. 43 cuspidata Ek.f.&Th. 219 K.Sch. 211
Levieria Becc. 263, 275 distincta Miers 223 Madhuca burckiana (Koord.)
acuminata (F.v.M.) Perkins inornata Miers 223 Lamk 593
277, 279 kunstleri King 233 Magnolia L. 564, 566, 567,
beccariana Perkins 277, 280*, microphylla Miq. 219 568
281* monilifera Burk. 221 sect. Gwillimia 566, 569

forbesii Perkins 281, 282 nativitatisRidl. 218 sect. Lirianthe 565

281
laxiflora Perkins 279, oblongaHk.f. & Th. 160, sect. Liriopsis Baillon 598
montana Becc. 277, 279, 165, 166, 223 sect. Maingola 565, 567
281,282 scandens Lour. 220*, 223 sect. Manglietia (Bl.) Baillon
montana (non Becc.) Kaneh. selwynii (F.v.M.) Bailey 589
& Hatus. 277 219 sect. Talauma Baillon 568
nitens Perkins 277, 278 sumatrana Scheff. 198 sect. Theorodon 569
orientalis Philipson 277, 278 triandra (Colebr.) Hk.f. & Th. subg. Magnolia 564-567,
parvifolia A.C. Smith 277 185 570
rudolfii Perkins 278 velutina Hk.f. & Th. 160, sect. Maingola Dandy 569,
scandens Philipson 277, 279 223 570
schlechteri Perkins 278, 279, var. glabrescens King 223 subg. Talauma (Juss.) Pierre
281 Limnanthaceae 639 564-567, 576
squarrosa Perkins 276*, 277, Linaceae 29, 607-619, 621, sect. Aromadendron (Bl.)

278 622, 629, 630, 639 Noot. 566, 569, 576

urophylla Perkins 279, 281 subfam. Hugonioideae 607, sect. Blumiana Bl. 565,
Levieria Kosterm. p.p. 263 608, 609 566, 569, 581
Libocedrus Endl. 337, 339, 340, subfam. Ixonanthoideae 621 subg. Talauma Pierre 568
342, 347, 443, 444 map subfam. Linoideae 607, 609 subg. Yulania 565
1989] Index to scientific plant names 733

(Magnolia) (Magnolia) Malulucban 35

aeqidnoctialis Dandy 57 maingayi (non King) Ridl. Malvanae 151
angaiensis Blco 585 577 Manglietia Bl. 564-568, 589
ashtoniiDandy ex Ncx)t. 570, mariusjacobsia NooL 581. calophylla Dandy 589, 591
576, 578*, 579 588 candollii (Bl.) WaU. 582
ashtonii Noot. 579 nilida563 celebica Miq. 582
beccariana (Agoslini) Nc»L nutans (Dandy) H. Keng 577 dolichogyna Dandy ex NooL
571 odoratissima Reinw. ex Bl. 589, 590*, 591
betongensis (Craib) H. Keng. 583 dolichogyna Dandy ex
585 pachyphylla Dandy 583 Cockbum 591
bintuluensis (Agoslini) Noot. paenetalauma 566 glauca Bl. 589, 590*
570, 576, 577 pahangensis Noot. 570. 576, var. glauca 589, 590
blumei Pranil 589 579 var. lanuginosa Dandy
bomeensis Noot. 570, 576, pealiana (non King) K. & V. 591
579, 580* 571 var. sumatrana Dandy
candollii (Bl.) H. Keng 569, pealii 571 589, 591
581, 582 persuaveolens Dandy 581, glauca (non Bl.) King 591
var. angaiensis (Blco) 587 g/auca (non Bl.) Ridl. 591
Noot. 582, 585 ssp. persuaveolens 587 glauca aucL 585
var. beccarii (Ridl.) Nool. ssp. rigida Nool. 587 lanuginosa (Dandy) Nool.
582, 586 var. pubescens NooL 589, 591
var. candollii 582, 584* 588
587. macklottii Korlh. 571
var. obovaia (Korih.) var. rigida 587 macklottii aucl. 589
Nooi. 582, 585 phaulantha Dandy ex Noot. minahassae K. & V. 593
var. singapurensis (Ridl.) 570. 571. 576 oortii Korlh. 577
Nool. 582, 586 Philippine nsis Parmeni. 604 oortii (non Korlh.) Miq. 577,
carsonii Dandy ex Nool. 570, plumierii Schwartz 568 589
571 pumila aucl. 582. 583 oortii aucL 589
var. carsonii 572*, 573, rumphii Spr. 582 pilosa Parmeni. 590. 591
576 sarawakensis (Agoslini) sabahensis Dandy ex Nool.
var. drymifolia Nool. Noot. 581, 588 589. 591. 592*
573, 574* singapurensis (Ridl.) H. Keng sabahensis Dandy ex
carsonii Dandy ex Cockbum 586 Cockbum 591
571, 572 splendens Reinw. ex Bl. 582 scortechinii King 60
coco (Lour.) DC. 569 sprengeri 567 sebassa King 582
decandollti 5S\, 5^2 stellata 562 singalanensis Agoslini 590
denudata 562 uvariafolia Dandy ex sumatrana Miq. 589, 591
drymifolia Dandy ex Cockbum 574 Manglieliastrum 563
Cockbum 571, 573 uvariifoliaDandy ex Noot. Martyniaceae, see Pedaliaccac
elegans (HI.) H. Keng 570. 570. 571. 574, 575* Malthaea Bl. 255, 258, 262,
576. 577 villosa (Miq.) H. Keng 581. 263, 288, 319, 321 map
forbesii King 582 588 calophylla Perkins 323
fragrans Reinw. ex Bl. virginiana L. 568 charlacea Merr. 321 map,
583 vrieseana (Miq.) Baillon ex 322*. 323
giganlifolia (Miq.) Nool. Pierre 596 heierophylla Quis. & Merr.
581, 586 Magnoliaceae 261. 561-605 321 map, 322*. 323.324
grandi flora L. 569 subfam. Liriodendroidcac 561. intermedia Merr. 321 map,
javanica K. & V. 571 564 322*, 323, 324
kachirachirai 563 subfam. Magnolioideae 561. laiifolia Perkins 323
kunsllen King 582 563. 564. 567. 568 philippinensis Perkins 326
lasia Noot. 581. 587 tribe Magnolieae 561, pincholiana Perkins 323
lilufera (L.) Baillon 582 568 pubescens Merr. ex Perkins
macklottii (Korlh.) Dandy tribe Michelicac Law Yuh- 321 map. 322*. 323
569. 570. 571 wu 561. 568. 593 roemcri Perkins 290
var. bctcariana (Agoslini) Magnolialcs 62, 64, 261 sancta Bl. 320*. 321 map.
N(x)l. 571 Maingaya malayana Oliver 7 1 322*. 323
var. macklottii 571 Malania 1 var. mindanaoensis
maingayi King 570. 571, Maipighiaceae 5: 125-145, Perkins 323
573 566; 6: 960; 10: 458 var. venulosa Perkins 323
734 Flora Malesiana [ser. I, vol. 104

(Malthaea) (Meliosma) (Meliosma)

vidalii Perkins 321 map, ferruginea Bl. 711 multiflora 709
322*. 323, 324 ferruginea (non Bl.) Backer & myriantha Sieb. & Zucc. 681
williamsii Perkins 323 Bakh.f. 709 nervosa K. & V. 704, 705
Matlhiola incana (L.) R.Br. 545 ferruginea (non Bl.) Koord. nitida Bl. 702
Meiogyne virgata (BI.) Miq. 716 709 var. cerasiformis Bl. 702
Melientha Pierre 31-34, 38, 40 ferruginea (non Bl.) Merr. & var. splendens BI. 702
map Perry 711 var. tridenta Bl. 702

acuminata Merr. 39 floribunda Bl. 709 pannosa Hand.-Mazz. 701

suavis Pierre 34, 39* fruticosa Bl. 701 patens Hemsl. ex Forb. &
ssp. macrocarpa Hiepko glauca Bl. 709 Hemsl. 699
40 map var. floribunda (Bl.) paucinervia Merr. 710
40* map
ssp. suavis K. & V. 709 pedicellata K. & V. 695
Meliosma Bl. 163, 679-682, glomerulaia Rchd. & Wils. pedicellata (non K. & V.)
690, 691*, 692* 699 Merr. 697
subg. Kingsboroughia grandifolia Lecomte 713 pendula Merr. 712
(Liebm.)Beus. 691*. harmandiana Pierre 699 petiolaris Miq. 715
694 hirsuta Bl. 694, 703 map, philippinensis Merr. & Perry
sect.Hendersonia Beus. 705, 715 702
692*, 694 humilis Merr. & Perry pinnata (Roxb.) Maxim. 694,
sect.Kingsboroughia 713 707
692*, 694 lanceolata Bl. 694, 703 map, ssp. amottiana (Wight)
subg. Meliosma 691*, 693, 704, 715 Beus. 707, 708 map,
694 var. chariacea Bl. 704 709
sect. Lorenzanea (Liebm.) var. genuina Hochr. 704 ssp. ferruginea (Bl.) Beus.
Beus. 694 var. lanceolata 705 707, 708 map, 709,
sect. Meliosma 692*, 693 f. lanceolata 705 710
subsect. Pinnatae f. nervosa (K. & V.) ssp. humilis (Merr. &
(Warb.) Beus. 693, Beus. 694, 705 Perry) Beus. 707, 708
694 var. membranacea Bl. 704 map, 713
subsect. Simplices var. obliqua Bl. 704 ssp. macrophylla (Merr.)
(Warb.) Beus. 693, var. obliqua (non Bl.) Beus. 706*, 707, 708
694 Koord. 711 map, 709, 711
ser. Curvinervia var. pendula Bl. 704 ssp. pendula (Merr.) Beus.
Beus. 693 var. polyptera (Miq.) Beus. 707, 708 map, 709,
ser. Rectinervia 703 map, 705 712
Beus. 693 /anc//b//aHk.f. 701 ssp. ridleyi (King) Beus.
acutrdnatissima Merr. 712 latifoliaKxfMlM 707, 708 map, 709,
angulata Bl. 690, 698 laurina BIT \5 710
apoensis Elmer 709 lepidota Bl. 694, 695 ssp. sylvatica (Elmer)
arnoitiana Walp. 709 ssp. dolichomischa (Vidal) Beus. 707, 708 map,
bartlettUM&n. 710 Beus. 695, 696* 709, 712
bonlocensis Merr. 697 ssp. kinabaluensis Beus. pinnata Koord. 702
brachyboirys Merr. 713 695, 697 polyptera Miq. 704, 705
cannarioides Elmer 709 ssp. lepidota 695 pungens (Wall, ex W. & A.)
celebica Warb. ex Dihm ssp. squamulata (Hance) Walp. 702
715 Beus. 697 pungens auct. 699
conferiiflora Merr. & Perry ssp. vulcanica (Merr.) reticulata Merr. 712
710 Beus. 695, 697 ridleyi King 710
confusa Bl. 702 levis King 704 rigida Sieb. & Zucc. 699
var. laxior Baker f. 713 loheri Merr. 701 var. angustifolia Miq.
costata CvSod. 701 luzonensis l(y) 699
cuspidata Bl. 702 luzonica 709 var. patens Cufod. 699
diepenhorstii Valet. 702 macgregorii Merr. 712 rufo-pilosa Hend. 694, 703
dolichomischa Vidal 696 macrocarpa Elmer 7 1 map, 714
elegans Rm. 110 macrophylla Merr. 711 sambucina Miq. 709
ellipticaHk.f. 701 megaloboirys Merr. 711 sarawakensis 694
elmeri Merr. 702 monophylla Merr. 701 schlechteri Merr. & Perry
evrardii Gagnep. 701 monophylla Ridl. 696 713
1989] Index to scientific plant names 735

