FLORA
MALESIANA
SERIES I - SpERMATOPHYTA
Flowering Plants
Vol. 10, part 4
DEDICATION TO BLUME
ADDENDA
INDEX TO REVISED FAMILIES
Aceraceae
4
•:3,
592
Dichapctalaceae
5:
305;
6:
941
Opiliaccae
10:
31
Aclinidiaceac s.s.
4:
37
Dillcniaccae
4:
141;
7:
824
Oxalidaceae
7: ]
151,
829
Aixoaccae
4:
267
Dioscoreaceae
4:
293
Papaveraceae
5:
114
Alismaiaccae 5:
317;
6:
915
Dipsacaceae
4:
290
Passifloraceae
7:
405
Alseuosmiaceae
10:
335
Dipterocaipaceae
9:
237
Pedaliaceae
4:
216;
7:
829
Amaranihaceae
Droscraccae
Pentaphragm atace ac
4:
517
4: 69, 593; 6:
915;
8:
549
4: 377;
5:
557;
9:
562
Pcniaphylacaccae
5:
121
Anacardiaceae
8:
395
Elaeagnaceae
10:
151
Philydraceae
4:
5
Ancistrocladaceae
4:
8
Elatinaceae
4:
203
Phyiolaccaceae
4:
228
Aponogctonaceae 4
I: 11;
7:
213
Epacridaceae
6:
422
Pinaceae
10:
447
Araliaceae-I
9
: 1,
553
Ericaceae
6: 469,
943
Pittosporaccac
6:
960;
5:
345
Araucariaceae
10:
419
8: 549;
9:
562;
10:
716
Plumbaginaceae
4:
107
Arislolochiaceae
10:
53
Erythroxylaceae
5:
543;
8:
549
Podocarpaceae
10:
351
Balanophoraceae
Fagaceae
7:
265;
9:
563
Podostemaceae
4
i:65;
6:
963
7:
783;
8:
549
Hacourtiaceae
5:
1,
565
Polemoniaceae
4:
195
Basellaceae
5:
300
6: 943;
7:
827;
9:
563
Polygalaceae
10:
455
Bat(id)aceae
5:
414
Flagellaceae
4:
245;
9:
564
Pontederiaceae
4:
255
Betulaccae 5:
207;
6:
917
Geraniaceae
6:
445;
9:
565
Portulacaceae
7:
121
Bignoniaceae 8:
114;
9:
554
Gnciaceae
4:
336;
6:
944
Primulaceae
6:
173
Bixaceae s.s.
4:
239
Gonystylaceae
4:
349
Proteaceae
5:
147
Burmanniaceae
Goodcniaceae
5: :
335,
567
Punicaceae
4:
226
4: 13
,592:
; 9:
554
6: 949;
7:
827;
9:
566
Rcsiionaceae
5:
416
Burseraceae
5: 209,
567
Hacmodoraceae
5:
111;
10:
717
Rhizophoraceae
6:917; 7:
820;
9:
555
Haloragaceae
7::
?39,
828
5: 429;
6:
965;
8:
550
Butomaceae
5: 1
118,
556
Hamamelidaceae
5:
363
Sabiaccae
10:
351
Byblidaceae
7:
135
Hippocrateaceae
6:
389
SaUcaceae
5:
107
Calliirichaceae
4:
251
Hydrocharitaceae
5:
381
Salvadoraceae
4:
224
Campanulaceae
6: 952; 7: 828;
9:
:566;
10:
717
Sarcosperma(ta)ccae
4:
32
6: 107,
928;
9:
556
Hydrophyllaceae
4:
207
Saururaceae
4:
47
Cannab(in)aceac
4:
222
Hypcriaceae
8: 1;
10:
717
Scyphosiegiaceac 5
:29';
'; 6:
967
Cappar(id)accae 6:61;
7:
822
Icacinaceae
7: 1;
9:
566
Simaroubaceae
6:
193,
968
Caprifoliaceae
Iiidaccae
i
1:77;
10:
717
Sonneratiaceae
4: 175; 6
:928
; 9:
556
Ixonanihaceae
10:
621
4: 280,
513;
6:
973
Cardioptcridaceae
7:
93
Juglandaccac
6:
143
Sparganiaceae
4:
233;
10:
718
Cclastraceae 6: 227, :
389,
930
Juncaccae
4:
:210;
9:
566
Sphcnocleaceae
4:
27
Ccntrolepidaceae
5:
421
Juncaginaceae
4:
57
Sphcnostemonaceae
10:
145
Ceratophyllaceae
4:
41
Labiatae
8:
:301;
9:
566
Stackhousiaceae
4:
35
Chcnopodiaccae
Lccaccae
7:
755
Staphylcaceac
6:
49
4:99,594; 6:
932;
9:
557
Lemnaccae
7:
219
Stylidiaccae
4:
529;
6:
976
Chloranthaceae
10:
123
Lcntibulariaceae
8:
275
Styracaceae
4:
49;
9:
568
Chr>'sobalanaceae
10:
635
Liliaccae-I
9:
189
Symplocaceae
aeihraceae
7:
139
Linaccae
10:
607
8: 205;
9:
569;
10:
718
Cochlospermaceae
4:
61
Loganiaccae
Taccaceae
7:
806
Combrctaccae
6: 293,
953;
9:
567
Taxaceac
10:
347
4:533; 5:
564;
6:
932
Lophopyxidaceae
7:
89
Thymelaeaceae
Conifc rales
10:
337
Magnoliaccae
10:
561
4:349; 6:
: 1,
976;
7:
830
Connaraceae
Malpighiaccae
5:
125
Trapaccae
4:
43
5:495; 6:
933;
9:
557
Manyniaceae
4:
216
Trigoniaccae
4:
59
Convolvulaceac
4: :
388,
599
Menispcrmaccae
10:
157
Trimcniaceae
10:
327
5: 558; 6: 936; 7
:823
; 9:
558
Monimiaceae
10:
255
Triuridaccae
10:
109
Comaccae
8:
85
Moringaccae
4:
45
Tumeraceac
4:
235
Corynocarpaceae 4
:262
; 5:
557
Myoporaceae
4:
265
Typhaccae
4:
242;
6:
982
Crassulaceae 4
: 197
; 9:
558
Myricaccae
4:
276
Ulmaceae
8:
31
Crucifcrae
10:
541
Najadaceac
6:
157
Umbcllifcrae
4:
113,
595
Cryptcroniaceae
8:
187
Nyctaginaccae
6:
450
5:555; 6:983;
7
:830
; 9:
569
Clcnolophonaceae
10:
629
Nyssaccac
4:
29
Valcrianaccac
4:
253
Cupressaceae
10:
442
Ochnaccac
7:
97
Violaceae 7: 179,
831;
10:
720
Cypcraccae 7:
435;
9:
107
Olacaccac
10:
21,
717
Xyridaccae 4: 366,
598;
9:
571
Daiiscaccae
4:
382
Onagraccac
8:
98
Zygophyllaceae
4:
64
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 I SYNONYMY I AND DISTRIBUTION /
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 9 K 4 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
the publishing programmes of
D. Reidel, Martinus Nijhoff, Dr W. Junk en MTP Press.
Sold and distributed in the U.S.A. and Canada
by Kluwer Academic Publishers,
101 Philip Drive, Norwell, MA 02061, U.S.A.
In all other countries, sold and distributed
by Kluwer Academic Publishers Group,
P.O. Box 322, 3300 AH Dordrecht, The Netherlands.
Publication dates
Part 1 20 Jan. 1984
Part 2 21 May 1986
Part 3 19 Oct. 1986
Part 4 3 Aug. 1989
All Rights Reserved
© 1989 Kluwer Academic Publishers
No part of the material protected by this copyright notice may be reproduced or utihzed in any form or by
any means, electronic or mechanical, including photocopying, recording, or by any information storage and
retrieval system, without written permission from the copyright owner.
Printed in the Netherlands
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 regret biographers frequently do not give sufficient attention to personality and motiva-
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 before his death in May 1976 the author of this Dedication and former Editor of Flora Male-
siana, Professor C.G.G.J. van STthsis, finished the text of the manuscript. He had the intention to use this
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.
When in 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,
financially and 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.
Shortly after his arrival in Java, on 1 1 January 1819, Blume was appointed deputy-director un-
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 at Buitenzorg (Bogor), enlarged for this purpose, in the Botanic Gardens.
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 1 824 he made a large tour of inspec-
tion in the company 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.
The writing of the Bijdragen itself was a tremendous task, let alone the research incorporated
in them, 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: 1 825), issued in part simultaneously with fascicle 6 of the Bij-
(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.
In 1823 Blume published the first Catalogus van ... 's-Lands Plantentuin te Buitenzorg (A).
In the listing were many manuscript names of Reinwardt under the latter's name. Blume himself
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.
In September 1825 the Governor-General permitted him a two-years stay in the Netherlands,
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 having been lost by shipwreck, for instance several of
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 II - a book containing drawings of Kuhl & van Hasselt (on Asclepiadaceae,
Orchidaceae, etc.); it is a further proof that Blume did not have these documents (D: 1).
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 1 828 - when
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 of Blume in Brussels, he reported to I). J. van Hwijck (1786- 1855), administra-
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 in Brussels, in 1828, under authority of Blume and
his adjunct. Dr. J.B. Fischer. J.G.S. van Breda (C: 1827-1829), then professor at Ghent and
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-
(12)
Dedication
ty, but was subjected immediately to the Ministry of the Interior. That Ministry drafted an In-
struction for the director, effective from 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
material. Moreover, there was the problem of frequent insect damage once plants are dried.
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 at Leiden he gave special attention and instruction.
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
years of exploration in the East Indies, but in this category it was only P.W. Korthals who per-
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.
(13)
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
(14)
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 the Governor-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 essential aspect of a prosperous colony. In this respect it is significant that he named
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. I do not know if van Haii 's pleading led to any further action,
but it shows that / lora Javae had supporters.
(15)
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 Piper aceae, 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 writing with his sharp pen an acid comment
in Rumphia (1847 or 1849?) on Junghuhn's so-called Lycopodium 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 (B: 1859) Blume complained that the Government had not
contributed to its financing; obviously Blume, who was a man of means, had taken the risk of
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
diseases among 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 cholera was 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
(18)
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.
I will now proceed with some remarks on Blume's personality and his motives, as an addition
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
at Zurich (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. With his 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,
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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.'.
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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 topographical and 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 imperial is.
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).
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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, 1 cannot evaluate it.
For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, I (1976) 236.
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.
(25)
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 contribution is 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.
In: H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen 6: 211-268.
The same as the preceding entry.
1831 - Eenige woorden over de redding van het Rijks Herbarium door Dr. J.B. Fischer.
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
Aspidistra, als mede de beschrijving eener nieuwe soort van dit laatste geslacht.
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.
(27)
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.
(28)
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 in Nusa Kambangan I. (S. Central Java) and claimed
this to be the largest flower, superseding Nelumbntm.
(29)
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, 1 (1976) 240.
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
in Taxon 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-
(30)
Dedication
Japanese and three species are described of Broussonetia (2 new). Furthermore there is a sec-
tion 'synonymic de quelques plantes peu connues', concerning species and 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
I'archipel Indien et du Japon.
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.
(31)
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. 1853. Bonplandia 1: 228. - Blume was in Berlin and offered (obviously at a meet-
ing) fibres of Boehmeria tenacissima Bl. which he said had a great durability and could possibly
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-
dition that it should not be incorporated in the Rijksherbarium.
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.
(32)
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. 4: 454-455.
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.
GiJZEN, A. 1938. 's-Rijks Museum van Natuurlijke Historie, 1820-1915: 100-101. Rotterdam.
- 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, F. 1837. Brief aan C.L. Blume vanuit Djocjakarta, 2 Febr. 1837. Alg. Konst- en Let-
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.
1853. Java, zijnegedaante, zijn plantentooi, en inwendigebouw, 1: 183-186. 2nd Dutch ed.
Kalkman, C. 1979. The Rijksherbarium, past and present. Blumea 25: 13-26, especially p. 14.
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 25: 57-77, especially pp. 60-62. - Blume's endeavours.
1980. The publication of Blume's Tabellen en Platen voor de Javaansche Orchideeen.
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
(34)
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.
1950. History of Malesian botany. 29 pp., unpublished. - Typed copies of letters to and
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,
the facilities for and availability to other botanists, loans, the making of a catalogue of the col-
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.
1 828. Ueber den Charakter der Vegetation auf den Inseln des Indischen Archipels. Ein Vor-
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 1 862 or 1 863 . Copies were sent
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, 1 photo.
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 institutes, effective onwards of December 1st, 1850.
A most peculiar stipulation was in art. 18: anybody could claim to receive duplicates from the
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-
mitted him to devote himself to the study of nature. In 1 840 he was charged with making investiga-
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. 1 12 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. Why he never
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 of Reinwardt 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, 1 863), Blume had a very large library, the total
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 last century concerned with the biology of the Indies were scientists with the German nationali-
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. (C: 1977) in their biography of J.C.P. Arckenhausen reproduced a letter
(in the 'Rijksarchief, The Hague) from Blume to the Minister of the Interior at The Hague (d.d.
27 Dec. 1 832), in which he pleads for the second time for a permanent position of Arckenhausen.
Blume mentioned that he had 1 500 drawings, mostly from Latour, made in Java. These drawings
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 did not defend himself more openly and publicly is not clear. It is of course
a fact that one cannot well oppose rumours without published evidence. He was clearly not a very
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 I am inclined to believe Blume felt it below his dignity to expose
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 (7 April): Foreign corresponding member of the Institut de France, Paris; France.
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.
(40)
ABBREVIATIONS AND SIGNS
ace. = according
Ak. Bis. = Aklan Bisaya (Philip, language)
Alf. Cel. = Alfurese Celebes (language)
alt. = altitude
Anat. = . Anatomy
.Ap. = .Apayao (Philip, language)
app. = appendix, appendices
appr. = approximate
Apr. = .April
Arch. = .Archipelago
atl. = atlas
auci. div. = auciores diversi; various authors
auct(t). mat. = auciores malayenses; authors dealing
with Malesian flora
auctflj. plur. - auciores plures; several authors
.Aug. = . August
Bag. = Bagobo (Philip, language)
basionym = original name of the type specimen; its
epithet remains permanently attached to the taxon
which is typified by it provided it is of the same
rank.
Bg. = Buginese (language)
Brk. = Bikol (Philip, language)
Bil. = Bila-an (Philip, language)
Bill. = Billiton
Bis. = Bisaya (Philip, language)
Bon. = Bont6k (Philip, language)
Born. = Borneo
Bt = Bukit; mountain
Bug. = Buginese (language)
Buic. = Bukidnon (Philip, language)
c. = circiier, about
C. Bis. = Cebu Bisaya (Philip, language)
cf. - confer, compare
Chab. = Chabecano (Philip, language)
citations = see references
cm = centimetre
c.n. =see comb, now
comb, now =combinaiio nova; new combination
CS = cross-section or transversal section of an organ
c.s. =cum suis; with collaboration
cum fig. = including the figure
cur. =curanie: edited by
D (after a vernacular name) = Dutch
Daj. = Dyak (language)
d.b.h. = diameter at breast height
D.E.I. = Dutch East Indies
descr. added behind a reference = means that this
contains a valid description
diam. = diameter
Distr. (as an item) = Distribution
Distr. (with a geographical name) = District
ditio = the same, see do
Div. = Division, or Divide
div. =diversus (masc); various
do = diiio (Ital.); the same
Dum. = Dumagal (Philip. language)
dupl. = duplicate
E = east (after degrees: eastern longitude)
E (after a vernacular name) = English
Ecol. = Ecology
cd. = edited; edition; editor
e.g. = exempli gratia; for example
elah. = elaboravit; revised
emfend). = emendavit; emended
em(erg). cd. = emergency edition
Engl. = English
etc.. dc. =el cetera; and (the) other things
ex audi. = ex auciores; according to authors
excl. =exclusus (masc); excluding, exclusive of
ex descr. = known to the author only from the de-
scription
/. (before a plant name) = forma; form
/. (after a personal name) =filius; the son
f . (in citations) = figure
fam. = family
Feb(r). = February
fide = according to
fig. = figure
f]. =flore, floret (floruit); (with) fiower, flowering
For. Serv. = Forest Service
fr. =fructu, fructescit; (with) fruit, fruiting
Fr. (after a vernacular name) = French
G. = Gunung (Malay); mountain
Gad. = Gaddang (Philip, language)
gen. = genus; genus
genus delendum = genus to be rejected
Germ. = German
geronl. = Old World
haud = noi, not at all
holotype = the specimen on which the original de-
scription was actually based or so designated by
the original author
homonym = a name which duplicates the name of an
earlier described taxon (of the same rank) but
which is based on a different type species or type
specimen; all later homonyms are nomenclaturally
illegitimate, unless conserved
I. = Island
ibfid). = ibidem; the same, in the same place
Ibn. = Ibanag (Philip, language)
ic. = icon, icones; plate, plates
ic. inedit. = icon ineditum, icones inedila; inedited
plate(s)
id. =idem; the same
i.e. =id est; that is
If. = Ifugao (Philip, language)
Ig. = Igorot (Philip, language)
Ilg. = Ilong6t (Philip, language)
Ilk. = Iloko (Philip, language)
in adnol. = in adnotatione; in note, in annotation
incl. =inclusus (masc); including, inclusive(ly)
indet. = indetermined
Indr. = Indragiri (in Central Sumatra)
inedit. =inediius (masc); inedited
in herb. = in herbario; in the herbarium
in lilt. = in lilteris; communicated by letter
in sched. = in schedula; on a herbarium sheet
in sice. =in sicco; in a dried state
in syn. = in synonymis; in synonymy
Is. = Islands
Is. (after a vernacular name) = Isinai (Philip, lan-
guage)
Ism. = Isamal (Philip, language)
isotype = a duplicate of the holotype; in arboreous
plants isotypes have often been collected from a
single tree, shrub, or liana from which the holo-
type was also derived
Iv. = Ivatan (Philip, language)
J(av). = Javanese (language)
Jan. = January
Jr = Junior
KIg. = Kalinga (Philip, language)
Kul. = Kuliman (Philip, language)
Kuy. = Kuy6non (Philip, language)
Lamp. = Lampong Districts (in S. Sumatra)
(41)
Flora Malesiana
Lan. = Lanao (Philip, language)
lang. = language
/.c. =loco cilalo; compare reference
lectotype = the specimen selected a posteriori from
the authentic elements on which the taxon was
based when no holotype was designated or when
the holotype is lost
livr. = livraison, part
lice. =l.c. (plur.)
LS = longitudinal or lengthwise section of an organ
m = metre
M = Malay (language)
Mag. = Magindanao (Philip, language)
Mak. = Makassar, Macassar (in SW. Celebes)
Mai. = Malay(an)
Mai. Pen. = Malay Peninsula
Mand. = Mandaya (Philip, language)
Mang. = Mangyan (Philip, language)
Mar. = March
Mbo = Man6bo (Philip, language)
Md. = Madurese (language)
Minangk. = Minangkabau (a Sumatran language)
min. part. =pro minore parte; for the smaller part
mm = milimetre
Mng. = Mangguangan (Philip, language)
Morph. = Morphology
ms(c), MS(S) = manuscript(s)
Mt(s) = Mount(ains)
n. = numero; number
N = North (after degrees: northern latitude); or New
{e.g. in N. Guinea)
NE. = northeast
nee = not
rieerl. - Netherlands, Netherlands edition
Neg. = Negrito (Philip, language)
N.E.I. = Netherlands East Indies
neotype = the specimen designated to serve as no-
menclatural type when no authentic specimens
have existed or when they have been lost; a neotype
retains its status as the new type as long as no auth-
entic elements are recovered and as long as it can
be shown to be satisfactory in accordance with the
original description or figure of the taxon
N.G. = New Guinea
N.I. = Netherlands Indies
no = numero; number
nom. =nomen; name (on\y) = nomen nudum
nom. al. = nomen aliorum; name used by other
authors
nom. alt(ern). = nomen alternativum; alternative
name
nom. cons(erv). = nomen conservandum, nomina
conservanda; generic name(s) conserved by the In-
ternational Rules of Botanical Nomenclature
nom. fam. cons. = nomen familiarum conservan-
dum; conserved family name
nom. gen. cons. = see nomen conservandum
nom. gen. cons. prop. = nomen genericum conser-
vandum propositum; generic name proposed for
conservation
nom. illeg(it). = nomen illegitimum; illegitimate
name
nom. leg(it). = nomen legitimum; legitimate name
nom. nov. = nomen novum; new name
nom. nud. = nomen nudum; name published without
description and without reference to previous pub-
lications
nom. rejdc.) = nomen rejiciendum; name rejected by
the International Rules of Botanical Nomenclature
nom. seminudum = a name which is provided with
some unessential notes or details which cannot be
considered to represent a sufficient description
which is, according to the International Rules of
Botanical Nomenclature, compulsory for valid
publication of the name of a taxon
nom. suhnudum = nomen seminudum
nom. superfl. = a name superfluous when it was pub-
lished; in most cases it is a name based on the same
type as an other earlier specific name
non followed by author's name and year, not placed
in parentheses, and put at the end of a citation =
means that this author has published the same
name mentioned in the citation independently.
These names (combinations) are therefore homo-
nyms.
Compare 56b line 5-4 from bottom. The same can
happen with generic names.
{non followed by abbreviation of author's name) be-
fore a reference (citation) headed by an other
author's name = means that the second author has
misinterpreted the taxon of the first author.
Compare p. 419a under species 47 the synonym H.
celebica. Diels misapplied the name H. celebica as
earlier described by Burck.
non al. = non aliorum; not of other authors
non vidi = not seen by the author
nov. =nova (femin.); new (species, variety, etc.)
Nov. = November
n.s. = new series
n. sp. = nova species; new species
n. (sp.) prov. = nomen (specificum) provisorium;
provisional new (specific) name
n.v. = non vidi; not seen
NW. = northwest
Oct. = October
op.cit. =opere citato; in the work cited
p. =pagina; page
P. = Pulau, Pulu (in Malay); Island
Pal(emb.) = Palembang
Pamp. = Pampangan (Philip, language)
Pang. = Pangasinan (Philip, language)
paratype = a specimen cited with the original descrip-
tion other than the holotype
part. alt. = for the other part
P. Bis. = Panay Bisaya (Philip, language)
P.I. = Philippine Islands
pi. = plate
plurim. = plurimus; most
p.p. =pro parte; partly
pr. max. p. =pro maxima parte; for the greater part
pro = as far as is concerned
prob. =probabiliter; probably
prop. = propositus; proposed
Prov. = Province
pr.p. =pro parte; partly
pt = part
quae est = which is
9«ooc^ basionym, syn., specimina, etc. =as far as the
basionym, synonym(s), specimen(s), etc. are con-
cerned
references = see for abbreviations the list in vol. 5,
pp. cxlv-clxv
Res. = Residency or Reserve
resp. = respective(ly)
(42)
Abbreviations and signs
S = south (after degrees: southern latitude)
S (after a vernacular name) = Sundanese (language)
Sbl. = Sambali (Philip, language)
SE. = southeast
sec. = secus; according to
sect. =sectio; section
sens. ampl. (ampliss.) = sensu amplo (amplissimo);
in a wider sense, in the widest sense
sens. lai. =sensu laio; in a wide sense
sens. sir. (sirictiss.) = sensu siricto (sirictissimo); in
the narrow sense, in the narrowest sense
Sept. = September
seq., seqq. =sequens, sequeniia; the following
ser. = series
S.I. =sensu lato; in a wide sense
S.-L. Bis. = Samar-Leyte Bisaya (Philip, language)
Sml. = Samal (Philip, language)
s.n.=sine numero: (specimen) without the collec-
tor's number
Sp. = Spanish (language)
sp(ec). = species; species
specim. = specimen(s)
sphalm. = sphalmaie; by error, erroneous
spp. = species; species (plural)
Sr = Senior
5.5. = see sens. sir.
ssp. = subspecies; subspecies
s.sir. = see sens. str.
Stat, now = status nova; proposed in a new rank
Sub. = Subanum (Philip, language)
subgfen). = subgenus; subgenus
subseci. =subsectio; subsection
subsp. = subspecies; subspecies
Sul. = Siilu (Philip, language)
Sum. E.C. = Sumatra East Coast
Sum. W.C. = Sumatra West Coast
Suppl. = Supplement
SW. = southwest
syn. = synonymum; synonym
synonyms = the names of taxa which have been re-
ferred to an earlier described taxon of the same
rank and with which they have been united on
taxonomical grounds or which are bound together
nomenclaturally
syniypes = the specimens used by the original author
when no holotype was designed or more specimens
were simultaneously designated as type
t. = tabula; plate
Tag. = Tagalog (Philip, language)
Tagb. = Tagbaniia (Philip, language)
Tagk. = Tagaka-6lo (Philip, language)
Tapan. = Tapanuli (in NW. Sumatra)
taxon = each entity throughout the hierarchic ranks
of the plant kingdom which can be described and
discriminated from other taxa of the same rank
Taxon. = Taxonomy
Tg = Tandjung (Malay); cape
Ting. = Tinggian (Philip, language)
Tir. = Tirurai (Philip, language)
transl. = translated
type = each taxon above the rank of a species is typi-
fied by a type belonging to a lower rank, for in-
stance a family by a genus, a genus in its turn by
a species; a species or infraspecific taxon is typified
by a specimen. Thenameof ata.xon is nomenclatu-
rally permanently attached to its type; from this it
cannot be inferred that the type always represents
botanically the most typical or average structure
found in the circumscription of the taxon.
type specimen = the specimen or other element to
which the name of a species or infraspecific taxon
is (nomenclaturally) permanently attached; botan-
ically a type specimen is a random specimen on
which the name was based by description. There-
fore, it does not need to represent the average or
most typical representative of a population. See
holotype, isotype, lectotype, syntype, paratype,
and neotype
typ. excl. = typo excluso; type excluded
typ. incl. = typo incluso; type included
typus = see type and type specimen
var. = varietas; variety
var. nov. = varietas nova; new variety
Vern. = Vernacular
vide = see
viz. = videlicet; namely
vol. = volume
W = west (after degrees: western longitude)
Yak. = Yakan (Philip, language)
± = about
& = and
= diameter
(^ =male (flower, etc.)
$ = female (flower, etc.)
5, <^ -bisexual (flower)
(cJ) (9) = dioecious with unisexual flowers
(c??) = monoecious with unisexual flowers
(<? 5) = polygamous
(??) = polygamous
oo = many
> = more than (in size, number, etc.)
< = less than (size, number, etc.)
X 2/5 = 2/5 of natural size
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-
tire; petioles often with 2 lateral glands. Stipules 2, minute and caducous to large
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-
tains full details of taxonomic history, morphology, anatomy, pollen, ecology and distribution
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 is predominantly a Neotropical genus (186 species there) with a single
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, as in Maranthes corymbosa, almost completely fills the tube.
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 species are difficult to observe. The grains are usually distinctly triangular
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 iricnoic and 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 too Httle is known from the chemistry of this
taxon to allow a sound chemotaxonomic discussion.
References: Coradin, Giannasi & Prance, Brittonia 37 (1985) 169-178; Grenand, Moretti
& Jacquemin, Pharmacopees traditionelles en Guyane, ed. Orstom, Paris (1987); Hegnauer,
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
All species of Aiuna and Maranthes have been included in Parinari. Despite their considerable
differences from 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.
In the original description of Parinari, which was based on P. campestris and P. montana from
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 earlier workers had implied that the bilocular ovary is a diagnostic character of
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 1: Petrocarya (correctly Parinari) included the African species P. excelsa and P.
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 1 838, but remained
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.
4. 'Stamens 15-26, the filaments hairy, exserted 1. Chrysobalanus
4. Stamens 7-10, the filaments glabrous, included 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
2. Stamens united into a unilateral ligule. Cotyledons only slightly ruminate 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, all fertile, inserted on margin of disk; filaments hairy,
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. 1 (1753) 513; rical, tomentose on exterior and interior. Calyx lobes
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 (1865) 606; Fritsch, Ann. Naturh. Mus. Wien
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. 1 (1775) 521, t. 208; DC. Prod. 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, 1 (1789) 160, nom. illeg. -
Angelesia Korth. Ned. Kruidk. Arch. 3 (1854) 384; Boerl. Handl. Fl. Ned.
Ind. 1 (1890) 424; Burk. Diet. (1935) 159; Corner, Wayside Trees (1940) 526;
Hutch. Gen. Flow. PI. 1 (1964) 191. - Trichocarya Miq. Fl. Ind. Bat. 1, 1
(1855) 358; ibid. 6 (1858) 1084, p.p. quoad T. splendens tantum. - Geobalanus
Small, Fl. Miami (1913) 80; Hutch. Gen. Flow. PI. 1 (1964) 191. - Coccomelia
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, pilose on 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.
9 (1972) 172. - Angelesia splendens Korth. Ned.
Kruidk. Arch. 3 (1854) 384; Boerl. & Koord. Ic.
Bog. 1, 4 (1901) 59, t. 96; Merr. Philip. J. Sc. 10
(1915) Bot. 307; Enum. Philip. PI. 2 (1923) 236; Cor-
ner, Wayside Trees (1940) 526; Browne, For. Trees
Sarawak & Brunei (1955) 307. - Licania angelesia
Blume, Melang. Bot. 2 (1855) 358. - Chrysobalanus
splendens Korth. ex Miq. F1. Ind. Bat. 1, 1 (1855)
358, in syn. - Parinarium fragile Teijsm. & Binn.
Cat. Hort. Bog. (1866) 253, nom. nud. - Parinarium
nitidum Hook. /. Fl. Brit. India 2 (1878) 310. -
Ferolia nitida {Hook, f.) Ridley, J. Str. Br. Roy. As.
Soc. n. 82 (1920) 183; Fl. Mai. Pen. 1 (1922) 671. -
Parinarium philippinense Elmer, Leafl. Philip. Bot.
10(1939)3809. - Fig. 2.
Tree to 25 m tall, the young branches sparsely Ian-
ate, soon glabrous. Stipules linear-lanceolate, to 3
mm long, caducous. Leaves 4-11 by 1.8-4.2 cm,
oblong, usually acuminate at apex, cuneate at base,
glabrous beneath; petioles 2-5 mm, canaliculate,
glabrous when mature. Inflorescence terminal and
axillary panicles of cymules, 1.5-14 cm long, the
rachis and branches grey-puberulous. Flowers c. 2
mm long. Receptacle campanulate, slightly swollen
to one side, grey-tomentellous on exterior, tomen-
tose within; pedicels c. 1 mm long. Calyx lobes acute,
tomentellous on both surfaces. Petals pubescent on
exterior. Stamens 7-10, slightly unilateral, the fila-
ments glabrous. Ovary at or near base of receptacle.
unilocular, pilose on exterior. Fruit ellipsoid, 1-1.3
cm long; epicarp smooth, glabrous; mesocarp thin,
fleshy; endocarp thin, hard, bony, breaking open by
longitudinal lines of weakness, tomentose within.
Distr. Thailand; in Malesia: Sumatra, Malay
Peninsula, W. Java, Borneo, Philippines. Fig. 3.
Ecol. Commonest in forest, including diptero-
carp forest, on hill slopes and ridges, but wide-rang-
ing in peat swamp, freshwater swamp forest, on sea-
shores, and in rocky places; 0-400(-800) m altitude.
Uses. The timber is strong, durable and resistant
to marine borers and is used for saltwater piles, rail-
road ties, etc. However, it is extremely hard to work
and requires special tools because of silica. The fruit
is edible but is not widely used.
Vern. Malay Peninsula: champrai, medang
merah, m. puteh, membatu, mempadang, merbatu
kechil; Borneo: piasau-piasau, Kedayan, gandulong,
Dusun, tampaluan, Sabah, sampaluan, Brunei,
buku-buku, bunga, djentihan burung, mauhi,
Kalimantan; Philippines: taguilom bay; amayan, ba-
lik, D.Bis., dagingan, dagingdingan, S.L.Bis.,
gapas, maralibus, Tagb.
2. Licania palawanensis Prance, Brittonia 31 (1979)
94.
Shrub, young branches sparsely puberulous soon
becoming glabrous. Stipules lanceolate, 1-2 mm
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,
emarginate, subcuneate at base, glabrous beneath;
petioles 1-3 mm long, c. 1.5 mm wide, lanate be-
coming glabrous with age, rugose. Inflorescences
panicles of cymules, 3-4 cm long, the rachis and
branches sparsely puberulous. Flowers c. 2 mm long.
Receptacle campanulate, slightly swollen to one side,
grey-tomentellous on exterior, tomentose within;
pedicels c. 1 mm long. Calyx lobes acute, tomentel-
lous on exterior, puberulous within. Petals puberu-
lous on exterior. Stamens 7, inserted around com-
plete circle, the filaments glabrous. Ovary inserted at
base of receptacle, lanate-pilose, unilocular. Fruit
(immature) ellipsoid, epicarp smooth, glabrous;
mcsocarp thin; endocarp thin, hard, bony, tomen-
tose within, breaking open by longitudinal lines of
weakness.
Distr. Malesia: Philippines (Palawan). Fig. 3.
Ecol. Confined to ultrabasic rock formation;
0-3(X) m altitude, including sea-shore forest.
V I.icania fuMcarpa (KosTkRM.) Prance, Brittonia
19 (1987) 3W). Hunga fusicarpa Kostfrm. Rcin-
wardtia 10(1985) 123.
Tree to 7 m tall, young branches puberulous, gla-
brescent, with small prominent round lenticels. Stip-
ules not seen. Leaves 5-10 by 1-4.5 cm, charta-
ceous, oblong to elliptic, acute to bluntly acuminate
at apex, cuneate at base, glabrous and glossy on both
surfaces, decurrent onto petiole; petioles 2-3 mm
long, rugose, puberulous becoming glabrous with
age. Inflorescences terminal and axillary panicles of
cymules, few-flowered, the rachis and branches
sparsely puberulous. Flowers c. 2 mm long. Recep-
tacle campanulate, grey-tomentellous on exterior,
tomentose within; pedicels 2-3 mm long. Calyx
lobes acute, narrow, tomentellous on exterior, pu-
berulous within. Petals not seen. Stamens persistent
beneath young fruit, 0.5-1 mm long, connate at
base. Fruit (2.5-)3-5 cm long, narrowly spindle-
shaped, narrowed at apex to a tip 2-3 mm long, nar-
rowed at base in stipe 5-10 mm long; epicarp
smooth, glabrous; mesocarp thin; endocarp hard,
bony, c. i mm thick, densely lanate within, without
lines of dehiscence.
Dislr. Malesia: E. Papua New Guinea (Milne
Bay Prov., [-erguson I., Morobc Prov.). Fig. 5.
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. ser. 2, 18 (1842) 208; Miq. F1. Ind. Bat. 1, 1 (1855)
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.
Ann. Sci. Nat. Bot. ser. 2, 18 (1842) 208; MiQ. Fl.
Ind. Bat. 1, 1 (1855) 359; Boerl. & Koord. Ic. Bog.
1, 4 (1901) 61, t. 97; Ridley, Fl. Mai. Pen. 1 (1922)
672; BuRK. Diet. (1935) 1693; Corner, Wayside
Trees (1940) 526; Browne, For. Trees Sarawak &
Brunei (1955) 308; Kochum. & Wyatt-Smith, Mai.
For. Rec. 17 (1964); Prance & Whitm. Tree Fl.
.Malaya 2 (1973) 331. - Embelia urophyl/a [Wall.
Cat. (1830) n. 2309, nom. nud.] ex A. DC. Trans.
Linn. Soc. 17 (1837) 131. - Diemenia racemosa
(KoRTH.) MiQ. Fl. Ind. Bat. 1, 1 (1855) 358. -
Licania diemenia Blu.me, Melang. Bot. 2 (1855) 10;
Hassk. Flora 41 (1858) 256, nom. illeg. -
Paraslemon spicatus Ridley, J. Str. Br. Roy. As.
Soc. n. 75 (1917) 29. - Fig. 4.
Tree to 40 m tall, or shrub, the young branches
glabrous, the trunk often buttressed. Stipules
triangular, c. 1 mm long, caducous. Leaves thinly co-
riaceous, narrowly oblong, 2.5-8 by 1.4-2.5 cm,
cuspidate acuminate at apex, the tip 5-15 mm,
cuneaic at base; midrib plane above, prominulous
beneath; primary veins 8- 1 1 pairs; petioles 4-5 mm
long, canaliculate, glabrous. Inflorescence of ax-
illary and rarely terminal racemes or occasionally
slightly branched, 4- 14 cm long, the rachis glabrous.
Flowers polygamo-dioecious, c. 1 .5 mm long. Recep-
tacle broadly cupuliform to flattened sauccr-shapcd,
glabrous on cxicrior, lomcntosc within; pedicels up
to 2 mm long. Calyx lobes acute, glabrous on exte-
rior. Petals 5. Stamens 2 fertile and 3 sterile stami-
nodcs opposite. Ovary inserted at base of receptacle,
pilose on exterior, unilocular. Style pilose at base,
glabrous above, the stigma trifid. fruit ellipsoid,
1-1.5 cm long; epicarp smooth, glabrous; mcsocarp
thin, hard; endocarp thin, hard, bony, glabrous
within, opening by 2 lateral plates.
Distr. Nicobar Islands; in Malesia: Malay Penin-
sula, Sumatra, Borneo. Fig. 5.
Ecol. Characteristic of peat swamp forest where
it is a common large tree, but wide ranging into
shorter, more open scrub forest.
Uses. The wood is hard to use because of the
silica content, but it is used locally for general con-
struction, posts, and as firewood.
Vern . Malay Peninsula: kelat, k. pasir, k. puteh,
nylas; Sumatra: galam tabanga, kayu gelang, malas,
meriawak; Borneo: mandailas, Brunei, Besaya, sem-
palawan, Brunei, tempalawan, Bajau, mengilas,
ngilas padang, obah, Sarawak.
Notes. The only record of this species from Java
(Blume s.n., L) is very dubious since the collector's
name was added later. It is probably either misla-
belled or from cultivated material. The only dif-
ference given between P. spicatus and P. urophyllus
is that the former is a shrub with sessile flowers.
Some forms of P. urophyllus have extremely short
pedicels and most sessile-flowered individuals are
recorded as being small trees. There is thus no reason
to maintain P. spicatus as a distinct species.
2. Paraslemon versleeghii Merr. & Pi:rrv, J. Arn.
Arb. 21 (1940) 197.
Tree to 40 m tall, the young branches sparsely
puberulous, soon glabrous. Stipules triangular, c. I
mm long, caducous. Leaves thinly coriaceous, nar-
rowly oblong. 5-9.5 by 1.8-3.7 cm, cuspidate acu-
minate at apex, the tip 7-15 mm long, cuncaie at
base; midrib plane above, prominulous beneath;
primary veins 8-12 pairs, inconspicuous, slightly
650
Flora Malesiana
[ser. I, vol. 104
"'^^
Fig. 5. Distribution of Licania fusicarpa (Kosterm.) Prance (diamond), Parastemon urophyllus (Wall, ex
A. DC.) A. DC. (dots), P. grandifructus Prance (squares), and P. versteeghii Merr. & Perry (triangles).
prominulous beneath; petioles 3-7 mm long, terete,
glabrescent. Inflorescence of axillary and terminal
racemes, 2-9 cm long, the rachis sparsely villous.
Flowers hermaphrodite, c. 1.5 mm long; pedicels
1.5-3 mm, sparsely villous. Receptacle broadly
cupuliform-flattened, sparsely villous on exterior,
tomentose within. Calyx lobes acute, with hirsute
margins. Petals 5, with hirsute margins. Stamens 5,
all fertile, opposite the petals in a complete circle.
Ovary inserted at base of receptacle, glabrous on ex-
terior except at base, unilocular. Style pilose at base,
glabrous above, the stigma with two large apical
lobes, the third reduced or missing. Fruit ellipsoid, c.
1.6 cm long; epicarp smooth, glabrous; mesocarp
thin, hard; endocarp thin, hard, bony, glabrous
within, opening by two lateral plates.
Distr. Malesia: Moluccas (Morotai), New
Guinea, and Admiralty Is. Fig. 5.
E c o 1 . Usually in well-drained hill forest up to 700 m
altitude, but also reported from secondary forest,
swampy (Campnosperma) forest and even beach
forest.
Vern. New Guinea: mangu, Tobelo, noeng,
Irian, gorsauw. Tor, gwarsau, Wainlag, sirebo,
sisero, Kemtoek, sosopi, Japen, telek, Mooi, sino-
ree, Biak.
3. Parastemon grandifructus Prance, Brittonia 39
(1987) 366.
Tree to 30 m tall, the young branches glabrous, the
trunk lightly buttressed to 1 m. Stipules caducous
(not seen). Leaves coriaceous, narrowly oblong,
5-8.5 by 1.8-3.2 cm, with long cuspidate acumen at
apex, the tip 10-16 mm long, cuneate at base,
glabrous on both surfaces; midrib prominent above,
prominulous or plane beneath; primary veins 5-6
pairs, prominulous above, plane beneath; petioles
5-8 mm long, glabrous, slightly canaliculate, slight-
ly swollen at base. Inflorescence of axillary and ter-
minal racemes, the rachis glabrous. Flowers seen on-
ly in fruiting specimens. Calyx lobes 5, acute,
glabrous on exterior, glabrous within except for a
few hairs around base. Receptacle glabrous on ex-
terior in fruiting condition. Style persistent below
fruits, the stigma bifid or trifid. Fruit ellipsoid,
2.3-3.5 by 1.3-1.5 cm, epicarp smooth, glabrous;
mesocarp thin, 0.25 mm; endocarp thin, hard, bony,
0.25 mm thick, glabrous within, opening by 2 lateral
plates 1.9-2 cm long.
Distr. Malesia: Borneo (Sarawak, Sabah). Fig.
5.
Ecol. Upland white sand areas.
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
652
Flora Malesiana
[ser. I, vol. 10^
Fig. 6. Hunga papuana (Baker f.) Prance. A. Habit, xO.5; B. flower, x7; C. flower section, x9; D. petal,
X 15; E. fruit, xO.5; F. ovary section, x 15. - H. novoguineensis Prance. G. Young fruit section, x 1; H.
habit, xO.5 (A-C Forbes 504, £)-F Womersley NGF 19307, G, H Hartley 12645).
1989]
Chrysobalanaceae (Prance)
653
.
.^X
£=» ^
""X
•i
%K
it^
!
A
\^
^^.
i
1 \
Fig. 7
(star),
. Distribution of Hunga longifolia Prance
H. novoguineensis Prance (dots), and H. pa-
puana (Baker/.) Prance (triangles).
bilocular, lanate-pilose externally. Style pilose at
base, glabrous above, stigma truncate. Fruit sagit-
tate-pyriform, unilocular, to 5 cm long, the upper
portion triangular, 2-3.5 cm long, the base narrowly
and abruptly tapered to a stipe 0.6-1 cm long;
epicarp smooth, glabrous; mesocarp thin, fleshy; en-
docarp thin, hard, bony, lanate-tomentose within.
Distr. Malesia: Papua New Guinea. Fig. 7.
Ecol. Oak forest; 500-l(X)0 m altitude.
3. Hunga longifolia Prance, Brittonia 31 (1979) 84.
- Fig. 8.
Tree 15 m tall, the young branches puberulous,
soon glabrous. Stipules linear-lanceolate, 5-6.5 mm
long, puberulous, subpersistent. Leave.s coriaceous,
oblong-lanceolate, 7-13 by 2-3.7 cm, acute to
bluntly acuminate at apex, cuneate at base, glabrous
on both surfaces; petioles 3-5 mm long, shallowly
canaliculate, lanate when young, glabrescent with
^A
vT Zokm
Fig. 8. Hunga longifolia Prance. A. Habit, xO.5; B. flower; C. petal; D. ovary section, all x20.
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- serted midway 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 (1775) 204; Hauman, Bull. Jard. Bot. Brux. 21
(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 panicle. Bracts and bracteoles eglandular, usually concealing flower buds
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-
ly prominent large lenticels; large tree of rain-forest and hills 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
lightly impressed on upper surface 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-
minal panicle 9-15 cm long, the rachis and branches
densely grey sericeous-tomentose; bracts and brac-
teoles ovate, acute, densely tomentellous on exterior,
glabrous within except near apex, caducous. Recep-
tacle campanulate, markedly gibbous, densely grey
tomentellous on exterior, 2-3 mm long; pedicels 1-3
mm long. Calyx lobes 2-3 mm long, narrowly ovate,
densely grey tomentose on exterior, tomentellous
within. Petals spathulate, 2 mm long, caducous. Fer-
tile stamens 7-8, base forming a conspicuous fused
ring with opposite tooth-like staminodes. Ovary
densely pilose. Style pilose, stigma truncate. Fruit
ovoid, 7-8 by 4.5-5.5 cm, exocarp densely len-
ticellate; mesocarp thin, fleshy; endocarp extremely
hard and thick (1 -8 cm thick), woody, granular, and
very irregularly ridged, with 2 small loculi in centre,
densely lanate within.
Distr. Malesia: North Borneo (Sabah). Fig. 9.
Ecol. Hillside forest, to 100 m altitude; forest
along rivers.
Vern. Berangan, Malay.
Fig. 9. Distribution of Parinari argenteo-sericea
Kosterm. (stars) and P. canarioides Kosterm.
(dots).
2. Parinari canarioides Kosterm. New & Crit. Mai.
PI. 3 (1955) 25, t. 12 (For. Dept. Bur. of Planning,
Bogor, Indonesia); Reinwardtia 7 (1965) 159,/. 2.
Trees to 60 m tall; trunk buttressed to 2.5 m high;
young branches sparsely puberulous, glabrescent,
lenticellate. Stipules linear, acute to 5 mm, hirsute,
early caducous, present on very young leaves only.
Leaves chartaceous, ovate, 5-9 by 2-4.3 cm, gla-
brous on both surfaces when mature, without sto-
matal crypts beneath, acuminate at apex, the tip
5-12 mm long, rounded to subcordate at base;
midrib lightly impressed above, prominent beneath,
sparsely pubescent when young; primary veins 7-11
pairs, plane to prominulous above, prominent be-
neath, arcuate; petioles 3-7 mm, glabrous when
mature, eglandular or with small rather inconspicu-
1989]
Chrysobalanaceae (Prance)
657
ous central glands. /n/Zore^cence^ dense-flowered ax-
illary panicles to 4.5 cm long, the rachis and branches
tomentose; bracts and bracteoles persistent, ovate,
puberulous on e.xterior, caducous. Receptacle cam-
panulate, 3 mm long, tomentose on exterior; pedicels
1-2 mm long; calyx lobes elliptic, concave, c. 2 mm,
acute, sparsely puberulous on exterior, densely to-
mentellous on interior. Petals elliptic, obtuse, 2 mm,
tapered to base. Fertile stamens 7-8. Fruit ellipsoid,
3.5-5 by 1 .5-2.5 cm; epicarp densely to sparsely len-
ticellate; mesocarp fleshy, 1 mm thick; endocarp 5
mm thick, hard, marbled, densely lanate within.
Distr. Malesia: Sumatra, Borneo, Sulawesi,
Philippines (Palawan). Fig. 9.
Ecol. Forest extending up to 800 m altitude.
Uses. The timber is much used, but of poor quali-
ty. Fruit edible, also eaten by pigs.
3. Parinari elmeri Merr. Univ. Calif. Publ. Bot. 15
(1929) 92; Kosterm. Reinwardtia 7 (1965) 161, f. 4;
Prance & Whitm. Tree Fl. Malaya 2 (1973) 335.
Trees to 32 m, without buttresses; the young bran-
ches densely tomentellous, glabrescent, obscurely
lenticellate. Stipules lanceolate, acute, to 18 mm long
by 3 mm broad at base, lateral, tomentellous, persis-
tent. Leaves oblong to oblong-lanceolate, 5-18 by
1 .5-7 cm, chartaceous to thinly coriaceous, glabrous
above, densely lanate pubescent beneath, without
stomatal cavities; acuminate at apex, the tip 5-13
mm long, subcuneate at base; midrib plane or slight-
ly impressed and pubescent above when young,
prominent beneath; primary veins 14-21 pairs,
prominent beneath, curved at margin; secondary
nerves more or less parallel forming ladder-like
reticulation; petioles 1.5-6 mm long, tomentellous.
