Tree Flora of Sabah and Sarawak, Volume I - ITTO

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Produced with the financial support of 

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Tree Flora of Sabah and Sarawak Editorial Committee - Chainnan: E. Soepadmo 

(Forest Research Institute Malaysia). Members: Abdul LatifJ Mohamed (Universiti 

Kebangsaan Malaysia), Ruth Kiew (Universiti Pertanian Malaysia), HS. Lee 

(Sarawak Forestry Department), K.M Wong (Sabah Forestry Department). Editorial 

assistant (Volume One): R.CK. Chung (Forest Research Institute Malaysia). 

Advisors: P.s. Ashton (Harvard University, U.S.A.), J Dransfield (Royal Botanic 

Gardens, Kew, u.K.), C Kalkman (Rijksherbarium, Leiden, the Netherlands), T.C 

Whitmore (University of Cambridge, u.K.).

TREE FLORA 

of 

SABAH AND SARA WAK 

Volume One 

edited by 

E. Soepadmo and K.M. Wong 

A joint publication of 

~ 

Sabah Forestry 

Department, Malaysia 

Forest Research 

Institute Malaysia 

1995 

~.A 

T 

Sarawak Forestry 

Department, Malaysia

Copyright © Forest Research Institute Malaysia (FRIM), 

Sabah Forestry Department, and Sarawak Forestry Department 1995. 

Forest Research Institute Malaysia (PRIM), Kepong, 52109 Kuala Lumpur, Malaysia 

. First published 1995 

Perpustakaan Negara Malaysia Cataloguing-in-Publication Data 

Tree flora of Sabah and Sarawak / edited by E. Soepadmo and 

K.M. Wong. 

Includes bibliography and index 

ISBN 983-9592-34-3 (V. 1) 

1. Botany-Sabah. 2. Botany-Sarawak. 3. Botany-Borneo. 

4. Botany-Classification. 5. Plants-Identification. I. Soepadmo, E. 

11. Wong, K.M. (Khoon Meng). 

581.095952 

Printed in Malaysia by Ampang Press Sdn. Bhd., Kuala Lumpur

CONTENTS 

1. Aceraceae (A. Noorsiha) 1 

2. Alangiaceae (A. Berhaman) 5 

3. Anisophylleaceae (KM Wong & L. Madani) 15 

4. Araucariaceae (PC Yii) 27 

5. Bignoniaceae (A. Berhaman) 33 

6. Burseraceae (KM Kochummen) 45 

7. Capparaceae (Deborah Kennard) 101 

8. Celastraceae (KM Kochummen) 107 

9. Chrysobalanaceae (Ghillean T Prance) 155 

10. Clethraceae (A. Berhaman) 181 

11. Connaraceae (Lesmy Tipot) 187 

12. Cornaceae (R.CK Chung) 199 

13. Datiscaceae (EJF Campbell-Gasis) 209 

14. Goodeniaceae (KM Wong) 213 

15. Hypericaceae (KM Wong) 219 

16. Illiciaceae (Richard MK Saunders) 227 

17. Juglandaceae (EJF Campbell-Gasis) 233 

18. Monimiaceae (PC Yii & Lesmy Tipot) 245 

19. Nyssaceae (P.C Yii) 253 

20. Ochnaceae (KM Kochummen) 257 

21. Olacaceae (Lesmy Tipot) 271 

22. Oxalidaceae (R. CX Chung) 287 

23. Pittosporaceae (John B. Sugau) 297 

24. Rhamnaceae (Carsten Schirarend) 305 

25. Rhizophoraceae (L. Madani & KM Wong) 321 

26. Rutaceae (David T Jones) 351 

27. Simaroubaceae (Julius Kulip & KM Wong) 421 

28. Sonneratiaceae (Othman Bojo) 443 

29. Staphyleaceae (J T Pereira) 453 

30. Styracaceae (R. Kiew) 463 

31. Trigoniaceae (KM. Wong) 467 

Abbreviations of Frequently Cited References 471 

Commonly Used Abbreviations for Localities 473 

Glossary 474 

Index to Scientific Names 487 

Index to Vernacular Names 506 

Foreword 

Acknowledgements 

INTRODUCTION 

Page 

Background to the Tree Flora of Sabah and Sarawak Project (E Soepadmo) XIII 

A Brief History of Botanical Collecting and Documentation in Borneo (KM Wong) XXI 

Biogeography and Ecology (PS. Ashton) XLIII 

THE TREE FLORA PROPER 

VII 

IX

FOREWORD 

Providing a foreword to the first volume of any flora is always a rare and pleasurable 

task, but at the same time a conscience-demanding one. In a region like Borneo, 

where the Malaysian states of Sabah and Sarawak are located, and where a complete 

documentation of the flora of anyone sizeable area is not available, a beginning volume of 

a new flora is an exciting endeavour. Yet the work to continue with further volumes must 

press on, and we can only congratulate ourselves within that first limit. 

The plant biodiversity of Sabah and Sarawak is a daunting topic. Estimates by specialists 

throughout the decades vary but are in the region of 10,000 species of vascular plants for 

the whole of Borneo. This is not likely to differ very much from the total for the habitatrich 

and topographically varied territories of Sabah and Sarawak. To engage enough 

specialists to carry out the immense responsibilities of preparing such cl flora is therefore 

conceivably a hugely expensive and time-consuming task. To divide the wqrk into workable 

components is an equally complicated affair: how does one select cetain families of 

importance, or determine which revisions should await more expert attentlOn at some later 

time? 

The Tree Flora of Sabah and Sarawak addresses those plant families that contain 

arborescent species for the mere fact that trees form the main framework of the rain forests 

and are of a fundamental ecological and conservation importance. The basic idea has 

always been to be able to identify these species, so that work in conservation, management 

and sustainable use of species and products have a sound scientific basis, without which no 

systematic applied work can proceed with ease. Our task is a major national effort with the 

Forest Research Institute Malaysia, the Sabah Forestry Department and the Sarawak 

Forestry Department as the key partners. We have the support of the universities and other 

research agencies in Malaysia, as well as key institutions involved with Southeast Asian 

botany within our region, in Australia, Japan, the United Kingdom, Europe and North 

America, without which would have made the work extremely tedious and time-consuming. 

The bulk of the funding for this enormous project comes from the Malaysian Government 

(especially through its funds for the !RP A, or the Intensification of Research by Priority 

Areas programme, disbursed to the participating institutions), the Overseas Development 

Administration (ODA) of the United Kingdom, and the International Tropical Timber 

Organization (ITTO). Such support has been crucial to the initiation of the Flora on an 

organized basis and will be the essential fuel for continuing work. The main testimony to 

the successful production of this first volume of the Flora, and the research now going on 

for further volumes, however, remains the local, regional and international cooperation 

among scientists, research managers and other resource people who have given time and 

importance to this project. During the projected ten-year period of the project, a new 

VII

generation of tropical botanists would have developed; in this, which is of paramount 

significance to the continuing and future undertaking of conservation and forestry 

management, the Tree Flora project would have served an enriching and catalystic role. 

We would like to congratulate Prof. Soepadmo the project leader, and the scientists 

involved in the project for their dedication and commitment, and acknowledge all national 

and international agencies for their support. A ten-year project is a long-term project which 

needs commitment and dedicated support and we hope that both elements will continue to 

be present to take this project to its ultimate successful end. 

Dato' Dr. SaHeh Mohd. Nor 

Director-General 

Forest Research Institute Malaysia 

Haji Awang Tengah bin Haji Awang Amin 

Director 

Sabah Forestry Department, Malaysia 

Datuk Leo Chai Chia Liang 

Director 

Sarawak Forestry Department, Malaysia 

VIII

ACKNOWLEDGEMENTS 

The successful initiation and continuation of the Tree Flora of Sabah and Sarawak 

project owes much to the financial and technical support provided by the Malaysian 

Government, the Overseas Development Administration (ODA) of the United 

Kingdom and the International Tropical Timber Organization (ITTO). In particular, the 

foresight and managerial skills of Dr. Jeff Burley and Dr. Philip Bacon (Oxford Forestry 

Institute), and Dato' Dr. B.C.y' Freezailah and Dr. Efransjah (ITTO) have been essential 

to the good progress made thus far. 

Numerous other institutions and individuals have also contributed invaluable advice, time 

and effort to this undertaking. Foremost in mind are our collaborating institutions in 

Malaysia (the Universiti Kebangsaan Malaysia, Universiti Malaya, Universiti Malaysia 

Sarawak, Universiti Pertanian Malaysia, Malaysian Agriculture Research and Development 

Institute, Sabah Parks Department) and abroad (the Royal Botanic Gardens, Kew, U.K.; the 

Oxford Forestry Institute, U.K.; the Rijksherbarium, Leiden, the Netherlands; the Royal 

Botanic Garden, Edinburgh, u.K.; the Herbarium Bogoriense, Indonesia; the Singapore 

Botanic Garden; the Arnold Arboretum of Harvard University, U.S.A; the Institute of 

Biological Sciences, University of Aarhus, Denmark; the Botanical Garden and Museum in 

Berlin-Dahlem, Germany). We extend our gratitude also to colleagues who have specially 

helped with facilities and research matters, at Kew (its director Prof. G.T. Prance, Prof. 

G.L.Lucas, Dr. J. Dransfield, M.J.E. Coode, Dr. D. Kirkup, Diane Bridson, P. Bygrave, N. 

Martland, AM. Smith), Leiden (its director Prof. P. Baas, Dr. M. Roos, Dr. C. Kalkman, 

Dr. J.F. Veldkamp, Dr. w.J.J.O. de Wilde, Dr. Ding Hou, Dr.M.M. van Balgooy, Dr. P. 

van Welzen, Dr. P. Kessler, Dr. E.F. de Vogel, Stans Kofman, H. Lut), Harvard (Prof. P.S. 

Ashton, Prof. P.F. Stevens), Oxford (Dr. P. Bacon, Dr. D. Filer, Prof. D.J. Mabberley), 

Vienna (Dr. C. Puff) and the C.S.I.R.O., Australia (Dr. M. Dallwitz, Dr. T.G. Hartley). The 

Keepers and Curators of the herbaria at Aarhus (A), the Arnold Arboretum (AA), Berlin 

Dahlem (B), Bogor (BO), British Museum (Natural History) (BM), Oxford (FHO), the 

Philippine National Herbarium (pNH), Sabah Parks, Sandakan (SAN), Singapore (SING), 

Universiti Kebangsaan Malaysia (UKMB), and the Universiti Pertanian Malaysia have 

assisted with loans of specimens, the provision of working facilities and information. 

At the Forest Research Institute Malaysia, we thank Dato' Dr. Salleh Mohd. Nor, Dr. Abd. 

Razak Mohd. Ali, L.C. Cheah, Dr. Roslan Ismail, Dr. H.T. Chan, Dr. N. Manokaran, Dr. 

L.G. Saw, K.M. Kochummen, R.C.K. Chung, Noorsiha Ayop, Lesmy Tipot, Rusea Go, Mat 

Asri Ngah Sanah, Kamamdin M. Saleh, Lucy Sigamoney V. Rajoo, Shahani Saad, 

Khartini Ahmad, and Jamaluddin Osman for assistance in many and different ways with 

the Project. Acknowledgement is also made here of the help provided in Sabah by past and 

present Directors of Forestry, Datuk M. Munang and Tuan Haji Awang Tengah bin Haji 

Amin, and colleagues from the Forest Research Centre, Sandakan (Anuar Mohamed, R.C. 

Ong, Y.F. Lee, A Berhaman, J.T. Pereira, J.B. Sugau, Pung Vui Lee, J. Tangah, L. 

Madani, Dewol Sundaling); and in Sarawak by its Director Datuk Leo Chai and colleagues 

from the Sarawak Forestry Department CH.S. Lee, Abang Mohtar, Runi S. Pungga, P.C. 

IX

TREE FLORA OF SABAH AND SARA W AK 

BACKGROUND TO THE TREE FLORA 

OF SABAH AND SARA WAK PROJECT 

E. Soepadmo 

Forest Research Institute Malaysia, 

Kepong, Malaysia 

Why a Tree Flora of Sabah and Sarawak? 

Borneo, the third largest island in the world, of which Sabah and Sarawak are parts, 

has been frequently acknowledged as one of the most important centres of plant 

diversity in the world. The island,/' which occupies a total land area of approximately 

740,000 sq. km, is conservatively, estimated to harbour 10,000-12,000 species of 

flowering plants, representing abour'5-6% of the world total (Merrill 1950; van Steenis . 

1950; Kiew 1984; Mat-Salleh et al. 1992). Of these, 40-50% are endemic to the island, and 

up to 80% of the endemic species in Borneo occur in Sabah and Sarawak. 

In certain localities in Sabah and Sarawak, where botanical exploration has been carried 

out more intensively, the species diversity is indeed extremely high. Beaman & Beaman 

(1990), for instance, have found that the flora of the Mt Kinabalu Park, Sabah, 

encompassing an area of about 700 sq. km, contains not less than 4,000 species of vascular 

plants in 180 families and 980 genera. 

For a number of economically important families and genera of flowering plants, Borneo 

(Sabah and Sarawak in particular), is also known as the centre of distribution and species 

diversit'j. Fer example, of the 386 known species of the Dipterocarpaceae, 291 species or 

about 75% are recorded from Borneo, of which 257 species or about 66% occur in Sabah 

and Sarawak. Of the 291 species occurring in Borneo, 156 or about 54% are endemic, of 

which 59 species or about 20% are restricted to Sabah and Sarawak (Ashton 1982). In the 

genus Durio (Bombacaceae), of the 30 species which have been described to date, 20 

species or about 66% are recorded from Borneo, with 16 species or about 53% occurring in 

Sabah and Sarawak (Kostermans 1958; 1990). Nine or about 45% of the 20 species known 

from Borneo are endemic, and all occur in Sabah and Sarawak. Similarly, the genus 

Mangifera (Anacardiaceae), comprising about 50 species, also has its centre of distribution 

and species diversity in Borneo, where 20 indigenous species have been recorded (Bompard 

& Kostermans 1992; Kostermans & Bompard 1993; Kochummen, pers. comm. 1995). Of 

these, 18 species are known from Sabah and Sarawak. 

For non-woody components of the flora, e.g., orchids and raffiesias, Borneo and Sabah and 

Sarawak in particular, also house the most number of species in the Malesian 

phytogeographic region. In this region, the orchid family or the Orchidaceae is known by 

XIII

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995) 

3,000--4,000 species, representing 12-16% ofthe entire flora (Chan et al. 1994). Of these, 

Lamb (1991) has estimated that 2,500-3,000 species are found in Borneo, equivalent to 

about 75% of the Malesian orchid flora. A great number of the Bornean species have been 

recorded from Sabah and Sarawak. The genus Rafflf!sia (Rafflesiaceae) with the largest 

flower in the world, also has its centre of species diversity in Borneo where 5-6 of the 14 

species known to date have been recorded, and all occur in Sabah and Sarawak (Meijer 

1984; Mat-Salleh 1991; Nais 1992). 

Other Old World tropical families which have their centres of distribution and species 

diversity in Borneo include the Anacardiaceae, Burseraceae, Celastraceae, Clusiaceae 

(Guttiferae; Calophyl/um), Euphorbiaceae, Fagaceae, Myrtaceae (Syzygium or Eugenia), 

Rhizophoraceae, and several others (Airy Shaw 1975; Ding Hou 1958, 1962, 1978; 

Kochummen 1995; Leenhouts 1956; Merrill & Perry 1939; Soepadmo 1972; Stevens 

1980). 

The presence of high species diversity in the natural forests of Sabah and Sarawak also 

means that there is a wealth of forest products to be harvested. There is no doubt that, in 

the past few decades, the harvesting and utilisation of these forest products, e.g., tropical 

hardwood timbers and rattans, has contributed significantly toward the socio-economic 

development of these two eastern Malaysian states. In recognising that of late, the 

exploitation and conservation ofbiodiversity in Sabah and Sarawak has become the focus of 

international attention and scrutiny, and the need to strike an acceptable balance between 

development and conservation of natural resources in these two states, it is imperative that 

up-to-date botanical inventories should be carried out without further delay. Such basic 

information is of paramount importance to the understanding of the availability, 

distribution, ecological and conservation requirements, and economic potential of the plant 

resources. Without such information, it will be extremely difficult if not impossible to 

develop and manage the available resources on a sustained basis. 