(Meliosma) (Menispermum) Microtinomiscium 158

simplicifolia (Roxb.) Walp. orbiculatum L. 231 Milling tonia Roxb. 690
681, 694, 697*, 698, ovalifolium Vahl ex Pers. arnoitiana Wighl 709
699 map iriandrumRoxh. 185 ferruginea Schull. & Schull.
ssp. fruticosa (Bl.) Beus. trilobum Thunb. 231 711
697*, 698, 699 map, tuber culalum Lamk 194 lanceolata Schull. & Schull.
700*, 701 Metasequoia 345 704
ssp. pungens (Wall, ex Michelia L. 562, 564-567. nitida Schull. & Schull. 702
W. & A.) Beus. 697*, 593, 595, 598, 605 pinnata Roxb. 707
698, 699*. 702 X alba DC. 601,605 pungens Wall, ex W. & A.
ssp. rigida (Sieb. & Zucc.) arfakiana Agoslini 596 702
Beus. 697*, 698, 699 beccariana Agoslini 571 sambucina Jungh. 709
map blumei Steud. 601 simplicifolia Roxb. 697, 698
ssp. simplicifolia 690, celebica Koord. 596 sumatrana Jack 702
697*, 698, 699 map champaca L. 562, 565, 598, Minquarlia 3
sumatrana (Jack) Walp. 694, 601 Miquelia 166
702, 703 map, 715 var. champaca 603 Mirtana loureiri (Pierre) Pierre
sylvatica Elmer 712 var. pubinervia (Bl.) Miq. 210
limorensis Bl. ex Blenk 715 603 Modeccopsis vaga Griff. 17
tongcalingii Elmer 711 champaca Miq. 603 Mollinedia 256
trichocarpa Merr. 710 champacca aucl. 596 acuminata F.v.M. 279
vulcanica Mcrr. 697 cumingii Merr. 604 coriacea (Bl.) Baill. 298
wallichii (non Planch, ex dollsopa auct. 589 sancta (Bl.) Baill. 323
Hk.f.)Koord. 711 ecicatrisaia Miq. 603 Molluginaceae, sec Aizoaceae
wightii Planch. 702 figo (Lour.) Spr. 599, 605 Monaria Konh. ex Val. 17
Meliosmaceae End!. 679, 680 forbesii Baker f. 596 Monimia 257-261
Memecylanthus Gilg & Schltr fuscala 562 oval folia 257
i

335 koordcrsiana Nooi. 601, Monimiaceae 145,255-326,

Meniscosta Bl. 682 602* 256 map, 327, 330
javanica Bl. 684 longiflora 605 subfam. AUierospcrmaloidcac
scandens Bl. ex Spr. 684 longifolia Bl. 605 255, 258-263
Menispermaceae 19, 157- var.racemosa Bl. 605 subfam. Horionioideae 260,
253, 305, 679, 680. mollis (Dandy) McLauchlin 261
681 596 subfam. Mollinedioideae 259,
iribc Coscinieac 161, 162, moniana Bl. 563, 601, 603, 261,263
164 604 map subfam. Monimioideae 259-
iribe Fibraureeae 162, 164, var. subvelulina Miq. 604 263, 277
168 montana aucl. 603 subfam. Siparunoideae 258-
tribe Menispermeae 161, 162, parviflora 604 261
164, 168 philippinensis (Parment.) tribe Hcdycaryeae 255, 259,
iribc Tiliacoreae 162, 164, Dandy 601,604 262, 263, 277
168 p////era Bakh.f. 603 tribe Mollincdieae 255, 262,
iribc Tinosporcae 158, 161, platyphylla Merr. 596 263, 277, 284 map, 326
162, 164, 167, 168, 189* pubinervia Bl. 603 tribe Monimieae 259
162
iribc Triclisieae rufinervis Bl. 603 tribe Trimcnicac 327
Mcnispcrmum 158, 165 salicifolia Agoslini 601, 604 Monnina 456
cocculus L. 213 scoriechinii (King) Dandy emarginaia458
crispum L. 194. 195 600*, 601 ejcce/.va(BI.)Spr. 501
dauhcum 165 spec. Slapf 587 longifolia {E\.)S\)v.5\A
fenesiralum Gaerin. 215 suaveolens Pers. 601 macrophylla Slcud. 514
flavescensLamk 210 sumatrae Dandy 604 vj/e/Z/na (Bl.) Spr. 514
flavum L. 210 isiampacca Bl. 603 Monoon incertum (Bl.) Miq.
glabrumBumn.f. 199,200 tsiampacca L. 596 515
glaucum Lamk 229 var. blumei Mor. 603 Monquarlia 4
japonicum TTiunb. 245 velutina Bl. 603 Moquilea Aubl. 645
lacunosum Lamk 213 Microcachrys338. 343. 351. sect. Cyclandrophora (Ha.ssk.)
limacia (DC.) Sprcng. 352 Endl. 665
223 Microcarpus 35) Morinjjaccae 4: 45—46; 5:

nuUabaricum Lamk 200 Microdcsmis Hk.f. 29 554; 6: 960

736 Flora Malesiana [ser. I, vol. 104

Moutabea Aubl. 456, 493 Natsiatum Rheede 213 (Olax)

Munnickia Bl. ex Rchb. 65 Ncmuaron 256 obtusa Bl. 7
Myoporaceae 4: 265—266; Neocarya Prance 642 psittacorum (Willd.) Vahl
568
9: macrophylla (Sabine) Prance 717
Myricaceae 4: 276-279 641 rosea Ridl. 9
Myristicaceae 605 Nephroia Lour. 231 scandens Roxb. 2, 7, 8*, 9,

Myrtales 151 elegans Ridl. 233 717

Myrtanae 1 5 sarmentosa Lour. 233 semiinfera Val. 9
Myrtiflorae 151 Nigrina Thunb. 129 stricta 2
spicala Thunb. 133 sumatrana Miq. 9, 50
spicifera Lamk 133 Onagraceae 8: 98-113
Nageia Gaertn. 337, 343, 347, Nouhuysia Laut. 145, 146 Ongokea 3, 4
353-355, 389, 391 map arfakensis (Gibbs) Steen. 149 Ophiocaryon Schomb. 163, 679,
sect. Nageia 390 novoguineensis (Gibbs) 680
sect. Polypodiopsis (Bertrand) Hatus. 147 Opilia Roxb. 4, 31-35,46
de Laub. 390, 394 pachyphylla (Gilg & Schltr) sect. Lepionurus (Bl.) Baill.

amara (Bl.) O.K. 387 Hatus. 147 sect. Opiliaslrum Baill. 35

beccarii (Pari.)Gordon 393 papuana Laut. 147 subg. Urobotrya (StapQ Engl.
blumei (Endl.) Gordon 393 pauciflora (A.C. Smitji) 41
cmot//i^/7 (Pari.) O.K. 381 Steen. 147 acuminata Wall. 44
discolor (Bl.) O.K. 400 Nyctaginaceae 6: 450-468; amentacea Roxb. 31-34, 47
falciformis (Pari.) O.K. 372 7: 829 map
leptosiachya (Bl.) O.K. 400 Nyssaceae 4: 29-31 celtidifolia (Guill.& Perr.)
maximus (de Laub.) de Laub. Endl. ex Walp. 33, 47
339, 394 map cumingiana Baill. 36
motleyi (Pari.) de Laub. 393, Ochanostachys Mast. 1-6, 12, fragrans Elmer 47
394 map 14 map javanica Miq. 47
neglecta (Bl.) O.K. 400 amentacea Mast. 5, 13*, 14 manillana Baill. 36
neriifolia (D. Don) O.K. 400 bancana (Becc.) Val. 14 pentitdis Bl. 47
polystachyus (R. Br. ex Ochnaceae 7: 97-119; 10: thorelii Gagn. 47
Endl.) O.K. 417 145, 621 tomentella (Oliv.) Engl. 47
/Mm/7;://(Bl.)F.v.M. 415 Ochthocosmus Benth. 621 Opiliaceae 2, 9, 29, 31-52
teysmannii (Miq.) O.K. 406 Octarillumfruticosum Lour. tribe Opilieae 34
thevetiaefolia (Bl.) F.v.M. 153 Osmoxylon sessiliflorum (Laut.)
417 Octoknema 3 Philipson 716
vitiensis (Seem.) O.K. 394 Octoknemaceae 33 Oxalidaceae 7:151-178,
map Odontocarya 158, 164 829; 10: 607
wallichiana (Presl) O.K. Odonlocaryioides 158
391, 392*, 393 map Olacaceae 1-29, 166, 629,
Najadaceae 6: 157—171 717 Pachydiscus Gilg & Schltr 335
Nallogia 35
Baill. tribe Agonandreae 34 Pachygone Miers 160, 165, 167,
gaudichaudiana Baill. 36 tribe Anacoloseae 3 168, 170, 172, 217
Nasturtium R.Br. 555 tribe Aptandreae 4 brachystachya (DC.) Miers
backeri Schulz 556 tribe Couleae 4 217
benghalense DC. 560 tribe Heisterieae 4 hebephylla Miers 217, 218
diffusum auct. 559 tribe Olaceae 4 leplostachya (DC.) Miers
heterophyllum Bl. 559 tribe Opilieae 34 217,218
homalospermum Schulz 557 tribeSchoepfieae 4 ovata (Poir.) Hk.f. & Th.
var. macrocarpum Schulz Olacales 33, 34 217, 218, 220*
557 Olacinea ignota 25 var. dasyphylla Miq. 217
hybospermum Schulz 559 Olacineae 29, 34 var. rolundifolia Miq.
indicum (L.) DC. Olax L. 1-5,6 217
vai.javana Bl. 559 sect. Triandrae Engl. 7 pubescens Benth. 217, 218
indicum auct. 557, 559 baticulin Blco717 zeylanica Sant. & Wagh
novo-guineense Gilli 557 benthamiana 3 218
officinale R.Bt. 555 imbricata Roxb. 7, 8 Pachylamax Dandy 561, 567,
palustre (L.) DC. 557 laxiflora Ridl. 9 568, 593
peekelii Schulz 557 muliiflora A. Rich, ex Baill. 8 praecalva Dandy 593, 594*
schlechieri Schulz 559 multiflora Ridl. 9 Paelae 493
1989] Index to scientific plant names 737