•"====^5;,
Fig. 10. Distribution of Pormor/e/fner/ Merr. (stars)
and P. parva Kosterm. (dots).
glandular, but glands often obscured. Inflorescences
of raceme-like reduced terminal and axillary panicles
or cymules, 1.7-3 cm long, the rachis and branches
densely brown tomentose; bracts and bracteoles
large, 2 mm long, ovate, persistent. Receptacle con-
ical, gibbous, to 3 mm long, brown-lanate on exte-
rior, pedicels 0.5-2 mm long. Calyx-lobes ovate,
acute, 2-3 mm long, lanate on exterior. Petals white,
oblong-ovate, 2-3 mm long, narrowed to base. Fer-
tile stamens 7-9 with tooth-like staminodes oppo-
site. Fruit oblong-ellipsoid, 6.7 by 3.7 cm; epicarp
sparingly lenticellate.
Distr. Malesia: Malay Peninsula, Borneo (Sara-
wak, Brunei, Sabah, NE. Kalimantan), Philippines
(Mindanao). Fig. 10.
Ecol. Lowland and hill forest to 900 m, including
areas on ultrabasic rock.
Uses. The wood is used for supports of Iban long
houses.
Vern. Borneo: resak, Iban.
4. Parinari parva Kosterm. Reinwardtia 7 (1965) 52,
f . 5; 162; Prance & Whitm. Tree Fl. Malaya 2 (1973)
335.
Tree to 15 m tall, bole often fluted at base, without
buttresses; the young branches densely tomentellous,
glabrescent, conspicuously lenticellate. Stipules lan-
ceolate, lateral, 13-37 mm long, up to 5 mm broad
at base, persistent, conspicuously reticulate and
densely tomentose on exterior. Leaves chartaceous,
oblong to elliptic, 11-28 by 5.5-11 cm, glabrous
above, densely lanate-arachnoid pubescent beneath,
the pubescence completely obscuring reticulate ner-
vation, but without stomatal crypts; finely acumi-
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, to 5 mm long, persistent.
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.
Distr. Malesia: Malay Peninsula (Kelantan,
Trengganu, Pahang, Johorc), Sumatra, Borneo. Fig.
10.
Ecol . Mostly on ridge tops and hillsides to 750 m
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,
with stomatal crypts filled with lanate pubescence
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
pairs, curved at margins; secondary nerves reticulate
slightly flattened, with a series of marginal glands at
veins on lower portion; petioles 5-10 mm long,
tomentellous, terete, with 2-4 prominent, conspic-
uous glands near mid point. Inflorescence of spread-
ing terminal and subterminal panicles, 5-11 cm
long, the rachis and branches rather sparsely grey-
brown tomentellous; bracts and bracteoles large,
ovate, 2.5-3 mm long, tomentose, caducous. Recep-
tacle campanulate, 2-3 mm long, tomentose on ex-
terior; pedicels 0.5-1 mm long. Calyx lobes triangu-
lar, acute, c. 1 mm long, tomentose on exterior,
tomentellous within. Petals 5, white, acute. Fertile
stamens 7-9, with tooth-like staminodes opposite,
lanate around base. Ovary villous. Style villous
lanate on lower portion, glabrous above; stigma
capitate. Fruit ovoid, epicarp sparingly lenticellate.
D i s t r . Australia (Queensland and Northern Ter-
ritory) and in Malesia: southern extreme of Papua
New Guinea and Irian Jaya. Fig. 11.
Ecol. Savanna, open forest, forest on rocky
areas in lowlands.
Vern. Papua New Guinea: warrem.
6. Parinari papuana C.T.White, J. Am. Arb. 31
(1950) 86. - Parinari nonda auct. non Benth.: Kos-
term. Reinwardtia 7 (1965) 170, p.p.
Large trees to 40 m tall, buttressed or unbut-
tressed, the young branches puberulous, soon gla-
brous, with clusters of large prominent Ienticels with
central slit. Stipules lanceolate, very early caducous.
Leaves thickly coriaceous to chartaceous, oblong,
4-18 by 1.5-6.5 cm, glabrous above, with stomatal
crypts filled with lanate pubescence beneath, acu-
minate at apex, the tip 3-10 mm long, rounded to
subcuneate at base; midrib plane or slightly impress-
ed and sparsely tomentellous when young above,
prominent beneath; primary veins 16-22 pairs,
Fig. 11. Distribution of Parinari nonda F.v.M. ex
Benth.
curved and anastomosing at margins; secondary
nerves reticulate, slightly flattened, with a series of
marginal glands at vein endings on lower portion;
petioles 2-8 mm long, tomentellous when young,
terete or slightly canaliculate, with 2 conspicuous or
sometimes obscure glands. Inflorescence of terminal
and subterminal panicles, 2-6 cm long, the rachis
and branches densely tomentose or tomentellous;
bracts and bracteoles large, ovate, 2-2.5 mm long,
tomentose on exterior; pedicels 0.2-1.5 mm long.
Calyx lobes triangular, acute, c. 1 mm long, tomen-
tose on exterior, tomentellous within. Petals 5,
white, acute. Fertile stamens 7-8, with tooth-like
staminodes opposite, lanate around base. Ovary
villous, style villous on lower portion, glabrous
above; stigma capitate. Fruit ovoid, 4-6 cm long;
epicarp sparingly to densely lenticellate; mesocarp
thin, fleshy; endocarp hard, thick, marbled, lanate
within.
KEY TO THE SUBSPECIES
1. Leaves coriaceous, 4-11 by 1.9-5 cm. Mature
fruit c. 6 cm long when dry. Montane
a. ssp. papuana
1. Leaves chartaceous, 7-18 by 2.5-7 cm. Lowland.
2. Fruit sparsely lenticellate, small, c. 4 cm long.
Leaf base subcuneate b. ssp. salomonense
2. Fruit densely lenticellate, large, c. 6.5 cm long.
Leaf base usually rounded c. ssp. whitei
a. ssp. papuana
Unbuttressed tree. Leaves thickly coriaceous,
4- 1 1 by 1 .9-5 cm, subcuneate at base. Mature fruit
c. 6 cm long when dry; densely lenticellate on ex-
terior.
Distr. Malesia: Northern, Central and Eastern
Papua New Guinea. Fig. 12.
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-
kuk. Tor, Anona.
b. ssp. salomonense (C.T.White) Prance, Brittonia
39 (1987) 369. - Parinari salomonense C.T.White,
J. Arn. Arb. 31 (1950) 87.
Buttressed tree. Leaves chartaceous, 7-12 by 3-7
cm, subcuneate at base. Mature fruit c. 4 cm long,
sparsely lenticellate on exterior.
Distr. Solomon Islands. Fig. 12.
Ecol . Lowland forest, hillsides and ridges to 300
m altitude.
Vern. Malmone, Kwara'ae, nakisi, one one,
sauialu, susui.
c. ssp. whitei Prance, Brittonia 39 (1987) 369.
Unbuttressed tree. Leaves chartaceous, 7-18 by
2.5-6.5 cm, rounded at base. Mature fruit c. 6.5 cm
long, densely lenticellate on exterior.
Distr. Malesia: West Irian and Papua New Gui-
nea along northern coast from extreme west to east.
Fig. 12.
Vern. Lowka, Manikiong, ogelel, Mooi.
7. Parinari oblont>ifolia Hook. /. Fl. Brit. India 2
(1878) 309; King, J. As. Soc. Beng. 66 (1897) 279;
RiDLKY. Agr. Bull. Str. & Fed. Mai. St. 1 (1902) 144;
Fl. Mai. Pen. I (1922) 668; Foxw. Mai. For. Rec. 3
(1927) 175; Corner, Wayside Trees (1940) 527;
Kosterm. Rcinwardtia 7 (1965) 165,/. 8; Prance &
Whitm. Tree Fl. Malaya 2 (1973) 335. - Ferolia
ohionfii/olia (HfXJK./.) O. Kt/c, Rev. (icn. PI. 1
(1891) 216. - Partnanum hurneense Mkrr. Univ.
Calif. Publ. Bot. 15 (1929) 93.
1 recs to 40 m tall, trunk low thick buttrcs.scd lo 2
m. the young branches minutely tomentellous, gla-
brescent, conspicuously prominently lenticellate.
Stipules ovate to lanceolate, acute, 3-5 mm, pilose
on exterior, early caducous. Leaves coriaceous, ellip-
tic to oblong, 14-23 by 4-9 cm, glabrous above,
with stomatal cavities filled with grey lanate pubes-
cence beneath, shortly acuminate at apex, the tip
3-13 mm long, rounded to subcordate at base;
midrib plane above, glabrous when mature except at
base, prominent, glabresceiit beneath; primary veins
23-35 pairs, erect, plane above, flattened and prom-
inent beneath; secondary veins prominulous and
parallel ± ladder-like beneath; petioles 9-17 mm
long, thick, tomentellous, when young, glabrescent,
eglandular or glandular. Inflorescences of large,
spreading terminal panicles, 10-21 cm long by 7-12
cm broad, the rachis and branches yellow-grey to-
mentellous; bracts and bracteoles ovate, 3 mm long,
early caducous. Receptacle campanulate, slightly
gibbous, 3 mm long, densely grey tomentose on ex-
terior; pedicels 1-3 mm long; calyx lobes ovate,
acute, 1.5-2 mm long, unequal, grey tomentose.
Petals white to bluish, lanceolate to spathulate, nar-
rowed towards base, c. 2 mm long, glabrous. Sta-
mens 8-10, with tooth-like staminodes opposite.
Ovary pilose; style glabrous; stigma truncate. Fruit
ellipsoid, 5-9 by 3-4 cm, epicarp den.sely len-
ticellate; mesocarp 1.5-2 mm thick; endocarp hard,
thick, marbled, 7-13 mm thick, fibrous, densely
lanate within.
Distr. Malesia: Malay Peninsula (S. Kelantan to
Johorc), Sumatra, Borneo (Sabah, Kalimantan).
Fig. 13.
Ecol . Lowland rain-forest and beside rivers or in
valleys extending lo 450 m altitude.
Vern. Malay Peninsula: hedara hulan, kemalau,
nwntelor, merhalu; dunfiun hukit, Malay; Borneo:
mankudar, mengkudu, Kalimantan.
660
Flora Malesiana
[ser. I, vol. 10^
Fig. 13. Distribution of Parinari oblongifolia
Hook./, (dots), P. gigantea Kosterm. (stars), and P.
metallica Kosterm. (triangles).
8. Parinari gigantea Kosterm. Reinwardtia 7 (1965)
182, f. 19.
Large trees to 40 m tall, trunk fluted at base, the
young branches densely lanate pubescent, glabres-
cent, with conspicuous small lenticels. Stipules lan-
ceolate, acute, to 25 mm long, caducous, membra-
neous, densely appressed tomentellous on exterior,
glabrous within. Leaver coriaceous, elliptic, 9-17 by
5-8 cm, glabrous above, with dense conspicuous
stomatal crypts beneath, bluntly acuminate at apex,
the tip 3-6 mm long, rounded at base; midrib plane
above, prominent and pilose, glabrescent beneath;
primary veins 20-28 pairs, slightly impressed on up-
per portion, prominulous on lower portion of upper
surface, straight, erect, parallel; secondary veins ±
parallel; petiole thick, 4-7 mm long, tomentellous
when young, with 2 small round glands on mid point
above. Flowers not seen. Infructescence axillary,
3-5 mm long. Fruit irregularly ellipsoid, 6.5 cm
long, 4 cm broad; epicarp densely lenticellate; meso-
carp fleshy; endocarp hard, bony, irregularly ribbed,
lanate within.
D i s t r . Malesia: Borneo (W. Kalimantan, Sabah).
Fig. 13.
Ecol. Lowland forest.
Vern. Lempong, Kalimantan.
9. Parinari metallica Kosterm. Reinwardtia 7 (1965)
49, f. 3; 160, f. 3.
Trees to 16 m tall, unbuttressed, the young bran-
ches appressed strigose, glabrescent, conspicuously
lenticellate. Stipules ovate-lanceolate, acute, 8-15
mm long, densely brown tomentose, membraneous,
early caducous. Leaves thickly coriaceous, elliptic,
8-17 by 4-9 cm, glabrous and shiny with metallic
sheen above when dry, with dense stomatal crypts
filled with hairs, apex rounded to shortly blunt acu-
minate, the tip 0-3 mm long, rounded or subcuneate
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
pilose. Style glabrous, equalling stamens; stigma
truncate. Fruit not seen.
Distr. Known only from Brunei, Sabah, and
Sarawak. Fig. 13.
Ecol. Forests on well-drained soil, hillsides,
50-300 m altitude.
10. Parinari prancei Kosterm. Reinwardtia 10(1985)
124.
Trees to 25 m tall, the young branches densely
brown lanate and pilose, lenticellate. Stipules
caducous (not seen). Leaves rigidly coriaceous, ellip-
tic, 9-21 by 6.5-12 cm, glabrous and shiny above
when mature, lanate when young, with conspicuous
stomatal crypts filled by lanate pubescence beneath,
broadly apiculate at apex, rounded to broadly sub-
cuneate at base; midrib ± plane above, prominent
beneath; primary veins 14-16 pairs, plane or slightly
impressed above, prominent beneath, arcuate near
margins, secondary venation parallel and forming a
ladder-like reticulum; petioles 10-12 mm long,
densely ferrugineous lanuginose when young, eglan-
dular. Inflorescences of axillary little branched small
panicles or racemes, to 3 cm long, the rachis and
branches densely appressed tomentellous; bracts and
bracteoles caducous. Receptacle campanulate-cupu-
liform, 3-4 mm long, appressed tomentellous on ex-
terior; pedicels 1.5 cm long; calyx lobes triangular,
acute, 1.5 mm long, tomentellous. F/-U/7 ellipsoid, c.
4 by 6 cm diam.; epicarp densely lenticellate; meso-
carp fleshy, 2 mm thick; endocarp woody, very hard
and thick, marbled, densely lanate within.
Distr. Malesia: E. Papua New Guinea (Milne
Bay Prov., Northern Prov.). Fig. 14.
Ecol. Lowland rain-forest to 400 m altitude.
11. Parinari rigida Kosterm. Reinwardtia 7 (1965)
53, f. 6a, b; 163. - Parinari ashtonii Kosterm. Rein-
wardtia 7 (1965) 53, f. 7; 164.
Trees to 30 m tall, unbuttressed, the young branch-
es tomentellous, glabrescent, inconspicuously len-
ticellate. S//pu/e5 caducous (not seen). Leaves rigidly
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
others, broadest below mid point, glabrous and
shiny above, sometimes slightly bullate, the lower
surface with stomatal crypts filled with pubescence,
with 2 glandular areas at junction of midrib and
petiole below, shortly and broadly acuminate at
apex, the tip 3-17 mm long, rounded or subcordate
at base; midrib plane or impressed for upper portion
above, prominent and appressed pilose beneath
when young; primary veins 13-20 pairs, slightly im-
pressed above, prominent beneath, slightly curved at
margins only; secondary venation flattened or
rounded, parallel; petioles thick, 3-10 mm long,
grey-pilose pubescent, rugose, with 2 small glands on
mid point of upper side. Inflorescences of narrow
terminal panicles to 13 cm long, the rachis and
branches tomentose; bracts and bracteoles lanceo-
late, to 2 mm long, early caducous. Receptacle cam-
panulate, slightly gibbous, 5 mm long, densely vil-
lous-tomentose on exterior; pedicels c. 1 mm long;
calyx lobes elongate triangular, 2-2.5 mm long.
Petals spathulate. Stamens 6-8. Ovary densely vil-
lous. Style equalling stamens; stigma capitate. Fruit
irregularly ellipsoid, 5 cm long, to 4 cm diam.,
tapered towards base almost into a stipe; epicarp
densely Icnticcllate; mcsocarp thin fleshy; cndocarp
thick, woody, marbled, lanatc within.
Distr. Malesia: S. Malay Peninsula, Sumatra,
Borneo (Sarawak, L. Kalimantan). Fig. 14.
Ecol. Heath and swamp forests, lowland forest;
0-1400 m altitude.
12. Parinari sumatrana (Jack) Benth. in Hook.,
Niger Fl. (1849) 335; Blume, Mus. Bot.Lugd.-Bat. 2
(1852) 97; MiQ. Fl. Ind. Bat. 1, 1 (1855) 353; ibid.
(1858) 1084; Suppl. Sumatra (1860) 115; ibid. (1861)
306; C.Muell. in Walp., Ann. 4 (1857) 644; Flora 41
(1858) 255; Hook. /. Fl. Brit. India 2 (1878) 309;
Miers, J. Linn. Soc. Bot. 17 (1879) 336; K. & V.
Bijdr. 5 (19(X)) 340, p.p. excl. P. costatum auct. non
Blume; Merr. J. Arn. Arb. 33 (1952) 239; Backer
& Bakh./. Fl. Java 1 (1964) 522, p.p. excl. P.
costatum; Kosterm. Reinwardtia 7 (1965) 176. -
Petrocarya sumatrana Jack, Mai. Misc. 2 (7) (1822)
67 (repr. Calc. J. Nat. Hist. 4 (1843) 165). - Lepido-
carpa ovalis (KoRTH.) Blume ex Miq. Fl. Ind. Bat. 1,
1 (1855) 353. - Ferolia sumatrana (Jack) O. Ktze,
Rev. Gen. PI. 1 (1891) 216. - Parinarium auct. non
Blume: Backer, Schoolfl. Java I (1911) 445, p.p. -
Fig. 15.
Trees to 30 m tall, without buttresses or small ones
to 50 cm; the young branches densely tomentellous,
glabrescent, lenticellate. Stipules oblong, to oblong-
ovate, 5 - 1 2 mm long by 3 - 5 mm wide at base, mem-
braneous, early caducous, pilose on exterior. Leaves
chartaccous to subcoriaccous, elliptic to oblong ellip-
tic, 7 14( 21) by 3-7.5 cm, obtuse to shortly broad
acuminate at apex, the tip up to 3 mm long, rounded
to subcordate at base; glabrous and shiny above,
with deep-set stomatal crypts beneath obscured by
dense caducous lanatc pubescence when young;
midrib plane to slightly impressed above, pilose
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
beneath; petioles 4-8 mm long, with 2 conspicuous
glands near middle, lightly canaliculate, glabrescent.
Inflorescence of short axillary panicles 2-6 cm long,
the rachis and branches brown tomentose; bracts and
bracteoles membraneous, tomentellous on exterior,
puberulous within, caducous. Receptacle conical-
campanulate, 3 mm long, densely pilose on exterior,
almost sessile. Petals spathulate, bluish. Calyx lobes
elongate-triangular, 2 mm long, acute, pilose on
both surfaces. Fertile stamens 8, unequal. Ovary
densely pilose. S/v/e glabrous, equalling stamens, the
stigma truncate. Fruit ellipsoid, 4 by 2.5 cm, epicarp
densely lenticellate; mesocarp 3-4 mm thick; en-
docarp marbled in cross section, hard, 5 mm thick,
densely lanate within.
Distr. Malesia: Sumatra, W. Java. Fig. 14.
Vern. Java: kanjere badak.
Note. This species is distinct from others by the
predominantly axillary inflorescences. The material
described as Lepidocarpa ovalis has much larger,
more pointed leaves than most of the collections, but
KosTERMANS is correct in placing that name in
synonymy under P. sumatrana.
13. Parinari costala (Korth.) Blume, Melang. Bot.
2 (1855) 10; Miq. FI. Ind. Bat. 1, 1 (1855) 354; ibid.
(1858) 1084; Suppl. Sumatra (1860) 115; C.Muell.
in Walp., Ann. 4 (1857) 644; Flora 41 (1858) 255;
Hook. /. Fl. Brit. India 2 (1878) 309; King, J. As.
Soc. Beng. 66 (1897) 277; Ridley, Agr. Bull. Str. &
Fed. Mai. St. 1 (1902) 145; Brandis, Indian Trees
(1906) 278; Burk. J. Str. Br. Roy. As. Soc. n. 73
(1916) 200; Merr. J. Str. Br. Roy. As. Soc. n. 76
(1917) 81; Enum. Born. PI. (1921) 290; Ridley, Fl.
Mal.Pen. I (1922) 666; Merr. Enum. Philip. Fl. PI.
2(1923)236; Burk. Diet. (1935) 1667; Heyne, Nutt.
PI. Ned. Ind. ed. 3 (1950) 697; Kosterm. Rein-
wardtia 7 (1965) 179, f. 17a, b; Prance & Whitm.
Tree Fl. Malaya 2(1973) 333.
For further synonyms, see under the subspecies.
key to the subspecies
I. Inflorescence and flowers densely ferrugineous
villous pubescent. Often at high altitudes
b. ssp. rubJKinosa
I . Inflorescence and flowers sparsely to densely grey
or brown appresscd pubescent. Lowlands.
2. Primary leaf veins 16-26 pairs; mature leaves
9- 15.7 cm long, oblong (index 2.3-3.65). Fruit
cxocarp usually densely vcrrucose
c. ssp. polyneura
2. Primary leaf veins 10-16 pairs; mature leaves
5 10.5 cm long, elliptic (index 1.7 2.7), rarely
oblong. Fruit cxocarp u.sually sparsely vcrrucose
a. ssp. coslata
a. ssp. costata. - Lepidocarpa coslata Korth. Ned.
Kruidk. Arch. 3 (1855) 387; MiQ. Fl. Ind. Bat. 1, 1
(1855) 354, in syn., sphalm. Lepidocarya costata. -
Ferolia costata (Korth.) O. Ktze, Rev. Gen. PI. 1
(1891)216.
Tree to 60 m tall, trunk buttressed up to 2 m, the
young branches densely appressed tomentellous,
glabrescent, with small conspicuous lenticels. Stip-
ules lanceolate, membranaceous, 3-7 mm long,
pilose on exterior, early caducous. Leaver coriaceous
or rigidly chartaceous, elliptic, subovate-elliptic to
oblong (leaf index 1.7-2.7), 5-10.5 by 1.8-4 cm,
glabrous above when mature but with sparse lanate
covering when very young, with stomatal cavities
filled with grey lanate pubescence beneath, acumi-
nate at apex, the tip 3-5 mm long, round to sub-
cuneate at base; midrib prominulous above, tomen-
tellous towards base, prominent beneath; primary
veins 10-16 pairs, arcuate, prominulous above,
prominent beneath; secondary veins rounded or only
slightly flattened; petioles 5-9 mm long, slender,
tomentellous when young, soon glabrous, usually
eglandular or with 2 inconspicuous median glands.
Inflorescences of predominantly axillary or terminal
few-flowered lax panicles to 8 cm long, the rachis and
branches appressed grey to brown appressed tomen-
tellous; bracts and bracteoles lanceolate, c. 2 mm
long, caducous. Receptacle campanulate, slightly
gibbous, grey-brown pubescent on exterior, 3-3.5
mm long; pedicels 0.5-1 mm long; calyx lobes ovate,
acute, 1.5-2 mm long, grey tomentellous on exte-
rior. Petals white, spathulate, 1.5-2 mm long, cadu-
cous, glabrous. Stamens 7-8, with small tooth-like
staminodes opposite, slightly unequal; style gla-
brous; stigma capitate. Fruit ellipsoid, to 3.5 by 4.5
cm; epicarp usually sparsely vcrrucose; mesocarp 2
mm, fleshy; endocarp hard, marbled, 3-5 mm thick,
fibrous, densely lanate within.
Distr. Malesia: Malay Peninsula, Sumatra,
Borneo, Philippines (Mindanao, Culion, Samar).
Fig. 16.
Ecol. Lowland forest, hillsides, ridges; altitude
up to 300 m.
Vern. Malay Peninsula: kemalau, mambatu,
merbatu; Borneo: augok, Piak, bugan, Iban.
b. ssp. rubii(inosa (Ridley) Prance, Brittonia 39
(1987) 368. Parinarium helferi Hook./. Fl. Brit.
India 2 (1878) 311, excl. syn. Parinarium .sumatra-
num sensu Kurz; Brandis, Indian Trees (1906) 278;
Kostkrm. Rcinwardtia 7 (1965) 175. - Parinari
ruhiginosa Ridley, J. Sir. Br. Roy. As. Soc. n. 75
(1917) 29; Fl. Mai. Pen. I (1922) 668; Foxw. Mai.
For. Rcc. 3 (1927) 175; Burk. Diet. (1935) 1667;
KosTiJRM. Rcinwardtia 7 (1965) 168. f. 10; Prance &
Whitm. Tree II. Malaya 2 (1973) 336. Parinarium
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.
Philip. J. Sc. 10 (1915) Bot. 309; Enum. Philip. Fl.
PI. 2 (1923) 235; Kosterm. Reinwardtia 7 (1965) 172,
/. 12.
Leaves 4-11.5 by 1.6-4.3 cm, oblong elliptic to
oblong lanceolate; primary veins 11-19 pairs; pet-
ioles 4-8 mm long, thickly tomentose. Inflorescence
dense to lax, ferrugineous villous pubescent. Fruit
exocarp sparingly lenticellate.
Distr. Burma; Malesia: Malay Peninsula, Bor-
neo (Sabah, Sarawak, Kalimantan), Philippines
(Mindanao, Bucas Grande I.). Fig. 16.
Ecol . In lower montane forests of Malay Penin-
sula and Borneo (750-1500 m) and lowland forests
of the Phihppines.
Vern. Merbatu, Malay (= Malesian standard
timber name for various genera); Borneo: mengku-
dur, Balikpapan.
c. ssp. polyneura (Miq.) Prance, Brittonia 39 (1987)
368. - Parinarium polyneurum Miq. Fl. Ind. Bat.,
Suppl. Sumatra (1860) 115; ibid. (1861) 306; Hook.
/. Fl. Brit. India 2 (1878) 309; King, J. As. Soc. Beng.
60 (1897) 278; K. & V. Bijdr. 3 (1901) 340; Kosterm.
Reinwardtia 7 (1965) 167, f. 9a, b; Prance & Whitm.
Tree Fl. Malaya 2 (1973) 336. - Ferolia polyneura
(Miq.) O. Ktze, Rev. Gen. PI. 1 (1891) 216.
Leaves 9-15.7 by 3.7-6.3 cm, oblong (index 2.3
-3.65); primary veins 16-26 pairs; petioles 3-7 mm
long, thick, tomentose. Inflorescence lax, inflores-
cence and flowers with grey appressed tomentellous
pubescence. Fruit exocarp usually densely verrucose.
Distr. Malesia: Malay Peninsula (Kelantan,
Perak, Pahang, Malacca), Singapore, Sumatra, Bor-
neo. Fig. 16.
Ecol. Lowland forest and occasionally in hills
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, 17 (1948) 461; Kosterm. Candollea 20 (1965) 118. - Moquilea sect.
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. 1 (1865) 607; Boerl. Handl. Fl. Ned. Ind. 1 (1890) 431, 424; Focke in E.
& 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 5, broadly ovate to lanceolate, tomentellous on both surfaces.
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,
Phil. Trans. Roy. Soc. Lond. 320 (1987) 132. -
Parinarium latifolium Hend. Card. Bull. Str. Settl.
7 (1933) 102, nom. illeg., non latifolium Exell. -
Parinari latifrons Kosterm. Reinwardtia 7 (1965) 54.
- Cyclandrophora latifolia (Hend.) Prance in
Kosterm., Candollea 20 (1965) 121. ~ Atuna latifolia
(Hend.) Kosterm. Reinwardtia 7 (1969) 421.
Small tree to 5 m tall, the young branches densely
lanate-tomentellous becoming glabrous, obscurely
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-
ceous on exterior, appressed puberulous within. Re-
ceptacle tube narrowly cylindrical, 7-11 mm long,
sericeous on exterior, sessile; calyx lobes lanceolate
to oblong-ovate, 5-10 mm long, unequal, densely
sericeous on exterior, tomentellous within. Petals
obovate narrowed to base, 10-11 mm long. Stamens
c. 20, inserted on faucal annulus 2 mm high with
tooth-like staminodes opposite, the filaments 10-12
mm long. Ovary densely strigose. Style slender, gla-
brous; stigma truncate. Fruit unknown.
Distr. Known only from Malay Peninsula on
Kedah-Perak border. Fig. 17.
Fig. 17. Distribution of Atuna latifrons (Kosterm.)
Prance & White (star) and A. cordata Cockburn
ex Prance (triangles).
1989]
Chrysobalanaceae (Prance)
667
2. Atuna nannodes (Kosterm.) Kosterm. Rein-
wardtia 7 (1969) 422; Prance & Whitm. Tree Fl.
Malaya 2 (1973) 325. - Parinari nannodes Kosterm.
Reinwardtia 7 (1965) 50, f. 4. - Cyclandrophora
nannodes (Kosterm.) Kosterm. & Prance, Can-
dollea 20 (1965) 122.
Trees to 20 m, usually smaller, unbuttressed; the
young branches sparsely appressed hirsutulous-
strigose, soon glabrous, obscurely lenticellate. Stip-
ules narrowly lanceolate, acute, 6-12 mm long,
strigose to glabrous, subpersistent. Leaves thinly co-
riaceous, oblong-lanceolate, 6.7-19 by 2.5-5.5 cm,
glabrous on both surfaces, sometimes slightly bullate
above, long slender acuminate at apex, the acumen
7-22 mm long, rounded at base; midrib promin-
ulous above, prominent beneath; primary veins
10-12 pairs, arcuate, prominulous on both surfaces
or sometimes prominent beneath; petioles 2-4 mm
long, glabrescent, eglandular, the lower part swollen,
usually curved. Inflorescences axillary racemes 3-7
cm long, the rachis densely sericeous-tomentellous;
bracts and bracteoles lanceolate, 3-7(-13)mm long,
persistent, sericeous. Receptacle cylindrical, 8-13
mm long, densely sericeous on exterior, sessile; calyx
lobes to 6 mm long, unequal, acute, sericeous on ex-
terior. Petals white, spathulate to ovate, 8-12 mm
long, narrowed to base. Stamens 18-20, black to
purple, the filaments 10-15 mm long, slightly unilat-
eral with tooth-like staminodes opposite. Style to 15
mm long, glabrous; stigma capitate. Ovary pilose.
Fruit ellipsoid, 3-4 by 1.5 cm, slightly tapered to
base, crustaceous verrucose on exterior; mesocarp
2-2.5 mm, fibrous, hard, endocarp thin.
D i s t r . Malesia: Malay Peninsula (Trengganu and
Pahang southward), Borneo (Sabah, Sarawak). Fig.
18.
Ecol. Well drained forests to 500 m altitude.
Vern. Merbatu, Malay.
Fig. 18. Distribution of>4/i/na/ion/io^e5 (Kosterm.)
Kostfrm. (iriangles), A. penanffiana (Kosterm.)
KosTtRM. (dots).
3. Atuna penangiana (Kosterm.) Kosterm. Rein-
wardtia 7 (1969) 422; Prance & Whitm. Tree Fl.
Malaya 2 (1973) 326. - Cyclandrophora penangiana
Kosterm. & Prance, Candollea 20 (1965) 124. -
Parinari asperula auct. non MiQ.: King, J. As. Soc.
Beng. 66 (1897) 2U, p.p.
Trees to 20 m tall, unbuttressed, the young branch-
lets glabrescent, obscurely lenticellate. Stipules lan-
ceolate, acute, to 7 mm long, glabrous, stiff, subper-
sistent. Leaves thinly subcoriaceous, oblong to
oblong lanceolate, 3.7-13 by 2-5.5 cm, glabrous on
both surfaces, acuminate at apex, the acumen 3-10
mm, cuneate at base; midrib flattened prominulous
above, prominent beneath; primary veins 10-13
pairs, arcuate, prominulous on both surfaces; pet-
ioles 3-5 mm long, eglandular, glabrescent, smooth,
not swollen or curved. Inflorescences axillary ra-
cemes 3-7 cm long, the rachis densely appressed
pilose; bracts and bracteoles sericeous to 10 mm
long, persistent. Receptacle cylindrical, 5-7 mm
long, sericeous pubescent on exterior; calyx lobes
acute, 4-5 mm long, slightly unequal. Stamens c. 20,
the filaments to 8 mm long with tooth-like stami-
nodes opposite. Style to 10 mm long, stigma capitate.
Ovary pilose. Fruit (immature) ellipsoid, epicarp
crustaceous, verrucose.
Distr. Malesia: Malay Peninsula (Penang,
Perak, Johore, Kelantan and Trengganu). Fig. 18.
Ecol. Well drained forests to 500 m altitude.
Vern. Membatu, Malay.
Note. The two species Atuna nannodes and A.
penangiana are hard to separate. The larger flowers
of A. nannodes seem consistent and the species gen-
erally has leaves with a much longer apex. These may
be one variable species.
4. Atuna cordata Cockburn ex Prance, Brittonia
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
mm long, conical to campanulatc, tomcntellous on
exterior, sessile; calyx lobes slightly unequal, tomcn-
tellous on both surfaces. Petals c. 7 mm long,
668
Flora Malesiana
[ser. I, vol. 10^
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
ide of ring, the filaments 10-12 mm long. Ovary
lensely pilose. Style slender, hirsutulous on lower
)ortion. Fruit 6 cm long, 5 cm wide, ovoid; epicarp
Tustaceous verrucose, mesocarp 5 mm thick, fi-
)rous, hard, endocarp thin.
Distr. Malesia: Borneo (Sabah). Fig. 17.
Ecol. Hill forests on ultrabasic rock.
>. .\tuna racemosa Rafin. Sylva Tellur. (1838) 153;
^ERR. Index Rafin. (1949) 136; Kosterm. Reinward-
ia7 (1969)422. - Fig. 19.
For further synonyms, see under the subspecies.
KEY TO THE SUBSPECIES
1. Leaves 10-25(-35) cm long, usually elliptic,
oblong or lanceolate but sometimes ovate, char-
taceous or thickly coriaceous, the apex long finely
acuminate, 6-25 mm long; petioles thick. Flowers
10-17 mm long. Medium to large trees often with
fluted bole a. ssp. racemosa
I. Leaves 4.5-12 cm long, usually ovate or oblong-
ovate, subcoriaceous or coriaceous, the apex
bluntly acuminate, 3-10 mm long; petioles thin.
Flowers 8-11 mm long. Large trees with cylin-
drical bole b. ssp. excelsa
I. ssp. racemosa. - Atunus alba Rumph. Herb.
\mb. 1 (1741) 171, t. 66, nan Atunus litorea Rumph.
4erb. Amb. 3 (1743) 96, t. 63. - Cydandrophora
\laberrima Hassk. Flora 25 (2), Beibl. 1 (1842) 47;
bid. 11 (1844) 583; Cat. Hort. Bog. (1844) 269. -
°arinari glaberrimum (Hassk.) Hassk. Tijd. Nat.
3es. Phys. 10(1843) 147; C.Muell. in Walp., Rep.
i (1845/46) 647; in Walp., Ann. 4 (1857) 645; Blume,
Vlus. Bot. Lugd.-Bat. 2 (1852) 98; Miq. F1. Ind. Bat.
I. 1 (1855) 355; K. & V. Bijdr. (1900) 338, incl. var.
anceolatum (Teijsm. & Binn.) K. & V., p.p. quoad
;pec. Java; Burk. Diet. (1935) 1696; Backer &
3akh./. FI. Java 1 (1964) 522. - Parinarium sca-
irum Hassk. Tijd. Nat. Ges. Phys. 10 (1843) 147,
lomen; Cat. Hort. Bog. (1844) 269, nomen; Flora 27
1844) 585; C.MuELL. in Walp., Rep. 5 (1845/46)
>47; in Walp., Ann. 4(1857)645; Blume. Mus. Bot.
Lugd.-Bat. 2 (1852) 95, p.p.; Mio. Fl. Ind. Bat. 1, 1
;i855) 354, t. 5; K. & V. Bijdr. 5 (1900) 337. p.p.;
Backer, School fl. Java (191 1)445; Ridley, Fl. Mai.
Pen. 1 (1922)669. ~ Parinarium lanceolalumJtusM.
k Bins. Cat. Hor(. Bog. (1854) 253. 255. nomen. -
Parmarium amhomense Teijsm. & Binn. I.e. 254,
nomen. - Parmarium margarata A.Gray. Bot.
kVilkcs U.S. Expl. Expcd. 1 (1854) 489. I. 55;
C.Muell. in Walp., Ann. 4 (1857) 646. - Parina-
num laurinum A.Gray. Bot. Wilkes U.S. Expl. Ex-
Xd. 1 (1854) 490. I. 55; C.Muell. in Walp.. Ann. 4
[1857)646; Merr. Philip. J. Sc. 10(1915) Hot. 210;
Kanehira. Bot. Mag. Tokyo 45 (1931)282. - Petro-
carya glaberrima (Hassk.) Miers. J. Linn. Soc. Bot.
17 (1879) 336. - Ferolia glaberrima (Hassk.) O.
Ktze. Rev. Gen. PI. 1 (1891) 216. - Ferolia scabra
(Hassk.) O. Ktze. I.e. 216. - Petrocarya scabra
(Hassk.) Miers, J. Linn. Soc. Bot. 17 (1897) 336. -
Parinarium elatum King, J. As. Soc. Beng. 66 (1897)
280; Ridley, Fl. Mai. Pen. 1 (1922) 669. - Parina-
rium hahlii Warb. Tropenpfl. 6 (1902) 370. - Pari-
narium mindanaense Perk. Fragm. Fl. Philip.
(1904) 119. - Parinarium curranii Merr. Philip. J.
Sc. 4 (1909) Bot. 264. - Parinarium warburgii Perk.
ex Merr. J. Str. Br. Roy. As. Soc. n. 76 (1917) 82.
- Cydandrophora elata (King) Kosterm. Candollea
20 (1965) 122. - Cydandrophora scabra (Hassk.)
Kosterm. I.e. 126. - Cydandrophora laurina
(Gray) Kosterm. I.e. 135. - A. elata (King)
Kosterm. Reinwardtia 7 (1969) 421; Prance &
Whitm. Tree Fl. Malaya 2 (1973) 324. - Atuna
scabra (Hassk.) Kosterm. Reinwardtia 7 (1969) 422.
Trees to 45 m tall, usually smaller, the bole often
fluted, young branches glabrous or appressed strig-
ose. Stipules lanceolate, stiff, to 20 mm long, acute,
glabrous to strigose. subpersistent. Leaves usually
chartaceous. more rarely stiffly coriaceous, broadly
ovate, elliptic, oblong or even lanceolate, 10-25
(-35) by 3.5- 1 1 cm, acuminate at apex, the acumen
6-25 mm long, rounded to subcordate at base,
glabrous on both surfaces when mature, sometimes
sparsely strigose beneath on lower portion when
young; midrib prominent on both surfaces; primary
veins 10-13 pairs, prominulous above, prominent
beneath, straight or arcuate; the venation conspicu-
ously papillose and often giving leaf a scabrous ap-
pearance; petioles thick, 3-7 mm long, glabrous or
pilose glabrescent . Inflorescences of axillary racemes
or little branched with up to 3 racemose branches on
short main peduncle, 5-15 cm long, the rachis
tomentellous to sericeous; bracts and bracteoles
ovate, acute, to 8 mm long, caducous. Receptacle
turbinate-campanulate, 5-10 mm long, tomentose
to sericeous on exterior; pedicels 0.5-1 mm long,
calyx lobes 4-7 mm long, ovate to ovate-oblong,
densely tomentellous on both surfaces. Petals ovate-
oblong, to 10 mm long, blue or white. Stamens
15-20, pale blue, to 15 mm long with tooth-like
staminodes opposite. Ovary densely villous. Style
equalling filaments, stigma small. Fruit ellipsoid to
subglobose. to 7.5 cm diam.; epicarp crustaceous
verrucose; mesocarp to 1 1 mm thick, endocarp thin,
1-3 mm, dcn.scly pilose within.
Distr. A wide range from Thailand to the
Pacific: Admiralty, Caroline, and Solomon Islands,
Fiji. Tonga. Samoa; in Malesia: Malay Peninsula
(Pcrak). Singapore. Sumatra, Borneo (Sarawak,
Brunei), Sulawesi. Philippines. Ambon. Ternate.
Ccram. New Ciuinca, New Hrilain. lig. 20.
670
Flora Malesiana
[ser. I, vol. 10
Fig. 20. Distribution of A tuna racemosa Rafin. ssp. racemosa (dots) and ssp. excelsa (Jack) Pranc
(triangles). Atuna racemosa ssp. racemosa also occurs in Tonga, Fiji, and Samoa outside the area shown i
the map.
E c o 1 . Usually occurring in well-drained lowland
or hill forest, up to 600 m altitude, but also found on
riverbanks, freshwater or brackish swamps and even
in mangrove.
Uses. The fruit (cotyledon) is grated and made
into a putty for caulking canoes, widely used in
Pacific islands. An oil is extracted from the seeds
used variously in different areas, e.g. to scent
coconut oil and for hairdressing. The leaves are used
to thatch the outside walls of houses in Fiji. The
wood is used locally for posts and poles, but is not of
good quality.
Vern . Jangong, membatu, Malay; kisokka, Jav.;
Borneo: belibu, senumpol, Iban, kukut, Sarawak,
merampangi, tatambu, Sabah, torog, Orang Sungei,
K'tangan; Sulawesi: lomo, Makassar; Philippines:
aluma, Ceb., botabon, butabul, getabon, Tagb.,
bolga, Bik., pantog-usa, Kuy., pinae, tabontaba,
takoutaban. Bis., tabong. Bag., tabon-tabon, C.
Bis., Bik., Mbo., samake, Bug.; New Guinea:
asikua, asista, Saki, bata-bata, koewao, Kwerba,
dela, Mooi, kan, Oriomo, low tukwa, lowtukwa,
Manikiong, mangosowai, Japen; New Britain: latita,
tita; New Georgia: y//", tavai, tita, Uso; Caroline Is.:
agaratim, ais, eis, eritem, grihing, Palau, adidi. Yap,
Solomon Is.: do-omu, oso, saia, tij, Kwara'ae; Fiji:
makita; Tonga: hea, seea; Samoa: ifi-ifi.
Note. KosTERMANS included Cyclandrophoi
glaberrima as a synonym of Atuna excelsa rath(
than where it is placed here. There seems little doul
based on the original description and herbariui
material at Leiden bearing Hasskarl's writing thi
C. glaberrima is equal to Atuna racemosa ssp. rac
mosa as defined here. The original description of <
glaberrima indicates leaves that are far too large f(
ssp. excelsa. Atuna excelsa was distinguished t
KosTERMANS by its coriaceous leaves and short pe
ioles. However, many sheets which he determined {
A. racemosa have equally short petioles and there
much variation in leaf texture. Therefore it is n<
possible to maintain A. elata. Similarly the distin(
tion of A. scabra was the scabrous texture of tl
leaves and their more lanceolate shape. Many colIe(
tions of /4. racemosa are equally scabrous {e.g. LA
52392 from New Guinea) and there is so much varii
tion in leaf shape that it would be quite impossible t
separate /I. scabra on that feature. This was alread
placed under Parinari glaberrimum by Backer an
Bakhuizen van den Brink (I.e. 1964).
b. ssp. excelsa (Jack) Prance, stat. nov. - Petroci
rya excelsa Jack, Mai. Misc. 2 (7) (1822) 68 [rep:
Hook. Comp. Bot. Mag. 1 (1836) 220; Calc. J. Na
Hist. 4 (1843) 164]; Walp. Rep. 2 (1843) 7. - Parint
1989]
Chrysobalanaceae (Prance)
671
rium jackianum Benth. in Hook., Niger Fl. (1849)
335; MiQ. Fl. Ind. Bat. 1, 1 (1855) 356; C.Muell. in
Walp., Ann. 4 (1857) 644; Hook./. Fl. Brit. India 2
(1878) 312. - Parinarium aspentlum MiQ. Fl. Ind.
Bat., Suppl. Sumatra (1860) \\5,nomen; ibid. (1861)
307, descr.; Hook. /. Fl. Brit. India 2 (1878) 310;
King, J. As. Soc. Beng. 66 (1897) 281; K. & V. Bijdr.
(1900) 337, p./?.; Ridley, Fl. Mai. Pen. 1 (1922)670.
- Ferolia asperula ^iQ.) O. Ktze, Rev. Gen. PI. 1
(1891) 216. - Ferolia jackiana /Benth.) O. Ktze, I.e.
- Parinarium spicatum King, J. As. Soc. Beng. 66
(1897) 279; Ridley, Fl. Mai. Pen. 1 (1922) 669. -
Parinarium maingayi King, J. As. Soc. Beng. 66
(1897) 280; Ridley, Fl. Mai. Pen. 1 (1922) 669. -
Parinarium villamilii Merr. Philip. J. Sc. 10 (1915)
Bot. 308; Enum. Philip. Fl. Pi. 2 (1923) 236. -
Cyclandrophora villamilii ^err.) Prance ex
Kosterm. Candollea 20 (1965) 126. - Cyclan-
drophora excelsa (Jack) Kosterm. I.e. 128. -
Cyelandrophora asperula (Miq.) Prance ex Kos-
term. I.e. 130. - Atuna villamilii (Merr.) Kosterm.
Reinwardtia 7 (1969) 422. - Aluna exeelsa (Jack)
Kosterm. I.e. All; Prance & Whitm. Tree Fl.
Malaya 2 (1973) 324. - Fig. 19.
Tree to 45 m tall, the trunk buttressed up to 2 m,
not fluted, the young branches sparsely strigose,
glabrescent, obscurely lenticellate. Stipules lanceo-
late, 8-15 mm long, acute, sparsely strigose, subper-
sistent. Leaves rigidly chartaceous to coriaceous,
ovate to oblong-ovate or less frequently oblong,
4.5-12 by 2-5 cm, acuminate at apex, the acumen
3-10 mm long, subcordate, rounded or subcuneate
at base, glabrous on both surfaces; midrib prominent
on both surfaces; primary veins 9-13 pairs, arcuate,
prominulous above, prominent beneath, the vena-
tion papillose giving a beaded appearance; petioles
slender, 3-6 mm long, puberulous, glabrescent or
glabrous. Infloreseenees of axillary racemes to 7.5
cm long, or little branched with 2 or more racemose
branches on short main peduncle, the rachis and
branches densely short sericeous; bracts and brac-
teoles oblong, e. 3 mm long, persistent. Reeeptacle
turbinate-campanulate, 4-7 long, sericeous on ex-
terior; calyx lobes ovate, equal, to 4 mm, sericeous
on exterior, tomentellous within. Petals white to
bluish white, oblong, to 5 mm long, caducous. Sta-
mens 13-18, to 8 mm long with tooth-like stamin-
odes opposite. Ovary pilose. Style glabrous, equal-
ling filaments, glabrous above, stigma small. Fruit
subglobose to slightly pyriform, 5-7 cm diam. or
5-7 by 3.5-4.5 cm; epicarp crustaceous, verrucose;
mesocarp fibrous, 5-8 mm thick, endocarp thin,
densely pilose within.
Distr. Malesia: Malay Peninsula (Kedah and
Trengganu southward), Sumatra, Java, Borneo, N.
Sulawesi. Fig. 20.
Ecol. Lowland forests on well drained soils ex-
tending to 750 m altitude on ridges and hillsides.
Vern. Malay Peninsula: kemalau utat, merbatu;
Sumatra: kemiling utan, klappa soepai, pelee kamb-
ing, salak; Borneo: membatu, Sabah, mahadiu, Ban-
djar, temalang.
Notes . Kostermans is probably correct in inter-
preting Petrocarya exeelsa Jack as the species
described here. The original description is quite
detailed and fits this taxon better than any other
Atuna.
Kostermans treated these two subspecies as
separate species. They were differentiated by small
characteristics of leaf shape, the acumen and the
base. While there do seem to be two elements involv-
ed in this complex, there is a complete graduation of
any single character such as leaf length, apex length,
petiole thickness, leaf shape or fiower size. Ssp. ex-
eelsa is much commoner in Sundaland and ssp.
raeemosa in the Sahul shelf and Pacific islands, but
the two subspecies have considerable geographical
overlap with ssp. raeemosa occurring sporadically on
the Malay Peninsula. Since all characters merge and
are only weakly correlated, these two species are
reduced to subspecies, a rank more in accord with
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. Cameroun 20, Fl. Gab. 24 (1978) 29. - Exitelia Blume,
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. 1 (1855) 356; ibid.