The need for a Flora of Borneo 

The flora of a given region provides an inventory of plant species occurring in that 

region, facilitates a means of species identification, and provides a source of 

information pertaining to up-to-date taxonomic and conservation status, 

distribution, ecological amplitude, and economic potential of the treated species. It is, 

therefore, unfortunate to note that despite the widespread national and international 

recognition of the great economic and conservation value of Bornean rain forests, and that 

botanical exploration and collection in Borneo has begun as early as 1822, and botanical 

accounts of its flora have appeared since 1894 (ef Wong 1995 for more details), to date this 

species-rich island has neither a comprehensive flora of any kind nor even a concise 

checklist of plant species. For this reason Kiew (1984) has highlighted the need and 

urgency of producing a simple "Flora of Borneo" in order to facilitate the implementation 

of sustainable management practices of forest resources of this species-rich island. She 

argued that by using Merrill's (1921) and Masamune's (1942) enumerations as a basis, -and 

by adopting a pragmatic approach, a concise "Identification Flora of Borneo" can be 

readied within 10 years. However, due mainly to the lack of or insufficient institutional and 

financial support, as well as shortage of qualified man-power, the proposed project never 

materialised. 

XIV

BACKGROUND (SOEPADMO) 

The Tree Flora of Sabah and Sarawak Project 

In recognising the urgent need of producing a "Flora of Sabah and Sarawak" of any 

kind, that recent collections (more than 200,000 numbers from Sabah and Sarawak up 

to 1990) necessitate revision and updating of taxonomic accounts in the Flora 

Malesiana itself, and that trained scientific personnel is now available at various national 

institutions, the Tree Flora of Sabah and Sarawak Project was officially launched by the 

Director-General of the Forest Research Insitute Malaysia on the 18th of November 1991. 

The modest lO-year project is executed jointly by the Forest Research Institute Malaysia 

(FRIM), the Sabah Forestry Department (FD Sabah), and the Sarawak Forestry Department 

(FD Sarawak) with the collaboration of other research institutions and universities. The 

main objectives of the project are: 

• To document and update the taxonomic status of all tree-species (taken as reaching at 

least 5 m in height, and 10 cm in diameter) native to Sabah and Sarawak. 

• To publish 8 volumes (each containing 300-400 species) of a concise Tree Flora of 

Sabah and Sarawak within 10 years. 

• To upgrade Malaysian capability and expertise in plant taxonomic research and the 

survey and documentation of tree diversity. 

• To develop and strengthen the management capability of herbaria and their related 

data bases in the three participating institutions (FRIM, FD Sabah and FD Sarawak). 

For the first 5 years, the project is jointly funded by the Malaysian Government, the 

Overseas Development Administration (ODA) of the United Kingdom, and the 

International Tropical Timber Organization (ITTO). 

To prepare the manuscripts of the estimated 109 families comprising about 3,000 treespecies, 

botanists of the following national and international institutions are taking part: 

Malaysia-Forest Research Institute Malaysia, FD Sabah, FD Sarawak, Universiti 

Kebangsaan Malaysia (UKM), Universiti Malaya (UM), Universiti Malaysia Sarawak 

(UNIMAS), Universiti Pertanian Malaysia (UPM), Mt Kinabalu Park, Malaysian 

Agricultural Research and Development Institute (MARDI), and the WWF-Malaysia. In all 

36 botanists are involved. 

Overseas-Institute of Botany, University of Vienna, Austria; University of Brunei 

Darussalam, Brunei; University of Aarhus, Denmark; Botanischer Garten und Botanisches 

Museum, Berlin, Germany; University of Hong Kong, Hong Kong; Rijksherbarium Leiden, 

the Netherlands; Singapore Botanic Garden, Singapore; Oxford Forestry Institute, Oxford, 

United Kingdom; Royal Botanical Gardens, Kew, United Kingdom; Arnold Arboretum, 

Harvard University, USA; University of Florida, USA; and Everglades National Park, 

Florida, USA. A total of 21 botanists are involved. 

XV


Coverage. Being an identification type of flora intended to be "user-friendly" to nonspecialist 

readers, the Tree Flora will be written in simple and easy-to-understand English. 

The use of highly technical botanical terms is, therefore, to be avoided. 

In the Tree Flora, all dicotyledonous trees, here defined as woody plants with the main 

upright stems measuring not less than 5 m tall and 10 cm diameter, will be treated in full. 

Non-tree and introduced tree species known only in cultivation will be given cursory 

treatment and annotated in the keys only. For each tree-taxon (family, genus, species), 

treatment will be confined to the following aspects--correct/accepted scientific name, 

major references, description of diagnostic characters, vernacular names (if applicable), 

distribution, ecology, uses, notes on taxonomy (if applicable), and key(s) to lower rank taxa. 

For each genus, at least one line-drawing depicting important vegetative and reproductive 

characters will be provided. 

Format. To standardise the format of manuscripts to be published in the Tree Flora, a set 

of guidelines has been prepared by the Editors (c! Soepadmo & Wong 1995). This 

guideline is obtainable from the project Secretariat at FRIM. Among the main guidelines 

which should be adhered to by contributors of manuscripts are the following: 

• Nomenclature-Names of families and genera will normally follow Brummitt (1992), 

but in cases of potential confusion and controversy, authors of revisions for the Tree 

Flora are requested to confirm the accepted names with the Chief Editor. 

• References--Only selected references most relevant to the taxonomy and distribution 

of the family, genus, and species and which are relevant to Borneo in general and 

Sabah and Sarawak in particular will be cited. Unless listed in the "Guidelines", book 

names should be cited in full. Names of journals and other serials are to be cited 

following van Steenis-Kruseman (1956). 

• Derivation of genus and species names--Language and meaning of root words to be 

indicated. 

• Basionym and synonym-Basionym to be given if it indicates a new taxonomic 

perspective of the taxon being treated. Only synonyms relevant to Borneo are to be 

included. 

• Typification-Whenever possible, citation of type specimen(s) of accepted species 

should be provided. 

• Description-Required for any family with more than one genus worldwide, for 

genera, and for species, and should account for the following aspects: habit; leaf 

arrangement and type; stipules; inflorescence type and position; flower sexuality, 

symmetry, merism, details of fusion of parts; fruit type; seed, embryo, cotyledons, 

endosperm, aril and others if very relevant. 

• Vernacular names--Only names commonly used in Sabah and Sarawak will be 

included. The dialect or language of each entry of vernacular name should be 

indicated. 

XVI


• Distribution-Number of genera and/or species is to be mentioned, followed by global 

and Malesian distribution, and followed by that in Sabah and Sarawak. If the taxon 

also occurs in Brunei and/or Kalimantan, it should be indicated following the 

distribution in Sabah and Sarawak. 

• Ecology-For a family or a genus to be given only if general trends can be 

summarised. For a species, information on forest type, soil/rock type, and altitudinal 

range is required. 

• Uses--For a family or a genus to be provided only if there are general or interesting 

uses. For a species as much information as possible should be given. 

• Taxonomy-To be provided only if there are taxonomic controversies regarding the 

taxon. Brief commentary could be given if this will clarify the taxonomic status of the 

taxon under consideration. 

• Key(s) to lower rank taxa-Bracket key structure is to be used. Couplets to be 

numbered once only, the individual leads in the couplet not further numbered. In 

constructing the keys, only diagnostic vegetative and/or reproductive characters are to 

be used. Non-tree taxa are to be annotated only. 

• Infraspecific taxa-In cases where two or more infraspecific taxa exist in Sabah and 

Sarawak, a full description of the species, including all variations known in Sabah and 

Sarawak, should be given in the species description. This should be followed 

immediately by a key to the taxa in which each infraspecific taxon is annotated. 

In cases where only one of the infraspecific taxa is known to occur in Sabah and 

Sarawak, the species description should follow the following format: species name and 

authority, derivation of species name, reference for the species; infraspecific taxon 

name and authority, reference, basionym, type specimen, synonyms, description of the 

sole infraspecific taxon which represents the species in Sabah and Sarawak, vernacular 

name(s), distribution, ecology, uses, and taxonomy. 

Other project activities 

Apart from preparing and publishing eight volumes of the Tree Flora of Sabah and 

Sarawak, the project also carries out a number of activities relevant to its overall 

objectives. These activities include: 

• Organisation of collecting expeditions to a number of botanically little-known localities 

in Sabah and Sarawak. 

• Establishment of a data base for specimens and taxonomic references using BRAHMS 

and related softwares. 

• Conducting workshops and specialised training on flora writing, documentation 

software, botanical illustration, editing, curation and management of herbarium 

specimens, etc. 

XVII


• Postgraduate training to upgrade Malaysian capability in plant taxonomic research and 

the inventory and documentation of plant/tree diversity. 

• Collaboration with other institutions and agencies with similar aims in the Malesian 

region. 

References 

Airy Shaw, H.K 1975. The Euphorbiaceae of Borneo. Kew Bulletin Additional Series IV. 

245p. 

Ashton, P.S. 1982. Dipterocarpaceae. Flora Malesiana 1, 9(2): 237-552. 

Beaman, lH. & RS. Beaman. 1990. Diversity and distribution patterns in the flora of 

Mount Kinabalu. In: P. Baas, K. Kalkman & R Geesink (eds.), The Plant Diversity of 

Malesia, pp. 147-160. Kluwer Academic Publisher, Dordrecht, the Netherlands. 

Bompard, lM. & AlG.H. Kostermans. 1992. The genus Mangifera in Borneo: Results of 

a IUCN-WWF/IBPGR Project. In: Ghazally Ismail, Mustedza Mohamed & Siraj Omar 

(eds.), Forest Biology and Conservation in Borneo, pp. 61-7l. Centre for Borneo Studies, 

Publication No. 2. 

Chan, c.L., A Lamb, P.S. Shim & II Wood. 1994. Orchids of Borneo. Vol. l. 

Introduction and a Selection of Species. Sabah Society & Royal Botanic Gardens, Kew. 

xviii + 402 p. 

Ding Hou. 1958. Rhizophoraceae. Flora Malesiana 1, 5(4): 429-493. 

Ding Hou. 1962. Celastraceae I. Flora Malesiana 1, 6(2): 277-291. 

Ding Hou. 1978. Anacardiaceae. Flora Malesiana 1, 8(3): 395-548. 

Kiew, R. 1984. Towards a Flora of Borneo. In: Ismail Sahid, Zainal Abidin A Hasan, A 

Latiff Mohamed & A Salam Babji (eds.), Research Priorities in Malaysian Biology, pp. 

73-80. Penerbit Universiti Kebangsaan Malaysian, Bangi, Malaysia. 

Kochummen, KM. 1995. Burseraceae. In: E. Soepadmo & KM. Wong (eds.), Tree Flora 

ofSabah and Sarawak, Vol. 1: 45-106. Celastraceae, ibid.:107-154. 

Kostermans, AlG.H. 1958. The genus Durio Adans. (Bombacaceae). Reinwardtia 4(3): 

47-153. 

Kostermans, AlG.H. 1990. Durio bukitrayaensis Kosterm. (Bombacaceae), a new species 

from Borneo. Botanica Helvetica 100/1: 29-3l. 

Kostermans, A.J.G.H. & lM. Bompard. 1993. The Mangoes. IBPGRlLinnean Society 

London, Academic Press; xvi + 233. 

Lamb, A 1991. Orchids of Sabah and Sarawak. In: R Kiew (ed.), The State of Nature 

Conservation in Malaysia, pp. 78-88. Malayan Nature Society & IDRC, Canada. 

XVIII


Leenhouts, P.W. 1956. Burseraceae. Flora Malesiana 1, 5 (2): 209-296. 

Masamune, G. 1942. Enumeratio Phanerogamarum Bornearum. 739 p. 

Mat-Salleh, K 1991. Raffiesia-Magnificent Flower of Sabah, 49 p. Borneo Publishing 

Company, Kota Kinabalu, Sabah. 

Mat-Salleh, K, J.H. Beaman & H. Beaman. 1992. Specimen database and their utilization 

for the Flora of Borneo. In: Ghazally Ismail, Murtedza Mohamed & Siraj Omar (eds.), 

Forest biology and Conservation in Borneo, pp. 117-137. Center for Borneo Studies, Publ. 

No. 2. 

Meijer, W. 1984. New species of Raffiesia (Rafflesiaceae). Blumea 30: 209-215. 

Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. J. Str. Br. Roy. As. 

Soc., Special Number. 637 p. 

Merrill, E.D. 1950. A brief survey of the present status of Bornean botany. Webbia 7: 309- 

324. 

Merrill, E.D. & L.M. Perry. 1939. The Myrtaceous genus Syzygium Gaertn. in Borneo. 

Mem. Am. Acad. Arts & Sci. 18(3): 135-202. 

Nais, J. 1992. Distribution, dispersal and some notes on Raffiesia around Kinabalu, 

Malaysia. In: Ghazally Ismail, Murtedza Mohamed & Siraj Omar (eds.), Forest Biology 

and Conservation in Borneo, pp. 97-108. Center for Borneo Studies, Publ. No. 2. 

Soepadmo, E. 1972. Fagaceae. Flora Malesiana 1, 7(2): 165-403. 

Soepadmo, E. & KM. Wong. 1995. Guide to Preparing and Editing Manuscripts. 31p. 

Forest Research Institute Malaysia, FD Sabah and FD Sarawak, Malaysia. 

Steenis, C.G.G.J. van. 1950. The delimitation of Malaysia and its main geographical 

division. Flora Malesiana 1, 1: LXX -LXXY. 

Steenis-Kruseman, M.J. van. 1956. Citation of serial and some books. Flora Malesiana 1, 

5(2): CXL V-CLXY. 

Stevens, P.F. 1980. A revision of the Old World species of Calophyllum (Guttiferae). J. 

Am. Arb. 61 (2 & 3): 117-699. 

Wong, KM. 1995. A brief history of botanical collecting and documentation in Borneo. In: 

E. Soepadmo & KM. Wong (eds.), Tree Flora of Sabah and Sarawak, 1: XXI-XLI. 

XIX


A BRIEF HISTORY OF BOTANICAL 

COLLECTING AND DOCUMENTATION 

IN BORNEO 

K.M. Wong 

Forest Research Centre, 

Sabah Forestry Department, 

Sandakan, Malaysia 

M 

Uch of the information on early botanical collections in Borneo is summarised, by 

collectors' names, in the great Cyclopaedia of Collectors for the Flora Malesiana 

(van Steenis-Kruseman 1950, 1958, 1974). The present account attempts to collate 

this information, together with some information on collectors beyond van Steenis 

Kruseman (1974) so that the progress of plant exploration in Borneo may be followed 

through each period and in as chronological an order as possible. Collectors of lesser 

significance, in that they have only gathered very few numbers, are not normally listed 

here. We give importance here to collections made up to only around the beginning of the 

1980s, as these are often less immediately known to unfamiliar botanists, because they are 

not held in herbaria in the region (through historical factors) or because they have rarely 

been referred to in the literature. Many collectors, though not all and not in every instance, 

since the 1950s, have distributed good sets of their collections to regional herbaria serving 

areas where the collections have been taken, and in general there is a better chance that the 

more recent collections made (especially as the 1970s and 1980s are approached) may now 

be found deposited in local herbaria and are easily accessible for study locally. The period 

beyond this is not covered in the present account as it can be expected that such modern 

collections will be easily found for reference in either the local herbaria (Bogor, Kepong, 

Kuala Lumpur, Sandakan, Sarawak, Singapore) or in the large botanical institutions abroad 

that have a long-standing association with the study of Bornean plants. 

Early collectors and documentation 

The earliest significant efforts at plant collecting in Borneo were those of the early to 

middle 19th century, and were motivated initially either by an interest to study or 

collect groups of horticultural interest, or by the interest of specialists in Europe, 

Java (where the Dutch administration of the Indonesian islands was based) and Malaya and 

Singapore (the seat of English administration in the region then) to document interesting 

and rare life forms of a little explored, faraway tropical land that was virtually terra 

incognito as far as scientific documentation is concerned. 

As early as 1822, George Muller, the Acting Resident in Dutch West Borneo, made 

explorations and plant collections in the Kapuas valley and Pontianak, and around Kutei 

XXI

TREE FLORA OF SABAH AND SARAW AK VOL. 1 (1995) 

and the Mahakam River until 1826, when he is thought to have been murdered near the 

upper Kapuas. Muller collected for Carl Ludwig Blume, Director of the Buitenzorg (Bogor) 

Botanic Garden around this time. 