Palmeria F.v.M. 255, 257-263, (Parabaena) (Parinari)

267 elmeri Diels 189*, 202 bicolor Merr. 664
acuminata Kaneh. & Halus. var. philippinensis campesu-is 641
270, 271 (Merr.) Yamamoto 203 canarioides Kosterm. 636,
angica Kaneh. & Hatus. 267, hirsuta (Becc.) Diels 193 655, 656 map
271 hirsuta (non Becc.) Diels 202 coriacea Benth. 641
arfakiana Becc. 262, 267, megalocarpa Merr. 158, costata (Korth.) Bl. 636, 656,
270 186*, 189*. 202 663
brassii Philipson 269, 271, myriaditha 203 ssp. costata 663, 664 map
272 myrianiha K. Sch. 203 ssp. polyneura (Miq.)
clemensae Philipson 268*, philippinensis Merr. 203 Prance 655, 663, 664
269, 273 sagiitaia Miers 189*, 201, map
dallmannensis Kaneh. & 202 ssp. rubiginosa (Ridl.)
Hatus. 272 scylophylla Diels 197 Prance 663, 664 map
fengeriana Perkins 269, 271 tuberculata Becc. 189*, 202, curaiellifolia641
gracilis Perkins 267, 269, 203 elmeri Merr. 655, 657 map
272 Paracryphiaceae 145 excelsa Sabine 639, 641
habbemensis A.C. Smith 272 Parakibara Philipson 255, 262, giganiea Kosterm. 636, 655,
hooglandii Philipson 267, 263, 284 map, 286 660 map
269 clavigera Philipson 286, glaberrima Hassk. 641
hypargyrea Perkins 269, 272 287* glaberrimiun (Hassk.) Hassk.
hypochrysea Perkins 275 Paramanglielia Hu & Cheng 669
incana A.C. Smith 269, 272 589 var.lanceolatum (T. & B.)
montana A.C. Smith 269, aromatica (Dandy) Hu & & V. 669
K.
272, 275 Cheng 589 griffithiana 64
myrianiha Perkins 270, 271 Paramichelia H.H. Hu 565, 567, heteropetala Scort. ex King
myrtifoUa Perkins 270 593, 599 675
paniculate Ridl. 269 baillonii (Pierre) Hu 599 insularum 639, 656
parvifolia Kaneh. & Halus. scoriechinii (King) Dandy jackiana Benth. 641, 671
276,271 601 latifrons Kosterm. 666
puberula A.C. Smith 270 Pararistolochia 53, 57, 59, 61, macrophylla Sabine 641
pulchra Perkins 270, 271 62,64 meiallica Kosterm. 656, 660
pulteana Perkins 272 Parasitaxus 343, 351, 354 map
scandens 272 Paraslemon A. DC. 635-638, montana 641
schoddei Philipson 269, 275 640, 642, 648 myriandra Merr. 675
warburgii Perkins 270, 27 grandifructus Prance 649, nannodes Kosterm. 667
womersleyi Philipson 269, 650 map nonda F.v.M. ex Bcnlh. 636,
273, 274* spicalus Ridl. 649 655, 658 map
Pandaccae 29 urophyllus (Wall, ex A. DC.) nonda auct. 658
Papaveraceae 5: 114-117 A. DC. 637, 648*. 649, oblongifolia Hk.f. 636, 655,
Papuacedrus Li 443 650 map 659, 660 map
arfakensis (Gibbs) Li 446 versteeghii Merr. & Perry papuana C.T.Whilc 655, 658
papuana (F.v.M.) 444
Li 637, 649, 650 map ssp. papuana 658, 659
445
torricellensis (Schllr) Li Parinari Aubl. 635, 636, 639- map
/'apuz/V/a Ridl. 541,547 642, 654 ssp. salomonense
laeteviridis P. Royen 548 seel. Exitelia (Bl.) C. Mucll. (C.T.White) Prance
mmutiflora Ridl. 548 673 658, 659 map
minutiflora auct. 549 sect.Neocarya DC. 64 ssp. whilci Prance 658,
Parabacna Micrs 158, 161, 162, sect. Pclrocarya 64 659 map
167, 168, 170. 171, 201 sect. Sarcoslegia Benth. 671 parva Koslcrm. 636, 655,
amplifolia Diels 193 subg. Euparinari 654 657 map
cincinnans (K. Sch.) Diels subg. Exitelia Bl. 673 polyandra 64
203 subg. Sarcostegia (Bcnlh.) prancci Koslcrm. 656, 660,
dcnudata Diels 189*. 202. Miq. 671 661 map
203 anamensis 656 rigida Kosierm. 656, 660,
. cchinocarpa Diels 189*. 201. argcnlco-sericea Kosierm. 661 map
202, 204 655, 656 map rubiginosa Ridl. 663
var. pubescens Yamamoto ashtonii Koslcrm. 660 salomonense C.T. White
204 asperula auci. 667 659
738 Flora Malesiana [ser. I, vol. 104

(Parinari) Parinarium auct. 661, 665 (Phyllocladus)

scabra Hassk. 641 Passiflora aurantia Forst. f. 717 hypophyllus Hk.f. 342.
senegalensis DC. 641 Passifloraceae 7: 405-434, 357*, 358*, 359*,
sumatrana (Jack) Benih. 829; 10: 166, 717 360*
641, 656, 661 map, Pedaliaceae 4: 216-221; 5: var. protracta Warb. 359
662* 557; 7: 829 major Pilger 359
wallichiana R. Br. 665 /'e/ae Adanson 493 protractus (Warb.) Pilger 359
Parinari auct. 665, 675 Pentaphragmataceae 4: 517- Phyllocosmus Klotzsch 621
Parinarium Juss. 654 528 Phytocrene 91, 166
sect. Cyclandrophora (Hassk.) Pentaphylacaceae 5: 121- /o/jm Merr. 219
C. Mucll. 665 124, 566 malacothrix Sleumer 227
sect. Neocarya DC. 654 Pentildis Zipp. ex Bl. 46 Phytolaccaceae 4: 228-232;
sect. Petrocarya DC. 654 Peponaster major Rumph. 94 5: 557

subg. ///Hk.f. 665 Pericampylus Miers 160, 169, Picea 338

subg. Cyclandrophora 170, 172, 227 Pierotia Bl. 622
(Hassk.) 31. 665 glaucus (Lamk) Merr. 165, lucida Bl. 625
subg. Macrocarya Miq. 228*, 229, 236 reticulata Bl. 626

665 incanus (Colebr.) Hk.f. & Pimela angustifolia Bl. 715

subg. Petrocarya (DC.) Miq. Th. 229 Pinaceae 343, 347, 447-453
654 lanuginosus (Bl.) Miq. Pinus L. 337-340, 342, 347,
amboinense T. & B. 669 229 447
asperulum Miq. 67 membranaceus Miers 229 subg. Diploxylon 449
borneense Merr. 659 Periomphale 335
Baill. dammara Lamb. 433, 434
corymbosum (Bl.) Miq. 673 papuana 336
Steen. finlaysoniana Bl. 45
costatum Bl. Peripetasma polyanthum Ridl. 452
insular is Endl.
var. rubiginosum Ridl. 253 kasya 452
Pari.