(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, 1 (1855) 356;
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
(1923) 235; Craib, Fl. Siam. Enum. 1 (1931) 563;
BuRK. Diet. (1935) 1695; Corner, Wayside Trees
(1940) 527; Backer & Bakh./. Fl. Java 1 (1964) 522.
- Parinarium mulliflorum (Korth.) Miq. Fl. Ind.
Bat. 1, 1 (1855) 356; ibid. (1858) 1084; Suppl.
Sumatra (1860) 115; ibid. (1861) 307; C.Muell. in
Walp., Ann. 4 (1857) 646. - Parinarium salicifolium
(Presl) Miq. Fl. Ind. Bat. 1 , 1 (1855) 357; C.Muell.
in Walp., Ann. 4 (1857) 646. - Maranthes speciosa
KoRTH. ex Miq. Fl. Ind. Bat. 1, 1 (1855) 357. -
Chrysobalanus ciliatus Korth. ex Miq. I.e. 357. -
Petrocarya griffithiana (Benth.) Miers, J. Linn.
Soc. Bot. 17 (1879) 336. - Parinarium racemosum
ViDAL, Cat. PI. Len. Silv. Cult. Manila (1880) 29. -
Ferolia griffithiana (Benth.) O. Ktze, Rev. Gen. PI.
1 (1891)216. - Ferolia corymbosa (BtvuE) O. Ktze,
I.e. 216. - Ferolia salicifolia (Presl) O. Ktze, I.e.
216. - Parinarium nitidum auct. non Benth.:
KooRD. Meded. Lands Planten Tuin Btzg 19 (1898)
448. - Polyalthia pulchrinervia Boerl. Cat. PI.
Hort. Bog. (1899) 20; Icon. Bog. 1 (1899) 106. -
Parinarium palauense Kanehira, Bot. Mag. Tokyo
45 (1931) 282;~ Fl. Micrones. (1933) 129; J. Dept.
Agr. Kyushu Imp. Univ. Fukuoka 4 (1934) 325. -
Fig. 21.
Small to large tree up to 40 m, sometimes flower-
ing when only a few metres high, trunk not but-
tressed or slightly enlarged at base. Stipules intrapet-
iolar, lanceolate, acute, 5-10 mm long, sparsely
pilose on exterior, glabrous within, early deciduous.
Leaves coriaceous, usually oblong-lanceolate to
oblong-elliptic, 6.5-14 by 2.5-8 cm, acuminate at
apex, the acumen 8-20(-30) mm long, cuneate at
base, glabrous when mature but often sparsely cadu-
cous arachnoid-lanate when young, usually with 2
conspicuous prominent glands at junction of petiole
and decurrent lower surface; primary veins 7-10
pairs, arcuate, prominulous on both surfaces; midrib
plane above, prominulous beneath; petioles 4-9 mm
long, glabrous when mature, flattened above. In-
florescences of flattened many-flowered corymbose
panicles, rachis and branches sparsely pilose,
glabrescent. Bracts and bracteoles ovate to lanceo-
late, sparsely pubescent, caducous. Receptacle tur-
binate, tapering into pedicels 2-4 mm long, grey
tomentose to glabrous on exterior, glabrous within,
calyx lobes fleshy, ovate to elliptic, obtuse, 2.5-4
mm long, unequal. Petals white tinged pink, gla-
brous, 3-6 mm long, caducous. Stamens 25-35 in-
serted in several rows on one side of throat, with
tooth-like staminodes opposite. Ovary bilocular,
densely lanate and villous. Style glabrous except at
base; stigma truncate. Fruit ellipsoid, 3-4 mm long,
1 .5-2 cm broad, tapered towards base; epicarp thin,
glabrous on exterior when mature, sometimes lanate
1989]
Chrysobalanaceae (Prance)
675
when young; endocarp hard, 5 mm thick, rough on
exterior; densely lanate within; bilocular usually with
seed in one locule only. Cotyledons plane-convex.
Disir. S. Thailand extending east to Solomon
and Caroline Islands and Australia (Queensland,
Northern Territory); in Malesia: Malay Peninsula,
Sumatra, Java, Borneo, Lesser Sunda Islands,
Sulawesi, Philippines, Moluccas, New Guinea, New
Britain and Admiralty Islands. Fig. 22.
Ecol. Common in coastal areas on rocky and
sandy hills and extremely inland up to 600 m altitude.
Also in gallery forest and in Australia on sand dunes
behind mangrove swamp. In Kalimantan the fruit is
eaten by many bird species, including hornbills and
fruit pigeons, which probably disperse the seed. The
seed is also scatter-hoarded by the squirrel Sun-
dasciurus hippurus. African species of Maranthes
are bat-pollinated.
Uses. Wood used for house-building and for
posts. Fruit edible.
Vern. Thailand: chi-kai-pen, chi-ot-pen, Korat;
Malay Peninsula: chana, lejin, merbatu, m. layang,
mujagon, sau hulan, sunko rimau; Sumatra: damor
lilis, kajiebatu, kaju baiu, Banka, kalek kureseng, k.
parada; Java: gesing, kituwat, solo, sulo, triwulan,
Huloh, Jav., taritik, l. monjet, Sund.; Borneo: bang-
kawang, bonsissian, Malay; bansisian, Sabah,
Tengara; nyalin laat, Sarawak; buenza, kajebabu,
kajoe kambang, kambang, potang, Kalimantan;
Sulawesi: kolaka; Tidore: latan, Aru Is.; Philip-
pines: almag, delebaybai, kaphangan, kolaka, kola-
sa, kulingan, malapiga, malapuyan, sampinit, tak-
dangan. Tag., aningat, binggas, caratacat,
kagemkena, karatakat. Ilk., arangan, Tagb., daka-
yau. Pang., bakoyan, tapas, P. Bis., bongog, dau,
mata-mata, sarangun, S.-L.Bis., dumaga, Kuy., ka-
gangan, kalakangon, ogat. Bag., kamuli tingan,
Pamp., lank angan, Lan., langog, Buk., lumaluas,
sigaadan, Mag., maluktik, Sul., salipungan, salutui,
Neg., bareraga, baril, Bik., C.Bis., laiusin, Bik.,
S.C.Bis., liusin, Sbl., Tag., Bik., sabongkaag. Ilk.,
Ting., tadiang manok. Ting., Tag.; New Guinea:
badigal, Wagu, djuramun, Kemtuk, jambuan,
Kaigorin, kaupen, Jal, kawol, kowot, Muyu, kwanu,
Maprik, lakan, luikoko. Bush Mekeo, marigag,
Sinai, mehlue, Bembi, morolee, mun, Dagu, naas,
ningua, njali, Nemo, njiwa, niwa, Sidei, paguh,
Timbunke, phu, Wasuk, watu, Karopa; Solomon
Is.: asikisiki, giza, mon warlu, morigag, now-wa-ru,
santalan; Bougainville: mon-warku, Kugumaru,
marigai, Siwai, Bouin; Palau Is.: apgau.
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,
676
Flora Malesiana
[ser. I, vol. 10^
f D K.TincL
Fig 23. Kostermanthus heteropetalus (Scortech. ex King) Prance. A. Habit; ^•^"°^^^^^f^'°"' f„f ^
petals; £. stamen; F. fruit; C. ovary section (>»-£, G Ogata KEP 105153, FMeijer SAN 34279).
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,
ovate, foliaceous, acute to acuminate, persistent to
subpersistent. Leaves coriaceous, usually elliptic-
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
on venation of both surfaces giving a bead-like ap-
pearance; midrib prominulous above, prominent
beneath; primary veins 6-10 pairs, arcuate, slender,
prominent beneath; petioles 6-12 mm long, some-
times lightly alate from decurrent leaf margins,
slightly flattened above, eglandular. Inflorescences
little-branched, to 10 cm long, the rachis and branch-
es lightly tomentellous; bracts and bracteoles ovate,
acute, to 3 mm long, caducous. Receptacle broadly
campanulate, 2-3 mm long, tomentose on both sur-
faces; calyx lobes fleshy, unequal, acute, to 7 mm
long, pilose on both surfaces, reflexed in open flow-
ers. Petals white tinged pink, fleshy, ellliptic, con-
cave, largest up to 15 mm long, tomentellous on ex-
terior, enveloping staminal ligule, the others much
smaller to 6 mm long. Stamens 25-30 united into a
unilateral ligule for 2/3 length, to 12 mm long,
glabrous; anthers pubescent. Ovary densely pilose.
.S7>'/^ densely appressed pilose, stigma truncate. Fruit
ovoid, unilocular 4 by 3 cm; epicarp glabrous, crus-
taceous; endocarp hard, thick. Cotyledons slightly
ruminate, 1.5 by 3 cm.
Distr. Malesia: Malay Peninsula, Sumatra,
Borneo, Sulawesi, Philippines (Mindanao). Fig.
24.
Ecol . From sea level up to 500 m altitude.
Fig. 24. Distribution of Kostermanthus heterope-
talus (Scortech. ex King) Prance (dots) and K.
malayanus (Kosterm.) Prance (star).
2. Kostermanthus malayanus (Kosterm.) Prance,
Brittonia 31 (1979) 94; Prance & White, Phil.
Trans. Roy. Soc. Lond. 320(1988) 152. - Acioa ma-
layana Kosterm. Reinwardtia 7 (1965) 13.
Small tree to 10 m; young branches glabrous, len-
ticellate. Stipules lanceolate, acute, glabrous, sub-
persistent, c. 5 mm long. Leaves chartaceous, ellip-
tic, 14-20 by 6.5-8.5 cm, acuminate at apex, the
acumen 4-10 mm long, cuneate at base, glabrous on
both surfaces; midrib slightly prominulous to plane
above, prominent beneath, with a pair of round
glands at base; primary veins 10-13 pairs, pro-
minulous above, prominent beneath; petioles 2-3
mm long, glabrous, slightly alate with decurrent leaf
margins. Inflorescence of subterminal racemes or lit-
tle branched, the rachis brown pilose pubescent;
bracts and bracteoles ovate, acute, to 3 mm long,
caducous. /?ecfp/flc/e slender cylindrical, 5 mm long,
sessile tomentose on exterior, densely tomentose
within; calyx lobes ovate, acute, 4-5 mm long,
densely tomentose on exterior, glabrous within ex-
cept at apex. Petals spalhulate, 6 mm long, clawed.
Stamens S- 10, united into a unilateral ligule for half
of length. Ovary densely pilose. Style pilose for most
of length. Fruit unknown.
Distr. Malesia: Malay Peninsula (Penang).
Known only from the type collection.
678
Flora Malesiana [ser. I, vol. 10^
Insufficiently known
Acioa percohacea Kosterm. Reinwardtia 7 (1965) 14. j j- . •• u ^ «•
This species was described from a single sterile collection from the Malay Penmsula, and distmgu.shed from
Kostermanthus heteropetalus Prance by its pubescent branches and caducous pubescent leaf undersurfaces.
It is impossible to evaluate until further material is collected, but almost certamly belongs withm K.
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 (4-)5, mostly opposite the sepals (rarely alternate:
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 leading botanists: Warburg (1895), von Wettstein (1911), Hutchinson (1926, 1973),
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 t he monographs of both authors in Biumea volumes 1 9 and 26, and provided with
an introduction.
(679)
680
Flora Malesiana [ser. I, vol. 10^
spermaceous scheme and mentioned that Radlkofer was not in favour of an affinity with Sapin-
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
opposition of calyx, corolla and stamens is a most unusual feature, but can probably be derived
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-
patible with a position near Menispermaceae, but that of Meliosma more similar with some
members of the Rosidae. There may be more arguments to accommodate Meliosma in a separate
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 1 (1862)413; Blume, Mus. Bot. Lugd. -Bat. 1 (1851) 369; Cronquist, An integrated system
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 (1936) 18; Melchior, Engler's Syllabus 2 (1964) 285; Takhtajan, Flowering plants: origin and
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 clusters common near the veins.
Young stems. Cork superficial. Cortex with stone cells in some species oi Meliosma. Pericyclic
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, solitary and 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
families, or alternatively to retain their tribal position in the same family. Anatomically Sabia is
quite distinct from 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-
tacyclic triterpenoids of the oleanene series and the absence of starch in seeds. The 3-acetates of
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 pairs, ascending to patent, curved to straight. Flowers bisexual, 5-
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 less superimposed, attached to the septum, hemi-anatropous. 'Drupe-
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-) I - 1 .5(- 1 .75) mm long 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- I mm long.
3. Cymes up to 2 cm, 1 4( 6)-nowcrcd; petals suborbicular to elliptic, 1.75 2.5 by 1.25-2 mm, obtuse
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 to 2(-3. 5) cm, (l-)2-10(-12)-flowered; petals oblong, 2.5-4(-4.5) by c. 1-1.5 mm, obtuse;
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(- 1 8) by 2-6(-8) cm; nerves (5-)6-8(-9) pairs; cymes either ar-
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)
35.
Evergreen, woody. Twigs glabrous to somewhat
pubescent; flowering twigs up to 2.5 mm diam., ±
lax-pubescent. Buds ovoid, acute; scales ± pubes-
cent, ciliolate. Leaver oblong, 5-8 by 2.5-3 cm, in-
dex 2-2.7, pergamentaceous, above glabrous or still
sparsely pubescent especially at the base and on
midrib, beneath laxly pubescent especially on midrib
and nerves; base acute, apex acute or short-acumi-
nate; nerves 6-7 pairs, patent, ± .straight to some-
what curved; petiole up to 1 .5 cm, glabrous to pubes-
cent. Cymes solitary, axillary up to 4.5 cm, up to
40-flowered, ± lax-pubescent; pedicels up to 4 mm;
bracteoles oblong to oblong-ovate, up to 0.8 mm,
pubescent, ciliolate. Sepals ovate to somewhat ellip-
tic, 0.8-1 by 0.5-0.75 mm, obtuse to acute, ±
pubescent, ciliolate. Petals oblong or oblong-ovate
to sublanceolate or ovate-lanceolate, 3.75-4 by
1-1.5 mm, acute to narrow-obtuse, subciliolate,
nerves up to 6, dark-coloured. Stamens 2.3-3 mm;
filament flattened, 1 .8-2.6 by 0.25-0.4 mm; anther
ellipsoid to oblong-ellipsoid, c. 0.4-0.6 mm,
upright. Disk crown-shaped; lobes very short or ab-
sent; ribs ± prominent. Pistil 2.75-3 mm; style
narrowly-conical to cylindrical, 2.25-2.5 mm, with
some hairs at the base; ovary somewhat globular to
subreniform, 0.5-0.6 by 0.6-0.8 mm, densely
pubescent. Drupelets not available.
Distr . Malesia: Singapore (Bt. Timah Res.), only
known from the type, collected in 1940.
Notes. In habit somewhat resembling S. par-
viflora but readily distinguished by floral characters.
On the label noted as a 'tree, 100 ft', but this is
suspected to be a wrong annotation or field observa-
tion or a wrong label.
2. Sabia javanica (Blume) Backer ex Chen, Sargen-
tia 3 (1943) 59; Backer & Bakh./. F1. Java 2 (1965)
144; VAN DE Water, Blumea 26 (1980) 39. -
Meniscosta javanica Blume, Bijdr. (1825) 29. -
Meniscosta scandens Blume ex Spreng. Syst. Veg. 4,
2 (1827) 114, nom. illeg.; Dietr. Syn. PI. 2 (1840)
923. - Sabia meniscosta Blume, Mus. Bot. Lugd.-
Bat. 1 (1851) 369, f. 44, nom. illeg., incl. var. firma
Blume, var. latifolia Blume et var. glabriuscula
Blume; Miq. F1. Ind. Bat. 1, 2 (1859) 6 1 8 ( 'me«/co5-
tay, Fl. Arch. Ind. (1870) 71; //j/c^. (1871) pi. 31, /nc/.
var. elliptica Miq.; Hook./. Fl. Brit. India 2 (1876)
3 ('menescortay. Backer, Schoolfl. Java (1911) 273;
KooRD. Exk. Fl. Java 2 (1912) 544. - Sabia elliptica
(Miq.) Miq. Sum. (1861) 203, 521. - Sabia javanica
^lume) Chen var. glabriuscula (Blume) Chen,
Sargentia 3 (1943)61.
Evergreen woody climber or scandent shrub, up to
10 m. Twigs glabrous; flowering twigs up to 5 mm
diam., glabrous or ± pubescent. Buds ovoid, up to
2 mm, acute; scales glabrous or with few hairs, ±
ciliolate. Leaves elliptic-oblong to sublanceolate,
6-19 by 2-8(-10) cm, index 2-3(-4), pergamen-
taceous to pergamentaceous-coriaceous, above and
beneath glabrous or with some hairs on midrib; base
acute to rounded, apex acute, acuminate; nerves
4-7(-8) pairs, patent, curved to straight; petiole up
to 2.5 cm, glabrous to sparsely pubescent, ± (fine-)
wrinkled. Cymes arranged in an axillary, up to 12 cm
long, glabrous to pubescent, thryrsoid inflorescence,
subtended by bracts or sometimes by small leaves
and then inflorescence up to 17 cm long; cymes up to
3 cm, forming a lax to dense cluster of 3-10(-12)
flowers, subglabrous to pubescent. Bracts ovate to
sublanceolate, up to 5 mm, subglabrous to more or
less pubescent, ± ciliolate; bracteoles as bracts but
1989]
Sabiaceae (van Beusekom & van de Water)
68S
smaller, or bracteoles minute or sepal-like and then
situated near calyx; pedicel up to 4 mm. Flowers
green to yellow or white. Sepals sometimes 6 (see
bracteoles), ± ovate or broad-ovate, 0.75- 1{- 1.25)
by 0.5-0. 8(-l) mm, acute to obtuse, ± pubescent,
ciliolate. Petals oblong, 2.5-3.5(-4) by 1-1.5 mm,
obtuse, nerves up to 5, often dark-coloured and then
conspicuous. Stamens (1-)1.25-1.5 mm; filament
± flattened, (0.75-)l-1.25 mm long, 0.25-0.5 mm
wide; anther globular to ellipsoid, 0.2-0.3 mm, in-
flexed. Disk crown-shaped; ribs sometimes faint or
absent. P/5/// 0.8-1.2 mm; style ± conical, 0.2-0.5
mm, much shoner than the adjacent side(s) of the
drupelet(s); ovary globular to subreniform, 0.5-0.6
by 0.5-0.7 mm, glabrous. Drupelets obovoid or
sometimes globular, ± compressed, 9-11 by 9-10
mm, without persistent petals and stamens at the
base; reticulate pattern often coarse and limited to
the margin. Embryo with somewhat undulated or
faintly folded cotyledons.
Distr. Malesia: Sumatra (East Coast Res., In-
dragiri, Lampongs), W. Java. In all c. 30 collections.
Ecol. Forests, at (20-)200-1500 m. Ft. fr.
Jan. -Dec.
Vern. Java: areuj bebentjojan, a. kahawatang,
a. katjapi, S.
Notes. Sabia javanica strongly resembles S.
pauciflora from the Philippines, the Moluccas, New
Guinea, and the Solomon Islands. It can be
distinguished from that species by its often more-
flowered cymes, its shorter style, and some other
slight differences. Since both species are geographi-
cally separated, it was also possible to combine them
into one species and give them the rank of subspecies.
Although the differences are rather small, 1 believe
that S. javanica and 5. pauciflora represent two dif-
ferent, well-delimited, but very closely related spe-
cies. Moreover, a reduction of both species to a single
one would increase the variability of several taxo-
nomic important characters, in consequence of
which the delimitation with some other related
species, like S. parvijlora and 5. racemosa, and
possibly also S. limoniacea, would become less
distinct. Finally, this might result into a far-going
lumping and a reduction of all these species to, say,
subspecies. Contrary to the situation in the extra-
Malesian species .S. campanuiata Wall., however, in
this case I believe that the differences between these
taxa have reached a higher level already, resulting in
the distinction of mutually closely related but ±
well-delimited species, each with its own specific
combination of characters.
In vegetative characters and in drupelets S.
javanica resembles .S. racemosa from Borneo. It can,
however, easily be distinguished from that species by
its morc-flowcred inflorescences and its floral
characters, es{>ccially its petals.
3. Sabia limoniacea Wall. [Cat. (1829) n. 1000,
nom. nud.] ex Hook./. & Th. Fl. Ind. 1 (1855) 210;
Walp. Ann. 4 (1857) 139; Benth. Fl. Hongk. (1861)
70; Hook./. Fl. Brit. India 2 (1876) 3; Kurz, J. As.
Soc. Beng. 45, ii (1876) 204, excl. syn. Sabia sp.
Griffith (= S. parvijlora ssp. parviflora); For. Fl.
Burma 1 (1877) 300 {'timonaceay, Forbes &
Hemsley, J. Linn. Soc. Bot. 23 (1886) 144; King, J.
As. Soc. Beng. 65, ii (1896) 454; Prain, Beng. PI. I
(1903) 246; Brandis, Indian Trees (1906) 194; Dunn
& Tutcher, Kew Bull. Add. Ser. 10 (1912) 68;
Ridley, Fl. Mai. Pen. 1 (1922) 513; Merr. Lingnan
Sc. J. 5 (1927) 19; Kanjilal c.s. Fl. Assam 1,2 (1936)
326; Chen, Sargentia 3 (1943) 56, f. 7; Biswas, PI.
Darj. Sikkim Himal. 1 (1966) 261; van de Water,
Blumea 26 (1980) 44, / 6b, 8. - Androglossum
reticulatum Champ, ex Benth. Hook. J. Bot. Kew
Gard. Misc. 4 (1852) 42; Benth. Fl. Hongk. (1861)
70; Chen, Sargentia 3 (1943) 58, non S. reticulata
Elmer (1909). - Sabia celastrinea Muell. in Walp.,
Ann. 6 (1865) 1269. - Sabia malabarica Bedd. Ic.
PI. Ind. Or. 1 (1874) 39. t. 177; HooK./ Fl. Brit. In-
dia 2 (1876) 2; Brandis, Indian Trees (1906) 194;
Gamble, Fl. Pres. Madras 1 (1918) 254; Chen,
Sargentia 3 (1943) 48. - Fig. 2, 3.
Evergreen woody climber, up to 10 m. Twigs
glabrous or sometimes sparsely pubescent; flowering
twigs up to 5 mm diam., glabrous to lax-pubescent.
Buds broad-ovoid to ovoid, up to 2.5 mm, acute;
scales (sub)glabrous, often ciliolate. Leaves oblong-
ovate to lanceolate, 4-18 by 1.5-6.5(-8) cm, index
2-4(-4.5), ± pergamentaceous-coriaceous, above
and beneath glabrous or with some hairs especially
on midrib; base acute to rounded, apex acute,
sometimes obtuse, acuminate or not; nerves 5-9
pairs, ± patent, sometimes somewhat ascending,
curved to straight; petiole up to 2.5 cm, glabrous to
lax-pubescent. Cymes either solitary, axillary,
subtended by small and often herbaceous leaves, or
when either the leaves are fallen or the cymes are
subtended by bracts arranged in an up to 1 5 cm long,
glabrous to ± lax-pubescent or tomentellous, race-
mose to thyrsoid inflorescence, cymes up to 2 cm,
l-4(-6)-nowered; pedicels up to 7 mm; bracts
oblong, up to 4 mm, glabrous to pubescent, ciliolate;
bracteoles ovate to oblong, up to 1 .75 mm, glabrous
to pubescent, ciliolate, often situated near calyx.
Flowers green to yellow or white. Sepals sometimes
6 or 7 (see bracteoles), broad-ovate to elliptic,
0.7- 1 .2(- 1 .5) by 0.6- I mm, acute to rounded, gla-
brous to ± pubescent, ciliolate. Petals suborbicular
to elliptic or ± obovate, 1.75-2.5 by 1.25-2 mm,
obtuse to rounded, sometimes broad-acute, nerves 5,
usually obscure. Stamens 1.5-2 mm; filament
somewhat flaticncd, 1 .25- 1 .75 by 0.3-0.4 mm; an-
ther ellipsoid, 0.25-0.35 mm, inflexed. Disk crown-
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
Fig. 3. Sabia limoniacea Hook./. & Thoms. a. fruit;
a', embryo, both xl.5 (a Poilane 24769; a'
POILANE 18918).
0.7-1.2 mm; style conical to cylindrical, 0.2-0.6
mm, much shorter than the adjacent side(s) of the
drupelet(s); ovary globular to subreniform, 0.5-0.6
by 0.5-0.8 mm, glabrous. Drupelets globular to
obovoid, strongly compressed, 11-14 by 10-13 mm,
red to blue or black when fresh, without persistent
petals and stamens at the base; reticulate pattern
usually faint or absent, sometimes more prominent
at the margin. Embryo with somewhat undulated
cotyledons.
Distr. Continental SE. Asia (throughout India,
Burma, Bangladesh, Thailand and Indochina to
China); in Malesia: Malay Peninsula (incl. also P.
Penang), Central Sumatra and Borneo (Sarawak), in
all 7 collections.
Ecol . Thickets and forest, 300-1200 m altitude.
FL Sept. -Jan.,//-. Dec. -April.
4. Sabia parvinora Wall, in Roxb., Fl. Ind. 2 (1824)
310; G.Do.M, Gen. Hist. 2 (1832) 69; Walp. Rep. 1
(1842) 557; Hook./ & Th. Fl. Ind. 1 (1855) 210;
Walp. Ann. 4 (1857) 139; Hook./ Fl. Brit. India 2
(1876) 2; Stapf, Trans. Linn. Soc. Lond. 4, 2 (1894)
142; Bra.ndis, Indian Trees (1906) 194; Lecomte, Fl.
Gen. I.-C. 2 (1908) 2, incl. var. harmandiana Le-
comte, Bull. Soc. Bot. Fr. 54 (1907) 674; Kanjilal
c.s. Fl. Assam 1, 2 (1936) 325; Chun, Sunyatsenia 4
(1940) 242; Merr. Brittonia 4 (1941) 112; Chen,
Sargentia 3 (1943) 64; Gagnep. & Vidal, Fl. Camb.
Laos, Vietnam 1 (1960) 16; Biswas, PI. Darj. Sikkim
Himal. 1 (1966) 261; Sen Gupta, Bull. Bot. Soc.
Bcng.22, ii(1968) 196; Hara. Fl. E. Himal. 2 (1971)
74; Sen Gupta, Rec. Bot. Surv. India 20, 2 (1973)65;
Hah A & Williams, Enum. Fl. PI. Nepal 2(1970) 100;
VAs de Water. Blumea 26 (1980) 48, f. 3c, 9. -
\al)ia harmandiana Pierre, Fl. For. Coch. 5 (1897)
pl 360B; Craib, Fl. Slam. Enum. I (1926) 340. -
Sahia phitippinensis Robins. Bull. Torrcy Bot. Club
' I9(J8) 70; .Merr. Enum. Philip. 2 (1923) 516;
•.. Sargentia 3 (1943) 67.
"■ a complete synonymy, see van de Water
'M
crgrccn climber or scandcnt shrub, up to 6 m.
'.'."> glabrous to laxly pubescent; flowering (wigs
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-
tenuate or not; apex acute, acuminate; nerves (5-)
6-9(-10) pairs, patent, straight or sometimes ±
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
spreading, lax, and with up to 35 or more flowers,
glabrous to sparsely pubescent; pedicels up to 1 cm;
bracts ovate to lanceolate, up to 2 mm or, when sub-
tending a cyme up to 6 mm, subglabrous to pubes-
cent, ciliolate; bracteoles as bracts. Flowers green to
yellow or white. Sepals broad-ovate to ovate,
0.7- 1 .5 by 0.5- 1 mm, acute to rounded, glabrous to
pubescent, ciliolate. Petals elliptic-oblong to lanceo-
late or sometimes oblong-ovate, 2-4(-4.5) by
0.7-1.3 mm, acute to obtuse; nerves up to 7, dark-
coloured or sometimes obscure. Stamens 1.2-2.25
(-2.5) mm; filament flattened, 0.9-2(-2.25) by
0.25-0.5 mm; anther ellipsoid to ovoid, 0.25-0.4
mm, often ± inflexed. Disk crown-shaped, usually
thin; lobes often distinct, relatively long and narrow,
sometimes short or margin of disk irregular; ribs
often faint or absent. Pistil l-2(-2.5) mm; style
either absent or obscure, or conical, (0.75-)l-1.5
(-1.75) mm, much shorter than the adjacent side(s)
of the drupelet(s); ovary globular to subreniform,
0.4-0.7 by 0.5-0.75 mm, glabrous. Drupelets glob-
ular to somewhat obovoid, ± compressed, 7-9 by
6-8 mm, green to red or blue when fresh, without
persistent petals and stamens at the base; reticulate
pattern rather fine, but often inconspicuous or ob-
scure. Embryo with faintly wrinkled cotyledons.
Distr. Widely ranging in SE. Asia from Nepal to
China; in Malesia: N. Borneo (Sabah) and the Philip-
pines (Luzon).
KEY TO the subspecies
I. Style normally developed, distinctly conical,
(0.75-)l-1.5(-1.75) mm long
a. ssp. parviflora
1. Style usually absent or obscure, the upper part of
the pistil carpel-like, sometimes normally devel-
oped and then up to 0.75 mm
b. ssp. philippinensis
a. .w/7. parviflora - Sabia parviflora Wall. - Sabia
harmandiana Pii.rri. .
leaves oblong, somclimcs oblong-ovatc to (sub)
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-
ing, lax, and with up to more than 35 flowers. Petals
oblong to lanceolate, sometimes oblong-ovate,
2.25-4(-4.5) by 0.7-1.25 mm. Style distinctly con-
ical, (0.75-)l-1.5(-1.75) mm long.
Distr. SE. Asia; in Malesia: Borneo (Sabah), 9
collections.
Ecol. Roadsides, in thickets, and in forests,
mainly 600-2000 m altitude. Fl. Jr. probably
throughout the year.
b. ssp. philippinensis (Robins.) van de Water,
Blumea 26 (1980) 50. - Sabia philippinensis Robins.
- Fig. 4.
Leaves oblong or oblong-ovate to lanceolate,
3-11 by 1-3.5 cm. Cymes 1.5-4.5 cm, 4-20-flow-
ered. Petals elliptic-oblong to sublanceolate, 2-3.5
by 1-1.25 mm. Style absent or obscure and often
carpel-like, sometimes normal-developed and then
up to 0.75 mm. Fruits not seen.
Fig. 4. Sabia parviflora ssp. philippinensis (Robin-
son) VAN DE Water, a. & b. disk and pistil showing
the absence of a style; c. a feebly developed one; all
X 12 (fl Ramos 26973; b Jacobs 7402; c Merrill
7708).
Distr. Malesia: Philippines (Luzon: Benguet
Prov.), 11 collections.
Ecol. Forests, 71000-2100 m. Fl. mainly
Febr. -April.
Vern. Baybayok, kopdas, uakal, udok, Ig.
Notes. Ssp. philippinensis can be distinguished
rather easily from ssp. parviflora by the absence of
a normally developed style. In all the specimens I
have seen (except one) the upper parts of the two
carpels of each flower are not connate with each
other and differentiated into a style as usual, but re-
main free and carpel-like, although the tip of each
carpel is sometimes slightly stigmatic. Moreover, the
margins of the upper part of a carpel are not fused,
so that the upper half of each carpel remains open.
Although this phenomenon is unique within the
genus, I have reduced S. philippinensis to a subspe-
cies of S. parviflora because it agrees very well with
that species in all other main characters.
Like in all Sabia species the leaves are dark above,
paler beneath, but in the present one the contrast is
especially conspicuous. In ssp. philippinensis the
pale margins and undersides of the leaves provide a
useful character to distinguish vegetative specimens
from those of S. pauciflora, another Philippine
species
5. Sabia pauciflora Blume, Mus. Bot. Lugd.-Bat. 1
(1851) 370; MiQ. Fl. Ind. Bat. 1, 2 (1859) 619; Fl.
Arch. Ind. (1870) 72; ibid. (1871) pi. 32; Chen,
Sargentia 3 (1943) 61; van de Water, Blumea 26
(1980) 51. - Sabia papuana Warb. in K.Sch. &
Laut., Fl. Deut. Schutzgeb. Sudsee (1900) 425. -
Sabia reticulata Elmer, Leafl. Philip. Bot. 2 (1909)
579; Merr. Enum. Philip. Fl. PI. 2 (1923) 516;
Chen. Sargentia 3 (1943) 62.
Evergreen woody climber or scandent shrub, up to
20 m. Twigs glabrous; flowering twigs up to 5 mm
diam., glabrous or sparsely pubescent. Buds ovoid,
up to 2.5 mm, acute; scales glabrous to pubescent,
(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-
brous to sparsely pubescent; bracts oblong to
lanceolate, up to 3.5 mm, subglabrous to somewhat
pubescent, (sub)ciliolate; bracteoles as bracts but
smaller, or minute, or sepal-like and then often
situated near calyx; pedicels up to c. 1 cm. Flowers
green to yellow or white. Sepals sometimes 6 (see
bracteoles), ovate to broad-ovate, 0.75-1.25 by
0.7-1 mm, acute to obtuse, glabrous to somewhat
pubescent, (sub)ciliolate. Petals oblong, sometimes
somewhat oblong-ovate, 2.5-4(-4.5) by (0.75-)
1-1.3(-1.5) mm, (narrow-)obtuse, sometimes sub-
ciliolate, nerves up to 5, sometimes dark-coloured
and then conspicuous. Stamens (l-)1.25-1.75(-2)
mm; filament flattened, (0.75-)l-1.5(-1.75) by
0.25-0.5 mm; anther globular to ellipsoid, 0.2-0.3
mm, inflexed. Disk crown-shaped; lobes often short
or irregular; ribs sometimes faint or absent. Pistil
1.3-1.7 mm; style conical, 0.6- 1 mm, much shorter
than the adjacent side(s) of the drupelet(s); ovary
globular to subreniform, 0.5-0.7 by 0.5-0.8 mm,
glabrous. Drupelets ± globular, sometimes some-
what obovoid, compressed, 7.5-11 by 8-10(-ll)
mm, white to red or dark-blue when fresh, without
persistent petals and stamens, reticulate pattern fine
to rather coarse, sometimes indistinct, limited to the
margin or not. Embryo with somewhat undulated or
faintly folded cotyledons.
1989]
Sabiaceae (van Beusekom & van de Water)
689
Distr. Malesia: Moluccas (Buru, Halmaheira,
Batjan), Philippines (Luzon, Negros, Mindanao),
New Guinea; Solomon Islands.
Ecol. Forests, from sea-level up to 2300 m. Fl.fr.
throughout the year.
Uses. Fresh leaves eaten against wound fever in
New Guinea.
Vern. Philippines: bungoi, dadabu. Bag.; New
Guinea: hambui, Poio, Enga lang., kubiakan,
Hagen-Chimbu, Yoowi dial., mongotya ka. North-
ern Prov., pehkuma, Mumuni, Orokaiva lang, pipi,
E. Highlands, pukhabu, S. Highlands.
Note. This species is closely related to S. javanica
from Java and Sumatra, but can be distinguished
from that species by its always few-flowered cymes,
its longer style, and its often ± globular drupelets (S.
javanica often obovoid).
6. Sabia racemosa Chen, Sargentia 3 (1943) 36, f. 2;
VAN DE Water, Blumea 26 (1980) 54.
Evergreen woody climber or scandent shrub, up to
6 m. Twigs glabrous; flowering twigs up to 4 mm
diam., glabrous or somewhat short-pubescent. Buds
ovoid, up to 1.5 mm acute; scales (sub)ciliolate or
not. Leaves ob\ong or somewhat oblong-ovate, 6-25
by 2-10 cm, index 2-3(-3.5), pergamentaceous,
glabrous or with some hairs on midrib, rarely
beneath all over sparsely short-pubescent; base acute
to rounded, apex acute, acuminate; nerves 4-8 (or 9)
pairs, ± patent, curved to straight; petiole up to 2.5
cm, glabrous or with some very short hairs. Cymes
arranged in an axillary, up to 8 cm long, glabrous to
puberulous or short-tomentellous, racemose to thyr-
soid inflorescence, subtended by bracts but often
bracts fallen or sometimes leaf-like, cymes up to 1
cm, I -4(-7)-nowered, glabrous to somewhat puber-
ulous or short-tomentellous; bracts ovate to oblong,
up to 3 mm, glabrous to somewhat pubescent,
(sub)ciliolate; bracteoles as bracts but usually smal-
ler, or minute and then often situated near calyx;
pedicels up to 4 mm. Flowers (pale-)green to yellow.
Sepals ± ovate to broad-ovate, 0.6-1.3 by 0.5-1
mm, acute to obtuse, glabrous to somewhat pubes-
cent, (sub)ciliolate. Petals elliptic-oblong to ovate-
lanceolate, 3.5-6.5 by(1.25-)l.5-2.5 mm, acute to
obtuse, or ± acuminate, or gradually narrowed,
nerves up to 7, thin but distinct. Stamens 1.2-2.2
mm; filament flattened, i -2 by 0.2-0.5 mm; anther
globular to ellipsoid, 0.2-0.3 mm, inflexed. Disk
crown-shaped; lobes sometimes very short or in-
distinct; ribs sometimes faint or absent. Pistil i - 1.5
mm; style ± conical, 0.5-0.9 mm, much shorter
than the adjacent side(s) of the drupclct(s); ovary
globular to subrcniform, 0.5-0.6 by 0.5-0.7 mm,
glabrous. Drupe/e/5 obovoid, ± compressed, 10-12
by (7- )8- 10 mm, while to pink or red when fresh,
without persistent petals and stamens al the base;
reticulate pattern faint to rather coarse, often limited
to the margin. Embryo with somewhat to very
wrinkled or folded cotyledons.
Distr. Malesia: Borneo.
Note. In vegetative characters and somewhat in
the fruit this species resembles S. javanica. It differs,
however, from that species in its inflorescence (few-
flowered cymes) and in its floral characters, especial-
ly the petals.
Since the fruiting collections oi ssp. racemosa bear
only immature or damaged fruit, the description of
the drupelets has mainly been based on the fruit of
ssp. kinabaluensis.
The two subspecies can easily be distinguished
from each other by the difference in the shape of
their petals. Since they can be distinguished from
each other only when flowers are available, the iden-
tification of most of the vegetative and fruiting
specimens has mainly been based on the locality from
where they have been collected.
KEY TO THE SUBSPECIES
1. Petals oblong-ovate to ovate-lanceolate, acute,
somewhat acuminate or tapering to the apex
a. ssp. racemosa
1. Petals elliptic-oblong to oblong, acute to obtuse
b. ssp. kinabaluensis
a. ssp. racemosa - Sabia racemosa Chen.
Sepals 0.6- 1.1 by 0.5- 1 mm. Petals oblong-ovate
to ovate-lanceolate, (3.5-)4.5-6.5 by (1.25-)1.5-
2.5 mm, acute, somewhat acuminate or tapering to
the apex. Pistil 1-1.2 mm; style 0.5-0.7 mm long.
Distr. Malesia: Borneo (Kalimantan), 7 collec-
tions.
Ecol. Low altitudes, up to 100 m. Fl. fr.
throughout the year.
b. ssp. kinabaluensis van de Water, Blumea 26
(1980) 55.
Sepals 0.9-1.3 by 0.6-1 mm. Petals elliptic-
oblong to oblong, 3.5-5 by 1.5-2.5 mm, acute to
obtuse. Pistil 1.2-1.5 mm high; style 0.6-0.9 mm
long.
Distr. Malesia: Borneo (Sabah: Mt Kinabalu),
15 collections.
Ecol. Forests, mainly al 800- 1 500 m altitude. Fl.
fr. throughout the year.
7. Sabia sumalruna Biumi., Mus. Bot. Lugd.-Bat. I
(1851) 370; Miy. II. Ind. Bat. 1, 2 (1859) 619; Fl.
Arch. Ind.(1870)72;;7>/ry. (187l)pl. 33; Kino, J. As.
Soc. Beng. 65, ii (1896) 454; Ridley, Fl. Mal. Pen.
1 (1922) 513; Chen, Sargentia 3 (1943) 39; van de
Water. Blumea 26 (1980) 56.
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.,
glabrous. Leaves elliptic to oblong, sometimes (sub)
lanceolate, (5-)7- 15(- 18) by (1 .5-)2.5-7(- 10) cm,
index 2-3(-4), pergamentaceous, above and be-
neath glabrous; base acute, apex acuminate to sub-
cuspidate; nerves 5-7 pairs, patent, curved to
straight; petiole up to 2 cm, glabrous. Flowers yel-
lowish-green to white, either solitary, sometimes 2 or
3 together, axillary, or arranged in a thyrsoid, ax-
illary, up to 6.5 cm long, glabrous inflorescence;
pedicels up to 2.5 cm, glabrous, with few small
budscales at the base when flowers solitary; bracts ±
oblong-ovate, up to 1 .5 mm long, glabrous, ciliolate;
bracteoles as bracts. Sepals broad-ovate to ovate,
1.25-1.75(-2) by (0.75-)l-1.75 mm, acute to ob-
tuse, glabrous, (sub)ciliolate or not. Petals oblong or
ovate-lanceolate, c. 6- 10 by 1 .5-2.5 mm, sometimes
the upper part somewhat channeled, tapering to the
apex, acute to narrow-obtuse, nerves obscure.
Stamens 2.5-1.5 mm; filament ± flattened, 3-7 by
0.4-0.75 mm; anther ellipsoid, 0.5-0.7 mm,
upright. Disk short-cylindrical, small, the upper part
not enclosing the base of the ovary and without
lobes; ribs ± prominent. Pistil 3.5-c. 7 mm; style
narrow-conical, 3-6 mm, ± half as long as the adja-
cent side(s) of the drupelet(s); ovary somewhat
globular to subreniform, 0.5-0.8 by 0.7-1 mm,
glabrous. Drupelets obovoid, somewhat compress-
ed, 11-13 by 8-9 mm, white to blue when fresh,
without persistent petals and stamens, reticulate pat-
tern absent, often more or less rugged on the outside.
Distr. Malesia: Sumatra (W. Coast Res., Palem-
bang), 7 collections.
Ecol. Forests, 60-1000 m altitude. Fl. May-
Aug.,//-. July-Sept., Febr.
Note. Only a few collections are available. For
that reason no buds and embryos could be described,
whereas the description of the flowers has partly
been based on rather young ones.
Excluded
Sabia densiflora Miq. Sum. (1861) 203, 520 =
Meliosma angulata Blume: K. & V. Bijdr. 9 (1903)
131 = Meliosma simplicifolia (Roxb.) Walp. ssp.
simplicifolia: van Beusekom, Blumea 19 (1971) 476;
Fl. Males. 10" (1989) 698 (this issue).
Sabia floribunda Miq. Sum. (1861) 203, 521 =
Meliosma angulata Blume: K. & V. Bijdr. 9 (1903)
131 = Meliosma simplicifolia (Roxb.) Walp. ssp.
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
Fig. 5. Flower of Meliosma. A. Semi-diagrammatical sketch of flower (subg. Meliosma) with opened outer
petals, but stamens still in bud position. B. Semi-diagrammatical length section of bud (subg. Meliosma). C.
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, rarely minute style, with simple or
somewhat bifid, minute stigma. Ovules 2 (or 1) in each cell, more or less
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 named fossils from the Tertiary are found widely distributed on the northern hemisphere, in
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 leaf or leaflet may be entire or dentate and is often variable. In saplings, watershoots and
seedlings the margin is mostly dentate. In species with pinnate leaves the size of the leaflets mostly increases
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 America Sect. Lorenzanea (Liebm.) Beus.
1 . Leaves pinnate, petiolule of terminal leaflet articulate with the rachis. Sepals mostly 4. Outer petals widely
imbricate, the largest one widely reniform, much wider than long, the smaller ones of irregular shape ±
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 species. SE. Asia and S. Mexico Sect. Kingsboroughia
5. Evergreen trees. Vascular bundle connecting pedicel and seed running freely in the mesocarp. One species.
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,
rarely (sub)globose and then smooth, 6-14 cm diam 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 1 (or 2) leaflets of which the short petiolule is always well markedly articulated
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;
Walp. Ann. 2 (1852) 224; MiQ. Fl. Ind. Bat. 1, 2
(1859) 614; Sum. (1861) 203; van Beusekom, Blumea
19(1971)451,/. 25. - Fig. 7.
For further synonyms, see under the subspecies;
for a complete synonymy, see van Beusekom (1971).
Evergreen shrub or tree, up to c. 15(-22) m. Flow-
ering twigs pubescent when young, glabrescent.
Leaves elliptic or obovate to lanceolate, 2-32 by
0.7-12(-18)cm, inde.\(1.2-)1.5-3(-4), atthebase
acute, at apex acute to caudate, rarely obtuse, usual-
ly entire, sometimes remotely spinously dentate to-
wards the apex, beneath sometimes pubescent on
midrib and nerves, without domatia; nerves 7-15
pairs, usually strongly ascending, petioles usually
rather long, 1-10 cm, 1/5-1/3 as long as the blade.
Panicles usually axillary and erect, widely to usually
narrowly pyramidal, 3-30(-200) cm, usually dense-
ly pubescent, bearing numerous solitary to crowded
flowers which are sometimes spicately arranged;
side-axes usually many, usually short, up to c. 15
(-40) cm, sometimes subtended by normal to small
leaves; bracts ovate to linear-lanceolate, up to c.
2(-6) mm, usually densely pubescent. Pedicels ab-
sent or present, up to c. 3(-5) mm. Mature buds
1.5-3 mm diam. Sepals (4) 5, (round-)ovate, sub-
equal, 1 -2 mm, or the outer 1 or 2 smaller, often one
lowered on the pedicel, all entire, ciliolate. Outer
petals glabrous. Inner petals ± lanceolate and entire,
or bifid, (0.6-)0.8-2.5 mm, glabrous or somewhat
ciliolate at margin or tip, when bifid never with a cen-
tral lobule. Filaments 0.7-1.5 mm. Ovary 0.5-1
mm, very exceptionally pubescent. Fruit (sub)glo-
bose, sometimes elliptic, when ripe 5-10 mm diam.;
endocarp globose to ellipsoid, 6-8(-9) mm diam.,
usually with a slightly elevated rather fine reticulum;
median keel distinct, more or less prominent; ventral
pore whether or not sunken but never spouted.
Distr. SE. & E. Asia; in Malesia (with 4
subspecies): Sumatra, Malay Peninsula, W. Java, N.
Borneo (Sabah), and the Philippines (Luzon, Min-
doro).
Ecol . In evergreen forests under tropical or sub-
tropical conditions, at medium to high altitudes; for
details, see under the subspecies.
Notes . Meliosma lepidota displays a rather wide
variation, especially in the ramification of its pan-
icles which covers almost the whole range of possibil-
ities found throughout Meliosma.
Within M. lepidota seven subspecies are recog-
nized, four of which in Malesia. The differences be-
tween them arc on the same level as in other sub-
species in Meliosma. Transitional forms between
these subspecies, however, occur in only a few cases,
which is logical since there is perfect geographical
isolation between most of them. Sec further the notes
under the subspecies.
key to the subspecies
1. Inner petals distinctly bifid.
2. Leaves 1 .5-2(-2.5) times as long as wide; petiole
1/4-2/3 as long as the blade. Panicles 5-15 cm.
Mature buds 2-2.5 mm diam. Endocarps ellip-
soid to obovoid d. ssp. kinabaluensis
2. Leaves (1.6-)2-3 times as long as wide; petiole
(l/6-)l/5-l/3 (-1/2) as long as the blade.