In 1836, Dutch botanist Pieter Willem Korthals, a member of the Commission for Natural 

Sciences for the Dutch East Indies who had explored parts of Java and Sumatra, collected 

in southeast Borneo, mainly in the Banjarmasin, Barito River and Mt Pamatton areas, 

together with Salomon Muller (a German member ofthe Commission) and Ludwig Horner 

(Swiss member of the Commission, who apparently did not collect plants on this 

expedition). Merrill (1921) and van Steenis-Kruseman (1950) point out it is likely that 

some of Korthals's Borneo records are probably of Sumatran or Javanese origin, owing to 

confusion in labelling after the collecting. Korthals's collections are principally at the 

Leiden herbarium. In 1842, the "United States Exploring Expedition" under the command 

of Charles Wilkes, which began its journey in 1838 and had travelled through the 

Philippines, visited the Banggi and Balabac Islands off the northernmost tip of Borneo, 

during which expedition botanists William Rich and lD. Brackenridge, with the help of 

zoologist C. Pickering, made some plant collections (now principally with the U.S. 

National Herbarium and the Gray Herbarium at Harvard). Between 1852 and 1855, 

Cornelis de Groot, Chief of the Mining Department of the Dutch East Indies collected in 

southeast and east Borneo, including the Banjarmasin, Tanjung Batu, Mahakam and 

Samarinda areas. 

Among the more significant early collectors of Bornean plants must count also the Rajah 

James Brooke, who had helped the Sultan of Brunei in suppressing a revolt in 1840 and 

was consequently given Sarawak to rule in the'following year. He.was also made Governor 

of Labuan and Consul General of Borneo in 1847, and collected some plant specimens 

which the Kew herbarium acquired in 1853-1855. Hugh Low, appointed by the Rajah 

Brooke first as his secretary in 1845 and later as Colonial Treasurer of Labuan between 

1848 and 1877, and who made the first documented climb by a European to the summit of 

Kinabalu in 1851, was well known even at the time for his interest in orchids. Low's 

successful third trip to Kinabalu in 1858, together with Spenser St. John, secretary to the 

Rajah Brooke, included the first collections from Marai Parai, the ultramafic western spur 

of the mountain. Low had also collected in the Kuching area, Mt Penrissen and Lawas. 

Low sent most of his orchid specimens to John Lindley at the London University and 

Joseph Hooker at Kew. 

Low's plant-collecting contemporaries included James Motley, a civil engineer at the 

Labuan coal mine during 1851-1854 and who was later (1854-1859) at Banjarmasin in 

south Borneo (Motley's Labuan collections were often sent to Kew by E. Barber Scott 

labelled "colI. Barber, Labuan", although his Kalimantan collections were distributed under 

his own name), and Thomas Lobb, who made an unsuccessful attempt to reach the summit 

of Kinabalu in 1856. Lobb, who was collecting plants of horticultural value for the firm of 

Messrs Veitch in England, also collected some herbarium specimens from Sarawak and 

Labuan. Lobb is thought to have intentionally falsified localities on some labels in order to 

increase the value of his collections for the firm of Veitch; a specimen labelled "Luzon" in 

one herbarium may be labelled "Borneo" or "Singapore" or "Java" in another (van Steenis 

Kruseman 1950). The well-known zoogeographer Alfred Russel Wall ace visited and 

XXII

BRIEF HISTORY (WONG) 

collected mainly ferns in Sarawak (Sarawak, Simunjon and Sadong Rivers) during 1854- 

1856, at the start of his travels through the Malay Archipelago. 

Willem Hendrik de Vriese, a botanist at Leiden at some time during this period, collected 

in Dutch West Borneo in 1860. Eduard von Martens, Custodian of the Berlin Zoological 

Museum, also made plant collections in Dutch West Borneo in 1863. Between 1865 and 

1867, Odoardo Beccari, the well known Italian naturalist, made many plant and animal 

collections in Sarawak; many of his collections were made from around Kuching and 

Matang, and the Sarawak River and Batang Lupar. He also collected from the Bintulu area 

and the Rejang valley, and was also briefly in Labuan, Brunei and the then Dutch West 

Borneo. His plant collections number around 20,000; he wrote many papers, especially on 

palms. Perhaps his most well known book is Nelle Foreste di Borneo (Wanderings in the 

Great Forests of Borneo) (Beccari 1902). Burtt (1964) points out that although Beccari 

ascended the Poi range, he did not climb Mt Poi (Poe or Pueh) as that name is used on 

modern maps, but a more south-easterly peak called Mt Berumput. Somewhat less 

illustrious than Beccari but nevertheless a collector who during 1866-1869 sent several 

cases of important plant specimens from Sarawak and Sambas in Dutch West Borneo to the 

herbarium at Buitenzorg was a man documented only as a coffee planter in Java and 

manager of an estate in Sarawak, called Martin. 

In 1874-1875, the well-known botanical explorer Johannes Elias Teijsmann (also Curator 

of the Buitenzorg Botanic Gardens in 1831-1869) explored Dutch West Borneo, making 

numerous collections in the Pontianak, Kapuas River, Kenepei River, Penein Mts, Biang 

Mts and other areas. His Borneo collections are mainly in the Bogor, Leiden, Kew and 

Florence herbaria. In 1881 ~ 1884 Friederich Grabowsky, a German zoologist, collected a 

few hundred plant specimens from southeast Borneo, mainly around the Banjarmasin and 

Kapuas River areas. His collections are principally at the Berlin and British Museum 

herbaria. During 1879-1880, the Danish explorer, ethnographer and zoological collector 

Carl Bock made an expedition to northeast and southeast Borneo, collecting also some 

plants along the Mahakam and Telen and in the Samarinda and Banjarmasin areas. 

Another important plant collector about this time was Frederick William Burbidge, a 

collector for the Veitch nurseries in England, who collected in Labuan and Kinabalu with 

Peter Veitch in 1877 and in the Sandakan area with William Pryer, the founder of 

Sandakan, in 1878. He also collected along the Lawas, Meropok, Limbang and Pandaruan 

rivers in the northern part of Sarawak. Burbidge's botanical plates and sketches from 

Borneo are housed in the British Museum of Natural History. In 1880, two other Veitch 

collectors, David Burke and Charles Curtis collected in Sarawak; Curtis later (1884) 

became the Superintendent of Gardens and Forests in Penang. 

The Hose family, comprising of the Reverend George Frederick Hose (Bishop of Singapore, 

Labuan and Sarawak, 1881-1908), his nephews Charles Hose (district officer in the 

Baram in 1884 and afterwards, Sibu) and Ernest Hose (a planter), and his daughter 

Gertrude Hose all collected plants in Sarawak. The Rev. Hose collected on Matang and 

(with Charles and Alfred Hart Everett, freshly retired Resident of Trusan and Hon'ble of 

the 4th Division) on Mt Dulit, and other places, taking especially grasses, sedges and ferns, 

XXIII


as his daughter did. Charles Hose made zoological studies as well as collected plants, in the 

Baram, Mt Dulit, Mt Mulud, Kayang, Rejang and Niah areas. Their collections were sent 

mainly to the Kew and Edinburgh herbaria. 

Between 1883 and 1885 when he was murdered, M. Fraser, Medical Officer of the Chartered 

Company in North Borneo, collected several hundred plant specimens from Kudat, the 

Marudu Bay, Balambangan and Banggi Islands, Gaya Island and Papar. These specimens 

are at Kew. Around 1885 and 1890, a Singapore schoolmaster, R. Hullett visited Sarawak 

and collected several hundred specimens, now mostly in the Singapore and Kew herbaria. 

In 1886, Ignatz F. Forstermann, in the employ of the English horticultural firm of Sander 

& Sons, collected ferns and orchids in Sarawak. John Whitehead, an ornithologist, collected a 

small number of plants during his famous expedition to Kinabalu in 1887-1888. 

In 1892, George D. Haviland, Curator of the Sarawak Museum at Kuching, visited Kinabalu 

(with his brother H.A. Haviland) and collected about 450 numbers on its upper slopes. Both 

Burbidge's and Haviland's collections were studied by OUo Stapf of the Royal Botanic 

Gardens, Kew, who wrote the first monograph on the Kinabalu flora (Stapf 1894), 

enumerating 360 species of flowering plants, ferns and bryophytes. Haviland also collected 

at Kuching, Mt Braang, Mt Penrissen, Limbang, Sibu, Ulu Tawaran, and (with Charles 

Hose) Mt Lambia. van Steenis-Kruseman (1954) has compiled a list of Haviland's 

collections in Borneo as then known. Edward Bartlett, a zoologist who also collected a 

small number of plant specimens in Sarawak around 1893, later succeeded Haviland in 

1895. Robert W.e. Shelford, who in 1897 was the Curator of the Sarawak Museum, 

collected Sarawak plants between 1897 andJ903, mainly in the Kuching and Trusan areas. 

Charles van de Leur Creagh, the Governor of North Borneo during 1888-1895, collected 

at least 1839 numbers on Kinabalu and also made collections from Pulau Gaya, Kimanis 

and Labuan on the west coast to Cowie Harbour, Tawau and Sandakan on the east coast 

(Meijer 1969). Johannes Waterstradt, Danish zoological collector, collected mainly 

orchids as well as insect specimens on his trips in Borneo. He went to the Padas River 

(North Borneo) in 1891, the mouth of the Lawas River and Kinabalu (1892-1893), Kudat, 

Banggi, Balambangan, Labuan , Kinabalu (N Borneo) and the Lawas and Limbang Rivers 

(Sarawak) (894), Kinabalu (1895), Brunei (1902), and again Kinabalu (1903, 1908 and 

1912). Much of his orchid collections before his return to Europe in 1904 went to the firm 

of Hugh Low in London. 

This was the time when the great Dutch explorer and ethnologist, Anton Willem 

Nieuwenhuis, made his expeditions into Dutch Borneo, exploring with German botanist 

Johann Gottfried Hallier (then an assistant at the Buitenzorg herbarium; he also used the 

initial 'H' for 'Hans') and Dutch geologist Gustaaf Frederik Molengraaff in 1893-1894 

the Mandai, Kenepai and Mendalam Rivers, the Kapuas delta and the Pontianak area; with 

Indonesian plant collector Jaheri (of the Buitenzorg Botanical Garden) in 1896-1897 the 

Mendalam and Mahakam areas; with Indonesian collectors Amdjah and Sakaran (of the 

Buitenzorg Botanical Garden) in 1898-1899 the Kapuas, Mahakam and Samarinda areas; 

and in 1899-1900 the Apo Kayan highlands. These collections (several thousand numbers) 

are principally at the Bogor and Leiden herbaria. Amdjah collected again in northeast 

Borneo in 1912. 

XXIV


Early 20th Century collectors 

H.N. Ridley, Director of Gardens and Forests, Straits Settlements during 1888- 

1900, and Director of the Singapore Botanic Garden during 1901-1912, visited 

and collected in Bau, Matang and Lundu in Sarawak in 1893. In 1897 he collected 

in Labuan, Kudat, Sandakan and the Labuk Bay area, and in Sarawak again. His 

subsequent visits to Sarawak were in 1903, 1905 and 1915. Ridley's collections are 

principally at the Kew and Singapore herbaria. John Hewitt, who became Curator of the 

Sarawak Museum during 1905-1908, collected plants mainly in the Kuching, Saribas, 

Baram and Limbang areas. 

The German orchid specialist Friedrich RR Schlechter collected in Labuan, Kudat, 

Sandakan in British North Borneo, and Banjarmasin, Balikpapan, Samarinda and the Kutei 

region in Dutch Borneo in 1901, and later in Kuching in Sarawak during 1906-1907. H. 

Witkamp, a mining engineer with the Royal Batavian Oil Company, made plant collections in 

Kutei district in east Borneo in 1905, 1907, 1910 and 1928. 

Around this time, Cecil 1. Brooks, a chemist with the Borneo Company which exploited 

gold in Sarawak, collected plants, mainly ferns, along the Sarawak River and on Mts 

Santubong and Poe, and around Bau and the Bungo Range, between 1907 and 1910. 

In 1908 the German botanist Hubert Winkler collected more than a·thousand numbers 

during his trip to east and southeast Borneo, covering the Banjarmasin, Barito River, 

Balikpapan, Samarinda and Mahakam areas. His collections are principally at the herbaria 

at Breslau, Berlin, Geneva and Manila (the last destroyed in the Second World War). 

During 1910-1914, the Dutch geologist Louis Martin Robert Rutten collected plants in 

east and northeast Borneo in the Balikpapan and Samarinda areas, along the Kayan and 

Rapah Rivers, and in the Sangkulirang Bay area. 

Lilian Suzette Gibbs, an English botanist interested in tropical alpine floras, made 

collections in the Beaufort area and the Tambunan plain, and also collected on Mount 

Kinabalu in 1910 (she collected on the Gurulau and Marai Parai spurs and ascended to the 

summit via Kamborangoh). Gibbs collected about 1000 numbers from North Borneo. Her 

paper on Kinabalu (Gibbs 1914) was the second monograph on the Kinabalu flora after 

Stapf's. The American botanist Frederick William Foxworthy, serving at the Bureau of 

Science in Manila (1906-1911), collected in the Lundu, Niah, Mt Poe, Mt Santubong and 

Kuching areas in Sarawak in 1908 and also climbed Kinabalu in 1910. John Coney 

Moulton, Curator of the Sarawak Museum during 1905-1915, collected widely in Sarawak 

between 1909 and 1920, and on Kinabalu in 19l3. Moulton's collections are largely in the 

Edinburgh, Kew and Singapore herbaria. Sarawak's first Conservator of Forests in 1919, 

John Phillips Mead, also collected plant specimens. Eric P. Mjoberg collected in Mt 

Murud, Mt Dulit, Mt Poe, the Bidi Caves, Mt Matang, Mt Penrissen and other areas in 

Sarawak when he was Director of the Sarawak Museum in 1922-1924, and afterwards in 

Dutch East Borneo and Southeast Borneo in 1925-1926. Forest Ranger D. Carroll collected 

several hundred specimens, mainly from the Kuching, Lundu and Sadong areas in Sarawak 

between 1922 and 1953. 

xxv


The Forest Research Institute, Buitenzorg began sending its collectors to Dutch 

(Indonesian) Borneo since about 1913. Johan Philip Pfeiffer, a Dutch chemical engineer, 

made botanical collections in 1913 in south and southeast Borneo (in the Sampit and 

Martapura areas), in 1915-1916 in northeast Borneo (in the Telok Selimau, Tanjung 

Mangkalihat and Berouw River areas) and again in 1918 in northeast Borneo (Telok 

Selimau); his collections are mainly at the Bogor and Leiden herbaria. 

10han Frederik Labohm, Forest Officer in southeast Borneo, collected many plant 

specimens in the Banjarmasin, Sampit, Balikpapan and Samarinda areas during 1916- 

1920. At this time (1917-1925), a Forest Overseer in the same district, Carel Nicolaas 

Johan Delmaar, also collected many plant specimens from the Kuala Kapuas area. 

Mohamad Dachlan, another overseer in southeast Borneo, too, made several hundred 

collections around the Banjarmasin, Martapura and Kuala Kapuas areas between 1918 and 

1939. Frederik Hendrik Hildebrand of the Buitenzorg Forest Research Institute made some 

small collections of plants in 1925 and 1928 in southeast Borneo. Another Forest Officer, 

Laurens Verhoef, also collected in southeast Borneo in 1928-1929. 

Joseph Clemens, a chaplain in the D.S. army, and his wife Mary Strong Clemens made 

substantial collections during several trips to Kinabalu between 1915 and 1917 (collecting 

at least 1,839 numbers) and between 1931 and 1933 (making more than 7,000 collections). 

Most of these have been sent to the Arnold Arboretum in the U.S.A. and the British 

Museum of Natural History. Their specimens were worked on by Merrill and others, and 

subsequently enumerated by H. Heine (1951, 1953). David L. Topping, a government 

official in the Philippines, collected more thall 500 fern specimens on Kinabalu together 

with Mary Clemens in 1915. A smaller collection of about a hundred numbers was also 

made on Kinabalu by George A.G. Haslam in 1916. 