663 Perrottetia alpestris (Bl.) Loes. kesiya Royle ex Gordon 341,

curranii Merr. 669 ssp. philippinensis (Vidal) 346,451, 452 map
elaium King 669 Ding Hou 716 khasia Engelmann 452
excelsum 654 Petalinia Becc. 12 khasyaWk.LASl
fragile T. & B. 646 bancana Becc. 14 var. insularis (Endl.)
grifhihianum Benth. 673 Petrocarya Schreb. 641, 654 Gaussen 452
hahlii Warb. 669 excelsa Jack 641, 670, khasyana Griff. 452
helferi Hk.f. 663 671 latteri Mason 451, 452
heieropelalum Scort. ex King glaberrima (Hassk.) Miers merkusiana Cooling &
677 669 Gaussen 45
jackianum Benth. 670 scabra (Hassk.) Miers 669 merkusii Jungh. & de Vriese
kunslleri King 677 sumatrana Jack 661 340, 341, 346,448*,
lanceolatum T. & B. 669 Petrocarya auct. 665 449*, 450*. 451 map
laiifolium Hend. 666 Petrosavia 109, 112 var. tonkinensis Chev.
laurinum A. Gray 669 Petrosaviaceae 109 451,452
maingayi King 671 Peumus 259, 260-262 sumatrana Jungh. 451
maranthes Bl. 673 boldus 260 sylvestris auct. 45
margarata A.Gray 669 Phanerodiscus 4 taeda auct. 452
mindanaeruse Perk. 669 Phelima Nor. 605 timorensis 452
mulliflorum (Korlh.) Miq. Pherosphaera 343 Piperaceae 127
674 Pherosphaeraceae 354 Piperales 127, 567
myriandrum Merr. 677 Philbomea Hall.f. 607-609, Piptocalyx Oliv. ex Bth. 255,
nitidum Hk. f. 646 614 261, 326, 327, 330
nitidum auct. 674 magnifolia (SiapO Hall.f. macrurus Gilg & Schltr 330,
palauense Kaneh. 674 614*, 615 map 332
philippinense Elmer 646 palawanica Hall.f. 615 moorei Oliv. 330
polyneurum Miq. 664 Philydraceae 4: 5-7; 7: 829 Pittosporaceae 5: 345-362;
racemosum Vidal 674 Phlebocalymna 145 6: 960; 7: 829; 9: 568;
salicifolium (Presl) Miq. 674 lobospora F.v.M. 149 10: 717
scabrum Hassk. 669 Phyllocladaceae 354 Pittosporum moluccanum
spicaium King 671 Phyllocladus L.C. Rich, ex (Lamk) Miq. 717
villamilii Merr. 671 Mirbel 337-343, 347, Platea 145
warburgii Perk. 669 354, 355, 357 map Plumbaginaceae 4: 107-112
1989] Index to scientific plant names 739

Podocarpaceae 338, 342, 343, (Podocarpus) (Podocarpus)

347, 351-419, 420 brassii Pilger 412, 413 map javanica sensu Merr. 379
Podocarpus rHerit. ex Persoon var. brassii 413 javanicus (Burm. f.) Merr. 453
337, 340, 343, 347, 351- var. humilis de Laub. 339, javanicus (non Burm. f.) Merr.
355, 395, 397 map 413, 414 377
sect. Acuminatus de Laub. brevifolius (Stapf) Foxw. junghuhniana Miq. 400
397, 398, 404 402,412,413 map. 414 kawaii Hayata 379
sect. Dacrycarpus Endl. 374 brevifolius (non StapO Foxw. koordersii Pilger ex K. & V.
sect Dacrydioideae Bennett 411 415
374 celebica (non Hemsl.) Warb. latifolia f. ternaiensis de Boer
sect.Dacrydium Bertrand 374 410 393
sect.Dammaroides Bennett celebicus Hemsl. 349 latifolius (non Thunb.) BI.
390 cincius Pilger 383 391
sect. Foliolatus de Laub. compacta Wasscher 383, laubenfelsii Tiong 399. 415.
397, 398, 399 445* 416 map
sect. Globulus de Laub. 397, confertus de Laub. 340, 341, ledermannii Pilger 405 map
398, 405 407, 408, 409 map leptophylla Wasscher 371
sect Gracilis de Laub. 398, costalis PresI 412, 413 map, leptostachya BI. 400
409 414 levis de Laub. 398. 400,
sect. Longifoliolatus de Laub. costalis (non Presl) Foxw. 403, 404 map
398, 407 410 lophatus de Laub. 410, 411
sect. Macrostachyus de Laub. crassigemmis de Laub. 353, map
398, 412 402, 412. 413 map lucienii 406
secL Nageia Endl. 390 cumingii Pari. 381 macrocarpus de Laub. 399,
sect. Polypodiopsis Bertrand cupressina sensu Lane-Poole 417, 418 map
394 379 maximus (de Laub.) Gaussen
sect. Polystachyus de Laub. cupressina R. Br. ex Mirbel 394
398, 399, 416 377 micropedunculatus de Laub.
secL Prumnopiiys (Philippi) cupressina Ridl. 379 339, 341. 405 map
Bertrand 384 var. curvulaMiq. 380 nakaii 406
sect. Rumphius de Laub. dacrydiifolia Wasscher 383 neglecta BI. 400
398, 399,406. 414 decipiens Gray 400 neriifolius D.Don 399. 400
sect Stachycarpus Endl. deflcxusRidl. 340, 341.400, map
384 403 map var. atjehensis Wasscher
sect. Sundacarpus Buchholz 400
discolor BI. 409
& Gray 385 dulcamara Seem. 387 var. bractcatus (BI.)
sect. Taxoideae Bennett 384 elaia R.Br. 453 Wasscher 408
subg. Foliolatus de Laub. eurhyncha Miq. 385, 387 var. brevifolius Stapf
397 fale ifor mis Pari. 372 414
subg. Stachycarpus (Endl.) filicifolius Gray 395 var. brevipes (BI.) Pilger
Engl. 384 gibbsii N.E. Gray 398, 407, 408
affinis410 408 map var. polyantha Wasscher
agalhifoliaB]. 391 glaucus Foxw. 409, 410. 400
amara BI. 385 411 map var. ridleyi Wasscher 419
archboldii N.E. Gray 399, globulus dc Laub. 406 map var. teysmannii (Miq.)
402, 403 map idenburgensis N.E. Gray 405 Wasscher 406
var. crassiramosus N.E. imbricatus BI. 376 Wasscher
var. timorcnsis
Gray 409 var. cumingii (Pari.) 401
archboldii (non N.E. Gray) Pilger 381 neriifolius D. Don in
Gausscn 412 var. curvula (Miq.) Lamb. 417
atjchcnsis (Was.schcr) de Wasscher 380 neriifolius (non D. Don)
Laub. 408 map. 409 var. kinabaluensis Stecn. 408
beccarii Pari. 393 Wasscher 38 novaccalcdoniac 403
blumei EnA\. 391 imbricatus scnsu Foxw. palcmffanica Miq. 453
bomccnsis dc Laub. 398, 379 papuanus Rki\. 379
400. 403 map imbricatus (non BI.) Gibbs papuiinus (non Ridl.) Pilger
bractcatus BI. 341,408,409 379. 381 383
map insularis dc Laub. 399. 402. papuanus (non Ridl.) Sicup.
var. brevipes BI. 408 404 map 380
740 Flora Malesiana [ser. I, vol. 104

(Podocarpus) (Polygala) (Polygala)

pcdunculala Bailey 387 arillalaBuch.-Ham. 461, linarifolia (non Willd.)

philippinensis Foxw. 415 467*. 469 Adema 481
piigcri Foxw. 402, 410. 411 arvensis Willd. 477, 478 linearifolia 48
map var. squarrosa Benth. 476 longifolia Poir. 460, 462,

pilgeri (non Foxw.) v. Roycn arvensis (non Willd.) Adema 474*

401 477 lutea460
polyanlha (Wasschcr) arvensis (non Willd.) Benth. luzoniensis Merr. 472
Gausscn 400 482 macrostachya Hassk. 481
polystachyus R.Br, ex Endl. brachislachyos 476 malesiana Adema 461, 463,
339-341, 417*, 418 brachystachya BI. 477, 480 464*
map brachystachya DC. 476, mariesii Hemsl. 465
var. rigidus Wasschcr 403 477 monspeliaca (non L.) Blco
polystachyus (non R.Br.) Li buchanani Buch.-Ham. ex 482
&Keng414 D. Don 474 monticola HBK 460
pscudobractealus dc Laub. buxiformis Hassk. 477 monticola (non HBK) Ridl.
342, 409 map cardiocarpa Kurz 461 463, , 467
ridleyi (Wasscher) N.E. Gray 464* oligophylla DC. 474
340, 341, 398,417,418 cardiocarpa (non Kurz) Ridl. orcotrephes Burtt 461, 467*
map, 419 463 paenea 457
405
ridleyi auct. chamaebuxus 460 palustris Lace 463
rotundus de Laub. 410, 411 chinensis (non L.) Benn. 477 paniculataL. 458, 460, 461,
map chinensis L. 458, 462, 477, 475*
rubcns de Laub. 399, 401*, 478*, 479 papuana (Steen.) Meijden
402 map var. brachystachya (Bl.) 461, 465, 466*
rumphii Bl. 403, 415,416 Benn. 481 persicariacfolia DC. 460,
map var. linearifolia (non 462, 473*
410
schlechleri Pilger Willd.) Chodat 481 polifolia Presl 462, 476,
solomoniensis 403 var. triftora (L.) Benn. 482 477*
spathoides dc Laub. 398, 400, ciliata L. 487 476
polyfolia
404 map 478
densiflora Bl. polygama 458
steupii Wasscher 380, 381 discolor Buch.-Ham. ex prostrata Willd. 482
map D. Don 474 pulchra Hassk. 468
teysmannii Miq. 406 map elongata WiUd. 477, 480 pyramidalis Lev. 474
ihevetiifolia B 1.417 elongata (non Willd.) Benn. rhinanthoides Benth. 460,
vitiensis Seem. 394 482 461, 480*
wallichianus Presl 391 criocephala Benth. 476 riukiuensis Ohwi 474
wangii Chang 410 eumekes Hassk. 481 rufa Span. 474
Podocarpus Labill. 356 exsquarrosa Adema 460, 4o2, septemnervia Merr. 474
Podostemaceae 4: 65-68; 476* sibirica L. 473
6: 963; 10: 717 fernandesiana Paiva 475 472
sibirica (non L.) Hassk.
Polemoniaceae 4: 195-196 furcata Roylc 463, 465, simadae Masam. 477
Polyalihia pulchrinervia Boerl. 488 simassan Miq. 468
674 glaucocarpa Ridl. 468 sumatrana Miq. 461, 467*,
Polycarpicae 165, 261 glaucoides L. 462, 477, 479, 468
Polygala L. 455-457, 459 480, 481* tatarinowii Regel 461, 462,
sect. Chamaebuxus DC. 455, var. triflora (L.) Trimen 464*, 488
460,461, 465, 467 482 telephoides Willd. 477,
sect. Melchiora (Stccn.) glomerata Lour. 478, 479 478
Meijdcn 461, 465 hondoensis Nakai 472 telephoides (non Willd.)
sect. Orihopolygala Chodat humilis Span. 481 W. & A. 476
469 isocarpa Chodat 463 tinctoria (non Vahl) Hassk.
sect.Polygala 460, 461, 469 japonica Houtt. 462, 472* 471
sect.Pseudoscmciocardium javana DC. 460, 461, 470*, tonkinensis Chodat 469
Adema 460,461, 462 471* toxoptera Turcz. 478
sect. Semeiocardium [non karensium Kurz 469 iranquebarica Mart. 478
(Zoll.) Hassk.] Chodat khasyana Hassk. 472 trichocolpa Chodat 469
462 leptalea DC. 474 triflora L. 462, 479, 481,
acicularis Oliv. 465 ///iar//b//a Willd. 477, 481 482*
1989] Index to scientific plant names 741