Panicles 3-30 cm. Mature buds 1 .5-2(-2.2) mm
diam. Endocarps long- to short -ellipsoid (always
distinctly higher than wide) ... .a. ssp. lepidota
1. Inner petals entire, usually lanceolate.
3. Inner petals 2.5 mm. Panicles distinctly axillary
or ramiflorous. Mature buds 2.5-3 mm diam.
b. ssp. dolichomischa
3. Inner petals 1-1.5 mm. Panicles terminal or
crowded at the end of the twigs, rarely distinctly
axillary. Mature buds 2-2.5 mm diam.
c. ssp. vulcanica
a. ssp. lepidota. - Meliosma lepidota Blume, Rum-
phia 3 (1849) 199; Walp. Ann. 2 (1852) 224; MiQ. Fl.
Ind. Bat. 1, 2 (1859) 614; Sum. (1861) 203; Illustr.
(1871) 73. - Meliosma pedicellata K. & V. Bijdr. 9
(1903) 134; Koord. Exk. Fl. Java 2 (1912) 545; Atlas
2 (1914) t. 379; Backer & Bakh./. Fl. Java 2 (1965)
144.
Leaves oblong, sometimes somewhat ovate-
oblong, rarely elliptic, 5-26 by (1.5-)2-12 cm, en-
tire, base acute, apex acute to caudate, glabrous
when mature; nerves 8-12(-14) pairs; petiole 1.5-6
cm. Panicles axiWary, rarely terminal or ramiflorous,
often several together near the end of a branch, 3-30
cm, rather poor and lax, ramified up to the 2nd
order; primary (essentially secondary!) side-axes
short, up to c. 6(-10) cm. Mature buds 1.5-2 mm
diam. Inner petals about halfway bifid, 0.7-1 mm;
lobes rather narrow. Endocarp obovoid to ellipsoid,
(8-)9-14 mm long, 5.5-8 mm diam., with or
without rather wide and feeble reticulum; median
keel distinct, more or less prominent, blunt to rather
sharp, at one or both ends running out into a ventral,
often somewhat beak-like processus; ventral pore
rather wide, somewhat sunken.
Distr. Malesia: Sumatra (not uncommon in
Aceh, Tapanuli, and West Coast), W. Java.
Ecol . Primary montane rain-forest; 900-26(X) m
altitude in Sumatra, 1050-1600 m in Java.
Field notes . Outer bark dark brown, finely cor-
ky, 0.5 mm; inner bark turning rcdbrown, 0.5 cm;
wood ochre with reddish stripes.
Vern . Sumatra: antuang, hontuang, Batak lang.,
Toba, kalompang bag^h, CJn. Talamau.
Note. .Ssp. lepidota is similar and probably most
closely related to the adjacent s.<!p. dolichomi.\cha
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.
longipes (Merr.) Beus. from Vietnam, from which
it can sometimes only be distinguished by the shape
of the endocarp.
b. ssp. dolichomischa (Vidal) Beus. Blumea 19
(1971) 458, f. 25. - Meliosma dolichomischa Vidal,
Not. Syst. 16 (1960) 304. - Meliosma monophylla
Ridley, J. Str. Br. Roy. As. Soc. n. 54 (1910) 40,
nom. illeg.,non Merr. (1909); Fl. Mai. Pen. 1 (1922)
514; Vidal, Not. Syst. 16 (1960) 306. - Fig. 7.
Leaves elliptic to oblong, 4-22 by 2-10 cm, en-
tire, base mostly attenuate, apex usually cuspidate,
glabrous or subglabrous, nerves 7-13 pairs, petiole
(l-)3-10 cm. Panicles axillary or ramiflorous,
solitary or a few together, 6-25 cm, rather poor and
lax, ramified up to the 2nd or 3rd order; primary (es-
sentially secondary!) side-axes up to c. 10 cm.
Mature buds 2.5-3 mm diam. Inner petals lanceo-
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.
Distr. Malesia: Malay Peninsula (Pahang:
Eraser's Hill, Cameron Highlands; Perak: Hermi-
tage Hill, once).
Ecol. Primary montane rain-forest, c. 1200-
1500 m altitude.
Field notes. Bark thick, red. Wood first white
when cut, darkening to orange-brown. Leaves
glaucous below.
c. ssp. vulcanica (Merr.) Beus. Blumea 19 (1971)
460. - Meliosma vulcanica Merr. Philip. J. Sc. 11
(1916) Hot. 15; Enum. Philip. Fl. PI. 2 (1923) 518. -
Machilus nervosa Merr. Philip. J. Sc. 4 (1909) Bot.
262; Enum. Philip. Fl. PI. 2 (1923) 189; Salvore &
Lagrimas, Philip. J. For. 4 (1941) 309; c/. Kosterm.
Reinwardtia 5 (1960) 377; Bibl. Laur. 1 (1964) 919.
- .Meliosma boniocensis Merr. Philip. J. Sc. 20
(1922) 403; Enum. Philip. Fl. PI. 2 (1923) 517;
Kosterm. Reinwardtia 5 (1960) 377.
Leaves obovate-oblong or oblong, 5-16 by 2-6
cm, base acute, apex acute to acuminate or
sometimes rounded, glabrous or subglabrous; nerves
8-11 pairs; petiole 2-4 cm. Panicles terminal,
sometimes axillary, 3-20 cm, rather profuse to poor,
ramified up to the 3rd or 4th order; primary side-axes
(mostly essentially primary!) up to c. 15 cm, usually
subtended by normal to reduced leaves. Mature buds
2-2.5 mm diam. Inner petals lanceolate, 1-1.5 mm,
entire, sometimes frayed at the tip. Endocarp subglo-
bosc, rather oblique, 6-7 mm diam., apart from a
few ribs smooth, median keel distinct, rather promi-
nent, at one end running out into a minute ventral
processus; ventral pore somewhat sunken.
Distr. Malesia: Philippines (Luzon, Mindoro).
Ecol. Primary rain-forest, low altitude up to c.
2000 m.
Note . Ssp. vulcanica is the only subspecies of M.
lepidota in which normal terminal panicles have been
found. In general habit it is more similar to certain
forms of ssp. longipes from Vietnam than to .ssp.
squamulata (Han* e) Bius. from Taiwan or to ssp.
kinabatuensis from Borneo, to which i( is obviously
less closely related.
d. ssp. kinabaluensis Beus. Blumea 19 (1971) 455. -
Meliosma pedicellata {nan K. & V.) Merr. & Perry,
J. Arn. Arb. 20 (1939) 356.
leaver elliptic, rarely oblong, 3-15 by 1.5-9 cm,
usually entire, base acute to rounded and somewhat
attenuate, cuspidate, above glabrous or ± pubescent
on the midrib, subglabrous beneath, usually with a
white waxy layer beneath, which gives a glaucous ap-
pearance; nerves 8-14 pairs; petiole 1-7.5 cm.
Panicles terminal or axillary, solitary or a few togeth-
er, 5-15 cm, rather poor and lax, ramified up to the
2nd (3rd) order; primary (essentially secondary!)
side-axes up to c. 8 cm. Mature buds 2-2.5 mm
diam. Inner petals halfway or somewhat less bifid,
1-1.2 mm; lobes rather narrow. Endocarp ±
obovoid, c. 8 mm long, c. 6 mm diam., with rather
wide and feeble reticulum; median keel only slightly
elevated, blunt, at one end running out into a minute
ventral processus; ventral pore wide, not sunken.
Distr. Malesia: Borneo (Mt Kinabalu).
Ecol. Montane forest, 1700-2700 m altitude.
Field notes. The lower surface of the leaves is
often said to be white to light grey; in herbarium
specimens indeed a whitish waxy layer can be ob-
served often. The general colour of the leaves is re-
ported to be glaucous.
Note. Ssp. kinabaluensis has a very low degree of
variability, a characteristic which is also found in
some other subspecies of M. lepidota. It is most
similar to ssp. dolichomischa from the Malay Penin-
sula and to ssp. lepidota from Sumatra, with which
it shares, amongst others, the more or less ellipsoid
endocarp; all other subspecies have (sub)globosc en-
docarps.
2. Meliosma simplicifolia (Roxb.) Waip. Rep. 1
(1842) 103; HA.ssK.Cat. Hon. Bog. (1844) 226; Miy.
Fl. Ind. Bat. 1, 2 (1859) 613; Sum. (1861) 203; van
BhUSEKOM, Blumea 19(1971)462, f. 28. Milling-
tonia simplicifolia Roxb. [Horl. Bcng. (1814) 3,
nomen] PI. Corom. 3 (1820) 50, t. 254. FIr. 8. 10.
For further synonyms, see under the subspecies;
for a complete synonymy, see van Beusekum ( 1 97 1 ).
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
cuneate, apex acute to acuminate, rarely caudate or
rounded, entire to spinously dentate, sometimes with
hairy domatia; nerves 7-25 pairs, ± ascending,
sometimes looped; petiole 0.5-6(-7) cm, 1/20-1/3
as long as the blade. Panicles terminal, very rarely
axillary, erect, lax to rather dense, widely to narrow-
ly pyramidal, (4-)10-50(-60) cm, usually profusely
branched up to the 2nd-4th order, bearing numer-
ous solitary to crowded or glomerulate flowers which
are usually spicately arranged; primary side-axes
usually many, up to c. 25 cm, often subtended by
leaves; bracts ovate to linear-lanceolate, up to c. 8
mm. Pedicels sometimes present, up to c. 3 mm.
Mature buds (1-) 1.5-3 mm diam. Sepals (4) 5,
sometimes by addition of empty bracts seemingly
more, up to 11(-13), (round-)ovate, equal or usually
more or less unequal, the inner ones 0.7-2 mm, the
outer one(s) smaller, often minute. Inner petals more
or less deeply bifid, 0.5-1.5 mm, with glabrous,
sometimes fimbriate or ciliolate lobes, never with a
central lobule. Filaments 0.5-1.5 mm. Ovary
0.5-0.7(-l) mm. Mature fruit (sub)globose, 4-10
mm diam.; endocarp globose to subglobose, often
depressed or oblique, 3-9 mm diam., with very
vague to very strong and prominent reticulum; me-
dian keel more or less prominent; ventral pore
somewhat sunken to somewhat spouted.
Distr. Continental SE. Asia (from Ceylon to
China, Taiwan and S. Japan); in Malesia: Sumatra,
Malay Peninsula, Borneo, Java, and Lesser Sunda
Islands. Fig. 9.
Ecol. Subtropical to tropical forests, under
various conditions, usually in mountains up to c.
3000 m, but also at sea-level. For details see under the
subspecies.
Note. Meliosma simplicifolia is a very variable
species, covering an enormous area in which it is
adapted to many different habitats. It can be divided
into eight well-marked subspecies, five of which cen-
tre in SW. Yunnan, and diverge over different parts
of the area.
KEY TO THE SUBSPECIES
1. Sepals (4-)5.
2. Panicles branched up to the 2nd (3rd) order,
nearly always (very) densely tomentose; primary
side-axes rarely subtended by leaves. Leaves
sparsely to densely but always distinctly pubes-
cent to tomentose, at least on midrib and nerves;
without domatia. Style c. 1.5-2 times as long as
the ovary b. ssp. rigida
2. Panicles branched up to the (2nd) 3rd or 4th
order, sparsely pubescent to moderately tomen-
tose; lower primary side-axes often subtended by
normal to small or reduced leaves. Leaves gla-
brous to densely pubescent, rarely tomentose,
with or without domatia. Style about as long as
the ovary or shorter.
3. Leaves with or without domatia; midrib on the
upper side of the full-grown leaf glabrous or
nearly so, more or less prominent, rarely flat.
Inner petals with entire lobes, which are
sometimes slightly fimbriate or ciliolate at the
very tips. Endocarps 3.5-5(-7) mm diam.
a. ssp. simplicifolia
3. Leaves with or without domatia; midrib on the
upper side of the full-grown leaf more or less
but distinctly pubescent, ± impressed to flat.
Inner petals usually with fimbriate, rarely entire
lobes which are rarely minutely ciliolate at the
very tips. Endocarps (4.5-)5.5-8 mm diam.
c. ssp. fruticosa
1. Sepals (8-)9-ll(-13). Leaves usually with do-
matia. Endocarps 3.5-5.5 mm diam.
d. ssp. pungens
a. ssp. simplicifolia. - Millingtonia simplicifolia
RoxB. [Hort. Beng. (1814) 3, nomen] PI. Corom. 3
(1820) 50, t. 254; Fl. Ind. 1 (1820) 103; Nees, Flora
8 (1825) 106; Griff. Not. PI. As. (1854) 162; Ic. PI.
As. (1854) t. 442; Anon. Ic. Roxb. 4 (1970) 40, t. 20;
VAN Beusekom, Blumea 19 (1971) 476. - Meliosma
simplicifolia Walp. Rep. 1 (1842) 103; Hassk. Cat.
Hort. Bog. (1844) 226; Thw. Enum. PI. Zeyl. (1858)
59; MiQ. Fl. Ind. Bat. 1, 2 (1859) 613; Sum. (1861)
203; Illustr. (1871) 73; Bedd. Fl. Sylv. 3 (1871) 77;
Brandis, For. Fl. (1874) 116; Hook./. Fl. Brit. India
2 (1876) 5; KuRZ, J. As. Soc. Beng. 45, ii (1876) 204;
Fl. Burma 1 (1877) 301; Trim. Fl. Ceyl. I (1893) 315;
Prain, Bengal PI. 1 (1903) 246; Brandis, Indian
Trees (1906) 194; Merr. Contr. Arn. Arb. 8 (1934)
95; Brittonia 4 (1941) 110; Vidal, Not. Syst. 16
(1960) 307. - Meliosma angulata Blume, Rumphia 3
(1849) 197; Walp. Ann. 2 (1852) 224; K. & V. Bijdr.
9 (1903) 131; Koord. Exk. Fl. Java 2 (1912) 545;
Atlas 2 (1914) t. 378; Baker/, in Rendle, J. Bot. 62
(1924) Suppl. 30; Vidal, Not. Syst. 16 (1960) 304. -
Sabia densiflora Miq. Sum. (1861) 203, 520. - Sabia
floribunda Miq. I.e. 203, 521; Kurz, J. As. Soc.
Beng. 39, ii (1870) 74. - Fig. 8d.
Leaves obovate-oblong to -lanceolate, up to c. 50
by 18 cm, base cuneate, apex acute to short-cuspi-
date, beneath often with domatia; nerves 8-23 pairs.
Panicles rather lax, 10-45 cm, branched up to the
3rd or 4th order; axes sparsely to densely pubescent
but never tomentose, the lower primary ones sub-
tended by leaves. Flowers more or less crowded to
solitary, (sub)sessile; mature buds 1.5-2 mm diam.
Sepals 5 (4). Inner petals 0.6-0.8 mm, usually over
halfway bifid, lobes more or less divergent, narrow,
glabrous, sometimes slightly fimbriate or ciliolate at
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-
ly triangular at ventral view, 3.5-5(-7) mm diam.,
with more or less prominent, rather coarse reticu-
lum; median keel usually very prominent, at one end
sometimes running out into a minute ventral pro-
cessus; ventral pore somewhat or not sunken, not
spouted.
Distr. Widely distributed in continental SE.
Asia; in Matesia: northern half of Sumatra, W. Java
(not found since Blume's time). Fig. 9.
Ecol. Primary and secondary evergreen forest,
from sea-level up to c. 1200<- 1500) m altitude. It is
often reported to occur along watercourses.
Vern. Sumatra: medanf; sungu, M, simulingga,
sumpa mana belawah, Karo, kayu gadis. West
Coast.
Note. A rather uniform, well recognizable sub-
species all over its area.
b. ssp. rigida (Sieb.&Zucc.)Beus. Blumca 19(1971)
473. Meliosma rigida Sieb. & Zurr. Abh. K.
(Bayer.) Ak. Wiss. M. Ph. Kl. Munchcn 4, 2 (1845)
153; Mio. Ann. Mus. Hot. Lugd.-Bat. 3 (1867) 93;
Cat. Mus. Bot. I (1870) 23, incl. var. angusli/ulia
MiQ., nomen; Maxim. Bot. Jahrb. 6 (1884) 60;
Forbes&Hemsley, J. Linn.Soc. Bot. 23 (1886) 145,
p.p., excl. M. pungens; Dunn, J. Linn. Soc. Bot. 38
(1908) 358; Hayata, Ic. PI. Formos. 1 (1911) 161
Dunn & Tutch. Kew Bull. Add. Ser. 10 (1912) 68
Chun, Sunyatsenia 1 (1933) 180; Hand.-Mazz
Beih. Bot. Centralbl. 52(1934) 166; Kaneh. Formos
Trees ed. 2 (1936) 416, f. 372; Cufod. Oest. Bot. Z
88 (1939) 267, incl. var. patens; Hara, Enum
Sperm. Japon. 3 (1954) 121; Making, III. Fl. Jap
(1954) 348, f. 1044; Walker, Imp. Trees Ryukyu Is
(1954)200, f. 121; How, Acta Phytotax. Sin. 3 (1955)
444; Gagnep. & Vidal, Fl. Camb. Laos & Vietnam
1 (1960) 47, in obs.; Liu, III. Lign. PI. Taiwan 2
(1962) 925, f. 762; Li, Woody Fl. Taiwan (1963) 503;
Ohwi, Fl. Japan (1965) 613. - Quercus jama-buwa
SiEB. in sched. ex Miq. Ann. Mus. Bot. Lugd.-Bat.
3 (1867) 93, nom. inval. - Meliosma pungens auct.
non (W. & A.) Walp.: Hook./. Fl. Brit. India 2
(1876) 4, p.p., quoad pi. Japon. - Melio.sma patens
Hkmsley ex Forbes &. Hemsley, J. Linn. Soc. Bot.
23 (1886) 145. Meliosma harmandiana Pierre,
II. For. Cochinch. 5 (1897) I. 360. - Meliosma
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.
10(1915) Bot. 38; Enum. Philip. Fl. PI. 2 (1923) 517.
- Meliosma pannosa Hand.-Mazz. Anz. Ak. Wiss.
Wien M.-N. Kl. 58 (1921) 179; How, Acta Phytotax.
Sin. 3 (1955) 442; Gagnep. & ViDAt, Fl. Camb. Laos
& Vietnam 1 (1960) 50, in obs., p.p. - Meliosma
costata CuFOD. Oest. Bot. Z. 88 (1939) 266; How,
Acta Phuotax. Sin. 3 (1955)444; Gagnep. & Vidai,
Fl. Camb. Laos & Vietnam 1 (1960) 45; Vidax, Not.
Syst. 16 (1960) 304. - Meliosma evrardii Gagnep.
Not. Syst. 14(1952) 212, p.p.
Leaves usually obovate-oblong to obovate-lanceo-
late, sometimes oblong to lanceolate, 4-25(-32) by
1.5-8(-ll) cm, base long-cuneate to acute, apex
acute to cuspidate, without domatia; nerves 7-19
pairs. Panicles lax to rather dense, 10-30 cm,
branched up to the 2nd (3rd) order; axes more or less
tomentose, sometimes woolly-pubescent, the pri-
mary ones only exceptionally subtended by small
leaves. Flowers more or less crowded, usually in
dense glomerules, sessile; mature buds 1.7-2.2 mm
diam. Sepals 5 (4). Inner petals 0.6-O.S mm, usually
less than halfway bifid, lobes hardly or not
divergent, rather narrow, fimbriate or ciliolate at the
tips. Style about (1 .5-)2 times as long as ovary. En-
docarps (sub)globose, not or not much oblique,
(3.5-)4-5 mm diam., with fine reticulum; median
keel blunt to rather sharp, hardly to distinctly promi-
nent, often at one end running out into a minute ven-
tral processus or tubercle; ventral pore not sunken,
often somewhat spouted.
Distr. Widely distributed in continental SE.
Asia, incl. China, Laos, S. Vietnam (only at Hue),
Taiwan (incl. Pescadores), Ryu Kyu Islands, Japan;
in Malesia: Philippines (Luzon: Mountain Provi-
nce). Fig. 9.
E c o I . In evergreen broad-leaved or laurophyllous
forests, on different soils; in dry as well as in wet
places; altitude usually 100-1000 m, but in Luzon
reported from 1200- 16(X) m.
Field notes. Bark grey, smooth. Branches
brown. Leaves lustrous green above, sometimes
glaucous beneath. Fruit blue-purple to purplish
black.
Vern. Philippines: gahatan. If., Luzon; lasuit,
Bondoc dial.
Notes. Ssp. rigida is variable in quite some
characters in its area outside Luzon, for instance in
the degree of pubescence, leaf shape, and dentation.
In continental SE. Asia the area of ssp. rigida bor-
ders on or overlaps the areas of five or six other
subspecies of M. simplici/olia, which substantially
adds to the chance of confusing them, several
specimens being hybrids, it is probable that these
subspecies are ecologically isolated to a large extent
and thus contact between them is prevented.
Quercus gilva var. prucera Blumf:, Mus. Bot.
Lugd.-Bat. 1 (1850) 306, was included in the synony-
my of M. rigida by Hara, I.e., but I found it to
belong to Quercus gilva Blume.
c. ssp. fruticosa (Blume) Beus. Blumea 19 (1971)
477, f. 28. - Meliosma fruticosa Blume, Rumphia 3
(1849) 198; Walp. Ann. 2 (1852) 224; MiQ. Fl. Ind.
Bat. 1,2 (1859) 614; Illustr. (1871) 73; K. & V. Bijdr.
9 (1903) 133; Koord. Exk. Fl. Java 2 (1912) 545. -
Meliosma elliptica Hook./. Fl. Brit. India 2 (1876) 5,
p.p., excl. Sabia floribunda Miq.; King, J. As. Soc.
Beng. 65, ii (1896) 456; Ridley, J. Str. Br. Roy. As.
Soc. n. 33 (1900) 67; Fl. Mai. Pen. 1 (1922) 514;
BuRK. & Hend. Card. Bull. S. S. 3 (1925) 364. -
Meliosma lancifolia Hook./. Fl. Brit. India 2 (1876)
5; King, J. As. Soc. Beng. 65, ii (1896) 456; Ridley,
Fl. Mai. Pen. 1 (1922) 514. - Meliosma monophylla
Merr. Philip. J. Sc. 4 (1909) Bot. 286; Enum. Philip.
Fl. PI. 2 (1923) 517; Vidal, Not. Syst. 16 (1960) 306.
- Fig. 8c, 10.
Leaves usually oblong to lanceolate, 5-40(-45) by
2-15 cm, base acute, apex acute to acuminate,
densely pubescent on midrib and sometimes on
nerves and lamina, beneath glabrous to tomentose,
sometimes with domatia; nerves 7-25 pairs. Panicles
usually lax, sometimes more dense, 10-50 cm,
branched up to the 3rd (4th) order; eixes pubescent to
short-tomentose, the lower primary ones subtended
by small leaves or not. Flowers more or less crowded
to solitary, (sub)sessile; mature buds 1.5-2 mm
diam. Sepals 5 (4). Inner petals c. 0.7 mm, about
halfway or somewhat less bifid; lobes divergent or
not, usually rather narrow, more or less fimbriate,
sometimes entire. Style about as long as ovary or
shorter. Endocarps globose or ± ellipsoid,
(4.5-)5.5-8 mm diam., with rather wide, coarse
reticulum; median keel prominent, at one end often
running out into a minute ventral processus; ventral
pore not or not much sunken, not spouted.
Distr. S. Peninsular Thailand (Sural) and
Taiwan; in Malesia: common in the Malay Peninsula
and Sumatra; W. Java, Lesser Sunda Islands (Sum-
bawa, Flores), Borneo (Central Sarawak, Kinabalu,
W. Kutai), and the Philippines (Luzon). Fig. 9.
Ecol. Primary rain-forest, on various soil types,
reported to occur on limestone, sand, volcanic loam,
and andesite; altitude from sea-level up to 24(X) m.
Field notes. Bark smooth, grey to brown, len-
ticellatc, paperthin. Inner bark pale brown to dark
brownrcd. Wood reddish to rcdbrown. Fruit yellow
to pale red when ripening, dark red to brown when
ripe.
Vern. (all once noted). Malay Peninsula: bua
palu, Sclangor, medang kerkulu, mengading, Malac-
ca; Sumatra: lelagan, Gajo lang., Aceh, kaju djarap,
k. gasir, k. si raga, Asahan, kaju ardong ardong,
Toba, kabung kabung, Tapanuli, masadih pajo.
702
Flora Malesiana
[ser. I, vol. 10^
Simalur, kendung, Palembang, redjang, Djambi;
Java: ki (iwu, Preanger; Flores: kaju sar, Philip-
pines: malaligas. Tag.
d. spp. pungens (Wall, ex W. & A.) Beus. Blumea
19 (1971) 466. - Millingtonia pungens Wall, ex
W. & A. Edinb. New Phil. J. 15 (1833) 178; Prod. 1
(1834) 115; Wight, Ic. 3 (1845) t. 964/3. - Meliosma
pungens (Wall, ex W. & A.) Walp. Rep. 1 (1842)
423; Ann. 1 (1848) 135; Thw. Enum. PI. Zeyl. (1858)
59; Bedd. F1. Sylv. 3 (1871) 77; ibid. t. 160; Merr.
Contr. Am. Arb. 8 (1934) 94; Vidal, Not. Syst. 16
( 1 960) 306. - Meliosma wightii Planch, ex Brandis,
For. Fl. (1874) 116;HooK./. Fl. Brit. India 2 (1876)
4. - Fig. 8a.
Leaves elliptic to oblong, sometimes lanceolate,
5-20(-30) by 2-8(-10) cm, without or with some
distant teeth, acute to rounded at the base, acute to
acuminate at the apex, usually distinctly pubescent
on midrib and sometimes on nerves above, sparsely
to moderately pubescent beneath especially on mid-
rib and nerves, usually with domatia; nerves 7-18
pairs. Panicles lax to dense, (5-) 10-55 cm, branched
up to the 2nd (3rd) order; axes rather coarse, densely
short-tomentose, the lower primary ones almost
always subtended by small leaves. Flowers crowded
in dense glomerules, sessile; mature buds 2-2.5 mm
diam. Sepals {8-)9-ll(-\i). Inner petals c. 1 mm,
slightly bifid; lobes divergent, wide, glabrous. Style
about as long as ovary. Endocarps (sub)globose,
often rather irregular, 3.5-5.5 mm diam., with
usually lax reticulum; median keel distinct but not
very prominent, not running out into a ventral pro-
cessus; ventral pore hardly or not sunken, not spout-
ed.
Distr. Sri Lanka and Deccan Peninsula; in
Malesia: N. Sumatra (Gajo Lands, Takengon), one
collection. Fig. 9.
Ecol. Mountain forest, 1500-2000 m altitude.
3. Meliosma sumatrana (Jack) Walp. Ann. 1 (1848)
135; MiQ. Fl. Ind. Bat. 1, 2 (1859) 617; Sum. (1861)
203; lUustr. (1871) 75; Hook./. Fl. Brit. India 2
(1876) 6; KooRD. Minah. (1898) 408; Suppl. Cel. 2
(1922) 7, t. 56; ibid. 2 (1922) 28; Merr. Enum. Born.
(1921) 363; Enum. Philip. Fl. PI. 2(1923) 518; Contr.
Arn. Arb. 8 (1934) 95; Merr. & Perry, J. Am. Arb.
20(1939) 357; van Beusekom, Blumea 19 (1971) 485.
- Millingtonia sumatrana Jack, Mai. Misc. 2 (7)
(1822) 30; Hook. J. Bot. 1 (1834) 378; Merr. J. Arn.
Arb. 33 (1952) 236. - Meliosma nitida Blume, Cat.
(1823) 32; Nees, Flora 8 (1825) 106; Hassk. Tijd.
Nat. Gesch. Phys. 10 (1843) 139; Cat. Hort. Bog.
(1844) 226; Blume, Rumphia 3 (1849) 202, t. 169,
incl. var. tridenta Blume, var. cerasiformis Blume et
var. splendens Blume; Walp. Ann. 2 (1852) 225;
Miq. Fl. Ind. Bat. 1, 2 (1859) 617; Sum. (1861) 203,
520; Illustr. (1871) 74; King, J. As. Soc. Beng. 65, ii
(1896) 457; K. & V. Bijdr. 9 (1903) 117; Koord. Exk.
Fl. Java 2 (1912) 546, f. 81; Atlas 2 (1914) 377;
Ridley, Fl. Mai. Pen. 1 (1922) 515; Baker/, in Ren-
dle, J. Bot. 62 (1924) Suppl. 30; Burk. & Hend.
Card. Bull. Str. Settl. 3 (1925) 364; Heyne, Nutt. PI.
(1927) 1002; Merr. & Perry, J. Arn. Arb. 20 (1939)
357; Backer & Bakh./. Fl. Java 2 (1965) 145. - Irina
integerrima Blume, Bijdr. (1825) 231, non Hassk.
PI. Jav. Rar. (1848) 284 ('Iriney, Walp. Rep. 1
(1849) 416; Blume, Rumphia 3 (1849) 202, in syn.
sub M. nitida. - Millingtonia nitida Schult. &
ScHULT. Syst. Veg. Mant. 3, add. 2 (1827) 250;
DiETR. Syn. PI. 1 (1839) 103. - Meliosma confusa
Blume, Rumphia 3 (1849) 200; Walp. Ann. 2 (1852)
225; Miq. Fl. Ind. Bat. 1, 2 (1859) 616; Sum. (1861)
203, 520; Illustr. (1871) 74. - Meliosma cuspidata
Blume, Rumphia 3 (1849) 202; Miq. FI. Ind. Bat. 1,
2 (1859) 617; Illustr. (1871) 74; Hall./ Meded.
Rijksherb. 1 (1910) 2; Merr. Enum. Born. (1921)
362. - Meliosma pinnata (non Walp.) Koord.
Minah. (1898)408. - Meliosma diepenhorstii Valet.
Ic. Bog. 2 (1904) 195, t. 150. - Meliosma elmeri
Merr. PI. Elm. Born. (1929) 177. - Meliosma
philippinensis Merr. & Perry, J. Arn. Arb. 20
(1939) 357.
Evergreen tree, up to 15-20(-25) m. Leaves
2-5(-6)-jugate; rachis terete, 6-50 cm, including
the up to c. 25(-30) cm long petiole, up to c. 10(- 15)
mm across, rarely slightly pubescent, usually with
distinctly swollen base; leaflets usually elliptic to
lanceolate, (3-)5-35(-50) by (1.5-)2.5-I5(-20)
cm, base cuneate to rounded, shortly narrowed into
the petiole, apex acuminate to caudate, usually en-
tire, beneath rarely more or less pubescent, without
domatia; midrib slightly prominent above; nerves
(5-)7-13(-19) pairs, ascending, nearly always
looped and joined; petiolules very short or up to c.
6 cm, usually distinctly swollen at the base especially
in older leaves. Panicles usually terminal, usually
narrowly, sometimes widely pyramidal, 7-50(-75)
cm, usually profusely branched up to the 4th order,
rather stiff and coarse, puberulous, bearing numer-
ous crowded flowers; primary side-axes usually
rather short, up to c. 30 cm, the lower ones excep-
tionally subtended by small to reduced leaves; bracts
ovate to narrowly triangular, up to c. 6 mm, ±
puberulous. Pedicels absent or short, up to c. 2 mm.
Mature buds (1 .5-)2-3(-3.5) mm diam. Sepals 5 or
4, ovate, unequal, the inner 3 or 4 c. 1-2 mm, the
outer 1 or 2 usually smaller, often minute, sometimes
lowered on the pedicel, sometimes puberulous out-
side, especially the outer ones, with entire or 2- or 3-
lobed, often ciliolate margin. Outer petals glabrous.
Inner petals elliptic to lanceolate or strap-shaped
with wide-truncate tip, (1.2-)1.5-2(-3) mm, acute
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-
soid, when ripe 1-3 cm diam., with rather thick
spongy to pulpy mesocarp; endocarp ellipsoid, some-
times nearly globose, 0.7-2 cm diam., with almost
smooth to somewhat lumpy surface, often with a few
faint to sharply prominent ribs; median keel distinct,
slightly elevated to sharply prominent, at one end
often running out into a more or less prominent curv-
ing, at the other end sometimes into a minute tuber-
cle; ventral pore mostly rather wide, usually some-
what sunken.
Distr. Malesia: Sumatra (incl. Nias, Batu &
Sipora Is., Banka), Malay Peninsula (incl. Penang
I.), Anambas Is., W. half of Java, throughout Bor-
neo, Sulawesi, and the Philippines (Mindanao,
Palawan). Common. Fig. II.
Lcol . Primary and secondary lowland and mon-
tane rain-forest, up to c. 2200 m altitude. Found on
various soils, fertile as well as infertile, in dry to wet
localities, in dense to open forests, by streams as well
as on hilltops and ridges.
Field notes. Often a crooked tree, irregularly
branched. Trunk sometimes with small buttresses.
Bark surface grey to brown, smooth, with lenticels.
often with shallow fissures, sometimes said to be
dimpled, patchy or scaly. Inner bark 0.5- 1 cm thick,
soft, fibrous, light yellow or dirty white, soon turn-
ing pink, brownish, reddish, or rusty after exposure.
Sapwood said to be whitish, yellowish, creamy
orange, or brownish. Sap without special smell or
taste. Leaves bright green on both sides. Flower col-
our varying from white, cream, or greenish, to partly
or entirely pinkish to red. Fruit first yellow, then
yellow with red to red when ripe; pulp white, turning
quickly blood-red on exposure, finally becoming
black, sweetish to tasteless.
Uses. The species was proposed by Koorders &
Valeton, I.e., for reforestation purposes. In Min-
danao the triturated bark and leaves are several times
reported to be in use as a medicine applied for
wounds, to soothe itchy skin or - charred and put in
water - against tympanites. It was also .said to be
used in agricultural rituals. The fruits are many times
reported to be edible.
Vern. Malay Peninsula: pa-on^, Saki name, and
mennadinfi hesar, both from Pahang, huah mata
ikan, Icmuan, Sclangor, pokoh haran, Negri Scm-
bilan, kaju kahwa kanlu, membuloh, pokoh gra6
704
Flora Malesiana
[ser. I, vol. 10^
jantan, p. mala gajah, p. paigigi, p. pinang plandog,
p. ravoa antoo, pudding utan, Malacca, pelanlu;
Sumatra: laon, si paturut, sringkut, Karo country;
kaju durung durung, k. ining ining, Tapianuli, tarn-
pa bussie, Priaman, marazat, Mt Kerinci; Java: ki
liwu (landuk) (bodas), S, ki huut, Udjong Kulon;
Borneo, Sarawak: buliliap, Kenyah dial., maiak,
Kayan dial., buiu manuk, Iban, bilonok, Dyak;
Sabah: bung lai, Sungai, gapas gapas, kapas kapas,
keriyan, Dusun, illulal, limpangol, tunjang, Murut;
SE. Borneo: tambalilin, tandao, Dyak, Tidung dial.;
djangkanggunung, Bandjar lang., Riom dial.;
Sulawesi: see Koord. Minah. (1898) 408; eng-
golokia, W. Toradja dial., pulu putu, situi, Tobela
lang., Malili, pobumengo, Gorontalo; dama, Torai
dial., Menado; Philippines: carabo-rabo, daborabo,
kadabudabo, karabu-rabu, magobaylung, mahag-
kol, yagabogan, Mbo, Buk., bentinguasay, gepulu,
Zamboanga, waat, Cebuano, Mt Apo, salalab,
Moro dial., garong, gimbingimbing. Sub., sumaga-
sa, Bag.
Notes. Meliosma sumatrana is very constant in
its discriminative characters (especially the promi-
nent midrib and entire inner petals), but there is
nevertheless some geographical variation, especially
in the northern part of Borneo (Sarawak, Sabah). As
general tendencies may be noticed that towards the
centre of the area leaflets and fruits increase in size
and dentate leaflets become more common. More-
over, the number of leaflet pairs decreases when the
leaflets are larger.
Sterile hybrids between M. sumatrana and M. pin-
nata ssp. ridieyi are rarely found (Sabah).
DocTERS VAN Leeuwen (Zoocecidia Neth. East
Indies, 1926, 339, f. 612) described a leaf-gall on a
specimen from Sulawesi. This type of galls (usually
ball-shaped, c. 4 mm, ending in a short mucro, and
surrounded by a calyx-like circumvallation) is rather
commonly met with in this species, not only in
specimens from Sulawesi, but also from Borneo,
Sumatra, and the Malay Peninsula. The galls do not
only occur on the lower surface of the leaflets, but
occur also on the upper surface, and on rachis and
petiolules, often very many crowded together.
4. Meliosma lanceolata Blume, Cat. (1823) 32; Nees,
Flora 8 (1825) 106; Hassk. Cat. Hort. Bog. (1844)
226; Blume, Rumphia 3 (1849) 200, t. \6i,p.p., incl.
var. pendula Blume, var. membranacea Blume, var.
chart acea Blume et var. obliqua Blume; Walp. Ann.
2 (1852) 224; Miq. F1. Ind. Bat. 1 , 2 (1859) 614; Sum.
(1861) 203, 520; lllustr. (1871) 74, p.p.; Hook./. F1.
Brit. India 2 (1876) 7; King, J. As. Soc. Beng. 65, ii
(1896) 458; Ridley, J. Str. Br. Roy. As. Soc. n. 33
(1900) 67; K. & V. Bijdr. 9 (1903) 125; Hall./. Med.
Rijksherb. 1 (1910) 2, in obs.; Koord. Exk. Fl. Java
2 (1912) 546; Merr. Enum. Born. (1921) 363;
Ridley, Fl. Mai. Pen. 1 (1922) 516, f. 51; Baker/.
in Rendle, J. Bot. 62 (1924) Suppl. 30; Craib, Fl.
Siam. Enum. 1 (1926) 340; Ridley, Kew Bull. (1926)
63; Merr. PI. Elm. Born. (1929) 176; Hochr. Can-
dollea 6 (1936) 467, incl. var. genuina Hochr.;
Backer & Bakh./ Fl. Java 2 (1965) 145; van
Beusekom, Blumea 19 (1971) 489. - Millingtonia
lanceolata Schult. & Schult. Syst. Veg. Mant. 3,
add. 2 (1827) 250; Dietr. Syn. PI. 1 (1839) 103. -
Meliosma polyptera Miq. Sum. (1861) 203, 520;
lllustr. (1871)73. - Meliosma levis KwG, J. As. Soc.
Beng. 65, ii (1896) 457; Ridley, Fl. Mai. Pen. 1
(1922) 515. - Meliosma nervosa K. & V. Bijdr. 9
(1903) 129; Koord. Exk. Fl. Java 2 (1912) 546; Atlas
2 (1914) t. 376; Fl. Tjibodas 2 (1923) 158; Merr. &
Perry, J. Arn. Arb. 20 (1939) 359, in obs.; Backer
& Bakh./ Fl. Java 2 (1965) 145.
Evergreen tree, up to c. 25(-30) m. Twigs often
with conspicuous leaf-scars. Leaves (3-)7-18(-25)-
jugate; rachis terete, (10-)30-100 cm, including the
5-30 cm long petiole, up to c. 8 mm diam., usually
with distinctly swollen base, usually ± lenticellate;
leaflets usually oblong to lanceolate, hardly or not
asymmetrical, 5-20 by 2-7 cm, not or only slightly
increasing in size towards the top of the leaf, often
the lowermost pairs much smaller, base usually acute
to rounded, apex acuminate to cuspidate, glabrous
to moderately pubescent, always without domatia;
midrib usually deeply impressed above; nerves 5-16
pairs, ascending, looped. Panicles terminal, nearly
always pendulous and lax, rarely erect (then also
small), pyramidal, usually large, (15-)50-150 cm
and profusely branched up to the 3rd order, ± pu-
bescent, bearing numerous glomerulate or crowded
flowers which are usually spicately arranged, the
glomerules often with regular space; main axis terete,
often bent down abruptly at the base; primary side-
axes many, usually long, up to c. 90 cm, never
subtended by leaves; bracts ovate to narrowly
triangular, up to c. 5 mm, ± pubescent. Pedicels ab-
sent, up to c. 1 mm. Mature buds 1.5-2 mm diam.
Sepals 5 (4), ovate, more or less unequal, the inner 3
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
glabrous, and with an entire margin. Outer petals
1.5-2 mm. Inner petals about halfway bifid, c. 0.6
mm, with ciliolate, rarely glabrous lobes, usually
with a minute central lobule. Filaments c. 1 mm.
Ovary (0.5-)0.7(-l) mm, usually densely, some-
times sparsely pubescent, rarely glabrous. Fruit
(sub)globose, when ripe 7-10 mm diam.; endocarp
subglobose, often somewhat depressed to applanate
at the ventral side, usually strongly obUque, (5-)6-9
mm diam., with usually distinct, rather coarse, most-
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
small to minute ventral processus or tubercle; ventral
pore not or not much sunken.
D i s t r . Nicobar Is. , extreme South of Peninsular
Thailand; in Sfalesia: Sumatra (incl. Simalur, Batu,
and Banka Is.), W. Java, Borneo (northern halO-
Not uncommon, scarce in Borneo. Fig. 11.
Ecol. Primary and often secondary forests, at
low and medium altitudes, occasionally ascending to
1500 m,/. nervosa to 2900 m, on various soil types.
Field notes. Outer bark grey to brown, rather
smooth, later with longitudinal cracks, thin, often
lenticellate. Inner bark 0.5-1 cm, several times said
to be (light) red, orange brown, or redbrown, also
dirty white and then turning rusty after e.xposure.
Wood soft, white or pale yellow to light yellow
brown. Crown low, irregular and lax, with few usual-
ly crooked branches. The conspicuous large leaves
are rather crowded at the end of the twigs. Leaflets
when young red-brownish. Flowers white or yellow-
ish to pink or red (sometimes different colours in the
same panicle). Fruits, first dirty red, then bluish black
when ripe.
Vern. Malay Peninsula: medang siri, Malacca;
Sumatra: kabung kabung (blumut), Batak lang.,
Simelungun dial., bulung manuk, Batak lang., Karo
dial., sondang, sonlang, Timor on N. Sumatra, kaju
buluk hudjan, Lampong, angke foluh pajo, silaora,
sarin silo bulung, tulun surin or seulang (pajo), t.
lungke ali, Simalur I.; W. Java: ki tiwu, S, often
used as well for M. pinnata and M. sumatrana (also
with the addition lalaki, mindi bodas or persawon),
suren ieuweung, S. See also under var. lanceolata f.
nervosa and var. polyptera.
Notes. Meliosma lanceolata is generally very
well characterized by its large pendulous panicles and
its long leaves with many usually lanceolate leaflets.
Nevertheless it shows a wide variation especially in
number but also in shape and size of the leaflets and
the panicles. On the islands west of Sumatra
(Simalur, Nias, Batu) specimens are found with nor-
mal inflorescences but only 3-5-pinnate leaves, and
elliptic, sometimes subrotund, large leaflets. Transi-
tions to this extreme are common. There is another
deviating form, however, which takes a separate
position. It has many small, mostly lanceolate leaf-
lets which otherwise do not differ from those of M.
lanceolata. Also the panicles agree with that species.
In view of the wide variability in the leaves of M.
lanceolata, I prefer to include it here and I have
reduced it to a variety. The varieties and forms can
be distmguished as follows:
a. var. lanceolata.
f.cavcs(3 )6 18 jugate, with up IOC. lOOcm long
rachis (including the petiole); leaflets elliptic to
lanceolate, mcdium-si/cd to large, 5 20( 25) by
(2-)2.5-7(-10) cm, index (l.5-)2-5(-6), without
or with teeth, glabrous or pubescent.
Notes. In the lowland parts of its area var.
lanceolata is nearly always very constant in the main
characters. Mainly at higher elevations, however,
forms occur which deviate considerably, often to
such an extent that it is very difficult to separate them
from less typical forms of the otherwise well distinct
M. pinnata ssp. ferruginea and ssp. ridleyi; in a few
cases, especially when the material is incomplete, this
can only be done by a specialist who is thoroughly ac-
quainted with habitus and variability of both species.
For instance, a form with erect, unusually short
panicles (sometimes only 15 cm long) and other de-
viating characters may be met with. It occurs mainly
in the montane zone; transitional forms are found
lower, and these show a more or less gradual fading
of typical lanceolata characters. Specimens of this
mountain form have been described from Java by
KooRDERS & Valeton, I.c, as M. nervosa. In my
opinion this species should be reduced to the rank of
a form only; see below.
forma lanceolata.
Leaves (3-)6-18-jugate, with elliptic to lanceo-
late, glabrous to pubescent leaflets. Panicles pen-
dulous, usually much longer than 50 cm. Inner petals
ciliolate. Ovary pubescent.
/ormo nervosa (K.&V.)Beus. Blumea 19(1971)493.
- M. nervosa K. & V., vide supra.
Leaves not more than 8(-10)-jugate, with usually
elliptic glabrous leaflets. Panicles erect, shorter than
c. 50cm, minimum length c. 15 cm. Inner petal most-
ly glabrous. Ovary pubescent to glabrous.
Distr. Malesia: Sumatra (G. Leuser, G. Talak-
mau), W. Java.
Ecol. Mountain forest, 1300-2900 m altitude.
The tree can reach a height of 30 m by I m diam.
Vern. Java: ki Ijermeh badak, ki tjermeh beu-
reum, S.
b. var. polyptera (Miq.) Beus. Blumea 19 (1971) 492.
- M. polyptera Miq., vide supra.
Leaves 12-25-jugate, with at most 50 cm long
rachis (including the petiole); leaflets oblong to
linear-lanceolate, small, 4-11 by 1-2 cm, entire,
glabrous.
Distr. Malesia: Sumatra (Asahan, W. Coast).
Fig. II.
Ecol. At low altitudes.
Vern. Sumatra: badar badar, Lubuk Alung, tan-
dikat batu, Priaman, simarpapahu, Mula Padang.
5. Meliosma hirsuta Blumh, Rumphia 3 (1849) 200;
Waip. Ann. 2 (1852) 225; Mio. Fl. Ind. Bat. I, 2
(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,
Blumea 19 (1971) 493.
Evergreen small tree, c. 5 m. Leaves 15-20-(or
probably more-)jugate; rachis 50-100 cm including
the 10-20 cm long petiole, up to c. 6 mm across,
more or less hirsute, usually with distinctly swollen
base, sometimes sparsely lenticellate; leaflets (sub)
sessile, those in medium and upper part of the leaf
linear-lanceolate, 10-20(-25) by (1.5-)2-3 cm, in-
dex 5-10, the lower ones (ovate-)lanceolate to ovate,
gradually decreasing in length towards the base of
the leaf, up to only c. 3 cm, base rounded to acute,
sometimes slightly oblique, apex acuminate to
caudate, with entire to remotely spinously dentate
margin, thin-chartaceous, above glabrous except for
some pubescence on the midrib, beneath moderately
to sparsely hirsute especially on midrib and nerves,
without domatia; midrib above flat to slightly im-
pressed; nerves widely apart, (5-)8-12 pairs, as-
cendmg, looped and joined into a distinct marginal
nerve situated at 2-4 mm from the margin; venation
distinct, wide, reticulate; petioles absent or up to c.
1 mm, terminal one often longer, up to c. 8 mm,
densely hirsute, not swollen at the base. Panicles and
flowers as in typical M. lanceolala, but sepals up to
c. 1.5 mm. Fruit as in M. lanceolala.
Distr. Malesia: Sumatra (West Coast: G. Malin-
tang), only one collection. Fig. 11.
Notes. This species was by Blume erroneously
recorded to occur in S. Borneo.
Meliosma hirsuta is doubtless very closely related
to M. lanceolala, but very well distinct by its leaf
characters.
See also Pimela angustifolia under the dubious
species.
6. Meliosma pinnata (Roxb.) Maxim. Bull. Ac. Imp.
Sc. St. Petersb. 12 (1867) 64; Melanges 6: 263. -
Millingtonia pinnata Roxb. F1. Ind. 1 (1820) 103. -
Fig. 12.
For further synonyms, see under the subspecies;
for a complete synonymy, see van Beusekom (1 97 1 :
494).
Evergreen, sometimes deciduous tree, small to up
to c. 42 m. Twigs often with conspicuous leaf-scars.
Leaves 2- II -jugate; rachis terete, (2-)5-40(-60)
cm, including the up to c. 15(-25) cm long petiole;
leaflets usually ovate, elliptic, or obovate to ovate-
oblong, sometimes lanceolate, often asymmetric.