The Chartered Company which administered B:-itish North Borneo formed its Forestry 

Department based in Sandakan, with American forester D.M. Matthews as the first 

conservator in 1914, who had an interest in Philippine dipterocarps. The North Borneo 

Forest Department started botanical collecting around 1915 (when its collector Aniceto 

Villamil was joined by Foxworthy to make the first collections for the Sandakan 

herbarium), and sent its dipterocarp specimens to the Bureau of Science in Manila for study 

by Foxworthy, who also served as advisor to the Forestry Department in North Borneo 

during 1915-1916. In 1916, the Forest Department had obtained collections representing 

537 species. In 1916 also, the first bulletin of the forest department, on the timbers and 

minor forest products of North Borneo, was published (Foxworthy 1916). Another 

American, Devillo D. Wood, transferred from the Bureau of Forestry in Manila in 1916 

and who became conservator in North Borneo in 1917, began encouraging active botanical 

collecting in the east coast lowlands. The Forestry Department maintained close links with 

its counterparts in the Philippines and sent plant specimens to the Bureau of Science in 

Manila for identification. In 1920 Maximo Ramos, a trained collector borrowed from the 

Bureau of Science in Manila, collected 1256 specimens in Sabah, mostly in the lowlands 

around Sandakan. By 1921 Wood's own collections had numbered at least 2555 and in 

1922 together with Jose Agama, a Filipino forest ranger who joined the British North 

Borneo forestry service in 1915, he collected on Balambangan Island off the north coast. In 

1922 he sent forest officers P. Castro and F. Melegrito to collect on Balambangan and the 

XXVI


nearby Banggi island. Other forest officers who took part in this collecting effort during 

this time were L. Apostol, P. Orolfo and Aniceto Villamil. These collections, and those of 

the Clemenses from Kinabalu and of Hose from Sarawak, were the principal basis of a 

number of botanical papers on the North Bornean flora by Elmer Drew Merrill, first a 

botanist and later the director at the Bureau of Science in Manila (Merrill 1916, 1917a, b, 

1918, 1922a, b, 1924, 1926). Merrill, who published a bibliographic enumeration of 

Bornean plants (Merrill 1921), also studied the collections of Adolph Daniel Edward 

Elmer, a botanist and collector for the Bureau of Science, who collected in the Sandakan 

and Labuk Bay area in 1921, and the Tawau area in 1922-1923 (about 1,523 numbers) 

(Merrill 1929). The zoologist c.B. K1oss, of the Federated Malay States Museum in 

Taiping, also visited the Sandakan area and Balambangan and Banggi Islands in 1927 and 

Kinabalu in 1928, and collected a few plant specimens. Merrill left for the University of 

California at Berkeley in 1924, where many duplicates from North Borneo (including those 

of Elmer) were then sent, and moved to the New York Botanical Garden in 1930. 

The German botanist Hans Winkler, Director of the Botanical Garden at Hamburg, 

collected in Dutch West and Central Borneo in 1924-1925; these collections are mainly in 

the Hamburg, New York, Leiden and Bogor herbaria. Frederik Hendrik Endert of the 

Buitenzorg Forest Research Institute made collecting expeditions to Dutch Borneo in 1925 

(around Kutei in northeast Borneo), 1928 (Tanah Blomboe in southeast Borneo) and 1938 

(Sanggau, Sambas and Paloh areas in West Borneo). The 1928 expedition was made 

together with Dirk Fok van Slooten, an assistant of the Buitenzorg herbarium. Endert's 

several thousand Borneo collections are principally at the Forest Research Institute in 

Bogor, while the 200-odd collections of van Slooten are at the Bogor herbarium. 

Between 1929 and 1932 Jan Pieter Schuitemaker, Forest Officer in Dutch West Borneo, 

made a number of collections there. The Dutch botanist, Oene Posthumus, collected in the 

Samarinda, Balikpapan and Tengarong areas in east Borneo in 1930; the collections, 

mostly of ferns, are mainly in the Bogor herbarium. In 1931-1933, the Assistant Resident 

of Dutch West Borneo, Louis Coomans de Ruiter likewise collected plant specimens there. 

Betje Polak, temporarily attached to the Buitenzorg Botanic Gardens and later the 

Buitenzorg Soil Science Institute, collected plant specimens in West Borneo (around 

Pontianak) in 1930, southeast Borneo (around Banjarmasin) in 1939 and West Borneo in 

1940. 

Richard Eric Holttum, Director of the Singapore Botanic Garden, made a collection of 

about 300 ferns, mosses and other plants on Kinabalu in 1931 together with the Clemenses. 

In 1932, also together with the Clemenses, Caetano Xavier Furtado (Assistant Curator in 

the Straits Settlements Gardens Department and a specialist in palms) collected on 

Kinabalu. In 1933, Cedric Errol Carr, a retired rubber planter in Malaya, collected about 

2,000 numbers (including 700 orchids) on Kinabalu. Carr's collections are in the 

Singapore herbarium. 

The next conservator in North Borneo, Henry George Keith, who assumed headship of the 

Forest Department in 1931, continued to encourage botanical collecting and also collected 

plant specimens himself after his arrival in North Borneo in 1925. Keith was interested in 

pursuing 'a foresters' flora and also compiled a preliminary list of North Borneo plant 

XXVII


names (Keith 1937); he based much of his work on identifications provided by H.K. Airy 

Shaw at the Kew herbarium. Many of the duplicates of collections at this time were 

despatched to the Kew, Singapore and Kepong herbaria, the last where dipterocarp specimens 

were identified by the Forest Botanist of Malaya, Colin Fraser Symington (who visited the 

Kabili-Sepilok Forest Reserve, Semporna and the Betotan and Segaliud areas in North 

Borneo, and Brunei in 1938). By 1940, the Sandakan Herbarium had amassed 10,803 

sheets. 

During the Oxford University Expedition to Sarawak of 1932-1933, its botanist, Paul 

Westmacott Richards, made about 2,800 collections, from Miri up the Baram to Marudi, 

the Dulit range, Bidi, Bau and Santubong. The specimens are principally at the Kew, 

Oxford, British Museum of Natural History, Singapore and Sarawak herbaria. Patrick M. 

Synge, who assisted Richards but fell ill, collected principally orchids and pitcher plants. 

At about this time, e.O. Flemmich, who was appointed State Forest Officer in Brunei 

made a substantial collection from trees in Brunei, which are mainly at the Kepong 

herbarium. 

In 1937, the American ornithological collector John Augustus Griswold collected over a 

hundred plants on Kinabalu during the Asiatic Primate Expedition of Harvard to Siam and 

Borneo. Expedition leader H.l Coolidge, who never went up Kinabalu, is sometimes 

erroneously credited as having collected Kinabalu plants with Griswold. 

In 1938, the Japanese Nobuhira Hanada collected specimens of woody climbers of the 

Menispermaceae mainly from the Kuching area. He was probably assisting botanist 

Y oshimatsu Yamamoto who was a specialist in the family and who collected in Tawau in 

North Borneo and Pontianak and Banjarmasin in West and South Borneo in 1939. 

In 1940, Pieter Buwalda of the Buitenzorg Forest Research Institute collected more than 

300 numbers from the Sampit region in south Borneo. 

Post-War Collectors 

When the Second World War intervened, Australian air raids on Sandakan in 

1944 destroyed the herbarium. Still with Keith as conservator, the herbarium's 

collection programme resumed in 1947, assisted by a return of some 1,000 

sheets of specimen duplicates from the Singapore Herbarium. In the same year Keith 

(1947) published his account of North Borneo timbers. Dipterocarp specimens, in 

particular, which had been sent to Kepong for study and identification by Symington, were, 

after his death in 1943, also sent to van Slooten in Bogor and later Amsterdam, until van 

Slooten also died in 1953. 

Java-born Dutchman Andre lG.H. Kostcrmans, of the Botanical Division of the 

Buitenzorg Forest Research Institute, made a number of expeditions to Borneo around this 

time: in 1948 and 1953 around Sampit in south Borneo; in 1951, 1952, 1953-1954, 1955 

and 1957 (and also later in 1963) in northeast and east Borneo; and in 1955 and 1956 in 

central Borneo. Kosterman's 1952 and 1953 trips were partly joined by Willem Mcijcr, an 

XXVIII


assistant in the Bogor Herbarium then, and later in 1955 a lecturer at the Faculty of 

Agriculture in Pajakumbuh in Sumatra before his repatriation in 1958. Anne Johnson, 

demonstrator in botany at the University of Malaya in Singapore, collected some Sarawak 

plants, mainly bryophytes and ferns, in 1951; these are found in the Singapore herbarium. 

James Sinclair, Curator of the Singapore Herbarium, collected around Santubong in 

Sarawak in 1949, on Kinabalu and around Sandakan in 1957, and in Sarawak and Brunei 

in 1960. In 1955, John William Purseglove, Director of the Singapore Botanic Garden, 

collected in the 1st Division of Sarawak (with Mohamad Shah of the Singapore 

Herbarium), and again in the Bako National Park in 1956 (with Hamish Boyd Gilliland of 

the University of Malaya and Mohamad Shah). George Alphonso, Curator of the Singapore 

Botanic Gardens, also collected in Sabah (on the Crocker range, near Tenom, Tambunan 

and Patau, and around Keningau) in 1959; his further trips to Borneo (1965 to Sarawak, 

with Samsuri Ahmad; 1966 to Kinabalu) did not yield many more herbarium specimens. 

Hsuan Keng, botany lecturer at the University of Malaya in Singapore, made a short 

collecting trip to Sarawak (Baram district) in 1961. 

In Sarawak, James Aidan Robb Anderson, who became Assistant Conservator during 

1951-1954 and Forest Research Officer in Kuching in 1955 before transfer to Brunei as 

State Forest Officer in 1956, and back to Sarawak later as Forest Research Officer, made 

substantial collections, especially of peat -swamp plants in the Rejang, Baram and 

Kayangeran areas (Sarawak) and Badas (Brunei). His collections are mainly at the Sarawak 

and Kepong herbaria. Anderson published a guide to the peat-swamp flora (Anderson 

1963). At this time Francis George Browne was the Conservator, and Browne also made 

some plant collections besides writing a compendium (Browne 1955) of common forest 

trees. Anderson's successor as Assistant Conservator, and later also State Forest Officer in 

Brunei, was a German forester, F.W.O. Eberhard Briinig, who collected especially the flora 

of podzolized soils in the 1st and 2nd Divisions in Sarawak and Brunei (Brunig 1968). 

During the Oxford University Expedition to Borneo, 1955-1956, led by G. Arnold, forester 

Gordon Harold Pickles collected plant specimens (lodged principally at Forest Herbarium, 

Oxford, Singapore and Kuching) in the Plieran Valley along the Rejang River, the Usun 

Apau Plateau, Mt Kalulong and along the Baram River. 

In 1952, Keith was succeeded as conservator in North Borneo by A.B. Walton from the 

Malayan forestry service, who continued to emphasize floristic inventory work. Dipterocarp 

collection and inventory continued to dominate botanical work based at the forest 

department when, upon the recommendation of John Wyatt-Smith (Forest Botanist of the 

Federation of Malaya, at Kepong), Geoffrey Howorth Spencer Wood was appointed the 

first Forest Botanist for North Borneo in 1954, when G.L. Carson succeeded Walton as 

conservator. In the same year Wyatt-Smith and Wood collected in North Borneo together, 

and afterwards Wood spearheaded the department's botanical collecting and began to 

specialize in the dipterocarps. After a year's home leave to Britain, Wood returned in 1957 

and went on a collecting trip to Brunei with P.S. Ashton, Brunei's newly arrived and first 

Forest Botanist. Wood died in Brunei from an accidental explosion in camp. Between 1948 

and 1958, John Kidman Cox of the North Borneo Agriculture Department collected mainly 

orchids there (including on Kinabalu); he made about 900 collections from North Borneo, 

which were sent to Singapore, Kew, the British Museum, Berkeley and Leiden. 

XXIX


Bertram Evelyn Smythies, State Forest Officer in Brunei in 1952-1959 and thereafter 

Conservator of Forests in Sarawak until 1964, when he joined the Royal Society Expedition 

to Kinabalu for a short period, also added substantially to the botanical collections. He 

published a simple guide to common Sarawak trees from 34 families (Smythies 1965). 

Peter Frederick Burgess, Deputy Conservator at S~mdakan during 1956-1965 (when North 

Borneo became Sabah), and who wrote an excellent account of the timbers (Burgess 1966), 

also made small numbers of collections, especially in the east coast areas. 

Timothy CharlesWhitmore collected in Sarawak (around Kuching), Brunei and North 

Borneo (Labuan, Lungmanis, Sandakan, Kalabakan, Tawau areas) in 1957 when principally 

studying dipterocarps in the field; he returned when serving as Colombo Plan botanist at 

Kepong to collect in Sarawak (at Semengoh) in 1966.- Whitmore's Borneo collections are 

mainly at Cambridge, Sandakan, Singapore and Leiden. During this period, Marius Jacobs 

of the Bogor herbarium made collecting trips to Sarawak, Brunei and Kinabalu in North 

Borneo in 1958, taking nearly 800 numbers. University of Malaya plant physiologist John 

Carrick collected in Sarawak (Bako) in 1959 (with Le. Enoch) and 1961; his collections, 

about 600, are in the university herbarium (Kuala Lumpur) and in the herbaria in 

Singapore and Kuching. 

A successor to Wood who would want to travel out to North Borneo was not easily found, 

and the job was finally given in 1959 to Meijer, who had just been repatriated from 

Sumatra. Meijer continued working on dipterocarps, building on manuscript notes left by 

Wood. Shortly after Meijer's arrival, the wooden Sandakan Herbarium building was razed 

by fire that spread from an adjacent veneer factory in 1961, when almost all its 15,000 

mounted specimens were destroyed. In 1961, Dan H. Nicholson, then a student at Cornell 

University, collected mainly aroids in Sarawak (Semengoh; Matang, Bau, Bako, Setapok). 

Peter Shaw Ashton, who left Brunei in 1961, became Forest Botanist in Sarawak between 

1962 and 1966. Although there were suggestions for a herbarium in the Sarawak Forest 

Department in 1947, a combined Sarawak Museum and Forest Department herbarium 

came into being only by 1959, and Sarawak Museum specimens (including Haviland 

collections) were then incorporated into a new herbarium in its own building in 1961. 

Some specimens collected by Haviland, Hose and the Clemenses from the Brunei, 

Sandakan and Singapore herbaria were also returned to the Sarawak herbarium. Ashton' s 

numerous collections of Brunei plants (as well as those of Hasan Pukul, with whom 

Ashton sometimes collected) and Sarawak plants are mainly at the Sarawak, Brunei, Kew 

and Kepong herbaria. He collected again in Brunei in 1965. Ashton wrote a manual to 

Brunei dipterocarps (Ashton 1964), published the same year as the one by Wood & Meijer 

(1964) for Sabah, followed by a supplement (Ashton 1968), and also completed by 1973 the 

manuscript for a book on 25 non-dipterocarp families in Sarawak, which publication was 

delayed until 1988 (Ashton 1988). Based at the Sarawak herbarium, collectors of 

importance include herbarium assistant Ilias Paie (from 1962, active until the 1980s), tree 

climber Banyeng anak Nyudong, together with Ardzi Arshid, Bojeng Sitam, Galau, and 

Rashid Taggoi. 

In Sandakan, the 1960s saw a new drive in collecting with herbarium assistants Leopold 

Madani (from 1960 and still collecting in the 1990s) and Aban Gibot (from 1962, active 

until the 1980s); there, others with substantial collections to their credit include George 

xxx


MikiI, J. Ampuria, James Ah Wing, David Brand, Jaswir Singh, Muin Chai, David 

Charington, Abdul Rahim, Masirom Rundi, and Henry Sinanggul. Through the massive 

recollection programme some 17,200 specimens (including about 13,600 post-1961 

collections) had been accessioned in 1964, when a new herbarium building was ready 

(Meijer 1964). Meijer, Aban and Madani also collected with both Royal Society expeditions 

to Kinabalu (1961, 1964), but used the SAN number series. 

Interest in the Kinabalu flora had not waned during this period. Although there were very 

few who collected frequently on the mountain apart from Forest Department staff, there 

was also some notable exploration carried out, such as the exploration by Sheila Collenette 

(formerly Iris Sheila Darnton) of Kinabalu from Kundasang over the Pinosuk Plateau in 

1960. Her collections, at first largely within the Kota Belud area where she was 

headquartered in 1954, and later more generally in North Borneo, number nearly 2,000 and 

are mainly with the herbaria of the British Museum of Natural History and at Leiden. 