(Polygala) Prunus 640 Radix puluronica Rumph.

lriner\aia Ham. ex Wall. 487 Pseudaleia imbricata (Roxb.) 97
iriphylla Buch.-Ham. ex Hassk. ex Valeton 9 Ranunculales 163, 680
D. Don ongistylis Hassk. ex RaphanusL. 541, 543,545,
var. glaucescens (non Valeton 9 546
Wall.) Benn. 463 Pleleocarpa Oliv. 29 caudaius L. 546
triphylla (non Buch.-Ham. ex Pierocarya griffithiana (Benth.) raphanistrum L. 547
D. Don) Royle 462, 539 Miers 674 sativus L. 541, 546
umbonaia Craib 463 Pteroneurum DC. 550 Reinwardlia 609
undulaia Roxb. 487 decurrens Bl. 554 Restionaceae 5: 416-420,
variabilis (non HBK) Hassk. javanicum Bl. 551 569
475 Ptychopeialum 2-4 Rhamnales 151
vaulhieri Chodat 460 Punicaceae 4: 226-227 Rhipogonum album R.Br. 253
venenosa Juss. ex Poir. 461, Pycnarrhena Miers ex Hk.f. Rhizophoraceae 5: 429-493;
467*, 468 & Th. 157, 160-162, 6: 965; 8: 550; 9: 568;
ssp. pulchra (Hassk.) 167-171. 172. 173, 178. 10: 253, 717
Sieen. 468 305 Rhopalopilia 31-33
ssp. venenosa Sleen. 469 auslraliana F.v.M. 175 pallens Pierre 33
var. eramosa O.K. 468 balabacensis Yamamoto Rorippa Scop. 541, 543, 545,
var. robusta Craib 468 175 555
veronicaefoliaAll balanensis Yamamoto 175 backeri (Schulz) Jonsell 541,
veronicea F.v.M. 472 borneensis Diels 1 75 555, 556
virgaia458 calocarpa (Kurz) Diels 174 benghalensis (DC.) Hara 555,
vulgaris L. 460 casianopsidifolia Yamamoto 560
vulgaris (non L.) Thunb. 472 177 dubia (Pers.) Hara 559
wallichiana Wight 474 celebica (Boerl.) Diels 175 heterophylla (Bl.) Williams
warburgii Chodat ex Warb. elliptica Diels 175 555, 559
477 fasciculaia (Miers) Diels hybosperma (Schulz)
watiersii Hance 465 174 Jonsell 55, 559
wightiana W. & A. 460, 462, ^ran^/5 K.Sch. &Uut. 175 indica (L.) Hicm
479* insignis (Hatus.) Forman var. apeiala (Lour.) Hochr.
Polygalaceae 455-539, 639 173, 174 559
tribe Moulabeae 455, 458 longifolia (Dccne ex Miq.) islandica (Ocd.) Borb. 557
tribe Polygaleae 458 Becc. 173, 174 micrantha (Roih) Jonsell 555,
tribe Xanihophylleae 458 lucida (T. & B.) Miq. 173, 560
Polygalales 457, 458 174, 178 nasturtium-aquaticum (L.)
Polyosma 145 manillensis Vidal 173, 175 Hayek 555, 556*
Pontederiaceae 4: 255-261; membranifolia Merr. 175 officinalis (R.Br.) P. Royen
5:557; 6: 964 merrillii Diels 175 556
Porotheca K.Sch. 187 moniana Back. 174, 178 palustris (L.) Becc. 555, 557
peliolaia K.Sch. 187 nogovuineensis Miq. 173, pcekelii (Schulz) P. Roycn
Portulacaceae 7: 121-123 175 555, 557, 558*
Primulaceae 6: 173-192, ozantha Diels 173, 176*, schlechteri (Schulz) P. Roycn
964 177, 179 559
Proteaceae 5: 147-206, 566; papuana Kanch. & Hatus. Rosaceae 638, 639
6: 965; 7: 830; 9: 568; 177 subfam. Chrysobalanoidcac
10: 151 plcniflora 173 639
Proteales 151 sayeri Diels 175 subfam. Neuradoidcae 639
Proicanae 151 lumcfacta Miers 159, 173, subfam. Prunoidcae 640
Proiium 123 175, 176* Rosalcs 639
Protoalhcrospcrmoxylon 256 Pyrolaccac 335 Roucheria 608
Prumnopitys Philippi 343, 347, Pyrrhosa horsfieldii (Bl.) Hassk. coniesliana Pierre 619
352, 353. 355. 384. 385 605 griffithiana Planch. 617
map Pyrularia 33 Ruiaccac 165
sect.Sundacarpus (Buchhol/ Ryparo.sa kunstlcri King 539
& Gray) dc Laub. 385
amara (Bl.) de Laub. 385. Qucrcus gilva Bl. 701
386*. 387 map. 388*. var. procera Bl.701 Sabah Mangtieiia Dandy ex
389* jama-buwa Sicb. 699 Mcijcr 591
742 Flora Malesiana [ser. I, vol. 104

Sabia Colebrooke 163, 166, (Salomonia) Sarcosperma 29

679-681, 682, 683 map cordata Wight 487 Sarcosperma{ta)ceae 4: 32-
campanulata Wall. 685 cy//uinca(Bl.)Kur/491 34; 6: 967; 10: 29
celastrinea Muell. 685 edentula DC. 487 Sarcostegia Benih. 641
densiflora Miq. 690. 698 elongata (Bl.) Kurz ex Koord. Sanima53. 57, 59,61-64
elliplica (Miq.) Miq. 684 490 henryi 62
crralica v. d. Water 683, 684 horneri Hassk. 488 Saururaceae 4: 47—48

floribunda Miq. 690, 698, longiciliata Kurz 488 Saxegothaea343,351,352

701 mariinii L^v. 488 Saxegothacaceae 354
harmandiana Pierre 687 oblongifolia DC. 487 Saxifragaceae 145
japonica 163 obovata Wight 487 Scaphocalyx parviflora Ridl.
javanica (Bl.) Backer ex Chen parasitica Griff. 490 716
684, 685 peiiolata D. Don 487 spathacea Ridl. 716
var. gtabriuscula (Bl.) ramosissima Turcz. 488 Schisandraceae 163, 679
Chen 684 rigida Hassk. 488 Schoepfia Schreb. 1-6, 27
limonacea 685 seguinii Lev. 488 fragrans Wall. 2, 27, 28*
limoniacca Wall, ex Hk.f. & sessiliflora 487 Schoepfiopsis Micrs 27
Th. 684, 685, 686*, 687* sessilifolia D.Don 487 fragrans (Wall, in Roxb.)
malabarica Bedd. 685 setoso-ciliata Hassk. 488 Miers 27
menescorla 684 stricta Sieb. & Zucc. 488 Sciadocarpus Hassk. 287
menicosla 684 subrotunda Hassk. 487 brongniartii Hassk. 298
meniscosia Bl. 684 trinervata Steud. 487 Sciadopitys 354
var. eltiptica Miq. 684 uncinata Hassk. 488 Sciaphila Bl. 110, 111 map,
\ai.firma Bl. 684 Salomonia auct. 488 112, 720
var. glabriuscida Bl. 684 Salvadoraceae 4: 224-225 sect. Hermaphroditantha
var. latifoUa Bl. 684 Sampacca O.K. 598 subsect. Polyandra 112
papuana Warb. 688 domestica IV alba Rumph. sect. Hexanihera 1 13

parviflora Wall. 683, 687 605 sect. Oliganthera

ssp. parviflora 685, 687 longifolia O.K. 605 subsect. Bilobatae 113
ssp. philippinensis monlana O.K. 603 subsect. Quadrilobatae
(Robins.) v.d. Water montana Rumph. 582 112, 113
687, 688* sylvestris Rumph. 596 subsect Trilobatae
var.harmandiana Lecomte velutina O.K. 603 113
687 Santalaceae 2, 3, 33-35, 52 affinis 'Qtcc. 113
pauciflora Bl. 683, 684, 688 Santalales 5. 34 andajensis Becc. 117
philippinensis Robins. 687, Sapindaceae 639, 679, 680, 681 arfakiana Becc. 109, 112,
688 Sapindales 163,458.639,681 113, 115*. 117
racemosa Chen 684, 689 Sapindus microcarpus W. & A. aster ias Ridl. 116
ssp. kinabaluensis 709 atroviolacea Schltr 118
v.d.Water 683, 689 Sapotaceae453, 593 australasicaWQmsX. 117
ssp.racemosa 689 Sarcandra Gardner 123. 125. brachystyla Schltr 1 16
Elmer 688
reticulata 126. 128, 129, 134, 144 buruensis 112
sumatrana Bl. 682, 683, 689 glabra (Thunb.) Nakai 124, clemensae Hemsl. 117
Sabiaceae 163, 166,679-715 127, 128, 134, 135*, conferta J.J. Smith 115
Salacia banlettii RidJ. 25 136 consimilis Bl. 113, 120
Salicaceae 5: 107-110 ssp. brachystachys (Bl.) corallophyton K.Sch. &
Salmonea 486 Verdcourt 136 Schltr 113, 119
Salomonia Lour. 455-457, var. brachystachys 136 var. gracilis Giesen 1 19
459. 486 var. melanocarpa (Ridl.) comiculata Becc. 112, 115,
sect. Epirixanthes (Bl.) Benn. Verdcourt 136 116, 117
488 hainanensis (Pei) Swamy & corniculata (non Becc.) Went
angulata Griff. 488 Bailey 136 117
aphylla Griff. 490 glabra auctt. mult. 136 crinita Becc. 117
arnottiana Miq. 488 Sarcodiscus Griff. 287 decipiens Backer 113
canarana Miq. 488 chloranthiformis Griff. 298 densiflora Schltr 112, 113,
cantoniensis Lour. 487 Sarcopetalum F.v.M. 157-161, 118, 119*
cantoniensis auct. 488 167, 169. 172, 224 dolichostyla Schlti 119
cavalereriei Lev. 488 harveyanum F.v.M. 159, erubescens (Champ.) Miers
ciliata (L.) DC. 487 224. 228* 114
1989] Index to scientific plant names 743