1.5-25 by l-IO cm, usually increasing in size to-
wards the top of the leaf, base usually acute to
rounded, rarely slightly emarginate, apex acuminate
to cuspidate, entire or dentate, usually slightly to
densely pubescent, often with domatia; midrib flat to
impressed above; nerves 3-15 pairs, ascending,
looped; petiolules up to 5 cm, terminal one usually
longest, not or not much swollen at the base. Panicles
terminal, erect, sometimes somewhat pendulous,
dense to lax, widely to narrowly pyramidal,
10-55(-70) cm, usually profusely branched up to
the 4th order, bearing numerous solitary to usually
crowded flowers; primary side-axes usually many, up
to 35(-60) cm, lower ones sometimes subtended by
small to reduced leaves; bracts ovate to narrowly
triangular, up to c. 5(-10) mm, more or less pubes-
cent. Pedicels absent or up to 3(-4) mm. Mature
buds (1.5-)2(-3) mm diam. Sepals 5 or 4, ovate,
unequal, the 3 or 4 inner ones 1-1.5 mm, the outer
1 or 2 usually smaller, often minute, sometimes
lowered on the pedicel, sometimes slightly keeled,
glabrous or pubescent outside, all entire, usually
ciliolate. Outer petals usually glabrous. Inner petals
more or less deeply bifid, (0.3-)0.6(-l) mm, gla-
brous, ciliolate or fimbriate at the tips, often with a
minute central lobule, often frayed at the tips.
Filaments c. 1 mm. Fruit (sub)globose to obovoid,
when ripe (3-)4-10(-ll) mm diam., with thin
mesocarp; endocarp (sub)globose, oblique or not,
(2.5-)3.5-9(-10) mm diam., with more or less pro-
minently reticulate surface; median keel usually
distinct and more or less prominent, at one end
sometimes running out into a small to minute pro-
cessus or tubercle, and sometimes curving outwards
at the other end; ventral pore usually rather narrow,
whether or not sunken.
Distr. Throughout SE. Asia, from Sri Lanka and
China to Japan; throughout Malesia as far as New
Guinea (incl. New Britain). Fig. 13.
E c o 1 . Forests under moist tropical to subtropical,
sometimes warm-temperate conditions, on various
soils, from sea-level up to c. 3000 m altitude.
Notes. Meliosma pinnata covers a very large area
in which it has developed a complex and wide varia-
tion pattern. It can be divided up into nine well-
marked subspecies. Four of these are widely distrib-
uted, whereas five have a limited distribution. The
first group, the subspecies arnottiana, ridleyi,
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-
species of the second group occur scattered at the
periphery of the area of M. pinnata; I consider them
secondary offsplits from the primary subspecies, viz.
ssp. pinnata and ssp. angustifolia (Merr.) Reus.
from ssp. arnottiana, and ssp. pendula, ssp. sylva-
tica, and ssp. humilis from ssp. macrophylla.
The areas of the secondary subspecies fall partly or
entirely within the area of the primary subspecies
from which they are derived, but they are ecological-
ly isolated from these, usually by preference for dif-
ferent altitudinal zones; transitional or hybrid forms
are sometimes found. The areas of the four primary
subspecies, on the other hand, all touch or only
slightly overlap mutually, but generally they are per-
fectly replacing, and usually there is also different
ecological preference. Due to the scarcity of collec-
tions from critical regions, especially Sumatra,
Borneo and Sulawesi, it is mostly not clear how the
relation is in contact zones. There is some evidence
that one or two mutually may behave as good
species, where one or two others may be connected
by transitional forms, but in general the evidence re-
quired is still wanting. In this respect the picture is
not so complete as it is in M. simplicifolia.
The type subspecies does not occur in Malesia.
KEY TO THE SUBSPECIES
1. Ovary glabrous or only with a few hairs. Sepals
and petals always glabrous.
2. Leaves 3-5-jugate; leaflets dentate (sometimes
only a few teeth), with domatia in the axils of the
nerves beneath which are sometimes obscured by
very dense tomentum of the leaf-blade
g. ssp. humilis
2. Leaves (3-)4-6(-7)-jugate; leaflets dentate or
not, without domatia, never with very dense
tomentum.
3. Leaflets entire, index (l-)1.5-3, mostly round-
ed or obtuse to truncate or emarginate at the
base. Medium-sized to large trees
d. ssp. macrophylla
3. Leaflets dentate (sometimes very sparsely), in-
dex (l-)1.5-4(-5), acute or rounded, obtuse,
truncate or emarginate at the base. Small to
medium-sized trees, rarely shrubs.
1989]
Sabiaceae (van Beusekom & van de Water)
709
4. Leaflets moderately to rather densely villous-
pubescent (often more or less glabrescent when
older), mostly (especially lower ones) rounded
to truncate at the base, index 1.5-3(-4). En-
docarps 6-7 mm diam., without ventral pro-
cessus. Above c. 1800 m alt. e. ssp. pendula
4. Leaflets sparsely to densely short-pubescent,
rarely subglabrous, mostly with acute base, in-
dex I.5-4(-5). Endocarps 5-7.5 mm diam.,
mostly with a small but distinct ventral pro-
cessus. Below c. 1000 m altitude.
5. Leaves (3-)4-6(-7)-jugate; leaflets acute at
the base f . ssp. sylvatica
5. Leaves 3-5(-6)-jugate; lateral leaflets
rounded to truncate at the base
d. ssp. macrophylla (Celebes form)
1. Ovary entirely, rarely partly, but always densely
pubescent. Sepals and petals glabrous or pubes-
cent.
6. Sepals and usually also outer petals moderately
to densely pubescent on the outside. Leaflets en-
tire, index (l-)1.5-3 c. ssp. ferruginea
6. Sepals and petals glabrous or rarely a few hairs
on the outer sepals only. Leaflets entire or den-
tate, index (1-)1 .5-4(-5).
7. Endocarps 4.5-9(-10) mm diam., usually with
more or less sunken ventral pore. Inner petals
with fimbriate or ciliolate, rarely glabrous
lobes. Leaflets never with domatia
b. ssp. ridleyi
7. Endocarps (2.5-)3-4.5 mm diam., not with
sunken ventral pore. Inner petals with usually
glabrous, sometimes at the tips ciliolate or
frayed lobes. Leaflets with or without domatia
a. ssp. arnottiana
a. ssp. arnottiana (Wight) Beus. Blumea 19 (1971)
499. - Sapindus ? microcarpus W. & A. Prod. 1
(1834) 112, nom. illeg., non R. & P. (1804); Wight,
III. Ind. Bot. 1 (1840) 142; Walp. Rep. 1 (1842)416,
423. - Millingtonia arnottiana Wight, III. Ind. Bot.
1 (1840) 144, t. 53. - iVellinglonia arnottiana Meiss .
PI. Vase. Gen. (Comm.) 2 (1840) 207, in nolo. -
Millingtonia samhucina Jungh. Tijd. Nat. Gesch
Phys. 8 (1841) 365. - Meliosma arnottiana Walp
Rep. I (1842) 423; Thw. Enum. PI. Zcyl. (1858) 59
Bedd. FI. Sylv. 3 (1871)77; ibid. t. 160; Hook./. FI
Brit. India 2 (1876) 6; Trim. FI. Ceyl. 1 (1893) 315
Bra.sdis, Indian Trees (1906) 195; Gamble, FI. Pres
.Madras I (1918) 256. - Meliosma glauca Blume
Rumphia 3 (1849) 200. I. I68B, nom. illeg. \ Walp
Ann. 2 (1852) 225; Hassk. Hon. Bog. I (1858) 140
MiO. M. Ind. Bat. I, 2 (1859) 615; K. & V. Bijdr. 9
(1903) 135. ind. var. floribunda (Biijmi.) K. & V.
•Hall./. Mcdcd. Rijkshcrb. I (1910) 2; KrK)Rn. Fxk
n. Java 2 (1912) 546; M. Tjibodas 2 (1923) 157
Baker/ in Rcndlc. J. Bot. 62 (1924) Suppl. 30.
Meliosma floribunda Blume, Rumphia 3 (1849) 200;
Walp. Ann. 2 (1852) 225; MiQ. FI. Ind. Bat. 1, 2
(1859) 615; lllustr. (1871) 74; K. & V. Bijdr. 9 (1903)
137; Hall./ Meded. Rijksherb. 1 (1910) 2; Koord.
Exk. FI. Java 2 (1912) 546. - Meliosma sambucina
MiQ. lllustr. (1871) 74; K. & V. Bijdr. 9 (1903) 137,
in obs. - Meliosma luzonensis Merr. Publ. Govt.
Lab. Philip. 29 (1905) 24; Elmer, Leafl. Philip. Bot.
2 (1908) 492, in obs. ('luzonica'); Merr. Enum.
Philip. FI. PI. 2 (1923) 517. - Meliosma multiflora
Merr. Publ. Govt. Lab. Philip. 29 (1905) 25; Enum.
Philip. FI. PI. 2 (1923) 517. - Melliosma ferruginea
(non Blume) Koord. Gedenkb. Jungh. (1910) 177. -
Meliosma apoensis Elmer, Leafl. Philip. Bot. 10
(1939) 3784, descr. angl. - Meliosma cannarioides
Elmer, Leafl. Philip. Bot. 10 (1939) 3785, descr.
angl. - Meliosma ferruginea (non Blume) Backer &
Bakh./ FI. Java 2 (1965) \45, p.p., quoad M. glauca
et floribunda.
Small to medium-sized, rarely big tree, up to c.
20(-30) m. Leaves (2-)3-7(-8)-jugate; leaflets
ovate to ovate-oblong, elliptic, or lanceolate, small
to up to c. 25 by 10 cm, index (l-)1.5-4(-5), acute
to truncate at base, entire or dentate, chartaceous to
coriaceous, often with domatia. Panicles erect,
spreading, lax to dense, lower primary side-axes
usually subtended by small or reduced leaves. Sepals
glabrous or the outer ones rarely with a few hairs.
Petals glabrous, inner ones sometimes a bit ciliolate
or frayed at the tips of the lobes. Ovary densely
pubescent, very rarely subglabrous. Endocarps (sub)
globose, not or not much depressed, hardly or not
oblique, (2.5-)3-4.5 mm diam., with distinct, more
or less prominent, fine reticulum, with slightly to
rather strongly prominent, blunt to rather sharp me-
dian keel which does not run out into a ventral pro-
cessus or tubercle; ventral pore not sunken, some-
times a bit elevated.
Dist r . Sri Lanka throughout SE. Asia to China,
S. Korea, Japan and Taiwan; in Malesia: N. Sumatra
(Karo), Malay Peninsula, W. & Central Java, Philip-
pines (Batan Is., Luzon, Mindanao). Rare in W.
Malesia. Fig. 13.
Ecol . Primary or secondary montane rain-forest,
600-2500 m altitude, on loamy or volcanic soils, also
on limestone if the climate is wet enough. At higher
altitudes the subspecies is deciduous. In Malesia but-
tresses are sometimes developed, up to 1.5 m high.
Field notes . Bark dark to light grey, smooth, in
old trees sometimes distantly shallowly fissured. In-
ner bark soft, fibrous, with 'fingers' tapering out-
wards into granular tissue, pale pinkish brown to dull
red or redbrown, also said to be white, and turning
salmon red on exposure. Wood light and soft,
fibrous, easily split, white, with large pores and
beautiful grain, with prominent rays, hcarlwood in
older trees striped reddish and white. Leaflets be-
710
Flora Malesiana
[ser. I, vol. \0^
neath pale green, often glaucous. Fruits said to be
reddish, green brown, or black when ripe.
Vern. Sumatra: kabung sillang bulung, Batak
lang.; Java: dangdur bulu, kawayang, ki sum, ki
tiwu lalaki, S; Philippines: adope. adupong,
aropong. bantinan, kamug, Ig., bae, If.
Notes. Attention should be given to the relation
between ssp. arnottiana and ssp. pendula in the
Philippines (for the relation to ssp. macrophylla, see
under that subspecies). In the mountains of Luzon
both subspecies have been collected, ssp. pendula
above 1800 m altitude and ssp. arnottiana from c.
800-900 m up to c. 2400 m. Locally, e.g. on Mt San-
to Thomas, they have been found together, but
doubtless intermediate specimens are not observed.
It is possible that in such localities these subspecies
mutually behave as species; population studies in the
field might yield more evidence with regard to this.
The same problem arises in W. Malesia, where 55/?.
arnottiana has been collected (rarely). In Sumatra
and in Java its relation to ssp. ferruginea is in-
teresting since there is an altitudinal zone of overlap
between both, though ssp. ferruginea generally oc-
curs lower than ssp. arnottiana. In Java the situation
is as follows: ssp. ferruginea is by far the most com-
mon of both, ssp. arnottiana having only been col-
lected on a few mountains. Of these it is only G. Sa-
lak and G. Gedeh where both subspecies have been
found. Only of G. Gedeh more detailed ecological
evidence is available: Koorders (FI. Tjibodas 2,
1923, 157) stated that ssp. ferruginea occurs at c.
1400 m altitude and that ssp. arnottiana ('M. glau-
ca') occupies a zone between 1800 and 2400 m, being
especially abundant at c. 2200 m. This does suggest
the existence of ecological differentiation, but since
ssp. arnottiana on other mountains also grows at
lower altitudes, the situation remains unclear.
b. spp. ridleyi (King) Beus. Blumea 19 (1971) 505. -
Meliosma ridleyiKwG, J. As. Soc. Beng. 65, ii (1896)
458; Ridley, J. Str. Br. Roy. As. Soc. n. 33 (1900)
67; FI. Mai. Pen. 1 (1922) 516. - Meliosma elegans
Ridley, J. Str. Br. Roy. As. Soc. n. 54 (1910) 40; FI.
Mai. Pen. I (1922) 515. - Meliosma paucinervia
Merr. Philip. J. Sc. 10(1915) Bot. 39; Enum. Philip.
FI. PI. 2 (1923) 518. - Meliosma trichocarpa Merr.
Pap. Mich. Ac. Sc. 24 (1938) 80, nom. illeg., non
Hand.-Mazz. (1934). - Meliosma bartlettii Merr.
& Perry, J. Am. Arb. 20 (1939) 356. - Meliosma
confertiflora Merr. & Perry, I.e. 359.
Shrub or tree, up to c. 20 m. Leaves 3-7-jugate;
leaflets oblong to lanceolate, small to usually me-
dium-sized, up to c. 20 by 6 cm, base acute, rarely
rounded, usually entire, densely villous to glabrous,
without domatia. Panicles erect, usually rather lax
and slender; lower primary side-axes mostly subtend-
ed by small leaves. Sepals and outer petals glabrous.
Inner petals with fimbriate or ciiiolate tips, rarely
glabrous. Ovary densely pubescent. Endocarps
subglobose to very depressed and oblique, 4.5-9
(- 10) mm diam., with vague to distinct, more or less
prominent, rather wide reticulum, with slightly to
strongly prominent, blunt to very sharp median keel
which often at one end runs out into a minute ventral
processus, the curving at the other end sometimes far
drawn out into a blunt beak; ventral pore hardly to
rather deeply sunken.
Distr. Malesia: Central Sumatra, Malay Penin-
sula, Borneo (Sarawak, Sabah, W. Kutai), Philip-
pines (Mindoro). Fig. 13.
Ecol . Primary and secondary rain-forest, both in
mixed dipterocarp and in heath forest, on various
soil types, from sea-level up to 1400 m altitude.
Field notes. Bark mostly smooth, sometimes
somewhat scaly or slightly fissured, grey to brown.
Inner bark fibrous, pinkish to red or redbrown,
turning brown after exposure. Young branches,
inflorescence-axes, and leaf-rachises are sometimes
(Singapore) covered with a dense layer of soft dark
reddish brown hairs. Sepals sometimes said to be
purple. Fruit often ± hairy ( 'trichocarpa'), once said
to be bright purple.
Vern . Sumatra: kaju rokkam, k. rube gala, k. si
hasur, k. si (mardjuhut) (nij manuk, Asahan, mo-
dang halimponan, Tapanuli.
Notes . Ssp. ridleyi is rather variable when com-
pared to the other subspecies of A/, pinnata, especial-
ly in number and dentation of leaflets, in the degree
of pubescence, and in shape and size of the endo-
carps. In the Malay Peninsula, for instance, a form
with few subglabrous and somewhat dentate leaflets
has been found ('M. elegans'), as well as a beautiful,
densely rufous-pubescent form with distinctly more
and entire leaflets ('M. ridleyi'). It is not astonishing
that such different plants have been described as
separate species; only by studying material from
Borneo it becomes clear that these extremes are con-
nected by a range of transitions. Another form from
Dallas (Kinabalu), which has rather condensed
panicles, has been described as M. confertiflora.
This again is merely a local form without any
systematical significance, as is M. paucinervia, with
very lax panicles, from Mindanao. Yet, in spite of
this variation, it is obvious that ssp. ridleyi is a natu-
ral unit, probably most closely related to the adjacent
ssp. arnottiana from which it differs least of all
subspecies, mainly in shape and size of the en-
docarps, but also in some less important characters;
an especially close resemblance has been observed
between ssp. ridleyi and some deviating specimens
from South Vietnam which have been tentatively in-
cluded in ssp. arnottiana. Furthermore, the area of
ssp. ridleyi borders on or somewhat overlaps the
areas of ssp. ferruginea and macrophylla. The rela-
1989]
Sabiaceae (van Beusekom & van de Water)
711
tion between ssp. ridleyi and these subspecies has
been discussed under ssp. macrophylla.
Finally, it should be noted that the area of ssp.
ridleyi fully overlaps that of M. sarawakensis; this is
not accidental, since the latter probably is a
derivative of ssp. ridleyi (see the note under M.
sarawakensis).
c. ssp. ferruginea (Blume) Beus. Blumea 19 (1971)
507. - Meliosma ferruginea Blume, Cat. (1823) 32,
non SiEB. & Zucc. e.x Hook./. (1876), nee Kurz ex
King (1896); Nees, Flora 8 (1825) 106; Hassk. Cat.
Hort. Bog. (1844) 226; Blume, Rumphia 3 (1849)
200; Walp. Ann. 2 (1852) 225; MiQ. Fl. Ind. Bat. 1,
2 (1859) 616; Illustr. (1871) 74; K. & V. Bijdr. 9
(1903) 121; KooRD. Exk. Fl. Java 2 (1912) 546; Atlas
2 (1914) t. 375; Fl. Tjibodas 2 (1923) 157; Backer &
Bakh./. Fl. Java 2 (1965) 145, p.p., exct. M. glauca
et Jloribunda. - Millingtonia ferruginea Schult. &
ScHULT. Syst. Veg. Mant. 3, add. 2 (1827) 250;
DiETR. Syn. PI. 1 (1839) 103.
Medium-sized to big tree, up to c. 42 m. Leaves
2-6(-7)-jugate; leaflets elliptic to oblong, basal ones
sometimes a bit ovate, upper ones sometimes ±
obovate, usually rather large, up to 25(-38) by
10(-18) cm, base rounded to truncate, sometimes
acute, entire, firmly coriaceous, pubescent, rarely
subglabrous, rarely with domatia. Panicles erect,
spreading, lax to rather dense; lower primary side-
axes usually subtended by small leaves. Sepals usual-
ly densely pubescent, rarely on the outside sparsely
so to subglabrous. Outer petals pubescent outside,
rarely glabrous. Inner petals with fimbriate or
ciliolate tips. Ovary partly or entirely but almost
always distinctly and densely pubescent, very rarely
nearly glabrous. Endocarps subglobose, often some-
what depressed and oblique, 3.5-5.5(-8) mm diam.,
with rather vague to distinct, ± prominent reticulum,
with usually very prominent, rather sharp median
keel which does not run out into a ventral processus or
tubercle; ventral pore not or not much sunken.
Distr. Malesia: N. & Central Sumatra, through-
out Java, and the Lesser Sunda Islands (Bali, Sum-
bawa, Flores, Timor), locally common, especially in
Java. Fig. 13.
Ecol. Rain-forest, preferably on fertile, often
volcanic soils, 250-1600 m altitude.
Field notes. Bole cylindrical, straight, some-
times crooked, at the base up to c. 2.5 m diam. Bark
on the surface grey to brown, smooth, sometimes a
bit peeling or shallowly fissured to (deeply) cracked,
about 0.7-1.5 cm thick, easily detachable. Inner
bark pale brown to brownrcd or orange, with
streaks, also said to be dirty while and turning orange
brown when exposed to the air as a result of the
discolouring of the initially colourless watery exuda-
tion. Wood soft, yellowish to pinkish white. Leaflets
pale greyish to glaucous green beneath. Fruits
brownred to black when ripe.
Uses . Advocated for reafforestation purposes by
KOORDERS.
Vern. Sumatra: sekapong, Takengon, sontang,
Simelungun, sihubung, Kerinci; Java: ki tiwu, ki
tjerme badak, S, gempong, gijubuk, gompong, J;
Lesser Sunda Is.: gempong, sambuk, Bali, mladja,
tanggo, tawu, Flores, Endeh lang., lohot, raok,
Flores, kaju mangkok, W. Sumbawa.
Note. Ssp. ferruginea is usually well recognizable
by its outside pubescent sepals and petals. However,
in N. Sumatra and the Lesser Sunda Islands
specimens occur in which these characters are im-
perfectly or not developed, and they may also lack
the pubescence on the ovary and may have almost
glabrous leaves. They are not easily identifiable and
may be confused with A/, lanceolata var. lanceolata
f. nervosa or with the closely related M. pinnata ssp.
macrophylla and ssp. ridleyi.
A. ssp. macrophylla (Merr.) Beus. Blumea 19(1971)
510. - Meliosma macrophylla Merr. Philip. J. So.
7 (1912) Bot. 294; Enum. Philip. Fl. PI. 2 (1923) 517.
- Meliosma lanceolata var. obliqua {non Blume)
KooRD. Minah. (1898) 408; Suppl. 2 (1922) 7, t. 55;
ibid. 3 (1922) 28. - Meliosma wallichii (non Planch.
ex Hook./.) Koord. Minah. (1898) 408. - Meliosma
tongcalingii Elmer, Leafl. Philip. Bot. 8 (1915)
2815. - Meliosma megalobotrys Mekr. Philip. J. So.
11(1916) Bot. 16; Enum. Philip. Fl. PI. 2 (1923) 517.
- Meliosma macrocarpa Elmer, Leafl. Philip. Bot.
10 (1939) 3786, descr. angl. - Meliosma ferruginea
(non Blume) Merr. & Perry, J. Arn. Arb. 20(1939)
356. - Fig. 12.
Medium-sized to large tree, up to c. 42 m. Leaves
(3-)5-9-jugate; leaflets elliptic to oblong to ovate-
oblong, medium-sized to rather large, up to c. 20 by
9 cm, base rounded or obtuse to truncate, entire,
rarely with a few teeth (Sulawesi), chartaceous to
firmly coriaceous, very sparsely to densely pubes-
cent, always without domatia. Panicles erect and
spreading, lax and slender to rather dense; lower
primary side-axes usually subtended by small leaves.
Sepals and petals glabrous. Ovary glabrous, rarely
with a few scattered hairs. Endocarps subglobose,
sometimes more obovoid or depressed, more or less
oblique, 3.5-5 mm diam., exceptionally 5-7.5 mm
diam. (Sulawesi), with vague to distinct and promi-
nent reticulum, with rather sharp and prominent me-
dian keel which at one end mostly runs out into a
small but distinct ventral processus or tubercle; ven-
tral pore .somewhat sunken.
Distr. Malesia: E. Borneo (E. Sandakan, Bcrao,
W. & E. Kutai, Tandjung), Sulawesi (Minahasa,
Malili), Moluccas (Halmahcra, Scram). Philippines
(Luzon, Lcytc, Mindanao. Palawan), throughout
712
Flora Malesiana
[ser. I, vol. lO-*
New Guinea (incl. New Britain). Fairly common in
most parts of the area. Fig. 13.
Ecol. Usually in primary, rarely in secondary
rain-forest, at low to medium altitudes; in Borneo
only collected below 100 m. in the other parts of the
area also higher, up to c. 1 100-1200 m, in W. New
Guinea once at 1800 m. in Borneo usually found in
lowland dipterocarp forests. Generally reported to
occur on clayish, loamy, or sandy clayish soils, also
on red earth, on volcanic soil, and on loamsoil on
limestone. It is rarely found in occasionally sub-
merged areas. Once (New Guinea) said to occur on
peaty soil, and there developing stiltroots.
Field notes. Bole mostly straight, cylindrical,
up to at least 1 m diam. at the base, usually develop-
ing 1.5-2.5 m high buttresses, sometimes without
buttresses, once observed stiltrooted. Bark grey to
brown, or patchy brown-white-grey, smooth, some-
times with shallow vertical cracks, not or little peel-
ing, with vertical rows of lenticels. Inner bark c. 1 cm
thick, soft, light brown or pink to brownred, inside
paler, sometimes said to be streaked with cream, with
some colourless sticky exudate (which is also said to
be redbrown!); it is said to be rapidly darkening upon
exposure or 'a bright orange-brown stain quickly ap-
pears between bark and sapwood.' Wood very light
and soft; sapwood white to pale pink or brown, when
fresh with bright brown sap streaks, heartwood ab-
sent or present, darker than the sapwood. The fruit
is said to be brown to black.
Vern. Borneo: surian, E. Kutai; Sulawesi: kaju-
saul-rintek, Tooelooe lang., papako, Tontemboan
lang., mumping, Tonsea lang., Hasan, Ratahan
lang.; Moluccas: bais, Seram; Philippines: arocong,
Ig., agosos, balilang-uak (a corruption of barilan ng
uak). Tag., morau, S.L.Bis., mungapong, Bik., ma-
gasorod. Bag.; New Guinea: sebotebuk, tubuk,
Mooi, serajema, Manikiong, marwaskeipi, Japen,
bagare, Kapauku lang., biedewon, iediewat, Muju,
morrotuno, waito, Wapi, frikipa, Orne lang.,
tapuha, Managalase, kufi, Kutubu, uliga, Madang,
kombowase, Waskuk, wagebi, Wagu.
Notes. Ssp. macrophylla is the most common
and widespread of the East Malesian subspecies
group, characterized by a glabrous ovary by which it
is readily distinguished from the West Malesian
subspecies. Within its large area a few other sub-
species occur, viz. ssp. pendula, sylvatica, and
humilis, which have much more limited areas and
probably represent offsplits from it. These three
subspecies are ecologically well isolated from ssp.
macrophylla.
In Borneo the area oissp. macrophylla is, as far as
can be judged from the available evidence, sharply
delimited against that of the West Malesian ssp.
ridleyi, which, moreover, appears to prefer a higher
altitudinal zone (only in Borneo ssp. macrophylla
seems to be restricted to lowland forests below c. 100
m altitude!). To the SW the area o^ ssp. macrophylla
borders on that oi ssp. ferruginea which inhabits the
Lesser Sunda Islands. The latter two subspecies are
huge trees, very similar in general habit, and some-
times they have been confused. Nevertheless, they
are usually well distinct, mainly by flower characters,
though in both subspecies there is a tendency to lose
some of these characters.
In the Philippines ssp. macrophylla is sympatric
with ssp. arnottiana, but they prefer different
altitudinal zones, the first being a lowland subspecies
not exceeding c. 900 m altitude, the latter being a
montane subspecies occurring from c. 800 up to c.
2400 m (once recorded from c. 600 m).
e.spp. pendula (Merr.) Beus. Blumea 19(1971)512.
- Meliosma pendula Merr. Publ. Govt. Lab.
Philip. 29 (1905) 25; Enum. Philip. Fl. PI. 2 (1923)
518. - Meliosma reticulata Merr. Philip. J. Sc. 5
(1910) Bot. 195; Enum. Philip. Fl. PI. 2 (1923) 518.
- Meliosma macgregorii Merr. Philip. J. Sc. 10
(1915) Bot. 37; Enum. Philip. Fl. PI. 2 (1923) 517.
Small to medium-sized tree, up to c. 20 m. Leaves
(3-)4-6-jugate; leaflets elliptic to oblong, rarely lan-
ceolate, up to 18(-20) by 7(-ll) cm, the lower ones
at the base nearly always rounded or (sub)truncate to
obtuse, the upper ones more or less acute, nearly
always distinctly dentate, villous-pubescent, ± gla-
brescent, without domatia. Panicles erect and
spreading to somewhat pendulous and rather flaccid,
slender and rather lax; lower primary side-axes most-
ly subtended by small leaves. Sepals and petals
glabrous. Ovary glabrous. Endocarps subglobose,
slightly oblique, 6-7 mm diam., with rather vague,
slightly elevated reticulum, with hardly to moderate-
ly prominent, blunt median keel, the latter not run-
ning out into a distinct ventral processus or at most
into a very minute tubercle; ventral pore hardly or
not sunken.
Distr. Malesia: Philippines (Luzon: Mountain
Prov.). Fig. 13.
Ecol. Montane rain-forest, 1800-2500 m alti-
tude. In mossy forest, in ravines as well as on expos-
ed ridges.
Field notes. Bark thick, checked. Wood soft,
said to be soon assuming an orange-brown colour.
Uses. The leaves are once said to be used for
smoking by the Igorots.
Vern. Anitap, Ig.
Note. Ssp. pendula replaces the lowland and
lower hill ssp. macrophylla at high elevations (in this
respect being comparable to ssp. humilis from New
Guinea).
f.ssp. sylvatica (Elmer) Beus. Blumea 19(1971)513.
- Meliosma sylvatica Elmer, Leafl. Philip. Bot.
1989]
Sabiaceae (van Beusekom & van de Water)
713
2 (1908) 492; Merr. Enum. Philip. Fl. PI. 2 (1923)
518. - Meliosma acuminatissima Merr. Philip. J.
Sc. 10 (1915) Bot. 36; Enum. Philip. Fl. PI. 2 (1923)
517. - Meliosma brachyboirys Merr. Philip. J. Sc.
12 (1917) Bot. 275; Enum. Philip. Fl. PI. 2 (1923)
517.
Slender shrub or treelet, up to c. 5 m. Leaves
(3-)4-6(-7)-jugate; leaflets elliptic to usually ob-
long or lanceolate, usually medium-sized, up to c. 18
by 6 cm, acute at the base, sparsely to rather closely,
always distinctly dentate, very sparsely to moderately
pubescent, without domatia. Panicles erect, spread-
ing, usually slender and rather la,\; primary side-axes
(mostly?) not subtended by small leaves. Sepals and
petals glabrous. Ovary glabrous. Endocarp subglo-
bose, often somewhat ellipsoid, more or less oblique,
5-6 mm diam., with distinct, more or less prominent
reticulum, with rather sharp and prominent median
keel which at one end runs out into a small but
distmct ventral processus; ventral pore somewhat
sunken.
Distr. Malesia: Sulawesi (Minahasa, Latimod-
jong Mis), Philippines (Luzon, Negros). Fig. 13.
Ecol . Lowland rain-forest, usually not above 750
m altitude, growing in the shrub layer.
Field notes. Slender, suberect or bent shrub or
treelet of a sparsely branched habit. Bark smooth,
grey and brown mottled. Wood white, soft, easily
breakable. Leaves once said to be light bluish green
beneath.
Note. Ssp. sylvatica is closely related to ssp.
macrophylla. The most striking difference between
them is found in their physiognomy, the former be-
ing a small undergrowth treelet, the latter a large
forest tree. The main systematical differences are
found in the dentation and the shape of the base of
the leaflets. These fit nicely in the spectre of
character combinations present in the subspecies of
M. pinnata, and it seems justified to consider ssp.
sylvatica a subspecies of that species, instead of a
separate species.
R. spp. humilis (Merr. & Perry) Beus. Blumea 19
(1971) 514. - Meliosma humilis Merr. & Perry, J.
Arn. Arb.20(1939)358;/*/Vy. 22 (1941) 263, //j 065.
- Meliosma schlechteri Merr. & Perry, J. Arn.
Arb. 22(1941) 262.
Small to medium-sized tree, up to c. 20 m. Leaves
3-5 jugate; leaflets elliptic to oblong, rarely short-
lanceolate, usually ralhcr small, up to c. 15(~24) by
6<-9) cm, the base acute, lower leaflets sometimes
rounded at the base, sparsely to rather closely den-
tate, beneath subglabrous to ralhcr densely pubes-
cent, sometimes densely villous-tomcntose, always
with more or less distinct domatia in the axils of the
nerves beneath (obscure in densely lomentose
leaflets). Panicles erect, spreading, mostly rather lax
and with slender axes; lower primary side-axes often
subtended by small leaves. Sepals and petals gla-
brous. Ovary glabrous. Endocarps subglobose,
somewhat depressed and rather oblique, 5.5-7.5
mm diam., with more or less vague, slightly elevated
reticulum, with very prominent, rather sharp median
keel which at one end runs out into a small but
distinct ventral processus; ventral pore somewhat
sunken.
Distr. Malesia: Papua New Guinea (Highlands
Provinces, Madang, Morobe, Milne Bay). Common.
Fig. 13.
Ecol. Montane rain-forests, 1000-30(X) m
altitude. Observed as an understorey tree in dense
Castanopsis-Nothofagus forest, on ridges as well as
on streambanks, but also often reported from several
kinds of disturbed forest, such as bamboo regrowth,
old garden land, transition between coniferous forest
and treefern grassland, and even from open grass-
land. Once reported from limestone ridge.
Field notes. Bark greybrown, smooth, with big
lenticels. Inner bark straw-coloured to pink, red, or
reddish brown (due to discolouring, as in other sub-
species?), exuding 'resin'. Wood white to light
brown, with conspicuous rays and clear growth
rings, said to be of moderate weight and hardness.
Petioles, peduncle, and pedicels purplish to red-
brown; buds reddish. Fruits dark red to black when
ripe.
Uses. Once said to be used as housing timber,
free from borers.
Vern. Mansalong, Finschhafen, kokopong,
Nako lang., E. Madang Prov., kass, Maring lang.,
mappam, Enga lang., W. Highl. Prov.
Notes. Ssp. humilis is closely allied to ssp. ma-
crophylla, mainly differing by its dentate leaflets
with domatia. In Papua New Guinea it replaces ssp.
macrophylla mainly found below 1000 m altitude (cf.
ssp. pendula from Luzon). It is remarkable that ssp.
humilis has as yet been collected, even rather abun-
dantly, only in Papua New Guinea and not in W.
New Guinea.
7. Meliosma sarawakensis Ridley, Kew Bull. (1933)
193; Merr. & Perry, J. Arn. Arb. 20 (1939) 359. -
Meliosma grandifolia Lecomte, Bull. Soc. Bot. Fr.
54(1909)676, now. ///e/?.. non Urban (1895); Merr.
Enum. Born. (1921) 362; van Beusekom. Blumea 19
( 1 97 1 ) 5 1 5 . - Meliosma confusa var. laxior Baker /.
in Rendle, J. Bot. 62 (1924) Suppl. 30. - Meliosma
lattfolia RitUEY. Kew Bull. (1933) 193; Merr. &
Perry, J. Arn. Arb. 20 (1939) 359. m obs.
Evergreen, small tree, up to c. 10 m. Leaves
2-3(-4)-jugate; rachis terete, 12-30 cm, including
the 6-15 cm long petiole, up to c. 5 mm diam..
densely short-tomcntosc. later ± glabresccni; leaf-
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
often more or less obovate to obovate-oblong,
sometimes ± asymmetrical, (2-)5-22 by (1.5-)
3-12 cm, mostly distinctly increasing in size towards
the top of the leaf, base acute to rounded, apex more
or less acuminate, sometimes subacute or cuspidate,
with entire to remotely spinously dentate margin,
chartaceous, moderately to rather densely pubescent
especially beneath and on midrib and nerves, often
partly giabrescent when older, never with domatia;
midrib more or less impressed above; nerves 6-12
pairs, ascending, usually looped; venation distinct,
reticulate; petiolules up to c. 1.5(-3) cm, terminal
one usually longest, tomentose. Panicles terminal,
usually more or less pendulous, flaccid, lax, narrow-
ly pyramidal, (20-)25-55 cm, not profusely
branched up to the 2nd or 3rd order, branches
spreading, ± flaccid, usually slender, densely
tomentose, bearing numerous flowers crowded in
dense spikes; primary side-axes few to rather many,
up to c. 25(-35) cm, the lower ones usually subtend-
ed by reduced leaves; bracts ovate to usually narrow-
ly triangular or linear-lanceolate, up to c. 4 mm,
densely pubescent. Pedicels (almost) absent. Mature
buds c. 2 mm diam. Sepals 5 (4), ovate to ovate-
lanceolate, the 3 or 4 inner ones 1-1.5 mm, the outer
1 or 2 usually much smaller, often minute, densely
pubescent on the outside, with entire margin. Outer
petals glabrous. Inner petals about halfway or some-
what less bifid, 0.5-0.7 mm, glabrous, sometimes
with a minute central lobule. Filaments c. 1 mm.
Ovary 0.5-0.7 mm, densely pubescent. Fruit
(sub)globose, when ripe 0.7-1 cm diam.; endocarp
depressed-globose, applanate at the ventral side,
strongly oblique, 6-7(-8) mm diam., with usually
distinct, more or less sharply prominent reticulum;
median keel sharp and very prominent, not at one
end running out into a ventral processus or tubercle,
at the other end rather far curving outwards; ventral
pore rather sunken.
Distr . Malesia: Sumatra (Asahan to Palembang)
and NW. Borneo (Sarawak, around Kuching and
Pontianak; common). Fig. 13.
Ecol. Lowland rain-forest, up to c. 800 m
altitude.
Field notes. The sepals are redbrown to purple.
Vern. Sumatra: kaju rube boras, Asahan; Bor-
neo: bulu manok, Iban name, Kuching.
Note. The closest affinity of M. sarawakensis is
doubtless with M. pinnata ssp. ridleyi to which it is
very similar in all characters (they even share the red
sepals). Only after some hesitation M. sarawakensis
is maintained as a separate species and not made a
subspecies of A/, pinnata. It would fit rather well into
that species but is distinguished from it by a wider
range of characters. The most important of its
characters are the 2- or 3-jugate leaves and the dense-
ly pubescent sepals. These characters indeed are also
found in M. pinnata ssp. ferruginea, but this subspe-
cies is quite different from M. sarawakensis in
various other aspects. An additional argument to the
specific status of M. sarawakensis is found in the fact
that it is found together with M. pinnata ssp. ridleyi
in the same area and at the same altitudes in Sarawak
(near Kuching) and in Sumatra (Asahan), without
any sign of hybridization.
8. Meliosma rufo-pilosa Hend. Card. Bull. Str.
Settl. 7 (1933) 96, t. 18; Merr. & Perry, J. Arn.
Arb. 20 (1939) 360; van Beusekom, Blumea 19
(1971) 517.
Evergreen rather large tree up to c. 30 m. Flower-
ing twigs terete, 5- 10 mm diam., stout, abruptly ter-
minating in a tuft of leaves and inflorescences, gla-
brous, often with many large conspicuous leaf-scars.
Leaves (6-)7-9-jugate; rachis terete, (13-)25-50
(-65) cm, including the (3.5-)6- 16 cm long petiole,
pubescent, not swollen at the base, hardly or not len-
ticellate; leaflets elliptic to oblong, sometimes ovate
(-oblong), 3- 15 by 2-6 cm, base obtuse to truncate,
apex acuminate, entire, glabrous or ± puberulous
on nerves above, (sub)glabrous beneath, pubescent
on nerves, always without domatia; midrib im-
pressed above; nerves 7-18 pairs, ascending, looped;
venation fine, very distinct, reticulate; petiolules 1-6
mm, densely pubescent. Panicles terminal, one or a
few crowded together at the end of a twig, erect,
rather dense to lax, pyramidal, 30-50 cm, including
the 0-20 cm long peduncle, profusely branched up to
the 4th order, branches spreading, ± flaccid, densely
pubescent, bearing numerous solitary flowers;
primary side-axes well-spaced, c. 8-15, up to c. 30
cm, not lenticellate, the lower ones never subtended
by small or reduced leaves; bracts narrowly triangu-
lar to lanceolate, up to c. 4 mm, densely pubescent.
Pedicels 1-3 mm, densely pubescent. Sepals (3) 4,
ovate, (sub)equal, c. 1.5-2 mm, the outer one often
much smaller, rarely minute, usually lowered on the
pedicel, glabrous or somewhat pubescent outside;
margin flimsy, more or less ciliolate, entire or some-
times with some coarse irregular teeth. Outer petals
c. 1.5 by 1.5-2.5 mm. Inner petals ligular, usually
somewhat widened towards the top, 0.7-1 mm; top
entire or with a shallow incision, blunt, minutely
ciliolate. Filaments 0.7-1 mm. Ovary 0.5-0.7 mm,
glabrous. Fruit globose, when ripe 1.5-2 cm diam.,
with moderately thick, fleshy mesocarp; endocarp
semiglobose, broad-ovate to subcordate at ventral
view, 11-13 mm long and wide, 7-8 mm high, with
relatively thin wall, with slightly lumpy surface,
especially lumpy and somewhat furrowed at the ven-
tral curving of the wall; median keel faint, hardly
elevated but at one end drawn out into a con-
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)
614; lllustr. (1871) 73; Merr. Enum. Born. (1921) 363; Hall./. Beih. Bot. Centralbl. 39, 2 (1921) 161;
KosTER.M. Bibl. Laur. 1 (1964) 951. - As was noted by Hallier/., I.e., the type specimens (S. Muller^.a;.,
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. 1 (1850)226. - Canariopsis angustifolia Blvme e.x Miq.
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 it should then be placed close to M. laneeolata and M. hirsuta.
At first sight it is very 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 the preceding volumes, it seemed useful to correct some errors which have crept into the text
of volumes 4- 1 as well as to add some additional data, new records and references to new species which came
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
9: 47b Osmoxylon sessiliflorum (Laut.) Philip-
SON.
Add to Distr.: Moluccas (Halmaheira), 2
coll.
Add to Vern.: saha-saha, Ternate lang.,
tele, Sahu lang.
Bignoniaceae
8: 142b Dolichandrone spathacea (L.f.) K. Sch.
Add to Distr.: A.G. Wells, in H.J. Teas
(ed.). Biology and ecology of mangroves
(1983) 61, map, records this species for the
first time from Australia: the northern tip
of Cape York Peninsula (2 localities).
Cardiopteridaceae
7: 93 Cardiopteris [Wall, ex] Royle.
Add to Distr.: The genus extends to the
Solomon Islands (Bougainville) and N.
Queensland; cf. Bailey, Queensl. Fl.
(1899) 251; Compreh. Cat. Queensl. Pi.
(1912) 93, f. 76.
Celastraceae
6: 257b Glyptopetalum loheri Merr.
Distr.: Philippines: add Palawan (Rids-
DALE 683).
6: 266a Lophopetalum beccarianum Pierre.
Add to Distr.: Central Sumatra (Djambi:
Vreeken-Buys 69).
6: 291 b Perrottetia alpestris (Bl.) Loes. ssp. philip-
pinensis (Vidal) Ding Hou.
Add to Distr. (and map): Lesser Sunda
Islands (Flores: Loeters 650, 1796).
Convolvulaceae
4: 434b Jacquemontia brown iana Ooststr.
Add the synonym: J. pannosa (R. Br.)
Mabberley, Bot. Macar. 6 (1980, '1978')
63.
Cyperaceae
7: 653a Cyperus diaphanus Schrader ex R. & S.
var. latespicatus (Boeck.) Kern.
Add to Distr.: Lesser Sunda Islands
(Flores: Schmutz 5767).
9: 173a Carex bilateralis Hayata.
Add to Distr.: SE. to E. Asia.
Dipterocarpaceae
9: 436b Hopea gracilis Miq. (under Excluded).
It was unfortunately overlooked that the
type at L (W. Central Sumatra, Padang,
Teijsmann HB 424) was already in 1968
referred by Ashton to Meiogyne and later
in the same year identified by F.H. Hilde-
BRAND as Meiogyne virgata (Bl.) Miq. {An-
nonaceae).
Ericaceae
6: 856b Vaccinium angulatum J.J.S. 1914, non
(Griff.) Theobald, 1883 = Vaccinium
commutatum Mabberley & Sleumer,
Taxon 34 (1985) 155, nom. nov.
Flacourtiaceae
5: 24b Hydnocarpus nana King.
Add to Distr.: NE. Sumatra (Besitang R.,
Sikundur For. Res., de Wilde & de Wilde-
Duyfjes 19540).
5: 35a Scaphocalyx spathacea Ridl.
Add the synonym: S. parviflora RrDL. etc.
5: 35b Reduce Scaphocalyx parviflora Ridl. to S.
spathacea Ridl.
5: 56a Homalium dasyanthum.
Change authorship: (Turcz.) Theob. in
Mason, Burma ed. Theob. 2 (1883) 451;
Warb. etc. (see Mabberley, Taxon 34,
1985, 155).
5: 98b Reduce Casearia pallida Craib. to C
flavovirens Bl.
5: l(X)b Casearia flavovirens ^i..
Add the synonym: C. pallida Craib, etc.
(716)
1989]
Addenda, corrigenda et emendanda
717
5; 113a
Add to Distr.: Thailand; in Malesia: Suma-
tra, Java, Lesser Sunda Islands (Bali).
Haemodoraceae
Haemodorum corymbosum Vahl should
be called Haemodorum coccineum R.Br.,
according to T.D. Macfarlane, Perth,
Australia.
Iridaceae
8: 83 The correct name for Gladiolus natalensis
(Ecklon) Reinw. ex Hook, should be
Gladiolus daienii Geel, Sen. Bot. fasc. 28
(1829). See Milliard & Burtt, Notes Roy.
Bot. Card. Edinb. 37 (1979) 297.
Olacaceae
Hamamelidaceae
5: 366 Maingaya malayana Oliver.
A marvellous colour photograph of this
very rare species (and genus), endemic in
Malaya and Penang, was given by Francis
Nc in Nature Malaysiana 7, no 4 (1982) 8;
the species is thought to be ± extinct and is
now grown at the Forest Research Institute,
Kepong.
Hydrocharitaceae
5: 390b. Blyxa aiiheriii Rich.
391 Add: var. echinosperma (Clarke) Cook &
LiJOND, Aquat. Bot. 15 (1983) 14.
Add the synonym: B. echinosperma
(Clarke) Hook./, elc.
5: 392a Blyxa leiosperma Koidz.
Reduced to B. japonica (Miq.) Maxim, ex
AscHERS. & Gl'rke var. japonica.
5: 392b Blyxa aliernifolia (Miq.) Hartog.
Reduced to B. japonica (Miq.) Maxim, ex
Aschers. & GuRKE var. alternifolia (Miq.)
Cook & Luond, Aquat. Bot. 15 (1983) 25.
5: 393a fl/vx^crvopoAj/co (Miq.) Maxim, ex Aschers.
& Gurke.
Add: var. japonica and var. aliernifolia
(Miq.) Cook. & Luond, Aquat. Bot. 15
(1983) 22 and 25 respectively.
Add the synonyms: B. leiosperma Koidz.
and B. alternifolia (Miq.) Hartog.
Hypericaceae
8: 18b Replace Hypericum uralum Bif h.-Ham.
ex D.Don by Hypericum henryi Lev. &
Vaniot ssp. hancockii Robson, Bull. Br.
Mus. Nat. Hist. Bot. 12(1985)261, map 22.
Distr. of var.: Continental SE. Asia (S.
China, Vietnam, Burma, Thailand).
Icacinaceae
7: 6a Cilronelta suaveolens (Hi.) Howard.
Add to Distr.: Lesser Sunda Islands
(Flores: Schmutz 5819).
10: 7a The correct name for Olax scandens Roxb.
should be Olax psittacorum (Willd.)
Vahl; see Almeida, J. Bomb. Nat. Hist.
Soc. 81 (1985) 742.
10: 9b E.\cluded, add:
Olax baliculin Blanco, F1. Filip. ed. 2
(1854) 589; ed. 3 (1877) 38; Kosterm. Bibl.
Laur. (1964) 1153 = Litsea baliculin
(Blanco) Kosterm. Bull. Bot. Surv. India
10(1968) 268 (Lauraceae).