University of Minnesota professor Ernst Cleveland Abbe and his wife Lucy B. Abbe 

collected during 1959-1960 in Kuching, Brunei and (accompanied by his research 

assistants Linn BogIe and Robert Bruce Kaul) on Kinabalu (which the Abbes revisited in 

1962) and in 1964 in Sarawak; their collections, many of oaks, are mainly in the herbaria 

at the Arnold Arboretum, Kew, Edinburgh, the University of Minnesota and Singapore. 

With the encouragement of Carson, Edred John Henry Corner from the University of 

Cambridge led two Royal Society expeditions to Kinabalu, in 1961 (Corner 1964) and 

1964. Corner was Assistant Director of the Singapore Botanic Garden during 1929-1941. 

The other principal botanical collectors on these expeditions, which represented an 

important advance in knowledge for one of the richest floras in the tropics, were John 

David Adam Stainton, a barrister-at-Iaw and assistant organizer of the expedition (in 

1961) and Chew Wee Lek (both expeditions), first botanist and later Keeper of the 

Singapore herbarium. Chew also collected in Sarawak in 1962 (Niah, Baram, Mt Mulu, Mt 

Api and Mt Benarat); 1963 (the Bau limestones, Mt Gading, Priam River, Mt Mentawa, Mt 

Maja), 1966 (Lambir, Marudi, Baram, Limbang, and the Tutoh and Melinau Rivers) and 

1967 (Mt Api, Mt Benarat, the Tiang Bekap limestones near Kuching, Mt Mentawa and Mt 

Maja). Corner had previously collected in Brunei and Sarawak (Bako National Park) in 

1959, and also collected in the vicinity of Kuching in Sarawak after the first expedition to 

Kinabalu in 1961. These collections are principally at the Kew herbarium; Chew's other 

collections were lodged mainly with the Singapore herbarium, and duplicated at the 

Leiden, Kew and the Arnold Arboretum herbaria. Kinabalu collections were also made 

during the 1964 expedition by Martin Edward Duncan Poore, Professor of Botany at the 

University of Malaya, and student Ho Coy Choke and deposited mainly in the University of 

Malaya herbarium in Kuala Lumpur. 

Brian Laurence Burtt and Patrick James Blythe Woods, botanists from the Royal Botanic 

Garden, Edinburgh, collected in Sarawak in 1962 (Kuching, Niah, Marudi, Mulu, Lambir, 

Kapit, Lundu, Po i) and 1967 (Kuching, Hose Mountains, Kelabit Highlands). The 

collections are mainly at the Edinburgh, Leiden and Sarawak herbaria. Swiss Hans Peter 

Fuchs, a Shell Company research palynologist, collected in Sarawak (Santubong, Semengoh, 

the Bau limestone, Baram, Marudi, Niah, Lambir), Brunei and Sabah (Kinabalu) in 1963; 

parts of his Kinabalu trips were with Meijer, Sheila Collenette and Hermann Otto Sleumer, 

XXXI


a Leiden botanist who continued to Sarawak (Mt Matang) to collect. Duplicates of Fuchs's 

collections are also at Leiden, Kew and Sarawak. Eduardo Quisumbing, Filipino orchidologist, 

collected in 1963 in Sarawak (Bako, Santubong, Semengoh, Bau limestones), Brunei 

and Sabah (Mt Kinabalu). In 1963, T.D. Pennington from Oxford University collected 

mainly Meliaceae from Sarawak (Lundu, Rejang River, Kapit) and Sabah (Sepilok, 

Kinabalu); these are mainly at the Forestry Herbarium, Oxford, Singapore, Kuching and 

Sandakan. 

The Japanese botanist Mitsuru Hotta collected, while on a Kyoto University expedition to 

Borneo with Minoru Hirano of the Osaka City University as leader, in Brunei (Temburong) and 

Sarawak (including the Tatau River and the Mt Mulu area, Limbang, Marudi) in 1963- 

1964 (Hotta 1964). About 10,000 collections were taken, deposited mainly at the Kyoto, 

Kuching, Leiden and Kew herbaria. This resulted in a whole series of papers (Hotta 1964, 

1965a, b, c, 1966a, b, 1967a,b; Iwatsuki 1965a,b; Tagawa 1965, 1967; Tagawa & Iwatsuki 

1966). 

Harold Emery Moore, Jr., Director of the Bailey Hortorium, collected palm specimens in 

Sarawak (Matang, Bako, the Bau limestones, Bintulu, Nyabau) in 1963-1964 and Sabah 

(Sandakan, Sepilok, Kimanis, Beaufort, Tenom, the Crocker Range, Kota Belud, Jesselton; 

with Meijer) in 1964; the collections are mainly at the Bailey Hortorium, Kew, Kuching 

and Sandakan herbaria. In Brunei'; ·Joannes Petrus van Niel, a Shell Company palynologist; 

,made herbarium collections between 1964 and 1971; these are at Leiden. 

Andries Kanis, a Dutch botanist stationed by the Dutch Ministry of Foreign Affairs at the 

Sandakan herbarium in 1965, and acting head of that herbarium for about nine months 

before returning to Leiden in 1966, collected about 250 numbers in Sabah (as North Borneo 

became known then); he also collected in Brunei and Sarawak (Baram, Lambir Hills, 

Nyabau and Segan Forest Reserves, Bintulu, Matang, Semengoh, Bako) in 1966. Bruce 

Weber, an American botanist, was attached to the Sabah Parks and Forest Department 

during 1965-1967 and collected some plants. In 1966, a Chinese botanist who came to be 

settled in the Netherlands and employed at the Rijksherbarium in Leiden, Ding Hou, also 

visited to collect in Sabah (mainly in the Sandakan and east coast districts, and on 

Kinabalu) and Sarawak (mainly the Kuching, Bau, Bintulu, Sibu and Nyabau areas). 

Thomas Kenneth Newell, a graduate student at the University of Hawaii, made a small 

collection, mainly of monocots, in Sabah (Sandakan, Jesselton, Tamparuli and Kundasang) 

in 1966; his collections are at the Bishop Museum, Sandakan, Paris, Leiden, Florence and 

Kew. Meijer left Sabah in 1968, when also Ken Ogata from the Meguro Forest Experiment 

Station in Tokyo collected a large number of herbarium vouchers together with wood 

samples from Sabah. Shigeo Kurata of the Japanese Insectivorous Society collected 

pitcher-plant specimens in Sabah (mainly Kinabalu) during a MindorolNorth Borneo 

expedition in 1967-1968. Others in Sabah collected actively during this period, including 

planter Jim B. Comber, who took mainly orchids and ferns, principally from the Tenom 

area and the Crocker Range; his collections were given to Kew. Engkik Soepadmo 

(University of Malaya) collected with Anderson in the Simanggang and Bau areas in 

Sarawak in 1961, the collections numbered by Anderson under the Sarawak herbarium 

series, and with Gordon Smith (University of Malaya) and Paul Chai (Sarawak herbarium) 

in the Kapit and Bau areas in Sarawak in 1969. 

XXXII


John Dransfield collected in Brunei and Sarawak (Bako, Matang, Bintulu) in 1968 (as a 

research student), and Kalimantan Timor (east Borneo: Balikpapan, Samarinda, Kutei) and 

Kalimantan Selatan (southeast Borneo: Djaro, and Mt Sarempaka; with Indonesian botanist 

Kuswata Kartawinata and Dutch botanist Eduard Ferdinand de Vogel) in 1971 (while a 

Colombo Plan botanist at Bogor); his collections were distributed mainly to Kew, Ithaca, 

Bogor, Leiden, Kepong and Singapore. W. Soegeng Reksodihardjo, botanist of the Bogor 

Herbarium, collected in the Kutei Nature Reserve in East Kalimantan in 1970; the 

collections are mainly at Bogor. 

Hotta collected again in Sabah in 1968-1969 with Shohei Kokawa, on Kinabalu and the 

Trus Madi area, and in the Tawau Hills and on Mt Silam. In 1969, Peter Francis 

Cockburn was appointed Forest Botanist at Sandakan and continued a statewide collecting 

drive. He left in 1977. David l\1:abberley from Oxford collected in Sabah in 1974. Around 

this time, there are some collections by Anthony Lamb, of the Sabah Agriculture .Qepartment, 

registered; his collections were mostly orchids and although between 1977 and 1981 he 

used Sandakan herbarium collecting numbers, the specimens are not always lodged with 

the herbarium there. Lamb continued collecting into the 1990s, in the later part using ~is 

own numbers; many of his collections were sent to Kew or Leiden. P.S. Shim, a plantation 

silviculturist in the Forest Department in Sandakan also collected (mostly ferns and 

orchids), using the herbarium's number series. Gary Shea, a Canadian University Service 

Overseas volunteer attached to the herbarium in Sandakan, also collected widely in Sabah 

during 1971-1972; Shea's collections are at Sandakan and distributed to Kew, Kepong and 

Leiden. Cockburn organized the Trees of Sabah project, which saw the publication of two 

volumes (Cockburn 1976, 1980) that covered, in a very superficial and cursory manner, 44 

angiosperrn families and two conifer families of tree. Shea assisted with the write-ups for 

several families in the first volume. K.K. Tiong was appointed Forest Botanist in Sandakan 

in 1977, and continued the collection in Sabah. 

Paul P.K. Chai became Forest Botanist in Sarawak in 1970, collecting widely throughout 

Sarawak. Ilias & Chai also collected with the 1964 Royal Society Expedition to Kinabalu in 

Sabah. Hans Peter Nooteboom, Leiden botanist, made collecting trips to Sabah (Mt Alab, 

Mt Lurnarku, Mt Trus Madi, Tambunan) in 1969, and Sabah (Kinabalu, Sandakan) and 

Sarawak (Bario, Mt Murud, Apo Batu Buli Range; with Paul Chai) in 1970. Nooteboom 

and Chai's collections number about 1600, distributed to the Sandakan, Kuching and 

Leiden herbaria. In 1971 Anderson returned to collect in the Mt Mulu area in Sarawak. 

Chai again collected in the Mulu area (Mt Api, Mt Benarat, Mt Buda, the Melinau Gorge) 

in 1975 together with B.L. Burtt. 

De Vogel collected again in Kalimantan Selatan (Jaro, Mt Sarempaka) in 1972 and 1973; 

his collections are principally at Bogor and Leiden. In 1972, University of Hull research 

assistant RJ. Morley made a small collection of plants on Kinabalu, distributed to the 

University of Hull, Kew and Leiden. In 1973, a New Zealander, David Wilson Ives, 

consultant with the N.Z. Colombo Plan aid to Indonesia's beef industry, collected plant 

specimens in Kalimantan Selatan (Tandjung, Tabanio River, Djilatan, Djurong); his 

specimens are at the Christchurch herbarium. 

Benjamin Clemens Stone from the University of Malaya, who had collected in Sabah and 

Sarawak in 1967, also collected in Sabah's Danum Valley in 1976 as a World Wildlife 

Fund (WWF) consultant botanist, on Gaya Island and on Balambangan Island in 1977 

XXXIII


(Stone 1980) in the same capacity. These several hundred collections were lodged with the 

Sandakan, Kuching and University of Malaya herbaria. Earlier in 1970, Sabah Forest 

Department ecologist lE.D. Fox had also collected on Balambangan (about 80 numbers). 

P.J. Martin and A. Clive Jermy of the British Museum (Natural History) collected on Mt 

Mulu and along the Melinau and Melinau Paku Rivers in Sarawak in 1976. 

In 1976, Hotta again collected in Sabah (Kota Kinabalu, Kinabalu, Ranau, Telupid, 

Sandakan, Madai) and Sarawak, together with another botanist Michio Tamura on yet 

another Kyoto University Expedition to Borneo (Kobayashi & Hotta 1978). Further Kyoto 

University expeditions followed (lwatsuki et al. 1983). In a 1978-1979 expedition to East 

Kalimantan (Sekatak lowlands near Tarakan, Samarinda, Sebulu, Tabang, Balikpapan, Mt 

Beratus) and South Kalimantan (Banjarmasin, Mt Besar), led by Kunio Iwatsuki and 

joined by G. Murata, Masahiro Kato, J.P. Mogea and K. Kartawinata, some 4,650 numbers 

were collected. Another expedition in 1980-1981 to East Kalimantan, led by H. Ogawa 

together with Kato and Kunihiko Ueda and joined by Dedy Darnaedi of the Bogor 

Herbarium, and which collected about 2,500 specimens, surveyed Sebulu, Tahang, the 

Menubar River, Mt Kongkat, Mt Kongbotak, and the limestone Mt Njapa and Mt Buntung. 

In a further expedition to East Kalimantan in 1981, led by Iwatsuki and joined by Kato, 

Motoharu Okamoto and Ueda, Darnaedi and Eko Baroto Walujo of the Bogor Herbarium, 

and Rob Geesink of the Rijksherbarium in Leiden, the Long Bawan region near the 

Sarawak border and the Balikpapan and Samarinda areas were explored, netting some 

4,750 numbers for the Japanese collectors. Kalimantan collections were distributed mainly 

to the Kyoto, Bogor and Leiden herbaria; Sabah specimens to the Kyoto, Sandakan and 

Leiden herbaria. Two other Japanese collectors, Suehiro and Harigae, are credited with 

Kinabalu collections made in 1979, deposited in the Kyoto herbarium (lwatsuki et al. 

1983). 

During the Royal Geographical Society Expedition to Mt Mulu in Sarawak in 1977-1978 

(Jermy & Kavanagh 1982; Jermy 1984), several botanists collected in the area'at different 

times, including: Chai, Anderson, George Argent (Royal Botanic Garden, Ediburgh), 

J.A.R. Kerby (Edinburgh), Ivan C. Nielsen (University of Aarhus in Denmark), Carlo K. 

Hansen (Copenhagen Botanical Museum), J. Dransfield, B.C. Stone, Ruth Kiew (Agricultural 

University of Malaysia), Barbara S. Parris (University of Cambridge), A. C. Jermy, A. 

Touw (Leiden), WaIter F.B. Julich (Leiden), B. Cop pins (Edinburgh) and N. Sammy (a 

Singapore-trained biologist working in Australia). 

Dransfield collected rattans in Sabah in 1979 while on secondment from the Royal Botanic 

Gardens, Kew, to the British Overseas Development Administration project in Sabah to 

carry out a rattan survey. He was accompanied by his wife, Soejatmi Dransfield, who 

collected bamboos. John and Soejatmi Dransfield were in Sarawak in 1981, collecting 

rattans during a similar survey there. 

Of check-lists and floras 

Undoubtedly, the account by Stapf (1894) of the Kinabalu plant life must be the 

earliest systematic account of the flora of anyone place in Borneo. Stapf 

enumerated 360 species of flowering plants, ferns and bryophytes. After Gibbs 

(1914), which updated this Kinabalu account, it was largely people like Merrill (see 

XXXIV

eferences here), Copeland (1917, 1935), Ames & Schweinfurth (1920), Fisher (1932), 

Christens en & Holttum (1934), and others who produced numerous papers on Bornean 

plant taxa. Hallier (1916) probably made the first attempt at a flora of Borneo (in German). 

Merrill's bibliographic enumeration of Bornean seed plants (Merrill 1921), which lists 

4,924 species, not including the 120 orchid species listed by Ames & Schweinfurth (1920), 

giving a total of 5,044 species (in 1,162 genera and 156 families), was strong motivation 

for continuing work. Merrill's 1921 estimate was a Bornean flora of some 9,boo flowering 

plants including 3,000 species of tree. This work is essentially uncritical, as is the 

subsequent enumeration by Masamune (1942), listing 7,201 species (1,310 genera, 165 

families) of seed plants, that reproduces Merrill's list with few changes and adds partial 

information from the intervening period. Masamune also published a list of Bornean 

pteridophytes (Masamune 1945), which lists 963 species of ferns and fern-allies in 118 

genera. 