(Sciaphila) (Scleropynim) Sonneratiaceae 4: 280-289,

flexuosa Giesen 118 pentandrum (Dennst.) 513; 5: 557; 6: 973
galiensis Schllr 115 Mabberley 52 Sparganiaceae 4: 233-234;
hermaphrodila Schltr 1 13 wallichianum (W. & A.) Am. 10: 718
hydrophila Schllr 120 52 Sparganium fallax Graebn. 718
inaequalis Schltr 1 18 Scorodocarpus Becc. 1-6, 14 simplex Huds.
inornaia Fetch 116 map, 15 f. simplex 718

long ipes Schltr 118 bomeensis (Baill.) Becc. 5. subglobosum Morong 718
maboroensis Schltr 1 16 15, 16* Spermabolus T. & B. 605
macra K.Sch.&Uut. 120 Scyphosiegia Stapf 326 fruticosa T. & B. 605
macro K.Sch. & Schltr Scyphostegiaceae 5: 297- 299; Sphaerocarya leprosa Dalz. 22
116 6: 967; 7: 830; 10: 326 Sphenocleaceae 4: 27-28
macra Schltr 120 Sccuridaca L. 455-459, 483 Sphenostemon Baill. 145, 146,
maculata Micrs 112, 113 atro-violacea Elmer 483, 484 326, 333
major Bccc. 116 bracieata Benn. 484 sect. Apetalae (Stecn.)
micranthera Giesen 112, 121 var. papuana F.v.M. 485 Steen. 147
mindanaensis G'xcscn 120 complicata auct. 484 sect. Sphenostemon 147
minuta Schltr 113 corymbosa Turcz. 484 scr. Apetalae Stecn. 147
moniicola K.Sch. & Schltr cumingii Hassk. 484 ser. Sphenostemon 146, 147
116 ecristata Kassau 483, 485* arfakensis (Gibbs) Steen. &
multiflora Giesen 112, 113, var. niiida Kassau 485 Erdtman 147, 149
120 inappendiculata Hassk. 483 lobosporus (F.v.M.) L.S.
nana BI. 113, 116, 117 ssp. corymbosa (Turcz.) Smith 147, 149
neo-caledonica Schltr 1 15 Meijden 484 oppositifolius Hiirl. 146
nu/an.y Giesen 118 ssp. inappendiculata 484 pachycladus 146
oligochaeie Schltr 1 15 paniculaia Roxb. 484 papuanus (Laul.) Stecn. &
papillosa Becc. 121 philippincnsis Chodat 483, Erdtman 147, 148*, 149
papuana Becc. 116 484 pauciflorum (A.C. Smith)
pilulifera Schllr 116 scandens Ham. ex Benih. Steen. & Erdtman 147,
pumila Giesen 1 14 484 148
purpurea 109 lavoyana Wall, ex Benn. 484 Sphenostemonaceae 145-149,
quadribuUifera J.J.Sm. 112, volubilis auct. 484 326, 333
114 yaoshannensis Hao 484 Siachycarpus (Endl.) Ticgh. 352,
reficxa Schlu 118 Selwynia F.v.M. 218 384
sccundiflora Thw. ex Bih. laurina F.v.M. 219 sect. Sundacarpus (Buchholz

112, 115*. 116, 120 Semeiocardium Zoll. 539 & Gray) Gausscn 385
slcmmermannii Fosb. & hamillonii Hassk. 463 amara Gausscn 387
Sachet 120 Semeiocardium (non Zoll.) Stackhousiaceae 4: 35-36
suhhermaphrodita J.J. Sm. Hassk. 459, 462 Staphyleaceae 6: 49-59
114 Shepherdia Null. 151 Stcganthcra Perkins 255-263,
sumairana Becc. 1 16 Simaroubaceae 6: 193-226, 288, 306, 321
tcnclla Bl. 112. 113, 115* 968; 10: 621, 622, 718 alpina Perkins 319
var. robusta Giesen 1 14 Sinapis 541 atepala Perkins 319
var. voigiii Giesen 1 14 alba L. 545 austral iana C.T.White 309.
torricellensis K.Sch. & Schllr juncea L. 546 318
114 imoriana DC. 546 brassii (A.C. Smith) Kaneh.
trichopoda Schltr 1 8 Siparuna 259 & Hatus. 315
tuber culata Giesen 112, 118 gilgiana 261 buergersiana Perkins 316
valida Giesen 1 18 guyanensis 261 chimbucnsis Philip.son 309,
versteegiana Went 1 16 Siparunaccae 261 318
viliensis A.C. Smith 118 Siph()n(Kl()n celasuincus Grill. crispula Perkins 316
wariana (Schltr) 539 cyclopcnsis Philipson 308.
Mccrcndonk 112, 117 Sisymbrium 541 543 , 310
werneri Schltr 1 16 amphibujm L. denlaliJ (Val.) Kanch. &
wmklcri Schlu 112. 113. var. pcduslre L. 557 Hatus. 308, 310
120 micranihum Rolh 560 clliptica A.C. Smith 278
Sclcropyrum 33 naslurtium-aqualirum L. 555 fasciculata Philipson 307*.
aurantiacum (Laut.& K.Sch.) Skaphium Miq. 493 308. 309
Pilgcr 52 anceatuin Miq. 527 fengcriana F'crkins 315
744 Flora Malesiana [ser. I, vol. 104

(Steganthcra) (Stcphania) (Stcphania)

forbesii Perkins 316 borneensis Yamamoto montana Diels 159, 243-

hcntyi Philipson 308, 309 249 245, 250, 253
hirsuta (Warb.) Perkins 262, capitata (Bl.) Spreng. 159, ncoguineensis Kundu & Guha
300, 308.309, 314*, 160, 161, 165, 167,236, 253
315, 318 243-245, 252*, 253 obvia Miers 252
hospilans (Becc.) Kaneh. & calosepala Diels 249, 250 prapaiensis Yamamoto 248
Hatus. 257, 258, 308- calosepala (non Diels) Merr. psilophylla (Presl) Forman

310 311, 313*. 319 251 243-245, 251

A.C. Smith 256,
ilicifolia 250
cauliflora Becc. 249, ramosii Diels 249
296, 309, 317*, 318 cincinnans K. Sch. 203 ramuliflora Miers 249
insculpta Perkins 308, 309, concinna Miers 247 reticulata Forman 244, 245,
319 corymbosa (Bl.) Walp. 243- 249
insignis Perkins 312 245, 249, 250 rotunda (non Lour.) Miq.
ledcrmannii (Perkins) Kaneh. corymbosa (non Bl.) Turcz. 248
& Hatus. 257, 308, 319 251 rotundifolia 248
moszkowskii (Perkins) dictyoneura Diels 160, 167, salomonum Diels 244, 245,
Kaneh. & Hatus. 257, 243, 244, 253 250
308, 309, 311 discolor (Bl.) Spreng. 245 tomentosa (Bl.) Spreng.

myrtifolia (A.C. Smith) var. hernandiifolia (Willd.) 229

Philipson 309, 313 Boerl. 245 truncata Yamamoto 252
oblongiflora Perkins 315, exigua Miers 247 venenosa 248
316 florulenta Becc. 251 venosa (Bl.) Spreng. 243-
oblongifolia 315 formanii Kundu & Guha 250, 245, 248
odontophylla Perkins 319 251 zippeliana Miq. 243-245,
oligantha (Perkins) Kaneh. & forsteri (DC.) A. Gray 247 251
Hatus. 308, 309, 315 glaucescens (Decne) Walp. Strakaea Presl 65
oligocarpella Kaneh. & 245 melastomaefolia Presl 78
Hatus. 291 grandiflora Forman 244, 248 Strombosia Bl. 1-6, 19
parvifolia (Perkins) Kaneh. & hallieri Diels 247 ceylanica Gardn. 20*, 21, 22,
Hatus. 309, 315 hernandiifolia (Willd.) Walp. 23 map
psychotrioides Perkins 319 245, 247 var. lucida (T. & B. ex
pycnoneura Perkins 319 var. discolor (Bl.) Miq. Valet.) Hochr. 22
riparia Kaneh. & Hatus. 316 245 var.membranacea (Bl.)

royenii Philipson 257, 308, var. genuina (Bl.) Miq. Hochr. 22

309, 310 245 var. sessilis Hochr. 22
salomonensis (Hemsl.) var. glabra (Bl.) Miq. 245 dubia Vidal 22
Philipson 308, 309, 311, hernandiifolia [non (Willd.) elmeri Salvosa 22
312* Walp.] Koord. 249 javanica Bl. 5, 20*, 21 map
schlechleri Perkins 278 hernandiifolia [non (Willd.) sumatrana Val. 21
var.
schumanniana Perkins 3 1 Walp.] Ridl. 248 javanica auct. 22
suberoso-alata Kosterm. 311 hernandiifolia [non (Willd.) latifoliaSlap{22,23
symplocoides Perkins 319 Walp.] Vidal 251 lucida T. & B. 22
thyrsiflora Perkins 315, 316 japonica (Thunb.) Miers 159, maingayi (Mast.) Whitm. 22
lorrlcellensis Perkins 316 165, 243-245, 245, membranacea (Bl.) Valet. 22
villosa Kaneh. & Hatus. 246*, 247 minor Elmer ex Merr. 22
284 var. discolor (Bl.) Forman mulliflora King 22
warburgii Perkins 315 247, 248 philippinensis (Baill.) Rolfe
Stcllatopollisbarghoomii 256 var. japonica 247 21, 22
Sicmorwaws fruiescens Bl. 25 var. timoriensis (DC.) philippinensis [non (Baill.)
membranaceus Bl. 22 Forman 247 Rolfe] Lam & Holthuis 29
Stcphania Lour. 157, 160-162, japonica [non (Thunb.) rapaneoides S. Moore 22
165, 167, 169, 171, 243 Miers] Miers 25 rotundifolia King 22
sect. Eustephania Diels 243, longifolia Becc. 252 Strombosiopsis 3, 4
244 mcnadonensis Diels ex Slropha Noronha 129
sect.Thamnoihyrsa Diels Koord.-Schum. 249 Stylidiaceae 4: 529-532;
243, 244 250
merrillii Diels 249, 5: 564; 6: 976
acuminaiissima (Bl.) Spreng. moluccana Forman 244, 245, Styracaceae 4: 49-56; 6:
252 250 976; 9: 568
1989] Index to scientific plant names 745