Merrill (Enum. Philip. Fl. PI. 2, 1923,
195) recognized it as a Litsea and said that
it might prove to belong to L. leytensis
Merr. Since no material is known to exist,
the matter will remain speculative.
Passifloraceae
7: 416a Passiflora aurantia Forst. /.
Add to Distr.: W. Central Celebes, near
Palu, at 200 m, Meijer 10172.
Pittosporaceae
5: 356b Pittosporum moluccanum (Lamk) Miq.
Add the synonym: Coffea angustifolia
RoxB.;c/. FoRMAN,Kew Bull. 38(1983)64.
Podostemaceae
6: 66b Cladopus nymani H. Moll.
Add to Distr.: Central Borneo, en route
from Sinar Baru to Ryan Ruwan, N of
Long Bawan, Krayan, 1 1 5° 45 ' E, 4° 5 ' N,
on rock in a rapid stream (Pa Raya), sub-
merged and sterile, 1150 m alt., M.
Okamoto C.S., s.n. (L, Osaka). The
material matches the figure on p. 66 of Fl.
Males. I, 4 almost exactly.
Rhizophoraceae
5: 461b Bruguiera gymnorrhiza (L.) Saviony.
Distr. : It is worthy of record that according
to an internal report by A.N. Gillkson
(C'SIRO, Canberra) this mangrove species
is found on the lower limestone terrace on
I
718
Flora Malesiana
[ser. I, vol. 10"^
Christmas I. (Indian Ocean), in a healthy
and self-maintaining community near the
treshwaler Hosnie's Springs al 40 m alt.,
and not tar from it a stand of a remnant ht-
toral forest of Heriliera littoralis Dryand.
in Ait., another mangrove species.
5 : 471a Ceriops decandra.
The authorship must be (Griff.)
Theobald; see Taxon 34 (1985) 154.
Simaroubaceae
6: 218b Ailanthus inlegrifolia LAMk.
Add to Distr. (and map): N. Queensland
(several localities) (L. Pedley, in litt.).
Sparganiaceae
4: 233 The correct name for Sparganium simplex
HuDS./. simplex as used by Backer should
be:
Sparganium fallax Graebn. in E. & P. Pfl.
R. 2 (1900) 15, t. 3H; Cook & Nicholls,
Bot. Helv. 96 (1986) 253; ibid. 97 (1987) 3,
t. 13a, 14a, map 14. - S. simplex auct. non
HuDS./. simplex: Backer, FI. Males. I, 4
(1951) 233, fig.
Basal leaves usually exceeding the stem,
(4-)5-10(-15) mm wide. Lowermost bract
20-35(-60) cm long, 1 -several times as
long as the inflorescence. Female heads
3-4(-6), usually supra-axillary, sometimes
extending beyond the next internode; ped-
uncle up to 3 cm long. Male heads 5-8 or
more. Female flowers: perianth segments at
least connate below, in fruit more than 0.5
times as long as the fruit; pedicel 1-3 mm
long. Male flowers: filaments up to 6 or
more mm long; anthers (0.8-)l -2(-2.2)
mm long. Fruits fusiform, sometimes con-
stricted around the middle, 5-6 mm long,
light brown, dull, tapering below into a
1.5-3 mm long pedicel. Endocarp c. 2 mm
wide.
Distr. E. India to Japan; in Malesia:
Sumatra, Papua New Guinea (up to 2000 m
alt.).
Sparganium subglobosum Morong, Bull.
Torrey Bot. Club 15 (1888) 81, t. 79, f. 1;
Cook & Nicholls, Bot. Helv. 97 (1987) 4,
t. 13b, 14b, map 15.
Basal leaves usually exceeding the in-
florescence, (l-)2-4(-9) mm wide. Low-
ermost bract up to 15 cm long, 0.5-1 (rare-
ly more) times as long as the inflorescence.
Female heads solitary, axillary or on lateral
branches, occasionally one head supra-
axillary, sessile, rarely with an up to 5 mm
long peduncle. Male heads solitary. Female
flowers: perianth segments free, in fruit
0.3-0.5 times as long as the fruit; pedicel
O-I mm long. Male flowers: filaments
2.5-3. 2(-3. 8) mm long, anthers 0.5-0.9
(-1) mm long. Fruits obovoid to almost
globose, yellowish to pale brown, shiny,
subsessile or with an up to 1 mm long, not
persistent pedicel. Endocarp 1.6-1.8 mm
wide.
Distr. India (once, in Khasia), Yunnan,
Vietnam to Manchuria and to Okinawa,
Australia, New Zealand; in Malesia: Papua
New Guinea.
Note. The identity of a collection from
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 1 5-22 cm long 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,
West Germany; in litt.) the following
amendments should be made:
10: 110 Family distribution map: extend the central
area to the west to include part of East
Africa; see Kew Bull. 36 (1982) 733.
Sciaphila Blume.
Add/change description: Anthers some-
times 4-celled (America). Endosperm pres-
ent; embryo small, orthotropous, undif-
7: 198
ferentiated, bitegmic, only in ripe seed the
inner integument wholly or largely sup-
pressed.
Violaceae
Add under Viola the following species:
Viola rheophila Okamoto, Bull. Osaka
Mus. Nat. Hist. no. 37 (1984) 4-15, 3 fig.
Distr. Malesia: Borneo (Kalimantan
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 in bold type denote main treatment; an asterisk behind a page number denotes the
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
Aceraceae 4: 3-4, 592; 6:
915; 7: 820; 8: 549
Acioa Aubl. 639
heieropeiala (ScorL ex
King) Kosierm. 677
malayana Kosterm. 677
pcrcoriacea Kosterm. 678
Acioa aucl. 675
Acmopyle 343, 352, 354
Actinidiaceae s.str.4: 37-39
Adenia 166
Afrocarpus 354
Afrolicania Mildbr. 645
Agathis Salisb. 339,341-343,
347, 354,420,421, 429,
430', 431 map
sect. Agathis 431
sect Macrobracteaiae Mcijer
Drees 431
sect. Microbracleatae Mcijer
Drccs431
sect. Prismatobractcatae
Meijer Drees 431, 440
sect. Rostrata dc Laub. 431,
439
alba Foxw. 433
fl/6flauct. 435,437
beccarii Warb. 433
beckingii Mcijer 435
bornccnsis Warb. 340, 34 1 ,
430*. 432*. 433, 434*
map
cclcbica (Koord.) Warb. 433.
434. 435 map
ssp. ceiebica Vcldk. &
Whitm. 435
(Agathis ceiebica)
ssp. flavescens (Ridl.)
Veldk. & Whitm. ex
Veldk. & de Laub. 438
dammara (Lamb.) Richard
433, 434
ssp. dammara aucl. 435,
437
ssp. dammara VJhilm. 433
ssp. flavescens (Ridl.)
Whitm. 438
dammara auct. 435, 437
endcrtii Mcijer Drees 439,
440* map
flavescens Ridl. 340, 341,
433, 438
hamii Meijer Drccs 435
kinabalucnsis de Laub. 433,
438 map, 439
labiilardieri Warb. 436 map,
440, 441*
latifotia Mcijer Drees 433
Icniicula de Laub. 433,
436*, 437 map
loranthifolia Salisb. 433
loranthifolia auct. 435
macrostachys Warb. 433
motleyi (Pari.) Diimmcr
393
orbicuia de Laub. 433, 437*
map
palmerstonii (F.v.M.) Bailey
442
philippincn.sis Warb. 433,
437. 438 map
regia Warb. 437
rhomboidales Warb. 433
(Agathis)
robusia (Moore) Bailey 442
map
ssp. nesophila Whitm.
436
var. nesophila auct. 442
var. robusia Whitm. 442
spathulata dc Laub. 433,
435, 436 map
Agonandra 31-34
raccmosa 33
Ailanthus intcgrifolia Larnk 718
Aizoaceae 4: 267-275; 6:
915
Alberlisia Becc. 160-162, 164,
167-169, 171, 179
crassa Forman 179, 181
mccistophylla Forman 181
mcgacarpa Diels ex Forman
159, 179, 181
papuana Becc. 166, 179,
180*, 181
Alc.imandra Dandy 565, 566.
567, 569
cathcariii (Hk.f. & Th.)
Diindy 569
Alismataceae 5: 317-334;
6: 915, 9: 553
Alismauilcs 109
Allantospcrmum Forman 621,
622
Alnus 338
Alpam Rhccde 65
Alscuosmia 335
AlMCUosmiaccac 335-336
Aisoinitra macrcxarj^a (Bi.)
Rocm. 29
(721)
722
Flora Malesiana
[ser. I, vol. 104
Aliingiacxccl.sa341
Amaranthaceae 4: 69-98,
593; 5: 554; 6: 915;
7: 820; 8: 549
Ambora ficus Tul. 326
Amborclla 261
Amborcllaccae261
Amorpha pedalis Blanco 488
Anacardiaceae 8: 395-548;
9: 553; 10: 679-681
Anaoirdium occidcntale 352
Anacolosa Bl. 1,3-6, 23
arbor ea K. & V. 25
cauliflora Sleum. 25
celebica Valet. 25
frutcscens (Bl.) Bl. 2, 24*,
25, 26* map
heptandra Maing. ex Mast. 25
luzoniensis Mcrr. 25
maingayi Mast. 22
papuana Schelle.nb. 25, 26
Zollinger I Baill. 25
Anamirta Colcbr. 157, 158,
160-162, 166-168, 170,
171, 211
cocculus (L.) Wight & Am.
159, 165. 166, 212*,
213, 214*
flavescens (Lamk) Miq. 210
jucunda Micrs 213
lemniscata Miers 210
loureiriPicTTC 210
luctuosa Miers 210
Anaxagorea A. St. Hil. 605
Ancistrocladaceae 4: 8-10;
5: 553
Ancistrocladus pentagynus
Warb. 612
Androglossum reliculaium
Champ, ex Benth. 685
AndrurisSc\\\\s\09, 110
andajensis (Becc.) Schltr 118
anisophylla Cicscn 112, 118
australasica (Hemsl.) Giesen
118
bur uensis J. J. Smith 118
celebica Schltr 118
clemensae (Hemsl.) Giesen
112, 118
clemensae (Hemsl.) Giesen
var. borneensis Giesen 118
crinila (Becc.) Schltr 1 17
elegans G'lescn 118
gracillima Giesen 1 17
javanica Giesen 118
loheri Giesen 117
nana (Bl.) Giesen 117
palawensis Tuyama 118
/ene//a Schltr 118
(Andruris)
viiiensis (A.C.Smith) Giesen
118
wariana Schltr 1 17
Angelesia Korth. 645
papuana Baker f. 65 1
splendens Korth. 646
Anisadcnia 609
Anisophyllaea gaudichaudiana
Baill. 253
Annonaceae 63, 64, 261, 605,
716
Anthobembix Perkins 306, 308
brassii A.C. Smith 315
dentatus \ a\Q.iox\ 310
hospilans (Becc.) Perkins 311
ledermannii Perkins 319
moszkowskii Perkins 3 1 1
myrtifolia A.C. Smith 313
oligantha Perkins 315
parvifolia Perkins 315
Antidesma ghacsembilla Gaertn.
52
pcntandrum (Blco) Merr. 52
Ami taxis Miers 172
calocarpa Kurz 174
cauliflora (Miers) Dicls 174
fasciculata Miers 174
longifolius (Decne ex Miq.)
Miers 174
Apama Lamk 62, 63, 65
affinis Weisse 79
brevipes Weisse 79
corymbosa (Griff.) Willd. ex
Soler. 78
macranlha Weisse 81
tomeniosa Engl, ex Soler.
55,79
var. lanuginosa (Hk.f.)
K. & G. 79
iAp/zy/Ze/a Champion 110
erubescensCUdsn^). 114
Aponogetonaceae 4: 11—12;
5: 553; 7: 213
Aptandra 3, 4
Aptandropsis 4
Aquifoliaceae 145
Aquilegia 165
Ardbidopsis 543
Arabis 54 1
Araliaceae 9: 1-105, 553;
10: 716
Araucaria Juss. 337, 339, 421,
423 map
sect. Araucaria 422, 423
sect. Bunya Wilde & Eames
423
sect. Columbea All
sect. Colymbea Endl. 422
(Araucaria)
sect. Eutacta Endl. 422, 423,
425
sect. Euiassa (Salisb.) Benth.
& Hook. 425
sect. Intermedia White 423
subg. Colymbea Antoine 423
subg. Eutacta (Link) Antoine
425
beccarii Warb. 427
bidwiiiii 420, 423
cunninghamii Ait. ex D. Don
340, 347, 425
var. papuana Laut. 426*,
427 map, 428*
cunninghamii auct. non Ait.
427
hunsteinii K.Sch. 339, 347,
420, 423, 424 map
klinkii Laut. 423, 424
schumanniana Warb. 424
Araucariaceae 338, 339, 343,
347, 354, 419-442
Araucariacites 338
Arcangelisia Becc. 158, 160,
161, 166-171, 209
flava(L.) Mcrr. 160, 210,
211
inclyta Becc. 210
lemniscata (Miers) Becc. 210
loureiri (Pierre) Dicls 210
tympanopoda (Laut. &
K. Sch.) Diels 210, 211
Archichlamydeae 127
Ardisia glabra (Thunb.) DC. 134
Aristolochia L. 53-55, 57-59,
61-64,83, 166
acuminata Lamk 94
barbata 55
brasiliensis Mart. & Zucc.
55,64
clematitis 59
coadunata Backer 57-59, 84,
86, 95, 96*
var. bosschai Backer 95
var. coadunata 95
crassinervia Schmidt 58, 82*,
85, 87, 104*
curtisii King 84, 86, 89
decandra Ding Hou 57-59,
64, 83, 85, 87, 100,
101*, 102*
deltantha F.v.M. 97
dictyophlebia Merr. & Perry
97
dielsiana Schmidt 57, 61, 64,
84, 85, 88, 105
elegans Mart. & Zucc. 55,
58,64
1989]
Index to scientific plant names
723
(Aristolochia)
engleriana Schmidt 85, 88,
"106
foveolata Merr. 55, 58, 85-
87, 91 map
gaudichaudii Duchartre 56*,
57, 84, 88, 107
gigantea Mart. & Zucc. 64
glaucifolia Ridl. 55, 85, 86,
93
gracilifoUa Schmidt 97
grandiflora Sw. 57, 59, 62,
64
griffiihii Duchartre 59, 91
hastata Jack 88
hastata (non Jack) Klotzsch
92
humilis Merr. 55, 85, 87,
103
imbricaia Mast. 103
indica L. 99
indica (non L.) BIco 94
indica (non L.) L. 97
jackii Steud. 55, 58, 59, 84,
86, 87, 88, 89*
japonica Miq. 94
kaempferi WiUd. 92
kankauensis Sasaki 92
kaoi Liu & Lai 91
klossh Ridl. 85, 86, 93
kwangsiensis Liang 91
lautcrbachiana Schmidt 108
ledermannii Schmidt 106,
107 map
Icuconeura 62
leytensis Merr. 84, 85, 87,
104
linnemannii Warb. 55, 85,
88, 107
longifolia Roxb. 94
macgregorii Merr. 55, 56*,
57, 85, 87, 103
maurorum 58
megalophylla K.Sch. 94
memhranacea Merr. 103
mindanaensis Warb. 94
minuiiflora Ridl. ex Gamble
55. 85-87, 93
var. dolahrata Gamble 93
moluccana Duchartre 94
momandul K. Sch. 58. 59,
82*, 85, 86, 88, 97
naviculilimba Ding Hou 85,
87. 100*
novogumcCnsis Schmidt 108
ornilhrjcephala 55
papillifolia Dmg Hou 58, 84.
87. 98*. 99
pcliata 59
(Aristolochia)
philippinensis Warb. 60*,
66, 84, 87, 102
piihecurus Ridl. 97
ramosii Merr. 92
ringens Vahl 55, 64
roxburghiana Klotzsch 94
6 angustifolia Duchartre
94
ssp. kankauensis (Sasaki)
Kitamura 92
roxburghiana (non Klotzsch)
Warb. 107
rumphii KosteleL?Jcy 85, 86,
97, 101*
samarensis Merr. 85, 87,
102
schlechteri Laut. 58, 59, 85,
88, 97, 105, 106, 107
sericeaBlco55, 57, 58, 61,
66, 85, 87, 103
singalangensis Korth. ex
Ding. Hou 61, 84, 86,
89, 90*
tagala Chamisso 54, 55, 56*,
57, 63, 85-88, 94
tagala (non Chamisso) Halus.
92
var. hankaoensis 92
var. kankauensis (Sasaki)
Yamazaki 92
timorensis Dccne 94
Lranstillifera Ding Hou 60*,
84, 87, 99
iripariiia Backer 88
ungulifolia Mast. 88
zollingeriana Miq. 57, 85,
86, 92
Aristolochiaceae 53-108,
126, 166
Aristolochiales 567
Aromadendron Bl. 567, 568,
576
ashionii Dandy ex Cockbum
579
borneensis Dandy ex Cock-
bum 579
elegans Bi. 577
var. glauc.a (Korth.)
Dandy 577
glaucum Korth. 577
nutans Dandy 577
oreadum (Diels) Kanch. &
Hatus. 583
Arorrtadendrum 568
Aromoracia rusticana G.M. &
Schcrb. 543
Asarum canadcn.sc 57
virginitum 62
Ascarina J.R. & G. Forst. 123-
129, 137, 140 map
sect- Ascarina 137, 139, 140
map
sect. Madascarina Leroy &
Jeremie 123, 137, 140
map
diffusa A.C. Smith 139,
141
lanceolata (non Hk.f.) auct.
141
lucida 124, 139
maheshwarii Swamy 139,
140, 141
philippinensis C.B. Rob.
138*, 139, 141
reticulata Merr. 139
rubricaulis 127
subsessilis Verdcourt 139,
141
Ascarinopsis 123, 128
Ascarinopsis Humb. & Capuron
123, 128, 137, 140 map
As hto n s Aromadendron Mcijcr
579
Asiphonia Griff. 65
piperiformis Griff. 78
sp. Griff. 78
Aspidocarya 162
dissitiflora Laut. & K. Sch.
197
hirsuta^cjcc. 193
kelidophylla Laut. &
K. Sch. 253
uvifcra Hk. f. & Th. 189*
Aihcrospcrmataceae 261
Athcrospermoxylon 256
Athrotaxis 339
Au-iaecarpum 158
Atroxima 456
Atuna Rafm. 636-642, 665
cordata Cockburn ex Prance
636, 666 map, 667
data (King) Kostcrm. 669
elliplica Ko.stcrm. 666
excelsa (Jack) Kostcrm. 636,
640,641,671
indica (Bald.) Kostcrm. 666
latifolia (Hcnd.) Kostcrm.
666
lalifrons (Kostcmi.) Prance &
While 666 map
nannodes (Kostcrm.)
Koslerm. 666. 667 map
pcnangiana (Kostcnn.)
Ko.stcrm. 666. 667 map
raccmo.sa Rafin. 666. 669
ssp. cxccl.sa (Jack) I'rancc
668*. 669. 670 map
724
Flora Malesiana
[ser. I,vol. 104
(Atuna racemosa)
ssp. racemosa 669, 670
map
scabra (Hassk.) Koslerm.
641,669
travancorica (Bedd.) Kosterm.
666
villamilii (Merr.) Kosterm.
671
Aiunus Rumph. 665
alba Rumph. 669
Austrocedrus Florin & Boutelje
443
Austrotaxus 339, 348
Badiera (non DC.) Hassk. 459,
467
pulchra (Hassk.) Hassk. 468
venenosa (Poir.) Hassk. 468
Bafodeya Prance 642
Balanophoraceae 7: 783—805;
8: 549; 9: 554
Balantium Desv. ex Buch.-Ham.
654
Balsaminaceae 539
Bania Becc. 183
thyrsiflora Becc. 183
Banisterodes O.K. 493
affine (Miq.) O.K. 503
ellipilcum (Korth. ex Miq.)
O.K. 530
excelsum (Bl.) O.K. 501
glaucum (Wall, ex Hassk.)
O.K. 527
griffiihii (Hk.f. ex Benn.)
O.K. 513
insigne (Benn.) O.K. 537
/on^//b//a(Bl.)O.K. 514
maingayi (Benn.) O.K. 524
obscurum (Benn.) O.K. 537
rufum (Benn.) O.K. 505,
507
stipitatum (Benn.) O.K. 535
vitellinum (Bl) O.K. 515
Bannisierioides 493
Barbarea vulgaris R. Br. 545
Bamharlia Gleason 493
Basellaceae 5: 300-304
Baiania Ualus. 172
insignis Hatus. 174
Baterium Miers 183, 184
validum Miers 1 84
Bat(id)aceae 5: 414-415; 6:
917
Benneiiiodendron leprosipes
(Clos) Merr. 715
Bcrberidaceae 163
Berbcridales 163
Betulaceae 5: 207-208; 6:
917
Bignoniaceae 8: 114-186;
9: 554; 10: 716
Bixaceae s.str. 4: 239-241
Bladhia glabra Thunb. 1 34
Blumia Nees 569
candollii (Bl.) Necs 582
Blyxa alternifolia (Miq.) Hanog
717
aubenii Rich. 717
var. echinosperma (Clarke)
Cook &Luond 717
echinosperma (Clarke) Hk.f.
717
japonica (Miq.) Maxim, ex
Aschers. & Gurke 717
var. alternifolia (Miq.)
Cook&Luond^n
var. japonica 717
leiosperma Koidz. 717
Boraginaceae 29
Borneo Aromadendron Meijer
579
Bracea King 29
Brachynema 3, 4
Braganlia Lour. 65
affinis Planch, ex Rolfe 79
blumii Lindl. 79
brevipes Merr. 79
corymbosa (non Griff.)
F.-Vill. 103
corymbosa Griff. 78
macranlha Boerl. 81
melastomaefolia Duchartre
78
tomentosa Bl. 79
var. lanuginosa Hk. f. 79
Brassica L. 541, 543, 545
besseriana Andrz. ex Trautv.
546
campestris L. 543
chinensis L. 543
integrifolia (West) Rupr. 546
juncea (L.) Czem. 544*, 545
napus L. 543
oleracea L. 543
rapa L. 543
rugosa Prain 543
Bredemeyera 456, 457, 465
sect. Melchiora Steen. 465
floribunda 457
papuana Steen. 465
Brewsiera M.J. Roemer 622,
625
crenaia M.J. Roemer 625
Brongniartia Bl. 287
coriacea Bl. 298
Brownetera L.C. Rich. 356
Bruguiera gymnorrhiza (L.)
Savigny 717
Bubbia 123
Burmanniaceae 4: 13-26,
592; 5: 553; 7: 820;
9: 554
Burseraceae 5: 209-296,
567; 6: 917; 7: 820;
9: 555; 10: 715
Butomaceae 5: 118-120,
566
Byblidaceae 7: 135-137
Callitrichaceae 4: 251-252
Callitris sp. 453
Callitroideae 444
Calocedrus 444
Camelina 541
Campanulaceae 6: 107—141,
928; 8: 549; 9: 556
Canariopsis anguslifolia Bl. ex
Miq. 715
Canarium angustifolium Miq.
715
Candjera 5 1
Cannabinaceae 4: 222-223
Cansiera 50, 52
zyziphifolia 50
Cansjera Juss. 31-35, 48, 49
map
grossularioides Blco 52
lanceolala Bth. 50
leplostachya Bth. 31, 33, 34,
48*, 49 map, 51*
leplostachya Koord. 35
malabarica Lamk 50
var. 6 50
manillana Bl. 36
monostachya (Willd.)
M. Roemer 50
parvifolia Kurz 48*, 49 map
penlandra Blco 52
polystachya (Willd.)
M. Roemer 50
rheedii J.F. Gmelin 31-33,
48*, 49 map, 50*
rheedii Blco 52
scandens Roxb. 9, 50
timorensis Decne 5 1
zizyphifolia Griff. 50, 51
Cappar(id)aceae 6: 61-105;
7: 822
Caprifoliaceae 4: 175-194;
6: 928; 9: 556; 10: 335
Capsella Medicus 541, 543,
545, 549
bursa-pastoris (L.) Medicus
550
1989]
Index to scientific plant names
725
Cardamine L. 541, 543, 545,
550
africana L. 541, 551
ssp. borbonica (Pers.)
Schulz
var. papuana Laul. 55 1
alligena O.E. Schulz 551,
553
borbonica Pers. 551
decurrens (Bl.) Z. & M. 554
flexuosa With. 551, 554
flexuosa auct. 554
hirsuta L. 551, 554
javanica (Bl.) Miq. 552
keysseri O.E. Schulz 551,
553
papuana (Laut.) O.E. Schulz
551, 552*
re ge liana Miq. 554
sublyrata Miq. 559
Cardiopteridaceae 7: 93—96;
10: 166, 253, 716
Cardioptcris Royle 166, 716
moluccana Bl. 19, 253
Carex bilateralis Hayata 716
Caricaceae 61
Carpolobia 456, 457
Carronia F.v.M. 158, 160, 162,
167, 168, 170, 172, 183
thyrsiflora (Becc.) Dicls
182*, 183
Carson's Magnolia Meijer 571,
573
Cascaria flavovirens Bl. 716
pallida Craib 716
Casuarinaccae 453
Cathedra 3, 4
Cayloniales 338
Caytonipollcniies 338
Celastraceae 6: 227-291,
389-421,930; 10:
539, 716
Cclaslrales 145, 629
Celasiranac 151
Ccliis paniculaia (Endl.) Planch.
29
Ccntrolepidaceae 5: 421-427
Ccphaiotaxaccac 343, 347
Ccphaloiaxus 354
celebica Warb. 349
mannii (non Hk.f.) F*rilzcl ex
Dicls 349
sumatrana Miq. 349
Ceramium Bl. 65
lomenlosum fil. 79
Ceratophyilaceae 4: 41-42
CV.ri<jps dctandra 7 1 8
ChamaehuxiLS (DC.) Spach 459,
467
(Chamaebuxus)
arillaia (D. Don) Hassk. 469
pulchra (Hassk.) Hassk. 468
venenosa (Poir.) Hassk. 468
sub var. elliptica Miq. 468
subvar. obovata Miq. 468
var. apiera Miq. 468
var. gracilis Miq. 468
var. minor Miq. 468
var. robusla Miq. 468
Champaca Adans. 598
lurbinata Nor. 583
Champereia Griff. 31-34, 35
cumingiana (Baill.) Merr. 36
gaudichaudiana (Baill.) Tiegh.
36
griffithii Planch, ex Kurz 36
lanceolata Merr. 36
manillana (Bl.) Merr. 31, 34,
35, 36*, 37*, 38 map, 40*
oblongifolia Merr. 36
perrolteiiana Baill. 52
platyphylla Merr. 36
Champereya 35
gnetocarpa Kurz 36
griffiihiana Planch, ex Kurz
36
Chandlera 158
Chasmanihera 164
Chaunochiton 3, 4
Cheirolepidaceae 338
Chenopodiaceae 4: 99—106,
594; 6: 932; 8: 549; 9:
557
Chlaenandra Miq. 158, 161,
162, 167, 168, 170, 171,
187
ovaia Miq. 159. 186*, 187,
189*
Chloranthaceae 123-144
Chloranihales 127
Chloranlhus Swariz 123-128,
129. 139. 143, 144
brachysiachys Bl.
var. melanocarpus Ridl.
136
brachysiachys scnsu Benlh.
134
chincnsis 124
denliculalus Cordemoy 1 34
elalior R.Br, ex Sims 131
creclus (Buch.-Ham.) Verde.
124. 127. 128. 130*.
131. 132*. 133*. 136
ereclusSwccl 130
fortunci 127
glabcr scnsu Backer &
Bakh.f. 136
glaber (Thunb.) Makino \?A
(Chloranthus)
hainanensis Pei 1 36
henryi Hemsley 130, 133
inconspicuus Svj2iT\z 133
inconspicuus (non Swartz)
Blco 131
indicus Wight 133
japonicus 127, 130
monander R. Br. ex Sims
134
multisiachys Pei 128
obtusif alius Miq. 133
officinalis Bl. 124, 127,
128, 131, 132
oldhamii Solms-Laub. 133
oldhamii (non Solms-Laub.)
Merr. & Quis. 133
philippinensis Merr. 133
salicifolius Presl 131
serratus 127
spicalus (Thunb.) Makino
127, 128, 130, 132
veriicillatusMen. 133
Chrysobalanaceae 635-678
tribe Chrysobalaneae 635
tribe Couepieae 636
tribe Hinelleae 636
tribe Parinarieae 635
Chrysobalanus L. 635-637,
639, 640, 642, 643
ciliatus Korth. ex Miq. 674
icaco L. 643, 644* map
splendens Korth. ex Miq.
646
Cissampelos L. 157, 158. 160-
162. 165. 167. 169-171,
234
convolvulacea Willd. 236
var. hirsuta (DC.) Hassk.
236
cumingiana Turcz. 236
discolor DC. 236
var. cardiophylla A. Gray
236
hernandifolia 245
hernandiifotia Willd. 245
insularis Makino 240
ovala Poir. 217
pareiraL. 165
p236
var. hirsuia (Buch. ex
DC.) Forman 235*.
236
var. orbiculaia (DC.) Miq.
236
var. peltata Schcff. 236
var. typica Diels 236
parcira (non L.) Ridl. 229
psilophylla Prcsl 25 1
726
Flora Malesiana
[ser. I, vol. 104
Citronella suaveolens (Bl.)
Howard 717
Cladopus nymani H. M611. 717
Classopollis 338
Clethraceae 7: 139-150
Clypea BI. 243
acuminalissima Bl. 252
capitata Bl. 252
corymbosa Bl. 249
discolor BI. 245
glaucescens Decne 245
tomentosa Bl. 229
venosa Bl. 248
Coccomelia Ridl. 645
CocculusDC. 157, 158, 161,
162, 165, 169, 231
angustifolius Hassk. 234
bantamensis Bl. 199
blumei Boerl. 223
brachysiachyus DC. 217
celebicus Boer). 175
cinereus Zoll. & Mor. 229
coriaceus Bl. 199
corymbosus Bl. 229
crispus [non (L.) DC] Hassk.
199
cuspidatus V^ all. 2\9
cynanchoides PresI 233
elegans (Ridl.) Ridl. 233
ferrandianus Gaudich. 233
flavescens (Lamk) DC. 210
flavicans Wall. 253
forsteri DC. 245
glaucescens BI. 230
^/aucwi (Lamk) DC. 229
incanus Colebr. 229
japonicus (Thunb.) DC. 245
var. limoriensis DC. 245
kunslleri King 231
lanuginosus Bl. 229
laurifolius DC. 160, 166,
169, 171,231,232*,
233, 234
var. angustifolius (Hassk.)
Boerl. 233
var. triplinervis Boerl. 233
leptostachyus DC. 217
limacia DC. 223
longifolius Decne ex Miq.
174
lucidus Teijsm. & Binn. 174
macrocarpus W. & A. 230
mollis Hk.f. & Th. 233
orbiculalus (L.) DC. 159,
160, 162, 170, 172,
231, 232*
ovalifolius (Vahl ex Pers.)
DC. 233
populifolius DC. 213
(Cocculus)
rimosus Bl. 207
sarmenlosus Dicls 233
var. stenophyllus Merr.
233
tr iandr us CoXchx. 185
/r//?o/'U5DC. 219, 233
/n7(3feu5 (Thunb.) DC. 159,
163, 165,231,233,234
velutinus Wall. 223
Cochlearia officinalis L. 543
Cochlospermaceae 4: 61—63
Coffea anguslifolia Roxb. 717
Columbea All, Al^
Colymbea Spreng. 422
Combretaceae 4: 533-589;
5: 564; 6: 932; 7: 823
Comesperma 456, 461, 465
sect. Prosthemosperma
F.v.M. 465
Coniferales 337-453
Connaraceae 5: 495-541;
6: 933; 7: 823; 8: 549;
9: 557
Convolvulaceae 4: 388-
512,599; 5: 558; 6:
936; 7: 823; 9: 558;
10: 29,716
Cordai tales 126
Cornaceae 8: 85-97
Coronopus 541
Coryneliales 353
Corynocarpaceae 4: 262—
264; 5: 557; 6: 941
Corytholobium Mart, ex Benth.
483
Coscinium Colebr. 160-162,
166-168, 170, 171, 215
blumeanum Micrs ex Hk.f. &
Th. 212*, 215, 216,
217
blumeanum (non Miers ex
Hk.f. &Th.) Miq. 216
var. epel latum Boerl. 216
fenestratum (Gaertn.) Colebr.
160, 165, 212*, 215,
217
var. macrophyllum
Yamamoto215
var. ovalifolium
Yamamoto215
maingayi Pierre 216
mangayi 216
miosepalum Diels 216
peltatum Merr. 216
wallichianum Miers 215
Couepia 640
Coula 3, 4
Crambe541,543
Crassulaceae 4: 197-202;
9: 558
Cri.spiloba Stccn. 335
Croialaria duboisii Lev. 469
Cruciferae 541-560
tribe Arabideae 543
tribe Brassiceae 543
tribe Heliophilcae 543
tribe Lepidicae 543
tribe Stcnopctaleae 543
Cryphaea Buch.-Ham. 129
erecta Buch.-Ham. 131, 132
Crypteroniaceae 8: 187-204
Cryplocarya reticulata BI. 715
Ctenolophon Oliver 29, 629
englerianus Mildbr. 630, 631
grandifolius Oliver 631
parvifolius Oliver 630, 631,
632*, 633* map
philippinensis Hall.f. 630,
631
Ctenolophonaceae 607, 609,
621,629-634
Cucurbitaceae 29, 166, 253
Cupressaceae 338, 339, 343,
347,442-447
Curupira 3, 4
Cyclandrophora Hassk. 642, 665
asperula (Miq.) Prance ex
Kosterm. 671
elata (King) Kosterm. 669
excelsa (Jack) Kosterm. 671
glaberrima Hassk. 669
lalifolia (Hend.) Prance 666
laurina (Gray) Kosterm. 669
nannodes (Kosterm.)
Kosterm. & Prance 667
penangiana Kosterm. &
Prance 667
scabra (Hassk.) Kosterm. 669
villamilii (Merr.) Prance ex
Kosterm. 671
Cyclea Am. ex Wight 160-162,
165, 167, 169, 170, 172,
237
acuminatissima Merr. 239
apoensis Yamamoto 24 1
atjehensis Forman 237, 238*,
239
barbata Miers 159, 136, 139,
242, 243, 253
barbata Craib 239
caudata Merr. 239
cauliilora Merr. 237, 241
ciliata Craib 242
elegans King 187, 237, 239
insularis (Makino) Hatus.
237, 240
var. luxurians Hatus. 240
1989]
Index to scientific plant names
727
(Cyclea)
kinabaluensis Forman 159,
237, 239
var. hispida Forman 239
korihalsii Diels ex Norman
241,242
laxiilora Miers 237, 239,
241, 242, 243
merrillii Diels 237, 240
peliata [non (Lamk) Hk. f. &
Th.] Becc. 241
var. arnouii Miers 241
peliata [non (Lamk) Hk. f. &
Th.] Miq. 163, 242
percgrina Miers 237, 239,
241. 242
robusia Becc. 237, 240
scyphigera Suesseng. &
Heine 239
forma anguslifolia
Suesseng. & Heine 239
tomcntosa 243
tonkinensis (non Gagnep.)
Yamamoto 239
wallichii Diels 242, 243
Cyclodiscus tomentosus
Klolzsch 79
Cyperaceae 7: 435—753,
823; 9: 107-187,560;
10: 716
C>perus diaphanus Schrader ex
R. & S.
var. latespicalus (Boeck.)
Kern 716
CyrillopsisKuhlm. 621,622
Dacrycarpus (Endl.) dc Laub.
337, 338,343, 347, 351,
353-355. 374, 376 map
cincius (Piiger) de Laub. 376,
383*, 384 map
compaclus (Was.scher) de
Laub. 376, 382* map,
383*. 384
cumingii (Pari.) de Laub.
376, 381 map
dacrydiifoUa (Was-schcr)
Gausscn 383
cxpansus dc Laub. 342. 376,
381, 382 map
imbncatas (Bl.) dc Laub.
340, 341. 376
var. turvulus (Miq.) dc
Laub. 340, 341. 377.
379 map, 380*
var. imbncatus 377 map
var. palulas dc l^ub. 377
map. 378*. 379
(Dacrycarpus imbncatus)
var. robustus de Laub.
377, 379 map
kawaii (Hayata) Gaussen
379
kinabaluensis (Wasscher) de
Laub. 376, 381 map
leptophylla (Wasscher)
Gaussen 371
steupii (Wasscher) de Laub.
340, 376, 380
steupii (non de Laub.) de
Laub. 379
Dacrydium Soland. ex Forsl. f.
337, 338, 342, 343, 347,
352-355, 360, 362 map
balansae 365
beccarii Pari. 362, 366, 367
map
var. rudens de Laub. 368
var. subelatum Comer 363
comosum Comer 363, 370,
371 map
cornwalliana de Laub. 340,
362, 364 map, 366*
cupressinum 365
datum (Roxb.) Wall, ex Hk.
341, 362, 363. 364 map
ericoides de Laub. 340,
341, 363, 371 map
falciforme (Pari.) Pilgcr 372
falciforme [non (Pari.) Pilgcr]
Foxw. ex Mcrr. 373
falciforme [non (Pari.) Pilgcr]
Laut. 373
gibbsiae Siapf 363. 369
gracilis de Laub. 363, 367
map
junghuhnii Miq. 363
leptophyllum (Wa.sschcr)
dcLaub. 363,371
magnum de Laub. 363, 368
map
medium dc Laub. 339-341,
363, 368 map
micropedunculatum 340
nidulum de Laub. 362. 365
map
var. araucarioides dc Laub.
366
novo-guincensc Gibbs 362,
364 map
pctlmatum de Laub. 340.
.34 1 . 362, 364 map
var. robusium dc Laub.
364
pterrii Hickcl 363
spathoidcs dc I-aub. 340, 363,
367 map. 368
(Dacrydium)
xanihandrum Piiger 363,
369*, 370* map
Dactyladenia 639
Dahuronia Scop. 645
Dammara Link 429
alba Rumph. ex Hassk. 435
var. alba Hassk. 435
var. celebica Hassk. 435
celebica Koord. 435
loranthifolia Link 433
motleyi Pari. 393
orienialis Lamb. 433
var. alba Knight ex Henkel
& Hochst. 435
var. orienialis Carr. 433
var. pallens Carr. 435
palmerstonii F.v.M. 442
robusia Moore ex F.v.M.
442
rumpfii auct. 437
rumphii Presl 435
Daphnandra Perkins 256, 261,
263, 265
aromadca 261
novoguineensis Perkins 266
perkinsiae Gilg & Diels 266
Daphne monosiachya Willd. 50
polysiachya Willd. 50
Datiscaceae 4: 382-387; 7:
823
Davisicarpum 158
Decussocarpus de Laub. 389
sect. Dammaroides (Bennett)
de Laub. 390
.sect. Decussocarpus de Laub.
394
maximus de Laub. 394
motleyi (Pari.) de Laub. 393
viiiensis (Seem.) dc Laub.
395
wallichianus (Presl) dc Laub.
393
Dcgcncriaceac 562
Dichapetalaceae 5: 305-316.
567; 6: 941; 7: 823
Diclidanthcra456,457,493
Diemenia Korth. 648
racerrwsa (Korlh.) Miq. 649
Dillcnia 123
Dilleniaceae 4: 141-174;
5: 557; 7: 824
Dimocarpus longan 159
Diogoa 3, 4
Dioscorca 166
aculcata (non L.) Zoll. 199
spiculala Bl. 199
slcnomcridora Pram ik Hurk.
253
728
Flora Malesiana
[ser. I, vol. 104
Dioscoreaceae 4: 293-335;
5: 557; 10: 253
Dioscorcophyllum cumminsii
165
Diploclisia Miers 158, 160,
169, 170, 172, 229
glaucescens (Bl.) Diels 230
glaucescens (non (King)
Diels) sensu Forman 23 1
kunstleri (King) Diels 228*,
230
macrocarpa (W. & A.) Miers
230
Dipsacaceae 4: 290-292;
5: 557
Dipterocarpaceae 9: 237—552;
561; 10: 665, 716
Dipierocarpus comutus Dyer 665
Discogyne Schitr 622
papuana Schitr 627
Dolichandrone spalhacea (L.f.)
K.Sch. 716
Dombeya Lamk. 422
Dor>'phora 261
aromatica 261
Drebbelia Zoll. 6
subarborescens Zoll. 7
Droseraceae 4: 377—381;
5: 557; 6: 943; 9: 562
Dryadodaphne S. Moore 255-
257, 262, 263
celastroides S. Moore 266
crassa Schodde 264*, 265,
266
novoguineensis (Perkins)
A.C. Smith 261, 265,
266, 267
ssp. novoguineensis 266,
267
var. macra Schodde
266
var. novoguineensis
266, 267
ssp. occidentalis
Schodde 265, 267
novoguineensis (Perkins)
A.C. Smith p.p. 265
Drymis-leaved Magnolia Meijer
571, 573
Dugoriia Scop. 654
Dulacia 4
Durandea Planch. 608, 609
jenkinsii (F.v.M.) Siapf 612
magnifolia Stapf 615
pallida K.Sch.eW
pentagyna (Warb.) K. Sch.
612
var. rolundaia (K. Sch.)
Laul. 612
{Durandea)
robinsonii (Merr.) Hall.f. 612
rolundaia K. Sch. 612
Elaeagnaceae 151-156
Elaeagnales 1 5 1
Elaeagnus Toum. ex L. 151,
155*
seel. Elaeagnus 152
sect. Sempervirentes Serv.
152
anguslifolia (non L.) Blco
154
arborea Roxb.
var. dendroidea
Schlechtend. 153.
conferta Roxb. 152, 153,
154*
ssp. dendroidea
(Schlechtend.) Serv.
153
ssp. euconferia Serv. 153
ssp. javanica (Bl.) Serv.
153
var. calcullensis Serv. 153
var. malaccensis Serv. 153
var. pallescens Serv. 156
var. sepientrionalis Serv.
153
var. silheiensis Serv. 153
cumingii Schlechtend. 154
ssp. perrollelii Serv. 154
ssp. philippinensis Serv.
154
dendroidea Sc\\\tz\\\sx\± 153
ferruginea A. Rich. 153
ssp. sumairana Serv. 153
var. airovirens Serv. 153
var. richardia Serv. 153
fruticosa (Lour.) Cheval.
153
gaudichaudiana Schlechtend.
153
javanica Bl. 153
latifoliaL. 152
laiifolia (non L.) A. Rich.
153
var. iriflora Schlechtend.
153
/7erro»e/H Schlechtend. 154
philippinensis Perrotlet 153
rigida Bl. 153
rosiraia Serv. 154
iriilora Roxb. 152, 153
ssp. obsoleia Serv. 153
ssp. polymorpha Serv.
153
ssp. rigida Serv. 153
(Elaeagnus triflora)
ssp. leiragonia Serv. 153
ssp. leiragonia (non Serv.)
Merr. & Perry 155
var. brcvilimbata 't Han
154*, 155
var. brevilimhalus 155
var. brevipes Serv. 153
var. longipes Serv. 153
var. triflora 154*
Zollinger i Serv. 154
Eiaeocarpus 123
Elatinaceae 4: 203-206
Elmerrillia Dandy 562, 564-
567, 593, 595
sect. P seudoaromadendron
Dandy 595
celebica (Koord.) Dandy 596
mollis Dandy 596, 598
ovalis (Miq.) Dandy 595,
596
papuana (Schitr) Dandy 595,
596
var. adpressa Dandy 596
var. glaberrima Dandy
596, 598
platyphylla (Merr.) Noot.
595, 596
pubescens (Merr.) Dandy 595,
596
sericea C.T.White 598
tsiampacca (L.) Dandy 595,
596
ssp. mollis (Dandy) Noot.
567, 598, 599*
ssp. tsiampacca 598
var. glaberrima (Dandy)
Noot. 595, 598
var. tsiampacca 597*,
598
vriesearuj (Miq.) Dandy 596
Elsoia Adans. 483
bracieala (Benn.) O.K. 484
corymbosa (Turcz.) O.K.
484
lavoyana (Benn.) O.K. 484
Embelia urophylla Wall, ex
A. DC. 649
Emmenanihus Hk.f. & Am.
622, 626
chinensis Hk.f. & Am. 626
Entosiphon Bedd. 665
Epacridaceae 6: 422-444,
943; 10: 335
Ephedra 338
Ephippiandra 258
Epicryanlhes 489
Epinetrum 164
Epirhizanlhes 489
1989]
Index to scientific plant names
729
Epirixanthes Bl. 455, 456, 459,
486, 488
aphylla (Griff.) Merr. 490
cylindricaBl. 457,490, 491
elongata Bl. 489, 490*
kinabaluensis Wendt 490,
491
linearis Bl. 490
pallida Wendt 490, 492
papuana J.J. Sm. 489, 490,
491
tenella Hk. f. 490
Epirixanihus 489
Epirizanthes 489
Epirrhizanihe 489
Epirrhizanlhes 489
Eriandra Royen & Sleen. 455,
457, 459, 492
fragrans Royen & Sieen. 493
Ericaceae 6: 469-914; 943;
7: 827; 8: 549; 9: 562;
10: 335, 716
Ericybe Roxb. 29
Eryihropalla 17
Erythropalum Bl. 1-6, 17, 166
grandifoliwn Elmer 17
scandens Bl. 17, 18*
var. abbrevialum Hochr. 17
iriandrum Quis. & Merr. 29
vagum (Griff.) Mast. 17
Erythroropalum 17
Erythroxylaceae 5: 543-552;
8: 549; 10: 607, 609, 622
Escalloniaceae 335
Euglypha 53, 61,63
Euphorbiaceae 29, 59
Euiacta cunninghamii (Ait.)
Link 426
Eutassa Salisb. 425
cunninghamii Spach 426
Evodia 123
Exellodendron Prance 642
coriacca 64 1
Exiielea Bl. 642
Exiteles 673
ExiteliaBl61\
corymbosa (Bl.) Bl. 673
muliiflora (Konh.) Walp.
673
Eystathes Lour. 493, 507
Fagaceae 7: 265-403; 8:
549; 9: 563
Faika Philipson 255. 262. 263.
284 map
villosa (Kanch. & Hatus.)
Philip.son 284. 285*.
286*
Falcalifolium de Laub.343, 347,
352, 354, 355, 371
angustum de Laub. 372, 373
map, 374
falciforme (Pari.) de Laub.
372, 373* map
gruezoi de Laub. 372, 373
map
papuanum de Laub. 372, 373
map
Fawceltia F.v.M. 164, 188
me rril liana (Diels) Yamamoto
193
Ferolia O.K. 655
asperula (Miq.) O.K. 671
corymbosa (Bl.) O.K. 674
cosiata (Konh.) O.K. 663
glaberrima (Hassk.) O.K.
669
griffithiana (Benth.) O.K.
674
jackiana (Benth.) O.K. 671
/jmda(Hk.f.)Ridl. 646
nonda (F.v.M. ex Benth.)
O.K. 658
oblongifolia (Hk.f.)O.K.
659
polyneura (Miq.) O.K. 664
salicifolia (Presl) O.K. 674
scabra (Hassk.) O.K. 669
sumatrana (Jack) O.K. 661
Fibraurea Lour. 157, 160-162,
166-168, 170, 171, 207
chloroleuca Miers 160,207
chloroleuca (non Miers) Merr.
184
elliplica Yamamoto 184
fasciculata Miers 209
haematocarpus Hk.f. & Th.
184
laxa Miers 209
recisa 207
lincloria Lour. 160, 165,
207, 208*
Fibraureopsis Yamamoto 1 83
smilacifolia Yamamoto 1 84
Ficus/7u/cAra Wall. 323
Fissipelalum Merr. 29
Flacourlia camptoceras Miq.
618
Flacourtiaceae 5: 1-106.565;
6: 943; 7: 827; 9: 563;
10: 333. 539, 715, 716
Flagellariaceae 4: 245-250;
5: 557; 9: 564
Folium lunalum minus Rumph.