To Masamune's enumeration, Merrill (1950) adds another 200 species of seed plants 

published by Ridley, Airy Shaw and others and not consulted by Masamune; estimating the 

size of the Bornean flora at 12,000 to 15,000 species (including seed plants, ferns and fernallies), 

he laments the lack of a descriptive flora. ' 

While specialists in the large western herbaria continued to work on a taxonomic basis, 

foresters' check-lists of trees and dipterocarp floras ("identification manuals") heralded a 

next phase of botanical development in Borneo, beginning with those of Wood & Agama 

(1956), Hasan & Ashton (1964), Wood & Meijer (1964) and Ashton (1964, 1968). van 

Steenis (1950) pointed out that Borneo had the largest number of endemic genera among 

islands in western Malesia and Airy Shaw (1975), who named many Bornean collections at 

the Kew herbarium and continued an interest in Bornean plants over a long period, 

considered Borneo to be " ... floristically and phytogeographically ... an area of prime 

importance--almost the 'hub' of Western Malesia ... " Forest floras and tree floras continued 

to be in the ambitions of the forest departments of North Borneo (after 1963, Sabah) and 

Sarawak, and early attempts include Ashton's manual of non-dipterocarp trees of Sarawak 

(revising only 25 families of tree, completed by 1973 but published only in 1988) and the 

two volumes of the "Trees of Sabah" (accounting for 46 families of tree: Cockburn 1976, 

1980). Anderson (1980) published a check-list of Sarawak trees and a guide to the 

Moraceae of Sarawak (Primack 1983) appeared to push on with what Ashton began, but the 

writing of such manuals in the same series was not continued. Kiew (1984) called for a 

concerted effort towards a flora of Borneo and in 1985, an attempt to continue the "Trees of 

Sabah" project by B.c. Stone, then of the Academy of Natural Sciences, Philadelphia, 

never materialised (Wong 1994). 

For Indonesian Borneo, local check-lists and floras were longer in coming (Whitmore, 

Tantra & Sutisna 1990a, b, c) and did not, in the case of some large and taxonomically 

complex families, attempt to list beyond a few common taxa. However, keys by Ashton to 

all known Bornean dipterocarp species and genera are given. One of the more recent 

accounts includes that by Kessler & Sidiyasa (1994), of the trees of the Balikpapan 

Samarinda area in East Kalimantan. 

Aside from the tree flora, modern identification manuals, amounting to floristic 

enumerations, of the rattans of Sabah and Sarawak (1. Dransfield 1984, 1992), and the 

bamboos of Sabah (S. Dransfield 1992) have been published. Modern compilations of the 

xxxv 

BRIEF HISTORY (WONG)

Ashton, P.S. 1964. Manual of the Dipterocarp Trees of Brunei State. Oxford University 

Press, London. xii + 242 p. 

129. 

Ashton, P.S. 1988 Manual of the Non-Dipterocarp Trees of Sarawak. Vol. 11. Dewan 

Bahasa dan Pustaka for Forest Department Sarawak, Kuching. xix + 490. 

Beccari, O. 1902. Nelle Foreste di Borneo. Firenze. 

Browne, F.G. 1955. Forest Trees of Sarawak and Brunei and their Products. Government 

Printing Office, Sarawak. 369 p + xviii. 

XXXVI 

Anderson, lA.R. 1963. The flora of the peat swamp forests of Sarawak and Brunei, 

including a catalogue of all recorded species of flowering plants, ferns and fern allies. 

Gard. Bull. Sing. 20: 131-228. 

Ames, O. & C. Schweinfurth. 1920. Orchids of Mount Kinabalu, British North Borneo. 

Orchidaceae 6 (I-XIV): 1-233. 

Airy Shaw, H.K. 1975. The Euphorbiaceae of Borneo. Kew Bulletin Additional Series IV. 

Current estimates of the Bornean vascular flora hover between 9,000 (Merrill's original 

estimate in 1921) and 15,000 (the upper limit of Merrill's subsequent estimate in 1950). 

This state of uncertainty surely reflects the existence of large gaps still in our knowledge. 

of Kinabalu's vascular plant flora, John Beaman and associates have published annotated 

In the meantime, the incredibly high plant diversity of Kinabalu, Borneo's highest 

mountain, continues to attract systematic study. As part of a modern botanical enumeration 

check-lists of the ferns (parris, Beaman & Beaman 1992) and orchids (Wood, Beaman & 

Beaman 1993). Farther south, a new inventory of the Brunei flora towards compiling a 

modern check-list, initiated in 1989 between the Brunei Forestry Department and the Royal 

Botanic Gardens, Kew, is yielding more iriformation of a very rich flora, with many 

novelties. 

orchids of Borneo (Chan et al. 1994, Vermeulen 1991) as well as a check-list for Borneo 

(Wood & Cribb 1994) have also been published. 

References 


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XXXVII


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its environment and biota being the results of the Royal Geographical Society/Sarawak 

XXXVIII


Government Expedition and Survey 1977-1978. Part I. Sarawak Mus. 1., Special Issue No. 

2. xxiv + 279 p. 

Keith, H.G. 1937. A Preliminary List of North Borneo Plant Names. North Borneo Forest 

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p. 

Kessler, P.J.A. & K. Sidiyasa. 1994 Trees of the Balikpapan-Samarinda area, East 

Kalimantan, Indonesia. Tropenbos Foundation, Wageningen. 446 p. 

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Masamune, G. 1945. Enumeratio Pteridophytarum Bornearum. ii + 124 p. 

Meijer, W. 1969. The contribution of Governor Charles Van de Leur Creagh to botanical 

exploration in Sabah, 1888-1895. Sabah Soc. 1. 5: 67-68. 

Merrill, E.D. 1916. Notes on the flora of Borneo. Philip. J. Sci. 11, Bot.: 49-100. 

Merrill, E.D. 1917a. Contributions to our knowledge of the flora of Borneo. 1. Str. Br. Roy. 

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Merrill, E.D. 1917b. Alabastra Borneensia. 1. Str. Br. Roy. As. Soc. 77: 189-247. 

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Merrill, E.D. 1922a. New or noteworthy Bornean plants (Part 1). 1. Str. Br. Roy. As. Soc. 

85: 151-201. 

Merrill, E.D. 1922b Additions to our knowledge of the Bornean flora. Philip. 1. Sci. 21 (6), 

Bot.: 515-534. 

Merrill, E.D. 1924. Plants from Banguey Island. Philip. 1. Sci. 24(1): 113-116. 

Merrill, E.D. 1926. The Flora of Banguey Island. Philip. 1. Sci. 24(3): 341-427. 

XXXIX


Merrill, E.D. 1929. Plantae Elmerianae Borneenses. Univ. of California Publications in 

Botany 15: 1-3 16. 

Merrill, E.D. 1950. A brief survey ofthe present status of Borneon botany. Webbia 7: 309- 

324. 

Parris, B.S., R.S. Beaman & lH. Beaman. 1992. The Plants of Mount Kinaba1u. I. Ferns 

and Fern Allies. Royal Botanic Gardens, Kew. 

Primack, RB. 1983. Forester's Guide to the Moraceae of Sarawak. Forest Department, 

Sarawak, Kuching. 140 p. 

Smythies, B.E. 1965. Common Sarawak Trees. Borneo Literature Bureau. 

Stapf, O. 1894. On the flora of Mount Kinaba1u in North Borneo. Trans. Linn. Soc. Lond., 

Bot. 4: 69-263. 

Steenis, c.G.G.J. van. 1950. The delimitation of Malaysia and its main geographical 

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Steenis-Kruseman, M.l van. 1950. Malaysian plant collectors and collections being a 

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Duplicated typescript: Flora Malesiana Foundation, Leiden. 

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Malays. Br. Roy. As. Soc. 53 (1): 68-89. 

Tagawa, M. 1965. Ferns of Borneo, collected by M. Hirano and M. Hotta; 3. Acta Phytotax. 

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Acta Phytotax. Geobot. 22: 87-94. 

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Kew & Toihaan Publ. Co., Malaysia. x + 342 p. 

XL

BRIEF HISTORY (WON G) 

Whitmore, T.C., LG.M. Tantra & u. Sutisna. 1990a. Tree Flora of Indonesia. Check List 

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Royal Botanic Gardens, Kew. 

Wood, J.J. & P.J. Cribb. 1994. A Checklist of the Orchids of Borneo. Royal Botanic 

Gardens, Kew. xii + 409 p. 

XLI


BIOGEOGRAPHY AND ECOLOGY 

P.S. Ashton 

The Arnold Arboretum, 

Harvard University, Cambridge, 

Massachusetts, U.S.A. 

The physical setting 

Botanists from Peninsular Malaysia who visit Sabah and Sarawak for the first time, 

and who are interested in floristic ecology, are startled to find that their traditional 

reference points are missing. The customary altitudinal sequence-lowland 

dipterocarp, hill dipterocarp, upper dipterocarp, (lower) montane oak and (upper) 

ericaceous forest-is either unrecognisable as in the case of these categories of dipterocarp 

forest, or does not consistently adhere to the expected altitudinal sequence. With closer 

familiarity, a new set of forest types correlated to soil becomes recognisable, which more or 

less obscures the altitudinal sequence. Furthermore, the varied coastline in the northwest, 

which is mostly bordered by vast peat swamps for which there is no peninsular equivalent, 

lacks any consistent coastal hill association. It was Symington (1943) who first put forward 

the floristic classification of Peninsular Malaysian forests which is still adhered to, but up 

to now no East Malaysian scheme has been proposed. Why is it that the floristic ecology of 

East Malaysian forests is so different? 

The hills of Peninsular Malaysia are thought to have remained above sea level since before 

the origin of the flowering plants, in the Jurassic era perhaps 150 million years ago, and the 

present landscape may be dated from perhaps the Cretaceous, c. 100 million years ago 

(Richardson 1947; Gobbett and Hutchinson 1973). Their soil cover is extraordinarily deep: 

as a rule, soils there exceed two meters over rotting rock through which occasional roots 

may penetrate, and sometimes much more. By contrast, in much of northern Borneo it is 

difficult to find soils as deep as one meter. This is because northern Borneo is at the 

southeastern edge of the ancient Laurasian continent and continues, even now, to 

experience coastal up- and downwarping, alternately sinking and accumulating sediments, 

then lifting and rapidly eroding, and thereby forming series of anti- and synclinally folded 

sedimentary rocks (Liechti et al. 1960). The result is a sharply dissected terrain of parallel, 

. narrow interior ridges and valleys, in which ridge-top, slope and valley bottom are sharply 

differentiated. This landscape is much more rugged than the peninsular main range, which 

is gentle by comparison and with a dendritic lattice of ridges. 

Each successive period of uplift and erosion has led to a further loss of clay minerals and 

nutrients, and to the creation of new hills to the north and east near the coast which each 

time become sandier and with poorer soils, and whose rocks are successively softer. These 

sandy, infertile soils are mostly yellow, but are so acidic that litter decomposition is slow 

"XLIII


and a mantle of peaty organic matter forms. They are known as humult ultisols. The 

organic layer impedes surface erosion' and, as a result, these young sandy hills alone 

quickly develop a mature rounded topography and remarkably deep soils. This contrasts 

with coastal clay hills which, where they occur, remain steep even if low, with clear signs 

of surface erosion; there, the only deep soils are formed by slumping downslope. 

Superimposed on this landscape in the lowlands, along the northwestern coast where the 

tectonic activity is greatest, are the imprints of changing sea levels during and after the ice 

ages: the current coastline, mostly a straight, sandy shore often bordered with Casuarina 

equisetifolia and its associates, is backed by a vast peat swamp which originated as an 

embayed mangrove when the sea level last declined, about 5,500 years ago. Behind again, 

and also perched on the hills which in places push to the coast between Bintulu and Kota 

Kinabalu, are extensive white sand terraces bearing giant podsol soils with deep, peaty 

organic surface layers (except where they have been burned); these were former coastlines 

during high sea levels, between the great ice ages. 

Inland, the predominating sedimentary hills are intruded by volcanic masses, granite, and 

limestone. The volcanic rocks, which are mainly tertiary, are as diverse as the sediments, 

from acid rhyolites yielding similar humult ultisols to the sandy coastal hills, through 

dacites and base-rich fertile basalts, to ultramafic rocks which carry diverse soil types and 

consequently floras, and which are widespread in northern and eastern Sabah but absent 

from Sarawak. The granites are young, rocky and covered with shallow soils. The 

outstanding example of a granite formation in this territory is the great dome of Kinabalu, 

the highest mountain between the Himalayas and New Guinea, which arose less than one 

million years ago. There are other, tertiary, granite mountains in West Sarawak. The 

several limestone massifs in eastern Sabah and northeastern and western Sarawak are 

generally of a pure dolomitic limestone which erodes only by solution, yielding no mineral 

soil. What soils there, are organic and surprisingly acidic. 

The floristic composition of forests is most markedly differentiated where the soils are 

limiting to plant growth and survival, by adverse water conditions or low nutrients. The 

combination of sharply defined topography, extraordinarily diverse rock substrates, and 

generally shallow, low nutrient soils in most of Sabah and Sarawak has led to a greatly 

diversified forest cover in which soils-correlated differentiation of forest types dominates 

over altitudinal and other influences. This is particularly so between Kota Kinabalu and the 

west coast of Borneo south to Pontianak in Kalimantan, where sandy sedimentary soils and 

the often deep and diversified coastal peat swamps create a diversified landscape quite 

different from those found elsewhere in the country. . 

Impact on taxonomy 

A 

notable result, familiar to the experienced field botanist in Sabah and Sarawak but 

a potential pitfall to the herbarium taxonomist, is a marked and complex pattern of 

small-scale variation in the characteristics of widespread species which occur in 

the many disjunct habitats of the region. Identification of countless species from ecological 

plots scattered through Brunei and Sarawak persuade us that finely distinguished species 

based on single specimens will rarely be maintained as knowledge increases, and that van 

Steenis' (1957) admonition to hold to a conservative and broad species concept may prove 

wise. 

XLIV

p 

< 

Kinabalu 

sandstone kerangas---f:!.....-------ift! 

Littoral forest 

Agathis borneensis 

phasic 

terrace kerangas 

Meters 

communities 

" " altitude 

Anisoptera /. Dryobalanops "', 

4000 

----..1--4- Tree line 

3000 

Forms prevalent on the northwest coast 

Forms prevalent 

on the east coast 

Montane elfin woodland (upper)-----'i'l- 

(organic mor soils) 

--~~-Montane oak-laurel 

2000 

forest (lower) 

(inorganic mull soils) 