Symplocaceae 8: 205-274; {Talaunw) Taxus L. 337, 342, 347, 348

9:569; 10: 718 longifolia (Bl.) Ridl. 582 map
Symplocos Jacq. luzoniensis Warb. ex Perkins baccata (non L.) Mast. 349
subg. Hopea Clarke 718 585 ssp. cuspidata
ambangensis Noot. 719 macrophylla 583
Bl. var. chinensis Pilger
cochinchinensis 719 magna Agostini 586 349
ssp. laurina 719 megalophylla Merr. 586 ssp. wallichiana (non
cosialifrucia Noot. 719 miqueliana Dandy 582 Zucc.) Pilger 349
hcrzogii 718 mutabilis Bl. 582 var. sinensis Henry 349
iliaspaiensis Noot. 719 var. acuminata Bl. 582 celebica (Wall.) Li 350
laeteviridis 719 var. acuminaiissima chinensis (Pilger) Rehd. 350
maliliensis 719 T. & B. 583 cuspidata (non Sieb.& Zucc.)
ophircnsis var. brevifolia T. & B. Kaneh. 350
ssp. cumingiana 719 583 cuspidata var. chinensis (Pil-
rayaeNoot. 718, 719 var. laiifolia T. & B. 583 ger) Rehd. & Wilson 350
riangensis Noot. 718, 719 var. leiocarpa T. & B. mairei (Lemee & Lev.) Hu &
vcrtJciliifolia 718 583 Liu 350
Synandrium exsertum 243 var. longifolia Bl.582 speciosa Florin 350
inclusum 243 var. sciagraphia 582 sumatrana (Miq.) de Laub.
var. splendens Bl. 582 349, 350*, 351 map
mutabilis auct. 585 wallichiana (non Zucc.)
Taccaceae 7: 806-819 oblanceolata Ridl. 585, 586 Foxw. 349
Talauma Juss. 563, 568 oblongata Merr. 585 wallichiana var. chinensis
sect. Aromadendron Nliq. obovaia Korth. 585 (Pilger) Horin 350
569, 576 obovata auct. 586 yunnanensis Cheng 350
angatensis (Blco) Vidal 585 oreadum Diels 583 Temstroemiaceae 327
aihlianiha Dandy 583 avails Miq. 596 Tetanosia Rich, ex M. Roemer
beccarii Ridl. 586 papuana Schltr 596 46
betongensis Craib 585 peninsular is Dandy 583 olacioides (W. & A.)
binluluensis Agostini 577 persuaveolens Dandy 587 M. Roemer 47
horneensis Merr. 583 plumierii (Schwartz) A. DC. Tctrastylidium 3,4
candollei auct. 585 568 Tetrasynandra 284 map
candollii Bl. 569,581,582 pubescens Merr. 596 Thalamia Spr. 356
var. laiifolia Bl. 582 pumita Bl. 582 Thalictrum 165
elegans (Bl.) Miq. 577 pumila auct. 583 Thcaceae 145
var. glauca (Korth.) rabiana Craib Thealcs 145
Parmcnt. 577 var. villosa (Miq.) Thlaspi541,543
elegans auct. 583 Parment. 588 Thottea Rottb. 53-55, 57-59,
elmeri Merr. ex Soderbcrg reticulata Merr. 583 61-64, 65
587 rubra Miq. 582 beccarii Ding Hou 67, 74
forbesii King 582 rumphii Bl. 582 bomeensis Val. 65, 66, 73
giganiifolia Miq. 586 sarawakensis Agostini 588 celebica Ding Hou 67, 78,
giganiifolia auct. 585 sclerophylla Dandy 585 80*
gilingensis Elmer 583 sebassa Miq. ex Dandy 582 corymbosa (Griff.) Ding Hou
var. glabra Dandy 583 singapurensis Ridl. 586 56*, 57, 59, 62, 67, 78,
var. roiundaia Dandy soembensis Dandy 583 82*
583 sumulrana Agostini 583 curvisemcn Ding Hou 61, 66,
glaucum (Korth.) Miq. 577 undulatifotia Agostini 583 73
gracilior Dandy 583 villariana Rolfc 583, 585 depcndens (Planch.) Klotzsch
grandiflora Merr. 585 villarii 583 56*, 57, 62, 67, 75
injlata Parment. 582 villosa Miq. 588 dinghoui 62
intonsa Dandy 588 f. celebica Miq. 603 grandillora Rottb. 59, 62. 66,
javanica Parment. 583 vrieseana Miq. 569, 596 68*, 69
kunstlen King 582 Tambouri.ssa 257, 258 hirsutaR\a\.8\
kuleinensis Agostini 586 ficus(Tul.) A.DC. 326 macTdnlha (Boerl.) Ding Hou
lanigeraH\^A.&.l\\.58V, Taxaceae 343, 346, 347-35 1 .
60*, 62. 65, 67. 81
lanigera auct. 586 3.54 macrophylla Bccc. 5'), 66, 71
levissinui Dandy 585 Taxaics .347 muluensis Ding Hou 56*,
Uliifcra {L.)O.Y.. 582 laxodiaccac 338, 343, 354 57, 67. 76*, 78
746 Flora Malesiana [ser. I, vol. 104

(Thotlca) (Tinospora) (Tinospora)

parviflora Ridl. 59, 62, 66, cclcbica Dicls 190, 193, 194 uliginosa Micrs 199

67, 80* cordifolia 190, 197 uliginosa (non Micrs) Hk.f.

paucifida Ding Hou 59, 62, coriacea (Bl.) Bcunndcex & Th. 198
67, 72*, 73, 81 Hcync 199 Trapaceae 4: 43-44; 6: 982
pcniiilobala Ding Hou 57, crispa(L.)Hk.f. &
Th. 160, Trcmandraccac 458
67. 76*, 77 165, 166, 190, 191, 194, Triccrcandra A. Gray 128
philippincnsis Quis. 67, 75, 195, 196*, 197, 200 Trichocarya Miq. 645, 648
82* cmpa [non(L.) Hk.f. &Th.] splendens 645
rcniloba Ding Hou 60*. 67, Dicls 199 Triclisia 165

77 curtisii Ridl. 253 Trigoniaceae 4: 58-60; 10:

rhizantha Bccc. 67, 75 dcniata 190 458

robusta Siccn. 66, 70*, 71 dissitiflora (Laut. & K.Sch.) Trimcnia Sccmann 145, 255,
siliquosa (Lamk) Ding Hou Dicls 159, 190, 191. 261, 326. 327, 329 map
81 192*. 197 arfakensis Gibbs 330, 331
sp. Ding Hou 83 glabra (Burm.f.) Mcrr. 159, bougainvilleensis (Rodcn-
sU'aalmanii Ding Hou 66, 69 165, 186*. 191. 192*, burg) A.C. Smith 332
sumau-ana (Mcrr.) Ding Hou 194, 195. 199. 200 grandifolia Warb. 333
67, 80*. 81 glandulosa Mcrr. 190. 194 macrura (Gilg & Schllr)
lomcntosa (Bl.) Ding Hou hasiala Elmer 193 Philipson 329 map, 330,
54, 57, 59, 62, 65, 79 havilandii Dicls 193,210 331, 332
iricornis Maingay 62. 67, hirsuia (Bccc.) Forman 190, moorei (Oliv. in Bth.)

74, 80* 193 Philipson 328, 329 map,

iriscrialis Ding Hou 55, 59, homoscpala Dicls 190, 200 330
66, 69 macrocarpa Dicls 191, 192*, myricoides Gilg & Schltr
Thuja 444 198, 199 330, 331
javanica Burm.f. 453 megalobotrys Laut. & ncocalcdonica 328, 329 map.
papuana (F.v.M.) Voss 444 K.Sch. 187 330
Thujoidcae 444 mcrrilliana Dicls 159, 190, papuana Ridl. 328*, 329
Thuya javanica 453 191, 191 map, 330, 331*
Thymelaeaceae 349-365;
4: minuliflora K. Sch. & Laut. weinmanniifolia Seem. 328,
6: 1-48,976; 7: 830; 227 329 map, 330-332
10: 151 negrotica Dicls 1 93 ssp. bougainvilleensis
Thymclacalcs 151 var. moniicola Yamamoto Rodenb. 328, 330, 332
Tiliacora Colcbr. 157, 160-168. 193 Trimeniaceae 145. 255. 261.
170, 172, 185 peekelii Dicls 197 327-333
triandra (Colcbr.) Dicls 162, polygonoides Dicls 200 Trimeria grandifolia (Hochst.)
166, 185. 187, 208* pseudo-crispa Bocrl. 199 Warb. 333
Tinomiscium Micrs 157-162, reticulata 199 Trimeriza Lindl. 65
166-170. 204 rumphii Bocrl. 194. 195. Trisiichocalyx F.v.M. 217
arfakianum (Bccc.) Dicls 200 Triuridaceae 109-121. 110
197 •
sagitlata 190 map, 720
elasticwn Bccc. 205 sinensis (Lour.) Mcrr. 190, Triuridalcs 109
javanicum Micrs 205 200 Triuris 110, 112
molle Dicls 205 smilacina Benth. 191, 196* Trochiscus Gilli 555
pctiolare Hk.f. & Th. 159, subcordata (Miq.) Dicls 159. macrocarpus Gilli 559
205. 206* 191, 192*, 200 Tropacolaccac 639
phlUppinense Dicls 205 sumau-ana (Schcff.) Bccc. Tsjeru-caniram Rhecde 49
phytocrenoides Kur/. ex 190, 191, 198, 199 Tsjerucaniram 49
T. & B. 205 var. hanadac Yamamoto Tsjerucanirum 49
pyrrhoboiryum Miq. 205 198 Tsoongiodendron 563. 564. 567,
Tinospora Micrs 19, 158-171, icijsmannii Bocrl. 191, 192*. 593
188 199 Tsuga 338, 343
andamanica Dicls 199 tinosporoidcs (F.v.M.) mairei Lcmee & L6v. 350
angusta 190 Forman 190, 192* Tuba baccifera Rumph. 213
arfakiana Bccc. 190, 191, trilobata Dicls 159, 190, flava Rumph. 210
197 191, 202 Turneraceae 4: 235—238
bacnzigeri Forman 159, 160, tuberculata (Lamk) Beumee Typhaceae 4: 242-244; 6:
190. 191. 195, 196*, 197 ex Hcync 194 982
1989] Index 10 scientific plant names 747

Ulmaceae 31-76, 551;

8: (Xanthophyllum) (Xanihophyllum)
9: 569; 29
10: subg. Coriaceum Meijden contracuim Meijden 499,
Umbelliferae 4: 113-140, 494, 528 532
595; 5; 555; 6: 983: subg. Exserium Meijden 494, cordaium Korth. ex Miq. 515
7: 830: 9: 569 533 f aequale Chodat 5 1
.