229
Frcnclopsis 338
Funis felleus Rumph. 194
Ganua palembanica (Miq.) v.d.
Assem & Kosterm. 453
sp. 453
Geobalanus Small 645
Geraniaceae 6: 445-449;
9: 565; 10: 607, 639
Geraniales 609, 639
Gesneriaceae 335
Gigantopteris 338
Gjellerupia Laut. 31-35, 45
papuana Laut. 45*, 46
Gladiolus dalenii Geel 717
natalensis (Ecklon) Reinw. ex
Hk. 717
Glossocalyx 258
Glyptopetalum loheri Merr. 716
Gnetaceae 4: 336-347; 6:
944
Goodeniaceae 5: 335-344;
567; 6: 949; 7: 827; 9:
566
Gordonia decandra Roxb. 626
Govantesia Llanos 35
malulucban Llanos 36
Grossulariaceae 335
Groutia Guill. & Perr. 46
cellidifolia Guill. & Perr. 47
Grymania Presl 67 1
salicifolia Presl 671, 673
Guatteria incerta Bl. 515
Guttiferae 145
Gymnostoma sp. 453
Haematocarpus Miers 160, 161,
167-169, 171, 183
complus Miers 1 84
incusus Miers 1 84
subpeltatus Merr. 182*. 184
ihomsonii Miers 184
validus (Miers) Bakh.f. ex
Forman 184
Haemodoraceae 5: 111-113;
10: 717
Haemodorum coccineum R. Br.
717
corymbosum Vahl 7 1 7
Halocarpus 354
Haloragaceae 7: 239-263,
828
Ilamamelidaceae 5: 363-
379; 6:952; 10: 341,717
Harmandia Pierre ex Baill. 1-6.
9. 10 map
kunstleri King 9
mckongensis Pierre ex Bail!.
9, 10*
Hcbcpctalum 608
Ucckelia nymanii K.Sch. 253
730
Flora Malesiana
[ser. I, vol. 104
Hedycarya 255, 259, 260
arborea 256
salomoncnsis Hemsl. 311
Hedycrea Schrcb. 645
Hcdyosmum Swarlz 123-129,
143. 144
arboresccns 125, 127
brasiliense 143
mexicanum 125
nutans (non Sw.) Merr. 144
orientale Merr. & Chun
142*, 144 map
sp. Sieen. 144
sumatranum 144
Hcisieria 3,4
micranlha 3
Hcnnecartia 257
Ilernandia kunslleri 9
Ueierosamara birmanica (O.K.)
Chodat 463
Heterotropa 53, 63
Hexastylis 53, 63
Hibbenia 123
Himantandraceae 562
Hippophae L. 151
Ilolopeira Miers 231
australis Miers 234
laurifolia (DC.) Miers 234
Holostylis 53,61,63
Homalium dasyanthum 716
Hopca gracilis Miq. 716
Horsfieldia iryaghedi (Gaertn.)
Warb. 605
Hortonia 258, 259
Hortoniaceae 261
Hugonia L. 608, 609
sect. Durandea (Planch.)
BaUIon609, 611
sect. Hugonia 611
costata Miq. 610*, 611, 612
map
jenkinsii F.v.M. 611, 612
map, 613*
pentagyna (Warb.) K.Sch.
612
robinsonii Merr. 612
sumatrana Miq. 618
Hugoniaceae 607, 608, 629
Humiria 629
Humiriaceae 607, 609, 629, 630
Hunga Pancher ex Prance 635-
637, 642, 643, 650
fusicarpa Kosterm. 647
longifolia Prance 651, 653*
map
novoguineensis Prance 651,
652*, 653 map
papuana (Baker f.) Prance
651, 652*. 653 map
Hyalisma 112
Hydnocarpus nana King 716
Hydrocaryaceae 4: 43-44
Hydrocharitaceae 5: 381—
413,569; 6: 952; 7:
828; 9: 566; 10: 112,
717
HydrophyUaceae 4: 207-209
Hypericaceae 8: 1-29; 10:
717
Hypericinea macrocarpa Wall.
626
Hypericum henryi L6v. & Vaniot
ssp. hancockii Robson
717
uralum Buch.-Ham. ex
D. Don 717
Hyperixanthes 489
Hypserpa Miers 157, 161, 167-
169, 172, 222, 218
borneensis (Miq.) Becc.
219
cu5p/Jafa(Hk.f. &Th.)
Miers 219
var. imcrophylla (Miq.)
Boerl. 219
heteromera Miers 219
jagorii'D'itXs 2\9
latifolia Miq. ex Diels 221
laurina (F.v.M.) Diels 218,
219
monilifera (Burk.) Diels
221
nandinifolia Yamamoto
219
nitida Miers 219, 220*
parvifolia Kaneh. & Hatus.
219
polyandra Becc. 219, 221
var. tomentosa Forman
221
praevaricaia Miers 219
propensa Miers 219
raapii Diels 221
selebica Becc. 221
selwynii F.v.M. 219
iriflora [non (DC.) Miers]
Miers 219
Hypsipodes Miq. 188
Iberis amara L. 543
umbellaia L. 543
Icacinaceae 7: 1-87, 828; 9:
566; 10: 29, 33, 91, 145,
166, 629, 717
Idenburgia Gibbs 145, 146,
326, 333
arfakensis Gibbs 149
{Idenburgia)
elaeocarpoides Gilg & Schllr
147
novoguineensis Gibbs 147
pachyphylla Gilg & Schltr
147
pauciflora A.C. Smith 147
Indorouchera HaU.f. 607-609,
615
contestiana (Pierre) Hall. f.
616*, 619 map
griffithiana (Planch.) Hall.f.
616*, 617*, 618* map
rhamnifoUaWaXU. 6\9
Iridaceae 8: 77-84; 10: 717
Irina integer rima Bl. 702
Irvingiaceae 621 , 622
Isomerocarpa A.C. Smith 263
novoguineensis (Perkins)
A.C. Smith p.p. 265, 266
Ixionanthes 622
Ixonanthaceae 607, 609, 621-
627, 629
Ixonanthes Jack 621, 622, 623*
sect. Brewstera (M.J.
Roemer) Hall.f. 621, 625
sect. Emmenanthes Hall.f.
626
sect. Ixonanthes 621, 625,
626
beccariiWalU.en
chinensis (Hk. f. & Am.)
Champ. 626
cochinchinensis Pierre 626
crassifolia Hall.f. 627
cuneata Miq. 625
dodecandra Griff. 625
grandiflora Hochr. 626
grandifolia Ridl. 627
hancei Pierre 626
icosandra Jack 624*, 625
var. cuneata Miq. 625
var. obovata Ridl. 625
khasiana Hk. f. 626
longipedunculata Merr.
627
lucida Bl. 625
multiflora Stapf ex Ridl.
626
obovata mi.i. 615
papuana (Schltr) Hub.Winkler
627
petiolaris Bl. 623, 625, 626,
627*
petiolaris (non Bl.) Hall.f.
627
philippinensis Elmer 626
reticulata Jack 625, 626
subdodecandra 625
1989]
Index to scientific plant names
731
Jackia longifolia 514
vitellina 514
Jacquemontia browniana Oosistr.
716
pannosa (R.Br.) Mabberley
716
Jakkia Bl. 493, 508
excelsam.5Q\
longifolia Bl 514
vitellina Bl. 514
Juglandaceae 6: 143-154,
953
Juncaceae 4: 210-215; 5:
557; 6: 953; 9: 566
Juncaginaceae 4: 57; 5: 554
Juniperus 345
chinensis 453
elata Roxb. 363
Juppia borneensis Merr. 253
Kairoa Philipson 255, 256, 258,
259, 262, 263, 284 map,
305
suberosa Philipson 304*,
305, 306*
Kibara Endl. 255, 257, 259,
260, 262, 263, 287, 308
angusiifolia Perkins 298
archboldiana A.C. Smith
257, 289, 291, 293
arucnsis Becc. 305
blumei Sleud. 298
borneensis Boerl. 305
buergersiana Perkins 305
bullata Philipson 289,
293
carrii Philipson 257, 290,
291, 297
chariacea Bl. 298
chimbuensis Philipson 290,
301
clemensiae Perkins 298
coriacea (Bl.) Tulasne 262,
290, 298
cuspidaia Bl. 298
depauperata Merr. 3(X)
dichasialis Suesscng. &
Heme 298
ellipsoidea Merr. 298
clmeri Perkins 305
clongata A.C. Smilh 290.
297
fcrox Philipson 257. 258.
289. 291. 292*
flagclliformis Philipson 290.
301
form icar urn Becc. 305
fragrans Philipson 290. 300
(Kibara)
fugax Philipson 290, 300,
301, 302*
grandifolia Merr. 298
hartleyi Philipson 289, 295,
296
hirsutaV^aih. 315
hospitans Becc. 311
inamoena Perkins 298
karengana Philipson 289,
293
katikii Philipson 290, 298
kosiermansii Philipson 289,
294
lalifolia Philipson 257, 289,
291
laurifolia A.C. Smilh 290.
300
leachii Philipson 289, 294
ledermannii Perkins 296
longipes Perkins 298
macrantha Philipson 288*.
289, 297, 300
macrocarpa Perkins 298
macrophylla Perkins 298
merrilliana Perkins 298
microphylla Perkins 289,
296
mollis Merr. 298
moluccana Perkins 288*,
289, 295, 296, 303
monticolia Perkins 290, 303
motleyi Perkins 298
myrtoidea Perkins 289, 291
ncriifolia Perkins 305
nitens Philipson 290, 299
novobrilanica Philipson 290,
301
oblongata Philipson 288*.
289, 295
oblusa Bl. 290, 300
oligocarpclla (Kaneh. &
Hatus.) Philipson 256,
289, 291
olivaeformis Becc. 305
papuana A.C. Smith 290.
299
pcrkinsiae K.Sch. & Laul.
305
rigidifolia A.C. Smilh 256.
258, 288, 290, 302
rocmeri (Perkins) Philip.son
289, 290
rossclcnsis Philipson 289.
295
roycnii Philipson 289, 295
serrulula Pcrkms 298
shungolcnsis Philipson 289.
296
(Kibara)
sleumeri Philipson 290, 303
stapfiana Perkins 298
streimannii Philipson 288,
289, 296
sudestensis Philipson 290,
303
symplocoides Perkins 289,
290, 303
leijsmanniana Perkins 296
timorensis Boerl. 305
tomeniosa Perkins 298
irichantha Perkins 298
versieeghii Philipson 289,
294
vidalii Perkins 298
vrieseana Perkins 296
warburgii Perkins 298
warenensis Kaneh. & Hatus.
289, 295
Kibaropsis 260
Kingsboroughia Liebm. 690
Kmeria 566
Korihalsella 353
dacrydii 353
Kosiermanthus I*rance 636-
639, 641, 642, 675
heteropetalus (Scort. ex King)
Prance 676*. 677 map,
678
malayanus (Kosterm.) Prance
677 map
Krameriaceae 457, 458
Labiatae 8: 301-394; 9:
566
Lagarosirohus Quinn 360, 361
Lardizabalaceae 163, 679
Larix 343
Lauraceac539,715,717
Lauralcs 127,327,330
Lautcrbachia Perkins 263. 284
map, 326
novoguineensis Perkins
325*. 326
Lavallea Baill. 19
ceylanica (Gardn.) Baill.
22
philippinensis Baill. 22
Leeaceae 7: 755-782
Legnephora Micrs 158. 165,
169, 170, 172, 225
acuta Forman 159. 225.
226*
microcarpa Forman 225.
226*. 227
minutidora (K.Sch.) Dicis
225. 226*. 227. 228*
732
Flora Malesiana
[ser. I, vol. 104
(Lcgnephwa)
moorei (F.v.M.) Miers 225,
226*
nyctericarpa Diels 227
philippinensis Forman 225,
226*
Lemnaceae 7: 219-237
Lentibulariaceae 8: 275-300
Lepidium L. 541,543,545,547
lacteviride (P. Royen)
Hewson 548, 549*
maccowagei Hews. 547,
549
minuliflorum (Ridl.) Hews.
548
sativum L. 547, 548
virginicum L. 547, 548
Lepidocarpa Korlh. 655
costata Korth. 663
ovalis (Korth.) Bl. ex Miq.
661
Lepidocarya costata 663
Lepidothamnus Phil. 360
Lepionurus Bl. 31-35, 42, 43
javanicus G. Don 43
oblongifolius (Griff.) Mast.
44
var. angustifoUus Roxb.
44
pubescens Ridl. 52
sylvestris Bl. 31, 33, 34, 43,
44* map, 46
var. lanceolata Val. 43
Leptonium Griff. 43
oblongifolium Griff. 43
Levieria Becc. 263, 275
acuminata (F.v.M.) Perkins
277, 279
beccariana Perkins 277, 280*,
281*
forbesii Perkins 281, 282
laxiflora Perkins 279, 281
montana Becc. 277, 279,
281,282
montana (non Becc.) Kaneh.
& Hatus. 277
nitens Perkins 277, 278
orientalis Philipson 277, 278
parvifolia A.C. Smith 277
rudolfii Perkins 278
scandens Philipson 277, 279
schlechteri Perkins 278, 279,
281
squarrosa Perkins 276*, 277,
278
urophylla Perkins 279, 281
Levieria Kosterm. p.p. 263
Libocedrus Endl. 337, 339, 340,
342, 347, 443, 444 map
(Libocedrus)
subg. Eulibocedrus Pilger
443
arfakensis Gibbs 446
papuana F.v.M. 342, 444,
445*
var. arfakensis (Gibbs)
de Laub. 446, 447 map
var. papuana 445, 446*,
447 map
torricellensis Schltr ex Laut.
444
Licania Aubl. 635-640, 642,
643, 645
angelesia Bl. 646
diemenia Bl. 649
elaeosperma 640
fusicarpa (Kosterm.) Prance
646, 647, 650 map
macrophylla 639
palawanensis Prance 646,
647 map
splendens (Korth.) Prance
640, 646, 647 map
Liliaceae 9: 189-235; 10:
109, 112, 166
Limacia Lour. 158, 160, 161,
167, 168, 170, 172, 221,
222
blumei (Boerl.) Diels 222,
223
borneensis Miq. 219
cerasifera Becc. 224
cerasifolia 224
cuspidata Ek.f.&Th. 219
distincta Miers 223
inornata Miers 223
kunstleri King 233
microphylla Miq. 219
monilifera Burk. 221
nativitatisRidl. 218
oblongaHk.f. & Th. 160,
165, 166, 223
scandens Lour. 220*, 223
selwynii (F.v.M.) Bailey
219
sumatrana Scheff. 198
triandra (Colebr.) Hk.f. & Th.
185
velutina Hk.f. & Th. 160,
223
var. glabrescens King 223
Limnanthaceae 639
Linaceae 29, 607-619, 621,
622, 629, 630, 639
subfam. Hugonioideae 607,
608, 609
subfam. Ixonanthoideae 621
subfam. Linoideae 607, 609
Linum 609
Liriodendron 562, 563, 565-
567, 605
liliiferum L. 582
tulipifera 567
Liriopsis Spach 598
fuscata (Andr.) Spach 598
Litsea baticulin (Blco) Kosterm.
717
insignis (Bl.) Boerl. 539
leytensis Merr. 717
Lobbia Planch. 65
de pendens Planch. 75
Lobularia maritima (L.) Desv.
545
Loganiaceae 6: 293-387,
953,960; 7: 828; 9:567
Longpistillate Manglieiia Meijer
591
Lophopetalum beccarianum
Pierre 716
Lophopyxidaceae 7: 89-91
Lophostylis Hochst. 483
javanica Miq. 484
Loranthaceae 2, 3, 34, 35
Macadamia 123
Macrococculus Becc. 157, 158,
160-162, 167-169, 171,
178
pomiferus Becc. 159, 178,
179, 180*
tympanopodus Laut. &
K.Sch. 211
Madhuca burckiana (Koord.)
Lamk 593
Magnolia L. 564, 566, 567,
568
sect. Gwillimia 566, 569
sect. Lirianthe 565
sect. Liriopsis Baillon 598
sect. Maingola 565, 567
sect. Manglietia (Bl.) Baillon
589
sect. Talauma Baillon 568
sect. Theorodon 569
subg. Magnolia 564-567,
570
sect. Maingola Dandy 569,
570
subg. Talauma (Juss.) Pierre
564-567, 576
sect. Aromadendron (Bl.)
Noot. 566, 569, 576
sect. Blumiana Bl. 565,
566, 569, 581
subg. Talauma Pierre 568
subg. Yulania 565
1989]
Index to scientific plant names
733
(Magnolia)
aeqidnoctialis Dandy 57 1
angaiensis Blco 585
ashtonii Dandy ex Ncx)t. 570,
576, 578*, 579
ashtonii Noot. 579
beccariana (Agoslini) Nc»L
571
betongensis (Craib) H. Keng.
585
bintuluensis (Agoslini) Noot.
570, 576, 577
blumei Pranil 589
bomeensis Noot. 570, 576,
579, 580*
candollii (Bl.) H. Keng 569,
581, 582
var. angaiensis (Blco)
Noot. 582, 585
var. beccarii (Ridl.) Nool.
582, 586
var. candollii 582, 584*
var. obovaia (Korih.)
Nooi. 582, 585
var. singapurensis (Ridl.)
Nool. 582, 586
carsonii Dandy ex Nool. 570,
571
var. carsonii 572*, 573,
576
var. drymifolia Nool.
573, 574*
carsonii Dandy ex Cockbum
571, 572
coco (Lour.) DC. 569
decandollti 5S\, 5^2
denudata 562
drymifolia Dandy ex
Cockbum 571, 573
elegans (HI.) H. Keng 570.
576. 577
forbesii King 582
fragrans Reinw. ex Bl.
583
giganlifolia (Miq.) Nool.
581, 586
grandi flora L. 569
javanica K. & V. 571
kachirachirai 563
kunsllen King 582
lasia Noot. 581. 587
lilufera (L.) Baillon 582
macklottii (Korlh.) Dandy
569. 570. 571
var. bctcariana (Agoslini)
N(x)l. 571
var. macklottii 571
maingayi King 570. 571,
573
(Magnolia)
maingayi (non King) Ridl.
577
mariusjacobsia NooL 581.
588
nilida 563
nutans (Dandy) H. Keng 577
odoratissima Reinw. ex Bl.
583
pachyphylla Dandy 583
paenetalauma 566
pahangensis Noot. 570. 576,
579
pealiana (non King) K. & V.
571
pealii 571
persuaveolens Dandy 581,
587
ssp. persuaveolens 587
ssp. rigida Nool. 587
var. pubescens NooL
587. 588
var. rigida 587
phaulantha Dandy ex Noot.
570. 571. 576
Philippine nsis Parmeni. 604
plumierii Schwartz 568
pumila aucl. 582. 583
rumphii Spr. 582
sarawakensis (Agoslini)
Noot. 581, 588
singapurensis (Ridl.) H. Keng
586
splendens Reinw. ex Bl. 582
sprengeri 567
stellata 562
uvariafolia Dandy ex
Cockbum 574
uvariifolia Dandy ex Noot.
570. 571. 574, 575*
villosa (Miq.) H. Keng 581.
588
virginiana L. 568
vrieseana (Miq.) Baillon ex
Pierre 596
Magnoliaceae 261. 561-605
subfam. Liriodendroidcac 561.
564
subfam. Magnolioideae 561.
563. 564. 567. 568
tribe Magnolieae 561,
568
tribe Michelicac Law Yuh-
wu 561. 568. 593
Magnolialcs 62, 64, 261
Maingaya malayana Oliver 7 1 7
Malania 1
Maipighiaceae 5: 125-145,
566; 6: 960; 10: 458
Malulucban 35
Malvanae 151
Manglietia Bl. 564-568, 589
calophylla Dandy 589, 591
candollii (Bl.) WaU. 582
celebica Miq. 582
dolichogyna Dandy ex NooL
589, 590*, 591
dolichogyna Dandy ex
Cockbum 591
glauca Bl. 589, 590*
var. glauca 589, 590
var. lanuginosa Dandy
591
var. sumatrana Dandy
589, 591
glauca (non Bl.) King 591
g/auca (non Bl.) Ridl. 591
glauca aucL 585
lanuginosa (Dandy) Nool.
589, 591
macklottii Korlh. 571
macklottii aucl. 589
minahassae K. & V. 593
oortii Korlh. 577
oortii (non Korlh.) Miq. 577,
589
oortii aucL 589
pilosa Parmeni. 590. 591
sabahensis Dandy ex Nool.
589. 591. 592*
sabahensis Dandy ex
Cockbum 591
scortechinii King 60 1
sebassa King 582
singalanensis Agoslini 590
sumatrana Miq. 589, 591
Manglieliastrum 563
Martyniaceae, see Pedaliaccac
Malthaea Bl. 255, 258, 262,
263, 288, 319, 321 map
calophylla Perkins 323
charlacea Merr. 321 map,
322*. 323
heierophylla Quis. & Merr.
321 map, 322*. 323.324
intermedia Merr. 321 map,
322*, 323, 324
laiifolia Perkins 323
philippinensis Perkins 326
pincholiana Perkins 323
pubescens Merr. ex Perkins
321 map. 322*. 323
roemcri Perkins 290
sancta Bl. 320*. 321 map.
322*. 323
var. mindanaoensis
Perkins 323
var. venulosa Perkins 323
734
Flora Malesiana
[ser. I, vol. 104
(Malthaea)
vidalii Perkins 321 map,
322*. 323, 324
williamsii Perkins 323
Matlhiola incana (L.) R.Br. 545
Meiogyne virgata (BI.) Miq. 716
Melientha Pierre 31-34, 38, 40
map
acuminata Merr. 39
suavis Pierre 34, 39*
ssp. macrocarpa Hiepko
40 map
ssp. suavis 40* map
Meliosma Bl. 163, 679-682,
690, 691*, 692*
subg. Kingsboroughia
(Liebm.)Beus. 691*.
694
sect. Hendersonia Beus.
692*, 694
sect. Kingsboroughia
692*, 694
subg. Meliosma 691*, 693,
694
sect. Lorenzanea (Liebm.)
Beus. 694
sect. Meliosma 692*, 693
subsect. Pinnatae
(Warb.) Beus. 693,
694
subsect. Simplices
(Warb.) Beus. 693,
694
ser. Curvinervia
Beus. 693
ser. Rectinervia
Beus. 693
acutrdnatissima Merr. 712
angulata Bl. 690, 698
apoensis Elmer 709
arnoitiana Walp. 709
bartlettUM&n. 710
bonlocensis Merr. 697
brachyboirys Merr. 713
cannarioides Elmer 709
celebica Warb. ex Dihm
715
conferiiflora Merr. & Perry
710
confusa Bl. 702
var. laxior Baker f. 713
costata CvSod. 701
cuspidata Bl. 702
diepenhorstii Valet. 702
dolichomischa Vidal 696
elegans Rm. 110
ellipticaHk.f. 701
elmeri Merr. 702
evrardii Gagnep. 701
(Meliosma)
ferruginea Bl. 711
ferruginea (non Bl.) Backer &
Bakh.f. 709
ferruginea (non Bl.) Koord.
709
ferruginea (non Bl.) Merr. &
Perry 711
floribunda Bl. 709
fruticosa Bl. 701
glauca Bl. 709
var. floribunda (Bl.)
K. & V. 709
glomerulaia Rchd. & Wils.
699
grandifolia Lecomte 713
harmandiana Pierre 699
hirsuta Bl. 694, 703 map,
705, 715
humilis Merr. & Perry
713
lanceolata Bl. 694, 703 map,
704, 715
var. chariacea Bl. 704
var. genuina Hochr. 704
var. lanceolata 705
f. lanceolata 705
f. nervosa (K. & V.)
Beus. 694, 705
var. membranacea Bl. 704
var. obliqua Bl. 704
var. obliqua (non Bl.)
Koord. 711
var. pendula Bl. 704
var. polyptera (Miq.) Beus.
703 map, 705
/anc//b//aHk.f. 701
latifoliaKxfMlM
laurina BIT \5
lepidota Bl. 694, 695
ssp. dolichomischa (Vidal)
Beus. 695, 696*
ssp. kinabaluensis Beus.
695, 697
ssp. lepidota 695
ssp. squamulata (Hance)
Beus. 697
ssp. vulcanica (Merr.)
Beus. 695, 697
levis King 704
loheri Merr. 701
luzonensis l(y)
luzonica 709
macgregorii Merr. 712
macrocarpa Elmer 7 1 1
macrophylla Merr. 711
megaloboirys Merr. 711
monophylla Merr. 701
monophylla Ridl. 696
(Meliosma)
multiflora 709
myriantha Sieb. & Zucc. 681
nervosa K. & V. 704, 705
nitida Bl. 702
var. cerasiformis Bl. 702
var. splendens BI. 702
var. tridenta Bl. 702
pannosa Hand.-Mazz. 701
patens Hemsl. ex Forb. &
Hemsl. 699
paucinervia Merr. 710
pedicellata K. & V. 695
pedicellata (non K. & V.)
Merr. 697
pendula Merr. 712
petiolaris Miq. 715
philippinensis Merr. & Perry
702
pinnata (Roxb.) Maxim. 694,
707
ssp. amottiana (Wight)
Beus. 707, 708 map,
709
ssp. ferruginea (Bl.) Beus.
707, 708 map, 709,
710
ssp. humilis (Merr. &
Perry) Beus. 707, 708
map, 713
ssp. macrophylla (Merr.)
Beus. 706*, 707, 708
map, 709, 711
ssp. pendula (Merr.) Beus.
707, 708 map, 709,
712
ssp. ridleyi (King) Beus.
707, 708 map, 709,
710
ssp. sylvatica (Elmer)
Beus. 707, 708 map,
709, 712
pinnata Koord. 702
polyptera Miq. 704, 705
pungens (Wall, ex W. & A.)
Walp. 702
pungens auct. 699
reticulata Merr. 712
ridleyi King 710
rigida Sieb. & Zucc. 699
var. angustifolia Miq.
699
var. patens Cufod. 699
rufo-pilosa Hend. 694, 703
map, 714
sambucina Miq. 709
sarawakensis 694
schlechteri Merr. & Perry
713
1989]
Index to scientific plant names
735
(Meliosma)
simplicifolia (Roxb.) Walp.
681, 694, 697*, 698,
699 map
ssp. fruticosa (Bl.) Beus.
697*, 698, 699 map,
700*, 701
ssp. pungens (Wall, ex
W. & A.) Beus. 697*,
698, 699*. 702
ssp. rigida (Sieb. & Zucc.)
Beus. 697*, 698, 699
map
ssp. simplicifolia 690,
697*, 698, 699 map
sumatrana (Jack) Walp. 694,
702, 703 map, 715
sylvatica Elmer 712
limorensis Bl. ex Blenk 715
tongcalingii Elmer 711
trichocarpa Merr. 710
vulcanica Mcrr. 697
wallichii (non Planch, ex
Hk.f.)Koord. 711
wightii Planch. 702
Meliosmaceae End!. 679, 680
Memecylanthus Gilg & Schltr
335
Meniscosta Bl. 682
javanica Bl. 684
scandens Bl. ex Spr. 684
Menispermaceae 19, 157-
253, 305, 679, 680.
681
iribc Coscinieac 161, 162,
164
iribe Fibraureeae 162, 164,
168
tribe Menispermeae 161, 162,
164, 168
iribc Tiliacoreae 162, 164,
168
iribc Tinosporcae 158, 161,
162, 164, 167, 168, 189*
iribc Triclisieae 162
Mcnispcrmum 158, 165
cocculus L. 213
crispum L. 194. 195
dauhcum 165
fenesiralum Gaerin. 215
flavescens Lamk 210
flavum L. 210
glabrumBumn.f. 199,200
glaucum Lamk 229
japonicum TTiunb. 245
lacunosum Lamk 213
limacia (DC.) Sprcng.
223
nuUabaricum Lamk 200
(Menispermum)
orbiculatum L. 231
ovalifolium Vahl ex Pers.
iriandrumRoxh. 185
trilobum Thunb. 231
tuber culalum Lamk 194
Metasequoia 345
Michelia L. 562, 564-567.
593, 595, 598, 605
X alba DC. 601,605
arfakiana Agoslini 596
beccariana Agoslini 571
blumei Steud. 601
celebica Koord. 596
champaca L. 562, 565, 598,
601
var. champaca 603
var. pubinervia (Bl.) Miq.
603
champaca Miq. 603
champacca aucl. 596
cumingii Merr. 604
dollsopa auct. 589
ecicatrisaia Miq. 603
figo (Lour.) Spr. 599, 605
forbesii Baker f. 596
fuscala 562
koordcrsiana Nooi. 601,
602*
longiflora 605
longifolia Bl. 605
var. racemosa Bl. 605
mollis (Dandy) McLauchlin
596
moniana Bl. 563, 601, 603,
604 map
var. subvelulina Miq. 604
montana aucl. 603
parviflora 604
philippinensis (Parment.)
Dandy 601,604
p////era Bakh.f. 603
platyphylla Merr. 596
pubinervia Bl. 603
rufinervis Bl. 603
salicifolia Agoslini 601, 604
scoriechinii (King) Dandy
600*, 601
spec. Slapf 587
suaveolens Pers. 601
sumatrae Dandy 604
isiampacca Bl. 603
tsiampacca L. 596
var. blumei Mor. 603
velutina Bl. 603
Microcachrys338. 343. 351.
352
Microcarpus 35)
Microdcsmis Hk.f. 29
Microtinomiscium 158
Milling tonia Roxb. 690
arnoitiana Wighl 709
ferruginea Schull. & Schull.
711
lanceolata Schull. & Schull.
704
nitida Schull. & Schull. 702
pinnata Roxb. 707
pungens Wall, ex W. & A.
702
sambucina Jungh. 709
simplicifolia Roxb. 697, 698
sumatrana Jack 702
Minquarlia 3
Miquelia 166
Mirtana loureiri (Pierre) Pierre
210
Modeccopsis vaga Griff. 17
Mollinedia 256
acuminata F.v.M. 279
coriacea (Bl.) Baill. 298
sancta (Bl.) Baill. 323
Molluginaceae, sec Aizoaceae
Monaria Konh. ex Val. 17
Monimia 257-261
oval i folia 257
Monimiaceae 145,255-326,
256 map, 327, 330
subfam. AUierospcrmaloidcac
255, 258-263
subfam. Horionioideae 260,
261
subfam. Mollinedioideae 259,
261,263
subfam. Monimioideae 259-
263, 277
subfam. Siparunoideae 258-
261
tribe Hcdycaryeae 255, 259,
262, 263, 277
tribe Mollincdieae 255, 262,
263, 277, 284 map, 326
tribe Monimieae 259
tribe Trimcnicac 327
Monnina 456
emarginaia458
ejcce/.va(BI.)Spr. 501
longifolia {E\.)S\)v.5\A
macrophylla Slcud. 514
vj/e/Z/na (Bl.) Spr. 514
Monoon incertum (Bl.) Miq.
515
Monquarlia 4
Moquilea Aubl. 645
sect. Cyclandrophora (Ha.ssk.)
Endl. 665
Morinjjaccae 4: 45—46; 5:
554; 6: 960
736
Flora Malesiana
[ser. I, vol. 104
Moutabea Aubl. 456, 493
Munnickia Bl. ex Rchb. 65
Myoporaceae 4: 265—266;
9: 568
Myricaceae 4: 276-279
Myristicaceae 605
Myrtales 151
Myrtanae 1 5 1
Myrtiflorae 151
Nageia Gaertn. 337, 343, 347,
353-355, 389, 391 map
sect. Nageia 390
sect. Polypodiopsis (Bertrand)
de Laub. 390, 394
amara (Bl.) O.K. 387
beccarii (Pari.) Gordon 393
blumei (Endl.) Gordon 393
cmot//i^/7 (Pari.) O.K. 381
discolor (Bl.) O.K. 400
falciformis (Pari.) O.K. 372
leptosiachya (Bl.) O.K. 400
maximus (de Laub.) de Laub.
339, 394 map
motleyi (Pari.) de Laub. 393,
394 map
neglecta (Bl.) O.K. 400
neriifolia (D. Don) O.K. 400
polystachyus (R. Br. ex
Endl.) O.K. 417
/■Mm/7;://(Bl.)F.v.M. 415
teysmannii (Miq.) O.K. 406
thevetiaefolia (Bl.) F.v.M.
417
vitiensis (Seem.) O.K. 394
map
wallichiana (Presl) O.K.
391, 392*, 393 map
Najadaceae 6: 157—171
Nallogia Baill. 35
gaudichaudiana Baill. 36
Nasturtium R.Br. 555
backeri Schulz 556
benghalense DC. 560
diffusum auct. 559
heterophyllum Bl. 559
homalospermum Schulz 557
var. macrocarpum Schulz
557
hybospermum Schulz 559
indicum (L.) DC.
vai.javana Bl. 559
indicum auct. 557, 559
novo-guineense Gilli 557
officinale R.Bt. 555
palustre (L.) DC. 557
peekelii Schulz 557
schlechieri Schulz 559
Natsiatum Rheede 213
Ncmuaron 256
Neocarya Prance 642
macrophylla (Sabine) Prance
641
Nephroia Lour. 231
elegans Ridl. 233
sarmentosa Lour. 233
Nigrina Thunb. 129
spicala Thunb. 133
spicifera Lamk 133
Nouhuysia Laut. 145, 146
arfakensis (Gibbs) Steen. 149
novoguineensis (Gibbs)
Hatus. 147
pachyphylla (Gilg & Schltr)
Hatus. 147
papuana Laut. 147
pauciflora (A.C. Smitji)
Steen. 147
Nyctaginaceae 6: 450-468;
7: 829
Nyssaceae 4: 29-31
Ochanostachys Mast. 1-6, 12,
14 map
amentacea Mast. 5, 13*, 14
bancana (Becc.) Val. 14
Ochnaceae 7: 97-119; 10:
145, 621
Ochthocosmus Benth. 621
Octarillumfruticosum Lour.
153
Octoknema 3
Octoknemaceae 33
Odontocarya 158, 164
Odonlocaryioides 158
Olacaceae 1-29, 166, 629,
717
tribe Agonandreae 34
tribe Anacoloseae 3
tribe Aptandreae 4
tribe Couleae 4
tribe Heisterieae 4
tribe Olaceae 4
tribe Opilieae 34
tribe Schoepfieae 4
Olacales 33, 34
Olacinea ignota 25
Olacineae 29, 34
Olax L. 1-5,6
sect. Triandrae Engl. 7
baticulin Blco717
benthamiana 3
imbricata Roxb. 7, 8
laxiflora Ridl. 9
muliiflora A. Rich, ex Baill. 8
multiflora Ridl. 9
(Olax)
obtusa Bl. 7
psittacorum (Willd.) Vahl
717
rosea Ridl. 9
scandens Roxb. 2, 7, 8*, 9,
717
semiinfera Val. 9
stricta 2
sumatrana Miq. 9, 50
Onagraceae 8: 98-113
Ongokea 3, 4
Ophiocaryon Schomb. 163, 679,
680
Opilia Roxb. 4, 31-35,46
sect. Lepionurus (Bl.) Baill.
sect. Opiliaslrum Baill. 35
subg. Urobotrya (StapQ Engl.
41
acuminata Wall. 44
amentacea Roxb. 31-34, 47
map
celtidifolia (Guill.& Perr.)
Endl. ex Walp. 33, 47
cumingiana Baill. 36
fragrans Elmer 47
javanica Miq. 47
manillana Baill. 36
pentitdis Bl. 47
thorelii Gagn. 47
tomentella (Oliv.) Engl. 47
Opiliaceae 2, 9, 29, 31-52
tribe Opilieae 34
Osmoxylon sessiliflorum (Laut.)
Philipson 716
Oxalidaceae 7: 151-178,
829; 10: 607
Pachydiscus Gilg & Schltr 335
Pachygone Miers 160, 165, 167,
168, 170, 172, 217
brachystachya (DC.) Miers
217
hebephylla Miers 217, 218
leplostachya (DC.) Miers
217,218
ovata (Poir.) Hk.f. & Th.
217, 218, 220*
var. dasyphylla Miq. 217
var. rolundifolia Miq.
217
pubescens Benth. 217, 218
zeylanica Sant. & Wagh
218
Pachylamax Dandy 561, 567,
568, 593
praecalva Dandy 593, 594*
Paelae 493
1989]
Index to scientific plant names
737
Palmeria F.v.M. 255, 257-263,
267
acuminata Kaneh. & Halus.
270, 271
angica Kaneh. & Hatus. 267,
271
arfakiana Becc. 262, 267,
270
brassii Philipson 269, 271,
272
clemensae Philipson 268*,
269, 273
dallmannensis Kaneh. &
Hatus. 272
fengeriana Perkins 269, 271
gracilis Perkins 267, 269,
272
habbemensis A.C. Smith 272
hooglandii Philipson 267,
269
hypargyrea Perkins 269, 272
hypochrysea Perkins 275
incana A.C. Smith 269, 272
montana A.C. Smith 269,
272, 275
myrianiha Perkins 270, 271
myrtifoUa Perkins 270
paniculate Ridl. 269
parvifolia Kaneh. & Halus.
276,271
puberula A.C. Smith 270
pulchra Perkins 270, 271
pulteana Perkins 272
scandens 272
schoddei Philipson 269, 275
warburgii Perkins 270, 27 1
womersleyi Philipson 269,
273, 274*
Pandaccae 29
Papaveraceae 5: 114-117
Papuacedrus Li 443
arfakensis (Gibbs) Li 446
papuana (F.v.M.) Li 444
torricellensis (Schllr) Li 445
/'apuz/V/a Ridl. 541,547
laeteviridis P. Royen 548
mmutiflora Ridl. 548
minutiflora auct. 549
Parabacna Micrs 158, 161, 162,
167, 168, 170. 171, 201
amplifolia Diels 193
cincinnans (K. Sch.) Diels
203
dcnudata Diels 189*. 202.
203
. cchinocarpa Diels 189*. 201.
202, 204
var. pubescens Yamamoto
204
(Parabaena)
elmeri Diels 189*, 202
var. philippinensis
(Merr.) Yamamoto 203
hirsuta (Becc.) Diels 193
hirsuta (non Becc.) Diels 202
megalocarpa Merr. 158,
186*, 189*. 202
myriaditha 203
myrianiha K. Sch. 203
philippinensis Merr. 203
sagiitaia Miers 189*, 201,
202
scylophylla Diels 197
tuberculata Becc. 189*, 202,
203
Paracryphiaceae 145
Parakibara Philipson 255, 262,
263, 284 map, 286
clavigera Philipson 286,
287*
Paramanglielia Hu & Cheng
589
aromatica (Dandy) Hu &
Cheng 589
Paramichelia H.H. Hu 565, 567,
593, 599
baillonii (Pierre) Hu 599
scoriechinii (King) Dandy
601
Pararistolochia 53, 57, 59, 61,
62,64
Parasitaxus 343, 351, 354
Paraslemon A. DC. 635-638,
640, 642, 648
grandifructus Prance 649,
650 map
spicalus Ridl. 649
urophyllus (Wall, ex A. DC.)
A. DC. 637, 648*. 649,
650 map
versteeghii Merr. & Perry
637, 649, 650 map
Parinari Aubl. 635, 636, 639-
642, 654
seel. Exitelia (Bl.) C. Mucll.
673
sect. Neocarya DC. 64 1
sect. Pclrocarya 64 1
sect. Sarcoslegia Benth. 671
subg. Euparinari 654
subg. Exitelia Bl. 673
subg. Sarcostegia (Bcnlh.)
Miq. 671
anamensis 656
argcnlco-sericea Kosierm.
655, 656 map
ashtonii Koslcrm. 660
asperula auci. 667
(Parinari)
bicolor Merr. 664
campesu-is 641
canarioides Kosterm. 636,
655, 656 map
coriacea Benth. 641
costata (Korth.) Bl. 636, 656,
663
ssp. costata 663, 664 map
ssp. polyneura (Miq.)
Prance 655, 663, 664
map
ssp. rubiginosa (Ridl.)
Prance 663, 664 map
curaiellifolia641
elmeri Merr. 655, 657 map
excelsa Sabine 639, 641
giganiea Kosterm. 636, 655,
660 map
glaberrima Hassk. 641
glaberrimiun (Hassk.) Hassk.
669
var. lanceolatum (T. & B.)
K. & V. 669
griffithiana 64 1
heteropetala Scort. ex King
675
insularum 639, 656
jackiana Benth. 641, 671
latifrons Kosterm. 666
macrophylla Sabine 641
meiallica Kosterm. 656, 660
map
montana 641
myriandra Merr. 675
nannodes Kosterm. 667
nonda F.v.M. ex Bcnlh. 636,
655, 658 map
nonda auct. 658
oblongifolia Hk.f. 636, 655,
659, 660 map
papuana C.T.Whilc 655, 658
ssp. papuana 658, 659
map
ssp. salomonense
(C.T.White) Prance
658, 659 map
ssp. whilci Prance 658,
659 map
parva Koslcrm. 636, 655,
657 map
polyandra 64 1
prancci Koslcrm. 656, 660,
661 map
rigida Kosierm. 656, 660,
661 map
rubiginosa Ridl. 663
salomonense C.T. White
659
738
Flora Malesiana
[ser. I, vol. 104
(Parinari)
scabra Hassk. 641
senegalensis DC. 641
sumatrana (Jack) Benih.
641, 656, 661 map,
662*
wallichiana R. Br. 665
Parinari auct. 665, 675
Parinarium Juss. 654
sect. Cyclandrophora (Hassk.)
C. Mucll. 665
sect. Neocarya DC. 654
sect. Petrocarya DC. 654
subg. ///Hk.f. 665
subg. Cyclandrophora
(Hassk.) 31. 665
subg. Macrocarya Miq.
665
subg. Petrocarya (DC.) Miq.
654
amboinense T. & B. 669
asperulum Miq. 67 1
borneense Merr. 659
corymbosum (Bl.) Miq. 673
costatum Bl.
var. rubiginosum Ridl.
663
curranii Merr. 669
elaium King 669
excelsum 654
fragile T. & B. 646
grifhihianum Benth. 673
hahlii Warb. 669
helferi Hk.f. 663
heieropelalum Scort. ex King
677
jackianum Benth. 670
kunslleri King 677
lanceolatum T. & B. 669
laiifolium Hend. 666
laurinum A. Gray 669
maingayi King 671
maranthes Bl. 673
margarata A.Gray 669
mindanaeruse Perk. 669
mulliflorum (Korlh.) Miq.
674
myriandrum Merr. 677
nitidum Hk. f. 646
nitidum auct. 674
palauense Kaneh. 674
philippinense Elmer 646
polyneurum Miq. 664
racemosum Vidal 674
salicifolium (Presl) Miq. 674
scabrum Hassk. 669
spicaium King 671
villamilii Merr. 671
warburgii Perk. 669
Parinarium auct. 661, 665
Passiflora aurantia Forst. f. 717
Passifloraceae 7: 405-434,
829; 10: 166, 717
Pedaliaceae 4: 216-221; 5:
557; 7: 829
/'e/ae Adanson 493
Pentaphragmataceae 4: 517-
528
Pentaphylacaceae 5: 121-
124, 566
Pentildis Zipp. ex Bl. 46
Peponaster major Rumph. 94
Pericampylus Miers 160, 169,
170, 172, 227
glaucus (Lamk) Merr. 165,
228*, 229, 236
incanus (Colebr.) Hk.f. &
Th. 229
lanuginosus (Bl.) Miq.
229
membranaceus Miers 229
Periomphale Baill. 335
papuana Steen. 336
Peripetasma polyanthum Ridl.
253
Perrottetia alpestris (Bl.) Loes.
ssp. philippinensis (Vidal)
Ding Hou 716
Petalinia Becc. 12
bancana Becc. 14
Petrocarya Schreb. 641, 654
excelsa Jack 641, 670,
671
glaberrima (Hassk.) Miers
669
scabra (Hassk.) Miers 669
sumatrana Jack 661
Petrocarya auct. 665
Petrosavia 109, 112
Petrosaviaceae 109
Peumus 259, 260-262
boldus 260
Phanerodiscus 4
Phelima Nor. 605
Pherosphaera 343
Pherosphaeraceae 354
Philbomea Hall.f. 607-609,
614
magnifolia (SiapO Hall.f.
614*, 615 map
palawanica Hall.f. 615
Philydraceae 4: 5-7; 7: 829
Phlebocalymna 145
lobospora F.v.M. 149
Phyllocladaceae 354
Phyllocladus L.C. Rich, ex
Mirbel 337-343, 347,
354, 355, 357 map
(Phyllocladus)
hypophyllus Hk.f. 342.
357*, 358*, 359*,
360*
var. protracta Warb. 359
major Pilger 359
protractus (Warb.) Pilger 359
Phyllocosmus Klotzsch 621
Phytocrene 91, 166
/o/jm Merr. 219
malacothrix Sleumer 227
Phytolaccaceae 4: 228-232;
5: 557
Picea 338
Pierotia Bl. 622
lucida Bl. 625
reticulata Bl. 626
Pimela angustifolia Bl. 715
Pinaceae 343, 347, 447-453
Pinus L. 337-340, 342, 347,
447
subg. Diploxylon 449
dammara Lamb. 433, 434
finlaysoniana Bl. 45 1
insular is Endl. 452
kasya Pari. 452
kesiya Royle ex Gordon 341,
346,451, 452 map
khasia Engelmann 452
khasyaWk.LASl
var. insularis (Endl.)
Gaussen 452
khasyana Griff. 452
latteri Mason 451, 452
merkusiana Cooling &
Gaussen 45 1
merkusii Jungh. & de Vriese
340, 341, 346,448*,
449*, 450*. 451 map
var. tonkinensis Chev.
451,452
sumatrana Jungh. 451
sylvestris auct. 45 1
taeda auct. 452
timorensis 452
Piperaceae 127
Piperales 127, 567
Piptocalyx Oliv. ex Bth. 255,
261, 326, 327, 330
macrurus Gilg & Schltr 330,
332
moorei Oliv. 330
Pittosporaceae 5: 345-362;
6: 960; 7: 829; 9: 568;
10: 717
Pittosporum moluccanum
(Lamk) Miq. 717
Platea 145
Plumbaginaceae 4: 107-112
1989]
Index to scientific plant names
739
Podocarpaceae 338, 342, 343,
347, 351-419, 420
Podocarpus rHerit. ex Persoon
337, 340, 343, 347, 351-
355, 395, 397 map
sect. Acuminatus de Laub.
397, 398, 404
sect. Dacrycarpus Endl. 374
sect Dacrydioideae Bennett
374
sect. Dacrydium Bertrand 374
sect. Dammaroides Bennett
390
sect. Foliolatus de Laub.
397, 398, 399
sect. Globulus de Laub. 397,
398, 405
sect Gracilis de Laub. 398,
409
sect. Longifoliolatus de Laub.
398, 407
sect. Macrostachyus de Laub.
398, 412
secL Nageia Endl. 390
sect. Polypodiopsis Bertrand
394
sect. Polystachyus de Laub.
398, 399, 416
secL Prumnopiiys (Philippi)
Bertrand 384
sect. Rumphius de Laub.
398, 399,406. 414
sect Stachycarpus Endl.
384
sect. Sundacarpus Buchholz
& Gray 385
sect. Taxoideae Bennett 384
subg. Foliolatus de Laub.
397
subg. Stachycarpus (Endl.)
Engl. 384
affinis410
agalhifoliaB]. 391
amara BI. 385
archboldii N.E. Gray 399,
402, 403 map
var. crassiramosus N.E.
Gray 409
archboldii (non N.E. Gray)
Gausscn 412
atjchcnsis (Was.schcr) de
Laub. 408 map. 409
beccarii Pari. 393
blumei EnA\. 391
bomccnsis dc Laub. 398,
400. 403 map
bractcatus BI. 341,408,409
map
var. brevipes BI. 408
(Podocarpus)
brassii Pilger 412, 413 map
var. brassii 413
var. humilis de Laub. 339,
413, 414
brevifolius (Stapf) Foxw.
402,412,413 map. 414
brevifolius (non StapO Foxw.
411
celebica (non Hemsl.) Warb.