Montane 

1000 

Peat swamp forest 

'"IIi!:-- Ultramafic 

Cotylelobium lanceolatum 

~~~~~~~~~;i~~~z;~~~~g~r~o~ss:i~ve~n~'~a~M~D;F~.~/;/o~; .. :J~:_>I·~____ ~ __ S~h~o:re~a~p~a~rv~~~o~l~ia~M~D~F~~~~la~n~c~e~o~la~t~a~M~D~F~='~'~"~'~~o 

- raw humus 

soil surface 

Increasing 

Dipterocarpus beccarii, D. crinitus, 

Shorea laevis etc. MDF 

Diagrammatic chart of the major jloristic associations in Sabah and Sarawakforests 

MDF stands for Mixed Dipterocarp Forest


Forest types 

The overall ecology of the main forest types has been well summarised by Whitmore 

(1984). Here, emPhasi.s is given to the major floristic associations. The lowland as well 

as mountainous forests of Sabah and Sarawak vary dramatically in stature and 

canopy structure, from less than 10 m tall with dense, even canopy to over 70 m, with 

scattered clumps of giant emergent dipterocarps or Koompassia, or sometimes lofty, dense, 

even canopy as in the alan (Shorea albida) peat swamps. However, forest stature and 

structure, which correlate with soil water availability, do not always correlate in Borneo 

with species composition, which correlates most closely with soil chemistry. Floristic 

variation is actually continuous, but there are also certain rather sharp environmental 

boundaries which differentiate major floristic associations. The following main lowland 

types can therefore be recognised: 

Mixed dipterocarp forest (MDF). Although dipterocarps decline in abundance and in 

number of species with altitude, and although there are some ten species which are 

confined to higher altitudes, the change in flora with altitude is continuous and no 

altitudinally zoned types can be recognised. Most strikingly seraya (Shorea curtisii), 

although scattered in the coastal hills of Sarawak and southwestern Sabah, is local and 

never dominant there, and is rare on the inland ridges. Therefore, there is no consistent 

coastal hill species association, nor is there a recognisable "hill dipterocarp forest." In 

Borneo, the upland forests on yellow-red soils which are dominated by dipterocarps in the 

emergent canopy are termed Mixed Dipterocarp Forests (MDF) to distinguish them from 

others, such as the S. albida-dominated peat swamp forests, in which one or only a few 

emergent dipterocarp species occur. The major floristic division in these forests is between 

Mixed Dipterocarp Forests in which either Dryobalanops lanceolata or Shorea parvifolia 

(often also with Shorea macroptera) are among the commonest species, and which are 

confined to clay-rich soils lacking an acid matted surface organic mat; and forests in which 

Anisoptera grossivenia and variously Dryobalanops aromatica and Dipterocarpus globosus, or 

in western Sarawak Shorea falcifera, are among the commonest emergent species, and 

which are confined to humult ultisols. Probably fewer than one quarter of species are 

shared between these two major types (cf. Ashton 1964: 34). Ther:e is also a widespread 

type on intermediate, sandy clay soils of the lowlands and also inland ridges, of which 

Dipterocarpus crinitus is a characteristic representative, and in which Shorea macroptera 

is also common. 

These major types vary geographically and with habitat, but they are almost always sharply 

distinguishable and separated in the landscape, whereas variation within them is continuous. 

Within the first type, giant forests in which Dryobalanops lanceolata, Shorea superb a, and 

Dipterocarpus caudiferus were abundant; and Octomeles sumatrana, Pterocymbium 

tubulatum and Pterospermum javanicum were characteristic pioneers, once clothed the' 

extensive fertile volcanic hills and sedimentary clay loams of lowland east Sabah as well as 

the lower basalt massifs and low clay hills of Sarawak. These forests were the first to be 

logged and their land converted to agriculture but some remain at Quoin Hill in Tawau, 

Bukit Mersing in Sarawak, and elsewhere. The Shorea parvifolia subtype is the prevailing 

forest of inland Sabah and Sarawak, on the shallower yellow clays and sandy clays which 

cover much of the inland sediments. 

XLVI

BIOGEOGRAPHY AND ECOLOGY (ASHTON) 

The most distinctive forest type on humult ultisols is that which is confined to its deepest 

soils, and therefore to low soft sandstone hills which are mostly but not always near the 

coast. The most extensive areas are in the Lambir Hills and the Andulau and Labi Hills of 

Brunei. Smaller islands occur at Beaufort Hill in southwest Sabah, in the Ulu Tutoh, Baram 

and Belait in Brunei, in Similajau, Nyabau and Segan forests in Bintulu, on the Arip 

rhyolite, in the northernmost hills between the lower Balingian and Mukah rivers, and in 

some small patches west of the Lupar including the foothills of Gunong Gaharu, and Bukit 

Undan in Bau. This type deserves special mention on account of the confinement of certain 

well known peninsular species there (including Dryobalanops aromatica and Shorea 

curtisii), because it appears to be the richest in species of all forest types in Malaysia, 

because of its remarkable endemism, and because it still remains unfamiliar to many 

foresters and biologists. The commoner dipterocarp species which are concentrated in or 

confined to it include Anisoptera grossivenia, Dipterocarpus globosus, D. lowii, D. rigidus, 

D. sarawakensis, Dryobalanops aromatica, Hopea beccariana, H. treubii, H. tenuinervula, 

Shorea acuta, S. crassa, S. curtisii, S. cuspidata, S. elliptica, S. Jalcifera, S. jlemmichii, S. 

geniculata, S. kunstleri, S. ladiana, S. laxa, S. ovata, S. rubella, and S. slootenii; species 

apparently confined to this type which appear in this first volume of the Tree Flora include 

Anisophylla Jerruginea, Canarium divergens, C. grandifolium, Santiria megaphylla, 

Sarawakodendron filamentosum, Parinari metallica, Allantospermum borneense (often the 

most abundant species, and very characteristic), and QuaSSia borneensis. 

A variety of other MDF, floristically related to this but generally poorer in endemics but 

more widespread in Sabah and Sarawak, occur on humult ultisols richer in clay, where up 

to 30% of the flora also occur in the S. parvifolia subtype. Dipterocarpus crinitus, Shorea 

macroptera and on ridges Dryobalanops beccarii distinguish this association. There is also 

a range of MDF on shallower, frequently more sandy soils, where there is an increasing 

heath (kerangas) forest element. 

Heath (kerangas) forest. These are forests, often but not invariably of low stature and 

lacking emergents, that are confined to organic white sand podsol s6ils. Their flora is very 

distinctive, with perhaps fewer than one half of species also occurring in MDF, usually on 

humult ultisol soils. Although Brunig's monograph (1974) provides extensive insight into 

the great floristic diversity of kerangas forest, the relationship between floristic and habitat 

variation remains even more poorly understood in this type of environment than is the case 

with MDF. Kerangas occurs in two principal habitats: on raised beach terraces where the 

podsols are generally deep except in the center and along the lower edges where they are 

often poorly drained; and on sandstone ridges and plateaux where they are generally 

shallow and often rocky. There appear to be floristic differences between-and to perhaps a 

lesser extent within-these habitats; lowland Agathis, for instance is almost entirely 

confined to terraces. Certain families of kerangas are also particularly well represented in 

upper montane forest ("elfin woodland"). They include Myrtaceae, Theaceae, and 

Podocarpaceae, and to some extent Ericaceae: Clusiaceae (Guttiferae) and Ebenaceae are 

also well represented in kerangas. " 

Peat swamps. There are extensive peat swamps up the northwestern coast, eastwards to 

Papar in Sabah. Anderson (1963) recognised six phasic communities from the margins to 

the centers of individual swamps, which he regarded as succeeding one another over the 

XLVII


time scale of the development of the dome of the raised bog. Anderson documented a total 

vascular flora of only 317 species of flowering plants, including several which are confined 

to certain districts and which otherwise are known from kerangas or MDF on humult 

ultisols. The endemic flora is small, most species being found also in kerangas. Notable is 

the tendency towards canopy dominance of Shorea albida and, towards the centers, Litsea 

palustris, Combretocarpus rotundatus and sometimes Dactylocladus stenostachys; but 

there are no subcanopy dominants. 

Other specialized lowland forests. There are major limestone formations in Bau District of 

western Sarawak, Mulu National Park, and in East Sabah, and also many minor exposures. 

The limestone flora is rich in endemics, particularly among herbaceous species; the general 

characteristics are similar to the limestone outcrops of Peninsular Malaysia. The highest 

limestone in Malaysia is at Gunong Benarat, Mulu NP, at 1,300 m, which bears upper 

montane elfin woodland on its jagged upper slopes. 

In several parts of Sabah, notably on the slopes of G. Kinabalu and in the northeast and 

east, are extensive exposures of ultramafic rocks. These exposures can bear freely draining 

acid soils with a surface organic layer. In this case the forest is "of more or less short stature 

with elements of both the MDF on humult ultisols, and kerangas. A small but important 

element of Bornean affinities are present, such as the strictly endemic Borneodendron 

aenigmaticum Airy-Shaw (Euphorbiaceae). In other areas, however, as at Kinabalu, the soil 

is a friable red-brown loam and the flora bears affinities with that found on basalt. 

The mangroves of Sabah and Sarawak do not compare in extent with those of the 

Peninsular west coast, but are noteworthy because they are far richer in species than any 

others in the world. The most extensive are at the mouths of the Kinabatangan River, and 

around the Bay of Brunei where some magnificent primary stands still survive. The floristic 

formations are similar to those of the Peninsula, as in most respects are those of the inland 

river banks and alluvium. 

Altitudinal zonation 

The sharp topography and bold ridge lines of the mainly sedimentary mountains of 

Sabah and Sarawak cast a stamp on vegetation differentiation which obscures the 

zonation recognisable in the Peninsula. Below 1,200 m the primary differentiation is 

between ridge and slope forests (cf. Ashton 1964: 66-70). Lower, broader shale ridges bear 

the tallest forests, but those with the fewest endemics. Higher ridges are variably supported 

by sandstone ·strata: Where the soils are deep, a variety of MDF on shallow humult ultisols 

or yellow clay types will be present; Dryobalanops beccarii, Shorea laevis and Shorea 

multiflora are characteristic species. The slopes bear S. parvifolia-type MDF with pockets 

of Dryobalanops lanceolata on gentle lower surfaces. On shallow soils, which increase on 

the ridges with the altitude, a distinct short-stature ridge forest is widespread between 800 

and 1,200 m, where Shoreaflaviflora, Vatica umbonata and V dulitensis are characteristic 

of dipterocarps, and Engelhardia spp. are common. Where sandstone is exposed, shortstature 

kerangas prevails, and this grades imperceptibly and continuously on sandstone 

substrates into "upper montane" ericaceous elfin forest; this retains many kerangas species, 

such as Eugenia (Syzygium) bankense and Calophyllum nodosum. Nevertheless, true upper. 

XLVIII


montane elements such as Weinmannia blumei, Leptospermumflavescens,Xanthomyrtus spp., 

Rhododendron quadrasianum, Drimys piperita, Dacrydium comosum and Phyllocladus 

hypophyllus can begin to appear as low as 600 m on the most exposed peaks and skeletal 

soils, but only become common above 1,300 m. 

On the slopes at about 1,300 m as in the Peninsula, there is an ill-defined transition on 

clay-rich soils to oak-laurel dominated "lower montane forest" that is at approximately the 

same altitude at which the upper montane elements become common on the sandstone 

ridges. Again, it is the differentiation between ridge and slope, and between soils with and 

without a surface organic horizon, which primarily correlate with forest type differentiation, 

although the slopes adjacent to exposed summit ridges do become clothed in organic soils 

and ericaceous forest. 

Historical biogeography 

This Tree Flora will for the first time provide a sound basis for historical 

biogeographic analysis of the northern Borneo flora. For the present, two provinces 

are recognised, Northwest Borneo from Pontianak in West Kalimantan northeast to 

the Crocker Range and Kota Kinabalu, and Sabah north and east of the Cracker Range. 

The Northwest Borneo Province has phytogeographic connections with the Riau Archipelago 

and eastern coastal Peninsular Malaysia, that is the Rim~ Pocket of Corner (Corner 1960, 

Ashton 1992). Within it, two subprovinces are recognisable, being apparently divided east 

and west of the Lupar Valleys in west Sarawak and the Kapuas Lakes in Kalimantan 

(Ashton 1972, 1992). The western subprovince contains a number of endemics, and also a 

number of Peninsular Malaysian species not found elsewhere in Borneo such as the 

dipterocarps Shorea dmyphylla, S. dealbata, S. Jalcifera, and S. resinosa, and, in this first 

volume of the Tree Flora, Dacryodes rubiginosa, Glyptopetalum quadrangulare, and 

Lophopetalum pachyphyllum. The many endemic species include the dipterocarps 

Dryobalanops Jusca, Shorea alutacea, S. bakoensis, S. cuspidata, S. elliptica, S. 

induplicata, S. lunduensis, S. pallidifolia, S. richetia, S. splendida, S. stenoptera, S. 

subcylindrica, Vatica compressa, V pedicellata and, in this volume, Alangium circulare, 

Haplolobus beccarii, H. inaequifolius, Anisophyllea rhomboidea, Ellipanthus beccarii var. 

beccarii, Mastixia glauca, Schuurmansiella angustifolia, Sarcotheca macrophylla, 

Glycosmis longisepala and Maclurodendron parviflorum. Likewise, a number of species of 

the Province are absent from this subprovince including, surprisingly, Dryobalanops 

aromatica and Shorea curtisii. 

Ecological plots provide evidence that the habitats which occur in ecological islands, 

notably the areas of the organic podsols and humult ultisols so characteristic of this 

Province but also the areas of limestone and ultramafic rocks, experience significani 

between-locality variation in the characters of many species, and also the apparent 

fortuitous absence of species from suitable localities. The lack, to date, of any record of the 

usually common humult ultisol species Upuna borneensis from the Lambir Hills is a 

striking example. 

The Northeastern Province is not so clearly subdivided, but it includes two habitats unique 

for all Borneo. Kinabalu, owing to its great height, is the home both of many endemics and, 

XLIX


on a smaller scale presumably owing to its youth, a regionally widespread upper montane 

and alpine element. The extensive ultramafic extrusions support a number of endemics, 

both in the lowlands and also on Kinabalu to which such notable species as Scaevola 

micrantha, S. chanii and Piltosporum linearifolium are confined. But the Province is 

notable in addition for its lowland endemic flora, which includes the dipterocarps 

Dryobalanops keithii, Hopea badiifolia, Parashorea tomentella, Shorea symingtonii, S. 

waltonii, all of S. parvifolia type MDF, and, in this volume, Dacryodes elmeri, Kokoona 

sabahana, Microtropis sabahensis, Parinari argento-sericea, Connarus agamae, Atuna 

cordata, Sarcotheca rubrinervis, Maclurodendron pubescens, Melicope jugosa, M sororioa, 

M subunifoliolata, Monanthocitrus oblanceolata, and Turpinia nitida. Also, a number of 

Philippine species including Dipterocarpus validus, Protium connarifolium, Pitlosporum 

resiniferum, Melicope bonwickii, M denhamii, Turpinia borneensis and Scaevola 

micrantha, and species such as Canarium asperum, C. decumanum and Kibara obtusa are 

only known from this part of Borneo. Several of these are confined to ultramafic 

substrates. 

The southward extent of this Province into East Kalimantan is as yet inadequately 

understood. There is limited ultramafic rock in Kalimantan but some species of the zonal 

clay soils, such as D. keithii, hardly reach Tidung while others, such as H. badiifolia, 

extend as far as Balikpapan. This variable extent of ranges is not surprising in the Shorea 

parvifolia type MDF on clay soils, which extends continuously over the hills of East and 

Central Borneo. A number of other Philippine species such, for instance, as the 

dipterocarps Parashorea malaanonan, Shorea almon and S. falciferoides, also extend 

westward into north and central Sarawak. 

Lastly, there is an element in the Shorea parvifolia type MDF, represented by such species 

as Dipterocarpus mundus, D. pachyphyllus, Hopea bullatifolia, H centipeda, H dasyrrhachis, 

H jluvialis, H megacarpa, Shorea agamii ssp. diminuta, S. asahi, S. collaris, S. iliasii, Vatica 