Uroboirya Stapf 31-35, 41 subg. Triadelphum Meijden curtisii King 514

sccL Lepionuroides Hiepko 494, 530 densiflorum Chodat 521, 522
42 map subg. Xanthophyllum 494, discolor Chodat 495, 520
sect. Urobotr>'a 42 500 520
ssp. discolor
floresensis Hiepko 42 map, secL Eystathes (Lour.) macranihum Meijden
ssp.
43,48 Meijden 500, 507 520
laiisquama (Gagn.) Hiepko subsect. Eystathes 507, ecarinatum Chodat 499, 539
32, 43, 42 map 526 ellipticum Korth. ex Miq.
longipes (Gagn.) Hiepko subsect. Jakkia (Bl.) 499, 530, 531*
42 map Meijden 507, 508 er>'lhrosiachyum Gagnep.
parv'ifloraHiepko 41*, 42 sect. Xanthophyllum 500 497, 511, 523
map, 43 acuminaiissimum Miq. 515 eiu-hynchum Miq. 497, 524.
siamensis Hiepko 34, 42 adenopodum Miq. 501 525
map, 43, 46 adenoius Miq. 497, 498, 511, ssp. eurhynchum 525
Uvaria-leaved Magnolia Meijer 515 maingayi (Hk. f. ex
ssp.
574 var.adenotus 516 A.W. Benn.) Meijden
var.Imeare Meijden 516 525
affine Korth. ex Miq. 498, excelsum Bl. 503
Vaccinium angulaium J.J. Sm. 500, 501, 503 var. affine (Miq.) Boerl.
716 B excelsa K. & V. 500 503
commuialum Mabberley & a genuina K. & V. 503 excelsum (Bl.) Miq. 500
Sleumer 716 var. adenopodum (Miq.) ferrugineum Meijden 498,
Valerianaceae 4; 253-254 K. & V. 501 503
Vallisneria 112 affine (non Miq.) K.Sch. &; flavescens (non Roxb.)
VanhalUa Schuli. 65 Hollr. 537 F.-Vill. 514
omenlosa J. A. &. J.H. affine (non Miq.) Koord. 5 1 flavescens Roxb. 498, 500,
Schultes 79 affine (non Miq.) Ridl. 528 5(W, 528
Rumph. 1
V'idara liuorea amoenum Chodat 499, 533 var. virens (Roxb.)
Okamoio 720
Viola rheophila amoenum (non Chodat) Keith A.W. Benn. 528
Violaceae 7: 179-212,831; 535 flavovirens Elmer 520
10: 720 ancolanum Miq. 498, 511 flavum Ridl. 505, 507
Violaria Post & O.K. 568, 569 f. angustifolia Miq. 5 1 floriferum Elmer 501
Viscaceae 353 angustigcmma Meijden 496, forbesii Baker 523
Vochysiaceae 458 519 glabrescens Ridl. 508
arsaiii C.E.C. Fischer 516 glandulosum Merr. 501
bcccarianum Chodat 495, glaucescens Miq. 539
Wcinmannia 123 522 glaucum Wall, ex Hassk. 527
Weltingionia arnottiana Mcisn. bomeense Miq. 495. 508 gracile Chodat 5 1

709 bractealum Chodat 518*. griffithiiHk.f.ex A.W. Benn.

Wilkiea F.v.M. 255-257, 262, 496, 519 496, 513, 514
263. 282, 288 brevipcs Meijden 495. 535, ssp. angustifolium (Ng)
forcmanii Philipson 282, 536* Meijdcn 513
283* brigittae Mcijdcn 496, 498, ssp. crecium Meijden 513.
hucgciiana 288 511, 512* 514
macrophylia 288 bullatum King 498. 501 var. angustifolium Ng 5 1

WitLslcmia F.v.M. 335 cclcbicum Meijdcn 499, 532 var. curiisu (King) Ng
papuana (Stccn.) Siccn. 336* ccraccifohum Mcijdcn 497. 513.515
WorcesierianihusMen. 29 517 var. montanum Ng 5 1 3,
chartaccum Mcijdcn 499, 514
535, 539 griffiihii (non A.W. Benn.)
Xanihophyllum Roxb. 455-459. ciirifoltum Chodat 530 Rolfc5I4
493 clovis (Slccn. ex Mcijdcn) hchecarpum Chodat 539
subg. Brunophyllum Mcijdcn Mcijdcn 496. 517 hctcrophyllum Mcijdcn 496,
494. 535 cockbumii Mcijdcn 494, 502 519
748 Flora Malesiana [ser. I, vol. 104

(Xanthophyllum) (Xanthophyllum) (Xanthophyllum)

heteropleurum Chodat 505, pallidum Ridl. 501 stipitatum A.W. Benn. 497,
507 paniculatum Miq. 514 499, 535
hildebrandii Meijden 498, papuanum Whitm. ex var. borneense Chodat 535

532 Meijden 457, 499, 537, var. glabrum Meijden 535

hookerianum King 514 538* var. nitidum Chodat 533
502
hosei Ridl. 498, parvifolium Meijden 496, var.pachyphyllum Chodat
hypoleucum Merr. 520 510 533
impressum Meijden 496, parvum Chodat 5 1 var. stipitatum 535
513 pauciflorum Meijden 496, subcoriaceum (Chodat)
inccrium (Bl.) Meijden 496, 509 Meijden 498, 509
515 pedicel latum Meijden 495, suberosum C.T.White 499,
insigne A.W. Benn. 536 522 533, 534*
kalimantanwn Meijden 539 penibukanense Heine 495, subglobosum Elmer 539
kingii Chodat 530 521 var. longifolium Elmer
korthalsianum Miq. 496, petiolatum Meijden 496, 517 539
520 philippinense Chodat 498, sulfureum 502
kunslleri King 514 510 sulphureum King 494, 502
laeve Meijden 499, 524 pseudoadenotus Meijden 495, sumatranum Miq. 51
lanceatum (Miq.) J.J. Sm. 521 tardicrescens Meijden 498,
494, 496, 527 pseudostipulaceum Merr. 5 1 510
lanceolatum 527 pseudoslipulaceum (non tenue Chodat 498, 508
latcriflorum Miq. 497, 527 Merr.) Meijer 519 tenuipetalum Meijden 498,
loheri Merr. 501 puberulum Ridl. 525 512
longifolium (Bl.) Dietr. 514 pubescens Meijden 495, trichocladum Chodat 495,
macranlhum Chodat ex Elmer 504 522
520 pulchrum King 495, 521, velutinum Chodat 495, 505
macrophyllum Baker 498, 522 venosum King 497, 525
507 ssp. pulchrum 521 verrucosum Chodat 524,
maingayi Hk. f ex
. ssp. siapfii (Chodat) 525
A.W. Benn. 524, 525 Meijden 522 vircns Roxb. 497, 528
malayanum Meijden 495, purpureum Ridl. 495, 522 virescens 528
526 ramiflorum Meijden 494, vitcllinum (Bl.) Dietr. 497,
microcarpum Chodat 527 495, 529*, 530 498, 514
molle Ridl. 522 reflexum Meijden 496, 519 ex
var. clovis Steen.
montanum Meijden 499, 532 resupinatum Meijden 498, Meijden 517
moniicolum Meijden 496, 504 wrayi King 497, 525
514 reticulatum Chodat 495, Ximenia L. 1-6, 10
muliiramosum Elmer 501 522 americana L. 2, 5, 11
ncglectum Meijden 497, 498, retinerve Meijden 497, 524 var. americana 11, 12*
509 robustum Chodat 515 borneensis Baill. 15
ngii Meijden 496, 526 var. elmeri Chodat 5 1 loranlhifolia Span. 1

nigricans Meijden 495, 508 rufum Benn. 494, 505, olacioides W.

A. 47 &
novoguineense Meijden 496, 506* Xylosma leprusipes Clos
526 sarawakense Chodat 503 715
obscurum A.W. Benn. 499, saraH'akensis 503 Xyridaceae 4:366 — 376,
536 schizocarpon Chodat 495, 598; 5: 557; 9: 571
ovatifolium Chodat 498, 508 504
palawancnse Elmer 497, 516 scortechinii King 536
palembanicum Miq. 5 1 spec. 504 Zanonia indica L. 253
palembanicum (non Miq.) spec. Anderson 530 Zizyphus liitorea Teysm. ex
Keith 509 spec. £ Ng 539 Hassk. 1

palembanicum (non Miq.) stapferi 521 Zonalapollenites 338

King 524. 525 5/apyz/ Chodat 521, 522 Zygophyllaceae 4: 64
 New York Botanical Garden Librar

3 5185 00275 8033

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 FLORA MALESIANA
under the Auspices of the Centre for Research
and Development in Biology, Bogor, Indonesia,
and the Rijksherbarium, Leiden, Holland,
executed by Foundation Flora Malesiana

Scientific Communications
concerning the Flora Malesiana should be addressed to
the General Editor, Dr. W.J.J.O. de Wilde
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