410
celebicus Hemsl. 349
cincius Pilger 383
compacta Wasscher 383,
445*
confertus de Laub. 340, 341,
407, 408, 409 map
costalis PresI 412, 413 map,
414
costalis (non Presl) Foxw.
410
crassigemmis de Laub. 353,
402, 412. 413 map
cumingii Pari. 381
cupressina sensu Lane-Poole
379
cupressina R. Br. ex Mirbel
377
cupressina Ridl. 379
var. curvula Miq. 380
dacrydiifolia Wasscher 383
decipiens Gray 400
deflcxusRidl. 340, 341.400,
403 map
discolor BI. 400
dulcamara Seem. 387
elaia R.Br. 453
eurhyncha Miq. 385, 387
f ale if or mis Pari. 372
filicifolius Gray 395
gibbsii N.E. Gray 398, 407,
408 map
glaucus Foxw. 409, 410.
411 map
globulus dc Laub. 406 map
idenburgensis N.E. Gray 405
imbricatus BI. 376
var. cumingii (Pari.)
Pilger 381
var. curvula (Miq.)
Wasscher 380
var. kinabaluensis
Wasscher 38 1
imbricatus scnsu Foxw.
379
imbricatus (non BI.) Gibbs
379. 381
insularis dc Laub. 399. 402.
404 map
(Podocarpus)
javanica sensu Merr. 379
javanicus (Burm. f.) Merr. 453
javanicus (non Burm. f.) Merr.
377
junghuhniana Miq. 400
kawaii Hayata 379
koordersii Pilger ex K. & V.
415
latifolia f. ternaiensis de Boer
393
latifolius (non Thunb.) BI.
391
laubenfelsii Tiong 399. 415.
416 map
ledermannii Pilger 405 map
leptophylla Wasscher 371
leptostachya BI. 400
levis de Laub. 398. 400,
403, 404 map
lophatus de Laub. 410, 411
map
lucienii 406
macrocarpus de Laub. 399,
417, 418 map
maximus (de Laub.) Gaussen
394
micropedunculatus de Laub.
339, 341. 405 map
nakaii 406
neglecta BI. 400
neriifolius D.Don 399. 400
map
var. atjehensis Wasscher
409
var. bractcatus (BI.)
Wasscher 408
var. brevifolius Stapf
414
var. brevipes (BI.) Pilger
408
var. polyantha Wasscher
400
var. ridleyi Wasscher 419
var. teysmannii (Miq.)
Wasscher 406
var. timorcnsis Wasscher
401
neriifolius D. Don in
Lamb. 417
neriifolius (non D. Don)
Stecn. 408
novaccalcdoniac 403
palcmffanica Miq. 453
papuanus Rki\. 379
papuiinus (non Ridl.) Pilger
383
papuanus (non Ridl.) Sicup.
380
740
Flora Malesiana
[ser. I, vol. 104
(Podocarpus)
pcdunculala Bailey 387
philippinensis Foxw. 415
piigcri Foxw. 402, 410. 411
map
pilgeri (non Foxw.) v. Roycn
401
polyanlha (Wasschcr)
Gausscn 400
polystachyus R.Br, ex Endl.
339-341, 417*, 418
map
var. rigidus Wasschcr 403
polystachyus (non R.Br.) Li
&Keng414
pscudobractealus dc Laub.
342, 409 map
ridleyi (Wasscher) N.E. Gray
340, 341, 398,417,418
map, 419
ridleyi auct. 405
rotundus de Laub. 410, 411
map
rubcns de Laub. 399, 401*,
402 map
rumphii Bl. 403, 415,416
map
schlechleri Pilger 410
solomoniensis 403
spathoides dc Laub. 398, 400,
404 map
steupii Wasscher 380, 381
map
teysmannii Miq. 406 map
ihevetiifolia B 1.417
vitiensis Seem. 394
wallichianus Presl 391
wangii Chang 410
Podocarpus Labill. 356
Podostemaceae 4: 65-68;
6: 963; 10: 717
Polemoniaceae 4: 195-196
Polyalihia pulchrinervia Boerl.
674
Polycarpicae 165, 261
Polygala L. 455-457, 459
sect. Chamaebuxus DC. 455,
460,461, 465, 467
sect. Melchiora (Stccn.)
Meijdcn 461, 465
sect. Orihopolygala Chodat
469
sect. Polygala 460, 461, 469
sect. Pseudoscmciocardium
Adema 460,461, 462
sect. Semeiocardium [non
(Zoll.) Hassk.] Chodat
462
acicularis Oliv. 465
(Polygala)
arillala Buch.-Ham. 461,
467*. 469
arvensis Willd. 477, 478
var. squarrosa Benth. 476
arvensis (non Willd.) Adema
477
arvensis (non Willd.) Benth.
482
brachislachyos 476
brachystachya BI. 477, 480
brachystachya DC. 476,
477
buchanani Buch.-Ham. ex
D. Don 474
buxiformis Hassk. 477
cardiocarpa Kurz 461 , 463,
464*
cardiocarpa (non Kurz) Ridl.
463
chamaebuxus 460
chinensis (non L.) Benn. 477
chinensis L. 458, 462, 477,
478*, 479
var. brachystachya (Bl.)
Benn. 481
var. linearifolia (non
Willd.) Chodat 481
var. triftora (L.) Benn. 482
ciliata L. 487
densiflora Bl. 478
discolor Buch.-Ham. ex
D. Don 474
elongata WiUd. 477, 480
elongata (non Willd.) Benn.
482
criocephala Benth. 476
eumekes Hassk. 481
exsquarrosa Adema 460, 4o2,
476*
fernandesiana Paiva 475
furcata Roylc 463, 465,
488
glaucocarpa Ridl. 468
glaucoides L. 462, 477, 479,
480, 481*
var. triflora (L.) Trimen
482
glomerata Lour. 478, 479
hondoensis Nakai 472
humilis Span. 481
isocarpa Chodat 463
japonica Houtt. 462, 472*
javana DC. 460, 461, 470*,
471*
karensium Kurz 469
khasyana Hassk. 472
leptalea DC. 474
///iar//b//a Willd. 477, 481
(Polygala)
linarifolia (non Willd.)
Adema 481
linearifolia 48 1
longifolia Poir. 460, 462,
474*
lutea 460
luzoniensis Merr. 472
macrostachya Hassk. 481
malesiana Adema 461, 463,
464*
mariesii Hemsl. 465
monspeliaca (non L.) Blco
482
monticola HBK 460
monticola (non HBK) Ridl.
467
oligophylla DC. 474
orcotrephes Burtt 461, 467*
paenea 457
palustris Lace 463
paniculataL. 458, 460, 461,
475*
papuana (Steen.) Meijden
461, 465, 466*
persicariacfolia DC. 460,
462, 473*
polifolia Presl 462, 476,
477*
polyfolia 476
polygama 458
prostrata Willd. 482
pulchra Hassk. 468
pyramidalis Lev. 474
rhinanthoides Benth. 460,
461, 480*
riukiuensis Ohwi 474
rufa Span. 474
septemnervia Merr. 474
sibirica L. 473
sibirica (non L.) Hassk. 472
simadae Masam. 477
simassan Miq. 468
sumatrana Miq. 461, 467*,
468
tatarinowii Regel 461, 462,
464*, 488
telephoides Willd. 477,
478
telephoides (non Willd.)
W. & A. 476
tinctoria (non Vahl) Hassk.
471
tonkinensis Chodat 469
toxoptera Turcz. 478
iranquebarica Mart. 478
trichocolpa Chodat 469
triflora L. 462, 479, 481,
482*
1989]
Index to scientific plant names
741
(Polygala)
lriner\aia Ham. ex Wall. 487
iriphylla Buch.-Ham. ex
D. Don
var. glaucescens (non
Wall.) Benn. 463
triphylla (non Buch.-Ham. ex
D. Don) Royle 462, 539
umbonaia Craib 463
undulaia Roxb. 487
variabilis (non HBK) Hassk.
475
vaulhieri Chodat 460
venenosa Juss. ex Poir. 461,
467*, 468
ssp. pulchra (Hassk.)
Sieen. 468
ssp. venenosa Sleen. 469
var. eramosa O.K. 468
var. robusta Craib 468
veronicaefolia All
veronicea F.v.M. 472
virgaia458
vulgaris L. 460
vulgaris (non L.) Thunb. 472
wallichiana Wight 474
warburgii Chodat ex Warb.
477
watiersii Hance 465
wightiana W. & A. 460, 462,
479*
Polygalaceae 455-539, 639
tribe Moulabeae 455, 458
tribe Polygaleae 458
tribe Xanihophylleae 458
Polygalales 457, 458
Polyosma 145
Pontederiaceae 4: 255-261;
5: 557; 6: 964
Porotheca K.Sch. 187
peliolaia K.Sch. 187
Portulacaceae 7: 121-123
Primulaceae 6: 173-192,
964
Proteaceae 5: 147-206, 566;
6: 965; 7: 830; 9: 568;
10: 151
Proteales 151
Proicanae 151
Proiium 123
Protoalhcrospcrmoxylon 256
Prumnopitys Philippi 343, 347,
352, 353. 355. 384. 385
map
sect. Sundacarpus (Buchhol/
& Gray) dc Laub. 385
amara (Bl.) de Laub. 385.
386*. 387 map. 388*.
389*
Prunus 640
Pseudaleia imbricata (Roxb.)
Hassk. ex Valeton 9
ongistylis Hassk. ex
Valeton 9
Pleleocarpa Oliv. 29
Pierocarya griffithiana (Benth.)
Miers 674
Pteroneurum DC. 550
decurrens Bl. 554
javanicum Bl. 551
Ptychopeialum 2-4
Punicaceae 4: 226-227
Pycnarrhena Miers ex Hk.f.
& Th. 157, 160-162,
167-171. 172. 173, 178.
305
auslraliana F.v.M. 175
balabacensis Yamamoto
175
balanensis Yamamoto 175
borneensis Diels 1 75
calocarpa (Kurz) Diels 174
casianopsidifolia Yamamoto
177
celebica (Boerl.) Diels 175
elliptica Diels 175
fasciculaia (Miers) Diels
174
^ran^/5 K.Sch. & Uut. 175
insignis (Hatus.) Forman
173, 174
longifolia (Dccne ex Miq.)
Becc. 173, 174
lucida (T. & B.) Miq. 173,
174, 178
manillensis Vidal 173, 175
membranifolia Merr. 175
merrillii Diels 175
moniana Back. 174, 178
nogovuineensis Miq. 173,
175
ozantha Diels 173, 176*,
177, 179
papuana Kanch. & Hatus.
177
plcniflora 173
sayeri Diels 175
lumcfacta Miers 159, 173,
175, 176*
Pyrolaccac 335
Pyrrhosa horsfieldii (Bl.) Hassk.
605
Pyrularia 33
Qucrcus gilva Bl. 701
var. procera Bl. 701
jama-buwa Sicb. 699
Radix puluronica Rumph.
97
Ranunculales 163, 680
RaphanusL. 541, 543,545,
546
caudaius L. 546
raphanistrum L. 547
sativus L. 541, 546
Reinwardlia 609
Restionaceae 5: 416-420,
569
Rhamnales 151
Rhipogonum album R.Br. 253
Rhizophoraceae 5: 429-493;
6: 965; 8: 550; 9: 568;
10: 253, 717
Rhopalopilia 31-33
pallens Pierre 33
Rorippa Scop. 541, 543, 545,
555
backeri (Schulz) Jonsell 541,
555, 556
benghalensis (DC.) Hara 555,
560
dubia (Pers.) Hara 559
heterophylla (Bl.) Williams
555, 559
hybosperma (Schulz)
Jonsell 55, 559
indica (L.) Hicm
var. apeiala (Lour.) Hochr.
559
islandica (Ocd.) Borb. 557
micrantha (Roih) Jonsell 555,
560
nasturtium-aquaticum (L.)
Hayek 555, 556*
officinalis (R.Br.) P. Royen
556
palustris (L.) Becc. 555, 557
pcekelii (Schulz) P. Roycn
555, 557, 558*
schlechteri (Schulz) P. Roycn
559
Rosaceae 638, 639
subfam. Chrysobalanoidcac
639
subfam. Neuradoidcae 639
subfam. Prunoidcae 640
Rosalcs 639
Roucheria 608
coniesliana Pierre 619
griffithiana Planch. 617
Ruiaccac 165
Ryparo.sa kunstlcri King 539
Sabah Mangtieiia Dandy ex
Mcijcr 591
742
Flora Malesiana
[ser. I, vol. 104
Sabia Colebrooke 163, 166,
679-681, 682, 683 map
campanulata Wall. 685
celastrinea Muell. 685
densiflora Miq. 690. 698
elliplica (Miq.) Miq. 684
crralica v. d. Water 683, 684
floribunda Miq. 690, 698,
701
harmandiana Pierre 687
japonica 163
javanica (Bl.) Backer ex Chen
684, 685
var. gtabriuscula (Bl.)
Chen 684
limonacea 685
limoniacca Wall, ex Hk.f. &
Th. 684, 685, 686*, 687*
malabarica Bedd. 685
menescorla 684
menicosla 684
meniscosia Bl. 684
var. eltiptica Miq. 684
\ai.firma Bl. 684
var. glabriuscida Bl. 684
var. latifoUa Bl. 684
papuana Warb. 688
parviflora Wall. 683, 687
ssp. parviflora 685, 687
ssp. philippinensis
(Robins.) v.d. Water
687, 688*
var. harmandiana Lecomte
687
pauciflora Bl. 683, 684, 688
philippinensis Robins. 687,
688
racemosa Chen 684, 689
ssp. kinabaluensis
v.d.Water 683, 689
ssp. racemosa 689
reticulata Elmer 688
sumatrana Bl. 682, 683, 689
Sabiaceae 163, 166,679-715
Salacia banlettii RidJ. 25
Salicaceae 5: 107-110
Salmonea 486
Salomonia Lour. 455-457,
459. 486
sect. Epirixanthes (Bl.) Benn.
488
angulata Griff. 488
aphylla Griff. 490
arnottiana Miq. 488
canarana Miq. 488
cantoniensis Lour. 487
cantoniensis auct. 488
cavalereriei Lev. 488
ciliata (L.) DC. 487
(Salomonia)
cordata Wight 487
cy//uinca(Bl.)Kur/491
edentula DC. 487
elongata (Bl.) Kurz ex Koord.
490
horneri Hassk. 488
longiciliata Kurz 488
mariinii L^v. 488
oblongifolia DC. 487
obovata Wight 487
parasitica Griff. 490
peiiolata D. Don 487
ramosissima Turcz. 488
rigida Hassk. 488
seguinii Lev. 488
sessiliflora 487
sessilifolia D.Don 487
setoso-ciliata Hassk. 488
stricta Sieb. & Zucc. 488
subrotunda Hassk. 487
trinervata Steud. 487
uncinata Hassk. 488
Salomonia auct. 488
Salvadoraceae 4: 224-225
Sampacca O.K. 598
domestica IV alba Rumph.
605
longifolia O.K. 605
monlana O.K. 603
montana Rumph. 582
sylvestris Rumph. 596
velutina O.K. 603
Santalaceae 2, 3, 33-35, 52
Santalales 5. 34
Sapindaceae 639, 679, 680, 681
Sapindales 163,458.639,681
Sapindus microcarpus W. & A.
709
Sapotaceae453, 593
Sarcandra Gardner 123. 125.
126. 128, 129, 134, 144
glabra (Thunb.) Nakai 124,
127, 128, 134, 135*,
136
ssp. brachystachys (Bl.)
Verdcourt 136
var. brachystachys 136
var. melanocarpa (Ridl.)
Verdcourt 136
hainanensis (Pei) Swamy &
Bailey 136
glabra auctt. mult. 136
Sarcodiscus Griff. 287
chloranthiformis Griff. 298
Sarcopetalum F.v.M. 157-161,
167, 169. 172, 224
harveyanum F.v.M. 159,
224. 228*
Sarcosperma 29
Sarcosperma{ta)ceae 4: 32-
34; 6: 967; 10: 29
Sarcostegia Benih. 641
Sanima53. 57, 59,61-64
henryi 62
Saururaceae 4: 47—48
Saxegothaea343,351,352
Saxegothacaceae 354
Saxifragaceae 145
Scaphocalyx parviflora Ridl.
716
spathacea Ridl. 716
Schisandraceae 163, 679
Schoepfia Schreb. 1-6, 27
fragrans Wall. 2, 27, 28*
Schoepfiopsis Micrs 27
fragrans (Wall, in Roxb.)
Miers 27
Sciadocarpus Hassk. 287
brongniartii Hassk. 298
Sciadopitys 354
Sciaphila Bl. 110, 111 map,
112, 720
sect. Hermaphroditantha
subsect. Polyandra 112
sect. Hexanihera 1 13
sect. Oliganthera
subsect. Bilobatae 113
subsect. Quadrilobatae
112, 113
subsect Trilobatae
113
affinis 'Qtcc. 113
andajensis Becc. 117
arfakiana Becc. 109, 112,
113, 115*. 117
aster ias Ridl. 116
atroviolacea Schltr 118
australasicaWQmsX. 117
brachystyla Schltr 1 16
buruensis 112
clemensae Hemsl. 117
conferta J.J. Smith 115
consimilis Bl. 113, 120
corallophyton K.Sch. &
Schltr 113, 119
var. gracilis Giesen 1 19
comiculata Becc. 112, 115,
116, 117
corniculata (non Becc.) Went
117
crinita Becc. 117
decipiens Backer 113
densiflora Schltr 112, 113,
118, 119*
dolichostyla Schlti 119
erubescens (Champ.) Miers
114
1989]
Index to scientific plant names
743
(Sciaphila)
flexuosa Giesen 118
galiensis Schllr 115
hermaphrodila Schltr 1 13
hydrophila Schllr 120
inaequalis Schltr 1 18
inornaia Fetch 116
long i pes Schltr 118
maboroensis Schltr 1 16
macra K.Sch.&Uut. 120
macro K.Sch. & Schltr
116
macra Schltr 120
maculata Micrs 112, 113
major Bccc. 116
micranthera Giesen 112, 121
mindanaensis G'xcscn 120
minuta Schltr 113
moniicola K.Sch. & Schltr
116
multiflora Giesen 112, 113,
120
nana BI. 113, 116, 117
neo-caledonica Schltr 1 15
nu/an.y Giesen 118
oligochaeie Schltr 1 15
papillosa Becc. 121
papuana Becc. 116
pilulifera Schllr 1 16
pumila Giesen 1 14
purpurea 109
quadribuUifera J.J.Sm. 112,
114
reficxa Schlu 118
sccundiflora Thw. ex Bih.
112, 115*. 116, 120
slcmmermannii Fosb. &
Sachet 120
suhhermaphrodita J.J. Sm.
114
sumairana Becc. 1 16
tcnclla Bl. 112. 113, 115*
var. robusta Giesen 1 14
var. voigiii Giesen 1 14
torricellensis K.Sch. & Schllr
114
trichopoda Schltr 1 1 8
tuber culata Giesen 112, 118
valida Giesen 1 18
versteegiana Went 1 16
viliensis A.C. Smith 118
wariana (Schltr)
Mccrcndonk 112, 117
werneri Schltr 1 16
wmklcri Schlu 112. 113.
120
Sclcropyrum 33
aurantiacum (Laut.& K.Sch.)
Pilgcr 52
(Scleropynim)
pentandrum (Dennst.)
Mabberley 52
wallichianum (W. & A.) Am.
52
Scorodocarpus Becc. 1-6, 14
map, 15
bomeensis (Baill.) Becc. 5.
15, 16*
Scyphosiegia Stapf 326
Scyphostegiaceae 5: 297- 299;
6: 967; 7: 830; 10: 326
Sccuridaca L. 455-459, 483
atro-violacea Elmer 483, 484
bracieata Benn. 484
var. papuana F.v.M. 485
complicata auct. 484
corymbosa Turcz. 484
cumingii Hassk. 484
ecristata Kassau 483, 485*
var. niiida Kassau 485
inappendiculata Hassk. 483
ssp. corymbosa (Turcz.)
Meijden 484
ssp. inappendiculata 484
paniculaia Roxb. 484
philippincnsis Chodat 483,
484
scandens Ham. ex Benih.
484
lavoyana Wall, ex Benn. 484
volubilis auct. 484
yaoshannensis Hao 484
Selwynia F.v.M. 218
laurina F.v.M. 219
Semeiocardium Zoll. 539
hamillonii Hassk. 463
Semeiocardium (non Zoll.)
Hassk. 459, 462
Shepherdia Null. 151
Simaroubaceae 6: 193-226,
968; 10: 621, 622, 718
Sinapis 541
alba L. 545
juncea L. 546
imoriana DC. 546
Siparuna 259
gilgiana 261
guyanensis 261
Siparunaccae 261
Siph()n(Kl()n celasuincus Grill.
539
Sisymbrium 541 , 543
amphibujm L.
var. pcduslre L. 557
micranihum Rolh 560
naslurtium-aqualirum L. 555
Skaphium Miq. 493
anceatuin Miq. 527
Sonneratiaceae 4: 280-289,
513; 5: 557; 6: 973
Sparganiaceae 4: 233-234;
10: 718
Sparganium fallax Graebn. 718
simplex Huds.
f. simplex 718
subglobosum Morong 718
Spermabolus T. & B. 605
fruticosa T. & B. 605
Sphaerocarya leprosa Dalz. 22
Sphenocleaceae 4: 27-28
Sphenostemon Baill. 145, 146,
326, 333
sect. Apetalae (Stecn.)
Steen. 147
sect. Sphenostemon 147
scr. Apetalae Stecn. 147
ser. Sphenostemon 146, 147
arfakensis (Gibbs) Steen. &
Erdtman 147, 149
lobosporus (F.v.M.) L.S.
Smith 147, 149
oppositifolius Hiirl. 146
pachycladus 146
papuanus (Laul.) Stecn. &
Erdtman 147, 148*, 149
pauciflorum (A.C. Smith)
Steen. & Erdtman 147,
148
Sphenostemonaceae 145-149,
326, 333
Siachycarpus (Endl.) Ticgh. 352,
384
sect. Sundacarpus (Buchholz
& Gray) Gausscn 385
amara Gausscn 387
Stackhousiaceae 4: 35-36
Staphyleaceae 6: 49-59
Stcganthcra Perkins 255-263,
288, 306, 321
alpina Perkins 319
atepala Perkins 319
austral iana C.T.White 309.
318
brassii (A.C. Smith) Kaneh.
& Hatus. 315
buergersiana Perkins 316
chimbucnsis Philip.son 309,
318
crispula Perkins 316
cyclopcnsis Philipson 308.
310
denlaliJ (Val.) Kanch. &
Hatus. 308, 310
clliptica A.C. Smith 278
fasciculata Philipson 307*.
308. 309
fengcriana F'crkins 315
744
Flora Malesiana
[ser. I, vol. 104
(Steganthcra)
forbesii Perkins 316
hcntyi Philipson 308, 309
hirsuta (Warb.) Perkins 262,
300, 308.309, 314*,
315, 318
hospilans (Becc.) Kaneh. &
Hatus. 257, 258, 308-
310 311, 313*. 319
ilicifolia A.C. Smith 256,
296, 309, 317*, 318
insculpta Perkins 308, 309,
319
insignis Perkins 312
ledcrmannii (Perkins) Kaneh.
& Hatus. 257, 308, 319
moszkowskii (Perkins)
Kaneh. & Hatus. 257,
308, 309, 311
myrtifolia (A.C. Smith)
Philipson 309, 313
oblongiflora Perkins 315,
316
oblongifolia 315
odontophylla Perkins 319
oligantha (Perkins) Kaneh. &
Hatus. 308, 309, 315
oligocarpella Kaneh. &
Hatus. 291
parvifolia (Perkins) Kaneh. &
Hatus. 309, 315
psychotrioides Perkins 319
pycnoneura Perkins 319
riparia Kaneh. & Hatus. 316
royenii Philipson 257, 308,
309, 310
salomonensis (Hemsl.)
Philipson 308, 309, 311,
312*
schlechleri Perkins 278
schumanniana Perkins 3 1 5
suberoso-alata Kosterm. 311
symplocoides Perkins 319
thyrsiflora Perkins 315, 316
lorrlcellensis Perkins 316
villosa Kaneh. & Hatus.
284
warburgii Perkins 315
Stcllatopollisbarghoomii 256
Sicmorwaws fruiescens Bl. 25
membranaceus Bl. 22
Stcphania Lour. 157, 160-162,
165, 167, 169, 171, 243
sect. Eustephania Diels 243,
244
sect. Thamnoihyrsa Diels
243, 244
acuminaiissima (Bl.) Spreng.
252
(Stcphania)
borneensis Yamamoto
249
capitata (Bl.) Spreng. 159,
160, 161, 165, 167,236,
243-245, 252*, 253
calosepala Diels 249, 250
calosepala (non Diels) Merr.
251
cauliflora Becc. 249, 250
cincinnans K. Sch. 203
concinna Miers 247
corymbosa (Bl.) Walp. 243-
245, 249, 250
corymbosa (non Bl.) Turcz.
251
dictyoneura Diels 160, 167,
243, 244, 253
discolor (Bl.) Spreng. 245
var. hernandiifolia (Willd.)
Boerl. 245
exigua Miers 247
florulenta Becc. 251
formanii Kundu & Guha 250,
251
forsteri (DC.) A. Gray 247
glaucescens (Decne) Walp.
245
grandiflora Forman 244, 248
hallieri Diels 247
hernandiifolia (Willd.) Walp.
245, 247
var. discolor (Bl.) Miq.
245
var. genuina (Bl.) Miq.
245
var. glabra (Bl.) Miq. 245
hernandiifolia [non (Willd.)
Walp.] Koord. 249
hernandiifolia [non (Willd.)
Walp.] Ridl. 248
hernandiifolia [non (Willd.)
Walp.] Vidal 251
japonica (Thunb.) Miers 159,
165, 243-245, 245,
246*, 247
var. discolor (Bl.) Forman
247, 248
var. japonica 247
var. timoriensis (DC.)
Forman 247
japonica [non (Thunb.)
Miers] Miers 25 1
longifolia Becc. 252
mcnadonensis Diels ex
Koord.-Schum. 249
merrillii Diels 249, 250
moluccana Forman 244, 245,
250
(Stcphania)
montana Diels 159, 243-
245, 250, 253
ncoguineensis Kundu & Guha
253
obvia Miers 252
prapaiensis Yamamoto 248
psilophylla (Presl) Forman
243-245, 251
ramosii Diels 249
ramuliflora Miers 249
reticulata Forman 244, 245,
249
rotunda (non Lour.) Miq.
248
rotundifolia 248
salomonum Diels 244, 245,
250
tomentosa (Bl.) Spreng.
229
truncata Yamamoto 252
venenosa 248
venosa (Bl.) Spreng. 243-
245, 248
zippeliana Miq. 243-245,
251
Strakaea Presl 65
melastomaefolia Presl 78
Strombosia Bl. 1-6, 19
ceylanica Gardn. 20*, 21, 22,
23 map
var. lucida (T. & B. ex
Valet.) Hochr. 22
var. membranacea (Bl.)
Hochr. 22
var. sessilis Hochr. 22
dubia Vidal 22
elmeri Salvosa 22
javanica Bl. 5, 20*, 21 map
var. sumatrana Val. 21
javanica auct. 22
latifoliaSlap{22,23
lucida T. & B. 22
maingayi (Mast.) Whitm. 22
membranacea (Bl.) Valet. 22
minor Elmer ex Merr. 22
mulliflora King 22
philippinensis (Baill.) Rolfe
21, 22
philippinensis [non (Baill.)
Rolfe] Lam & Holthuis 29
rapaneoides S. Moore 22
rotundifolia King 22
Strombosiopsis 3, 4
Slropha Noronha 129
Stylidiaceae 4: 529-532;
5: 564; 6: 976
Styracaceae 4: 49-56; 6:
976; 9: 568
1989]
Index to scientific plant names
745
Symplocaceae 8: 205-274;
9: 569; 10: 718
Symplocos Jacq.
subg. Hopea Clarke 718
ambangensis Noot. 719
cochinchinensis 719
ssp. laurina 719
cosialifrucia Noot. 719
hcrzogii 718
iliaspaiensis Noot. 719
laeteviridis 719
maliliensis 719
ophircnsis
ssp. cumingiana 719
rayaeNoot. 718, 719
riangensis Noot. 718, 719
vcrtJciliifolia 718
Synandrium exsertum 243
inclusum 243
Taccaceae 7: 806-819
Talauma Juss. 563, 568
sect. Aromadendron Nliq.
569, 576
angatensis (Blco) Vidal 585
aihlianiha Dandy 583
beccarii Ridl. 586
betongensis Craib 585
binluluensis Agostini 577
horneensis Merr. 583
candollei auct. 585
candollii Bl. 569,581,582
var. laiifolia Bl. 582
elegans (Bl.) Miq. 577
var. glauca (Korth.)
Parmcnt. 577
elegans auct. 583
elmeri Merr. ex Soderbcrg
587
forbesii King 582
giganiifolia Miq. 586
giganiifolia auct. 585
gilingensis Elmer 583
var. glabra Dandy 583
var. roiundaia Dandy
583
glaucum (Korth.) Miq. 577
gracilior Dandy 583
grandiflora Merr. 585
injlata Parment. 582
intonsa Dandy 588
javanica Parment. 583
kunstlen King 582
kuleinensis Agostini 586
lanigeraH\^A.&.l\\.58V,
lanigera auct. 586
levissinui Dandy 585
Uliifcra {L.)O.Y.. 582
{Talaunw)
longifolia (Bl.) Ridl. 582
luzoniensis Warb. ex Perkins
585
macrophylla Bl. 583
magna Agostini 586
megalophylla Merr. 586
miqueliana Dandy 582
mutabilis Bl. 582
var. acuminata Bl. 582
var. acuminaiissima
T. & B. 583
var. brevifolia T. & B.
583
var. laiifolia T. & B. 583
var. leiocarpa T. & B.
583
var. longifolia Bl. 582
var. sciagraphia 582
var. splendens Bl. 582
mutabilis auct. 585
oblanceolata Ridl. 585, 586
oblongata Merr. 585
obovaia Korth. 585
obovata auct. 586
oreadum Diels 583
avails Miq. 596
papuana Schltr 596
peninsular is Dandy 583
persuaveolens Dandy 587
plumierii (Schwartz) A. DC.
568
pubescens Merr. 596
pumita Bl. 582
pumila auct. 583
rabiana Craib
var. villosa (Miq.)
Parment. 588
reticulata Merr. 583
rubra Miq. 582
rumphii Bl. 582
sarawakensis Agostini 588
sclerophylla Dandy 585
sebassa Miq. ex Dandy 582
singapurensis Ridl. 586
soembensis Dandy 583
sumulrana Agostini 583
undulatifotia Agostini 583
villariana Rolfc 583, 585
villarii 583
villosa Miq. 588
f. celebica Miq. 603
vrieseana Miq. 569, 596
Tambouri.ssa 257, 258
ficus(Tul.) A.DC. 326
Taxaceae 343, 346, 347-35 1 .
3.54
Taxaics .347
laxodiaccac 338, 343, 354
Taxus L. 337, 342, 347, 348
map
baccata (non L.) Mast. 349
ssp. cuspidata
var. chinensis Pilger
349
ssp. wallichiana (non
Zucc.) Pilger 349
var. sinensis Henry 349
celebica (Wall.) Li 350
chinensis (Pilger) Rehd. 350
cuspidata (non Sieb.& Zucc.)
Kaneh. 350
cuspidata var. chinensis (Pil-
ger) Rehd. & Wilson 350
mairei (Lemee & Lev.) Hu &
Liu 350
speciosa Florin 350
sumatrana (Miq.) de Laub.
349, 350*, 351 map
wallichiana (non Zucc.)
Foxw. 349
wallichiana var. chinensis
(Pilger) Horin 350
yunnanensis Cheng 350
Temstroemiaceae 327
Tetanosia Rich, ex M. Roemer
46
olacioides (W. & A.)
M. Roemer 47
Tctrastylidium 3,4
Tetrasynandra 284 map
Thalamia Spr. 356
Thalictrum 165
Thcaceae 145
Thealcs 145
Thlaspi541,543
Thottea Rottb. 53-55, 57-59,
61-64, 65
beccarii Ding Hou 67, 74
bomeensis Val. 65, 66, 73
celebica Ding Hou 67, 78,
80*
corymbosa (Griff.) Ding Hou
56*, 57, 59, 62, 67, 78,
82*
curvisemcn Ding Hou 61, 66,
73
depcndens (Planch.) Klotzsch
56*, 57, 62, 67, 75
dinghoui 62
grandillora Rottb. 59, 62. 66,
68*, 69
hirsutaR\a\.8\
macTdnlha (Boerl.) Ding Hou
60*, 62. 65, 67. 81
macrophylla Bccc. 5'), 66, 71
muluensis Ding Hou 56*,
57, 67. 76*, 78
746
Flora Malesiana
[ser. I, vol. 104
(Thotlca)
parviflora Ridl. 59, 62, 66,
67, 80*
paucifida Ding Hou 59, 62,
67, 72*, 73, 81
pcniiilobala Ding Hou 57,
67. 76*, 77
philippincnsis Quis. 67, 75,
82*
rcniloba Ding Hou 60*. 67,
77
rhizantha Bccc. 67, 75
robusta Siccn. 66, 70*, 71
siliquosa (Lamk) Ding Hou
81
sp. Ding Hou 83
sU'aalmanii Ding Hou 66, 69
sumau-ana (Mcrr.) Ding Hou
67, 80*. 81
lomcntosa (Bl.) Ding Hou
54, 57, 59, 62, 65, 79
iricornis Maingay 62. 67,
74, 80*
iriscrialis Ding Hou 55, 59,
66, 69
Thuja 444
javanica Burm.f. 453
papuana (F.v.M.) Voss 444
Thujoidcae 444
Thuya javanica 453
Thymelaeaceae 4: 349-365;
6: 1-48,976; 7: 830;
10: 151
Thymclacalcs 151
Tiliacora Colcbr. 157, 160-168.
170, 172, 185
triandra (Colcbr.) Dicls 162,
166, 185. 187, 208*
Tinomiscium Micrs 157-162,
166-170. 204
arfakianum (Bccc.) Dicls
197 •
elasticwn Bccc. 205
javanicum Micrs 205
molle Dicls 205
pctiolare Hk.f. & Th. 159,
205. 206*
phlUppinense Dicls 205
phytocrenoides Kur/. ex
T. & B. 205
pyrrhoboiryum Miq. 205
Tinospora Micrs 19, 158-171,
188
andamanica Dicls 199
angusta 190
arfakiana Bccc. 190, 191,
197
bacnzigeri Forman 159, 160,
190. 191. 195, 196*, 197
(Tinospora)
cclcbica Dicls 190, 193, 194
cordifolia 190, 197
coriacea (Bl.) Bcunndcex
Hcync 199
crispa(L.)Hk.f. & Th. 160,
165, 166, 190, 191, 194,
195, 196*, 197, 200
cmpa [non(L.) Hk.f. &Th.]
Dicls 199
curtisii Ridl. 253
dcniata 190
dissitiflora (Laut. & K.Sch.)
Dicls 159, 190, 191.
192*. 197
glabra (Burm.f.) Mcrr. 159,
165, 186*. 191. 192*,
194, 195. 199. 200
glandulosa Mcrr. 190. 194
hasiala Elmer 193
havilandii Dicls 193,210
hirsuia (Bccc.) Forman 190,
193
homoscpala Dicls 190, 200
macrocarpa Dicls 191, 192*,
198, 199
megalobotrys Laut. &
K.Sch. 187
mcrrilliana Dicls 159, 190,
191, 191
minuliflora K. Sch. & Laut.
227
negrotica Dicls 1 93
var. moniicola Yamamoto
193
peekelii Dicls 197
polygonoides Dicls 200
pseudo-crispa Bocrl. 199
reticulata 199
rumphii Bocrl. 194. 195.
200
sagitlata 190
sinensis (Lour.) Mcrr. 190,
200
smilacina Benth. 191, 196*
subcordata (Miq.) Dicls 159.
191, 192*, 200
sumau-ana (Schcff.) Bccc.
190, 191, 198, 199
var. hanadac Yamamoto
198
icijsmannii Bocrl. 191, 192*.
199
tinosporoidcs (F.v.M.)
Forman 190, 192*
trilobata Dicls 159, 190,
191, 202
tuberculata (Lamk) Beumee
ex Hcync 194
(Tinospora)
uliginosa Micrs 199
uliginosa (non Micrs) Hk.f.
& Th. 198
Trapaceae 4: 43-44; 6: 982
Trcmandraccac 458
Triccrcandra A. Gray 128
Trichocarya Miq. 645, 648
splendens 645
Triclisia 165
Trigoniaceae 4: 58-60; 10:
458
Trimcnia Sccmann 145, 255,
261, 326. 327, 329 map
arfakensis Gibbs 330, 331
bougainvilleensis (Rodcn-
burg) A.C. Smith 332
grandifolia Warb. 333
macrura (Gilg & Schllr)
Philipson 329 map, 330,
331, 332
moorei (Oliv. in Bth.)
Philipson 328, 329 map,
330
myricoides Gilg & Schltr
330, 331
ncocalcdonica 328, 329 map.
330
papuana Ridl. 328*, 329
map, 330, 331*
weinmanniifolia Seem. 328,
329 map, 330-332
ssp. bougainvilleensis
Rodenb. 328, 330, 332
Trimeniaceae 145. 255. 261.
327-333
Trimeria grandifolia (Hochst.)
Warb. 333
Trimeriza Lindl. 65
Trisiichocalyx F.v.M. 217
Triuridaceae 109-121. 110
map, 720
Triuridalcs 109
Triuris 110, 112
Trochiscus Gilli 555
macrocarpus Gilli 559
Tropacolaccac 639
Tsjeru-caniram Rhecde 49
Tsjerucaniram 49
Tsjerucanirum 49
Tsoongiodendron 563. 564. 567,
593
Tsuga 338, 343
mairei Lcmee & L6v. 350
Tuba baccifera Rumph. 213
flava Rumph. 210
Turneraceae 4: 235—238
Typhaceae 4: 242-244; 6:
982
1989]
Index 10 scientific plant names
747
Ulmaceae 8: 31-76, 551;
9: 569; 10: 29
Umbelliferae 4: 113-140,
595; 5; 555; 6: 983:
7: 830: 9: 569
Uroboirya Stapf 31-35, 41
sccL Lepionuroides Hiepko
42 map
sect. Urobotr>'a 42
floresensis Hiepko 42 map,
43,48
laiisquama (Gagn.) Hiepko
32, 43, 42 map
longipes (Gagn.) Hiepko
42 map
parv'iflora Hiepko 41*, 42
map, 43
siamensis Hiepko 34, 42
map, 43, 46
Uvaria-leaved Magnolia Meijer
574
Vaccinium angulaium J.J. Sm.
716
commuialum Mabberley &
Sleumer 716
Valerianaceae 4; 253-254
Vallisneria 112
VanhalUa Schuli. 65
omenlosa J. A. &. J.H.
Schultes 79
V'idara liuorea Rumph. 1 1
Viola rheophila Okamoio 720
Violaceae 7: 179-212,831;
10: 720
Violaria Post & O.K. 568, 569
Viscaceae 353
Vochysiaceae 458
Wcinmannia 123
Weltingionia arnottiana Mcisn.
709
Wilkiea F.v.M. 255-257, 262,
263. 282, 288
forcmanii Philipson 282,
283*
hucgciiana 288
macrophylia 288
WitLslcmia F.v.M. 335
papuana (Stccn.) Siccn. 336*
Worcesierianihus Men. 29
Xanihophyllum Roxb. 455-459.
493
subg. Brunophyllum Mcijdcn
494. 535
(Xanthophyllum)
subg. Coriaceum Meijden
494, 528
subg. Exserium Meijden 494,
533
subg. Triadelphum Meijden
494, 530
subg. Xanthophyllum 494,
500
secL Eystathes (Lour.)
Meijden 500, 507
subsect. Eystathes 507,
526
subsect. Jakkia (Bl.)
Meijden 507, 508
sect. Xanthophyllum 500
acuminaiissimum Miq. 515
adenopodum Miq. 501
adenoius Miq. 497, 498, 511,
515
var. adenotus 516
var. Imeare Meijden 516
affine Korth. ex Miq. 498,
500, 501, 503
B excelsa K. & V. 500
a genuina K. & V. 503
var. adenopodum (Miq.)
K. & V. 501
affine (non Miq.) K.Sch. &;
Hollr. 537
affine (non Miq.) Koord. 5 1 2
affine (non Miq.) Ridl. 528
amoenum Chodat 499, 533
amoenum (non Chodat) Keith
535
ancolanum Miq. 498, 511
f. angustifolia Miq. 5 1 1
angustigcmma Meijden 496,
519
arsaiii C.E.C. Fischer 516
bcccarianum Chodat 495,
522
bomeense Miq. 495. 508
bractealum Chodat 518*.
496, 519
brevipcs Meijden 495. 535,
536*
brigittae Mcijdcn 496, 498,
511, 512*
bullatum King 498. 501
cclcbicum Meijdcn 499, 532
ccraccifohum Mcijdcn 497.
517
chartaccum Mcijdcn 499,
535, 539
ciirifoltum Chodat 530
clovis (Slccn. ex Mcijdcn)
Mcijdcn 496. 517
cockbumii Mcijdcn 494, 502
(Xanihophyllum)
contracuim Meijden 499,
532
cordaium Korth. ex Miq. 515
f . aequale Chodat 5 1 5
curtisii King 514
densiflorum Chodat 521, 522
discolor Chodat 495, 520
ssp. discolor 520
ssp. macranihum Meijden
520
ecarinatum Chodat 499, 539
ellipticum Korth. ex Miq.
499, 530, 531*
er>'lhrosiachyum Gagnep.
497, 511, 523
eiu-hynchum Miq. 497, 524.
525
ssp. eurhynchum 525
ssp. maingayi (Hk. f. ex
A.W. Benn.) Meijden
525
excelsum Bl. 503
var. affine (Miq.) Boerl.
503
excelsum (Bl.) Miq. 500
ferrugineum Meijden 498,
503
flavescens (non Roxb.)
F.-Vill. 514
flavescens Roxb. 498, 500,
5(W, 528
var. virens (Roxb.)
A.W. Benn. 528
flavovirens Elmer 520
flavum Ridl. 505, 507
floriferum Elmer 501
forbesii Baker 523
glabrescens Ridl. 508
glandulosum Merr. 501
glaucescens Miq. 539
glaucum Wall, ex Hassk. 527
gracile Chodat 5 1 3
griffithii Hk.f.ex A.W. Benn.
496, 513, 514
ssp. angustifolium (Ng)
Meijdcn 513
ssp. crecium Meijden 513.
514
var. angustifolium Ng 5 1 3
var. curiisu (King) Ng
513.515
var. montanum Ng 5 1 3,
514
griffiihii (non A.W. Benn.)
Rolfc5I4
hchecarpum Chodat 539
hctcrophyllum Mcijdcn 496,
519
748
Flora Malesiana
[ser. I, vol. 104
(Xanthophyllum)
heteropleurum Chodat 505,
507
hildebrandii Meijden 498,
532
hookerianum King 514
hosei Ridl. 498, 502
hypoleucum Merr. 520
impressum Meijden 496,
513
inccrium (Bl.) Meijden 496,
515
insigne A.W. Benn. 536
kalimantanwn Meijden 539
kingii Chodat 530
korthalsianum Miq. 496,
520
kunslleri King 514
laeve Meijden 499, 524
lanceatum (Miq.) J.J. Sm.
494, 496, 527
lanceolatum 527
latcriflorum Miq. 497, 527
loheri Merr. 501
longifolium (Bl.) Dietr. 514
macranlhum Chodat ex Elmer
520
macrophyllum Baker 498,
507
maingayi Hk. f . ex
A.W. Benn. 524, 525
malayanum Meijden 495,
526
microcarpum Chodat 527
molle Ridl. 522
montanum Meijden 499, 532
moniicolum Meijden 496,
514
muliiramosum Elmer 501
ncglectum Meijden 497, 498,
509
ngii Meijden 496, 526
nigricans Meijden 495, 508
novoguineense Meijden 496,
526
obscurum A.W. Benn. 499,
536
ovatifolium Chodat 498, 508
palawancnse Elmer 497, 516
palembanicum Miq. 5 1 1
palembanicum (non Miq.)
Keith 509
palembanicum (non Miq.)
King 524. 525
(Xanthophyllum)
pallidum Ridl. 501
paniculatum Miq. 514
papuanum Whitm. ex
Meijden 457, 499, 537,
538*
parvifolium Meijden 496,
510
parvum Chodat 5 1 3
pauciflorum Meijden 496,
509
pedicel latum Meijden 495,
522
penibukanense Heine 495,
521
petiolatum Meijden 496, 517
philippinense Chodat 498,
510
pseudoadenotus Meijden 495,
521
pseudostipulaceum Merr. 5 1 3
pseudoslipulaceum (non
Merr.) Meijer 519
puberulum Ridl. 525
pubescens Meijden 495,
504
pulchrum King 495, 521,
522
ssp. pulchrum 521
ssp. siapfii (Chodat)
Meijden 522
purpureum Ridl. 495, 522
ramiflorum Meijden 494,
495, 529*, 530
reflexum Meijden 496, 519
resupinatum Meijden 498,
504
reticulatum Chodat 495,
522
retinerve Meijden 497, 524
robustum Chodat 515
var. elmeri Chodat 5 1 5
rufum Benn. 494, 505,
506*
sarawakense Chodat 503
saraH'akensis 503
schizocarpon Chodat 495,
504
scortechinii King 536
spec. 504
spec. Anderson 530
spec. £■ Ng 539
stapferi 521
5/apyz/ Chodat 521, 522
(Xanthophyllum)
stipitatum A.W. Benn. 497,
499, 535
var. borneense Chodat 535
var. glabrum Meijden 535
var. nitidum Chodat 533
var. pachyphyllum Chodat
533
var. stipitatum 535
subcoriaceum (Chodat)
Meijden 498, 509
suberosum C.T.White 499,
533, 534*
subglobosum Elmer 539
var. longifolium Elmer
539
sulfureum 502
sulphureum King 494, 502
sumatranum Miq. 51 1
tardicrescens Meijden 498,
510
tenue Chodat 498, 508
tenuipetalum Meijden 498,
512
trichocladum Chodat 495,
522
velutinum Chodat 495, 505
venosum King 497, 525
verrucosum Chodat 524,
525
vircns Roxb. 497, 528
virescens 528
vitcllinum (Bl.) Dietr. 497,
498, 514
var. clovis Steen. ex
Meijden 517
wrayi King 497, 525
Ximenia L. 1-6, 10
americana L. 2, 5, 11
var. americana 11, 12*
borneensis Baill. 15
loranlhifolia Span. 1 1
olacioides W. & A. 47
Xylosma leprusipes Clos
715
Xyridaceae 4: 366 — 376,
598; 5: 557; 9: 571
Zanonia indica L. 253
Zizyphus liitorea Teysm. ex
Hassk. 1 1
Zonalapollenites 338
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
c/o Rijksherbarium, P.O. Box 9514
2300 RA Leiden, Netherlands
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9.
Volume 10.
Series I-Spermatophyta (Flowering Plants)
Cyclopaedia of collectors & collections. 1950. pp. clii + 639 (microfiche edition)
General chapters and revisions. 1948-1954. pp. ccix + 631 (microfiche edition)
Bibliography, specific delimitation & revisions. 1955-1958. pp. cccxlii + 596
Systematic revisions. 1960-1972. pp. 20+ 1023 (microfiche edition)
Systematic revisions. 1971-1976. pp. 18 + 876
Cyclopaedia of Collectors, Suppl. 2. Systematic revisions. 1974-1978. pp. 19 + 577
Systematic revisions. 1979-1983. pp. 48 + 600
Systematic revisions. 1984-1989. pp. 44 + 748
Series II - Pteridophyta (Ferns & Fern Allies)
Volume 1. Taxonomical Revision. 1959-1982. pp. (20) + xxiv + 600
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