endertii and V granulata, which have been found from innermost lowland Sarawak across to 

the DIu Barito, and which may eventually delineate a distinct central Bornean transmontane 

floristic province. 

References 

Anderson, J.A.R. 1963. The flora of the peat swamp forests of Sarawak and Brunei 

including a catalogue of all recorded species of flowering plants, ferns, and fern allies. 

Gard. Bull. Sing. 20: 131-228. 

Ashton, P.S. 1964. Ecological Studies in the Mixed Dipterocarp Forests of Brunei State. 

Oxford Forestry Memoirs 25. 

Ashton, P.S. 1972. The quarternary geomorphological history of western Malesia and 

lowland forest phytogeography. In P. & M. Ashton (eds.), Hull Geog. Dept. Misc. Series 

13. Transactions of the second Aberdeen-Hull symposium on Malesian ecology: The 

quarternary era in Malesia. Pp. 35-49 

L


Ashton, P.S. 1992. Plant conservation in the Malaysiau region. In S.K. Yap & S.W. Lee 

(eds.), In Harmony with Nature. Proceedings of the International Conference of Tropical 

Biodiversity. Kuala Lumpur: Malayan Nature Society. Pp. 86-93. 

Brunig, E.F. 1974. Ecological Studies of the Kerangas Forests of Sarawak and Brunei. 

Kuching: Borneo Literature .Bureau. 

Corner, E.lH. 1960. The Malayan flora. In RD. Purchon (ed.), Proceedings of the 

Centenary and Bicentenary Congress of Biology, Singapore. Pp. 21-24. 

Gobbett, D.l & C.S. Hutchinson (eds.). 1973. Geology ofthe Malay Peninsula. New York: 

Wiley. 

Liechti, P., F.W. Roe & N.S. Haile. 1960. The Geology of Sarawak, Brunei and the 

Western Part of North Borneo. British Borneo Geological Survey Department, Bulletin 3. 

Richardson, lA. 1947. An outline of the geomorphological evolution of British Malaya. 

Geol. Mag. 84: 129-144. 

Steenis, C.G.G.l van. 1957. Specific and infraspecific delimitation. Flora Malesiana 1, 5, 

1: CLXVII -CCXXIX. 

Symington, C.F. 1943. Foresters' Manual of Dipterocarps. Malayan Forest Records 16. 

Whitmore, T.c. 1984. Tropical Rain Forests of the Far East. (2nd edition). Oxford: 

Clarendon. . 

LI


ACERACEAE 

A. Noorsiha 



Koorders & Valeton, Bijdr. Booms. Java 9 (1903) 252; Bloembergen, FM 1,4 (1948) 3; Backer & 

BakhuizenJ, FJ 2 (1965) 143; Keng, OFMSP (1969) 416; Whitmore, TFM 2 (1973) 1; Anderson, 

CLTS (1980) 133; Cockburn, TS 2 (1980) 14; Ashton, 1v1NDTS 2 (1988) 1; Whitmore, Tantra & 

Sutisna, CLK 1 (1989) 11. 

Trees or shrubs. Buds with protective scales. Leaves opposite-decussate, simple, palmately 

or pinnately veined, without stipules. Flowers in fascicles, panicles, racemes or corymbs, 

regular, often unisexual; sepals and petals 4-5, rarely without petals; disc annular or 

lobed, or reduced to teeth, rarely absent, intra- or extra-staminal; stamens (or staminodes) 

(4-)8(-10), hypogynous or perigynous, filaments free; ovary often present in rudimentary 

form in male flower, in female flowers 2-locular, superior, compressed, styles 2, free or 

connate at base, ovules 2 in each locule, placentation axile. Fruit composed of 2 oneseeded, 

one-winged samaras. Seeds without endosperm; embryo with folded cotyledons. 

Distribution. Two genera (Acer and Dipteronia) with about 200-250 species distributed 

mainly in mixed deciduous forests of the northern temperate zone. Only one genus (Acer) 

with one species in western Malesia. 

Taxonomy. Closely allied to the Sapindaceae, from which it can be distinguished by its 

opposite/decussate simple leaves, samara-type fruit, and seed without endosperm. 

ACERL. 

(Latin name of the maple tree) 

Sp. PI. 1 (1753) 1054; Koorders & Valeton I.e. 253; B10embergen I.e. 3; Backer & BakhuizenJ I.e. 

143; Whitmore I.e. 1; Anderson I.e. 133; Cockbum I.e. 14; Ashton I.e. 1; Whitmore, Tantra & 

Sutisna I. e. 11. 

Deciduous trees or shrubs with smooth bark. Leaves pinnately veined, more or less 

distinctly 3-veined at the base, long-stalked. Flowers unisexual, male and female on the 

same or different trees, organised in terminal or axillary racemes, corymbs, or panicles; 

sepals and petals imbricate in buds. Other floral, fruit and seed characters as in the family. 

Distribution. As for the family. 

Ecology. The samaras are wind-dispersed. 

1


1'·0 

] 2cm 

, .. [ 

}cm 

B 

Fig. 1. Acer laurinum. A, leafy twig; B, leaf venation; C, terminal vegetative bud; D, inflorescence; 

E, male flower; F, female flower; G, fruit. (A-C from SAN 36157. D from Haviland 2092, E-F from 

Forman 106, G from SAN 124720.) 

2

ACERACEAE (NOORSIHA) 

Uses. Many temperate species yield valuable timbers and a few others produce sugar/maple 

syrup obtained by boring holes through the bark in February and March each year. A 

number of shrubby species make excellent ornamental plants because of their strikingly 

coloured (yellow, red and purple) aging foliage. The timber of the Malesian species is only 

occasionally used for light construction. 

Taxonomy. In a number of earlier publications (e.g., Blume, Rumphia 3 (1847) 193; 

Miquel, FI. Ind. Bat. 1,2(1859) 581; Bentham & Hookerj, Gen. PI. 1 (1867) 409; Hooker 

j, Fl. Brit:lnd. 1 (1875) 692) the genus was included in the Sapindaceae. 

Acer laurinum Hassk. Fig. 1. 

(Latin, laurinum = resembling Laurus; the blue-grey colour of the leaves) 

in Hoeven & de Vriese, Tijd. Nat. Gesch. & Phys. 10 (1843) 138; van Steenis, FM 1,4 (1954) 592; 

Backer & BakhuizenJ I.c. 143; Anderson I.c. 133; Cockbum l.c. 14; Delc;:ndick, Reinwardtia 9, 4 

(1980)395; Ashton I.c. 1; Whitmore, Tantra & Sutisna l.c. 1l. Type: Junghuhn, s.n., West Java, 

Megamendong (holotype L; isotype BO). Synonyms: A. javanicum Jungh. apud Hoeven & de Vriese, 

Monatsber. Verh. Ges. Erdk. Berlin 3 (1842) 96; A. niveum Blume, Jaarb. Kon. Ned Maatsch. Aanm. 

Tuin (1844) 84;A. caesiaefolium Blume, Rumphia 3 (1847) 193; A. philippinum Merr., Philip. Govt. 

Lab. Bull. 35 (1909) 36; A. curranii Merr., Philip. J. Sc. 4 (1909) 285; A. caesium (Reinw. ex 

Blume) Kosterm., Reinwardtia 7 (1965) 141 (based on Laurus caesia Reinw. ex Blume, Bijdr. 

(1826) 553); non A. caesium Wall. ex Brandis, For. Fl. NW. & C. Ind. (1874) 3). 

Medium-sized to large tree up to 50 ni tall; clear bole straight, cylindrical, up to 28 m tall 

and 70 cm diameter; buttresses about 1.5 m high and 2 m out, thin, spreading, slightly 

concave; crown dense, pale fresh green from below, more or less conical to hemispherical, 

deciduous to semi-evergreen. Bark initially smooth, greenish grey, becoming rich red to 

grey-brown and longitudinally fissured and flaky with age; inner bark c. 1.5 cm thick, 

firmly fibrous, yellow-brown to red-brown, somewhat mottled, sometimes laminated. 

Heartwood absent; sapwoodwhite to pale yellow, with rather distinct concentric growth 

rings. Twigs slender, terete or ribbed, drying nearly black, with numerous leaf-scars and 

minute lenticels. Buds numerous, c. 4 mm long, covered by 4-11 pairs of decussate, 

caducous scales of c. 2 mm long. Leaves thinly leathery, glabrous, elliptic to ovatelanceolate, 

7-23 x 3-6 cm, glaucous to grey beneath, upper surface dark green when fresh, 

drying dull red-brown; base rounded to obtuse, margin entire, apex acute to acuminatecaudate; 

midrib slender, slightly raised above, more or less prominent beneath; lateral veins 

4-9 pairs, very slender, with a pair at the base of the midrib; intercostal veins fine, 

reticulate; stalk 2-7.5 cm long, slender. Inflorescences in the axils of fallen leaves, 

paniculate or corymbose, 3-10 cm long, 30-50-flowered; bracts and bracteoles fairly welldeveloped. 

Male flowers: sepals and petals (3-)5, free, obliquely erect, respectively 2.5-3 

and 1.5-2.5 mm long; stamens (4-)6(-8), arranged in one whorl on a flat, glabrous to 

woolly disc, filaments c. 5 mm long, anthers c. 1 mm long; pistillode strongly reduced. 

Female flowers with sepals and petals as in the males; staminodes strongly reduced; ovary 

densely hairy, c. 2 mm across; styles 2, c. 1.5 mm long, stigmas 2, sessile. Fruits red, wings 4- 

7.5 x 1-2.5 cm, asymmetrical, narrowed towards the base, pubescent. Young seedling 

leaves coarsely and distantly toothed; sapling leaves whorled. 

Vernacular name. Sarawak-perdu (Malay). 

3


Distribution. Burma, possibly Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the 

Phillipines, Celebes and Lesser Sunda Islands. Rare in truly non-seasonal parts of 

Peninsular Malaysia, Sumatra and Borneo, but relatively common elsewhere. In Sabah 

uncommon, known only from 2 collections from Mt. Kinabalu (SAN 38438) and Nabawan 

(SAN 124720). In Sarawak, it has a very local distribution, but is frequent where it occurs, 

e.g., in the Hose Mts. (3rd Div.), Mt. Meluku (2nd Div.), and Usun Apau Highlands (4th 

Div.). 

Ecology. Apparently confined to soils of relatively high nutrient status, on igneous rocks at 

200-1500 m in the upper limits of Mixed Dipterocarp Forest and on granodiorite rocks in 

the oak-laurel lower montane forests around 1200-1600 m. Flowering has been recorded in 

April-August and fruiting in July-November. In Sabah, fruiting has been rarely recorded. 

4


ALANGIACEAE 

A. Berhaman 




Merrill, PEB (1929) 232; Masamune, EPB (1942) 517; Backer & BakhuizenJ, FJ 2 (1965) 159; 

Eyde, J. Am. Arb. 49 (1968) 167; Kochummen, TFM 1 (1972) 56; Anderson, CLTS (1980) 133; 

Cockburn, TS 2 (1980) 15; Ashton, MNDTS 2 (1988) 4; Whitmore, Tantra & Sutisna, CLK 1 (1989) 

1l. 

The family consists of only one genus, Alangium, distributed in tropical Africa, and E Asia, 

Malesia, eastern Australia and some South Pacific Islands. 

Taxonomy. Alangium has in the past been included in the Cornaceae. Eyde I.e. investigating 

the anatomy of the flower and discussing the relationships of AI€lngium, concluded that this 

genus was not closely related to the Cornaceae. Instead, he suggested that the alkaloid 

characteristics and floral morphology indicate that Alangium has its closest relatives in the 

Rubiaceae. The evidences have not been further examined and the most recent treatments 

accept the Alangiaceae as a monogeneric family, usually placed in the Cornales. 

ALANGIUM Lam. 

(after the Malabar plant name, alangi) 

kondolon (Dusun, Sabah), midong (lban, Sarawak) 

Encycl. Meth. Bot. 1 (1783) 174; Merrill, EB (1921) 459 (as part of Comaceae), I.e. (1929) 232; 

Bloembergen, Blumea 1,2 (1935) 241, Bull. Jard. Bot. Btzg. 3, 16 (1939) 139; Masamune I.e. 517; 

Kochummen I.e. 56; Anderson I.e. 133; Cockburn I.e. 15; Ashton I.e. 4; Whitmore, Tantra & Sutisna 

I.e. 12; Berhaman, Sandakania 4 (1994) 3l. 

Trees, more rarely shrubs or woody climbers, often with thin buttresses or stilt-roots. Bark 

smooth, thin, grey-brown, often with pale lichen patches; inner bark thick, yellowish 

brown. Leaves simple, alternate, entire, pinnately veined, or sometimes 3-5-veined at the 

base; stalk terete or slightly grooved or flattened on upper side; stipules none. Inflorescence 

an axillary eyme, few- to many-jlowered. Flowers bisexual, sessile or stalked (the 

stalks, if present, articulate); calyx-tube connate with the ovary wall, truncate or with 4-10 

teeth or lobes; petals 4-10, valvate, linear, becoming recurved, alternate with the calyxteeth, 

usually white; stamens as many as or 2-6-times the number of petals; anthers linear, 

dehiscing laterally or introrsely; disc intrastaminal, well-developed; ovary inferior, 1-2- 

5

TREE FLORA OF SABAH AND SARAWAK VOL. \ (\995) 

eelled, style single, usually as long as the corolla, ovule 1 in each locule, anatropous. Fruit 

a drupe, globose to ellipsoid and somewhat bilaterally compressed, often longitudinally 

ribbed, crowned by the persistent calyx-teeth and disc. Seed 1-2 per fruit with a copious 

endosperm; cotyledons foliaceous, flat, palmately veined at base; radicle straight. 

Distribution. 21 species, distributed as noted for the family. 10 species in Sabah and 

Sarawak, including A. kurzii Craib, a new record for Borneo. 

Ecology. Primary and secondary forests, lowlands to 1500 m. 

Taxonomy. Bloembergen I.e. (1939) recognised four sections, Angolam Baill. (which Eyde 

I.e. points out should be called section Alangium as it contains the type species), Marlea 

Baill., Rhytidandra Baill., and Conostigma Bloemb.; all except Rhytidandra are represented 

in Sabah and Sarawak. Eyde I.e. has found that differences in style and stigma structure, 

stamen number, the pattern of floral vasculature, pollen morphology, and fruit endocarp 

characteristics support B10embergen's designation of the four sections. 

Key to Alangium species 

1. Woody climbers ...................................................................................................................... 2 

2. Leaf coriaceous with domatia in the axils of main veins. Inflorescence almost sessile, 

with less than 15 flowers .............................................................................................. . 

Alangium sp. 1. 

Woody climber. Leaves obovate-elliptic, 12-15 x 4.5-6 cm, coriaceous; base 3- 

veined, cuneate; apex acuminate; lateral veins 4-5 pairs; stalk 1.2-1.5 cm long, 

glabrous. 

Only one specimen, SAN 79310, from Telupid in Sabah known, with young flower 

buds. 

Leaf thinly chartaceous, without domatia. Inflorescence with stalk up to 4 cm long, and 

17-30 flowers .......................................................................................... . 

A. scandens Bloemb. 

I.e. (1935) 264, I.e. (1939) 193; Masamune I.e. 518; Whitmore, Tantra & Sutisna I.e. 12; 

Berhaman I.e. 32. Type: Endert 4076, Sarawak, 4th Division, Long Petah (lectotype BO). 

Woody climber. Leaves ovate to ovate-elliptic, 8-15.5 x 3.5-6 cm, chartaceous to 

thinly coriaceous; base subcordate to rounded, apex acuminate; lateral veins 6-7 

pairs; stalk hairy, 1-1.5 cm long. 

Sumatra and Borneo. Hill to lower montane forest, common in Sabah. and 

Sarawak. 

3. Leafbase 3-5-veined ..................................... ................... .4 

Leaf base pinnately veined .......................... ... 9 

4. Leaves markedly obovate with narrowed base, or if elliptic and base rounded then 

lateral veins 6-7 pairs ........................................................................ 6. A .. longiflorum 

6

ALANGIACEAE (BERHAMAN) 

Leaves rounded to broadly ovate with rounded to cordate base, or if (rarely) elliptic and 

base narrowed then lateral veins 4-5 pairs ................................................................... 5 

5. Leaf length at least twice the width ................. . ...................... 2. A. griffithii 

Leaf length not reaching twice the width ....... . ......... 6 

6. Leaf-stalk densely hairy; lateral veins 9-12 pairs. Inflorescence-stalk less than 8 mm 

long. Flowers almost se~sile ........................................................................ 7. A. nobile 

Leaf-stalk glabrous to sparsely hairy; lateral veins less than 8 pairs. Inflorescence-stalk 

more than 10 mn: long. Flowers distinctly stalked ...................................................... 7 

7. Leaf-apex blunt to emarginate; lateral veins 4-5 pairs. Petals c. 28 mm long and 5 mm 

wide at base, covered by tiny stellate scales ............................................ 1. A. circulare 

Leaf-apex acute to acuminate; lateral veins 5-7 pairs. Petals less than 15 mm long and 

1-2 mm wide at base, covered by straight hairs .......................................................... 8 

8. Leaf lower surface velvety (seldom sparsely) hairy; midrib and lateral veins on upper 

surface densely hairy. Calyx-tube densely hairy, teeth to only about 0.5 mm long 

.................................................................................. 5. A. kurzii 

Leaf lower surface glabrous to sparsely hairy, never velvety; midrib and lateral veins 

on upper surface glabrous. Calyx-tube sparsely hairy, teeth more than 1 mm long 

....................................................................................................... 8. A. rotundifolium 

9. Plants of mixed swamp forest. Leaf-base typically asymmetric. Calyx-limb with distinct 

triangular teeth. Style thickly yellow hairy. Stamen filaments broadened and thickly 

woolly hairy at base .............................................................................. 3. A. havilandii 

Plants of lowland mixed dipterocarp forest. Leaf-base typically symmetric (rarely asyIp. 

metric). Calyx-limb subtruncate. Style sparsely white-hairy. Stamen filaments uniformly 

thick throughout, subglabrous to sparsely short-hairy at base ........... .4. A. javanicum 

1. Alangium circulare Stone & Kochummen 

(Latin, circularis = circular; the round-shaped leaves) 

Blumea 22 (1975) 219; Ashton I.e. 4. Type: Salleh ak Nantah S. 24325, Sarawak, 1st Division, 

Bukit Siol, Kuching (holotype SAR; isotypes A, BO, K, KEP, L, SAN, SING). 

Small to medium-sized tree to 15 m tall. Bark smooth, grey-brown; inn.er bark pale brown. 

Sapwood pale yellow. Twigs grey to brown. Leaves broadly ovate-elliptic-obovate to 

subcircular, 8.5-10.5 x 8-10 cm, coriaceous; base rounded to cordate, 3-veined, apex 

blunt to emarginate; midrib raised on both surfaces; lateral veins 4-5 pairs, slightly raised 

on both surfaces, glabrous; stalk glabrous, 7-10 mm long, 2-3 mm diameter, grooved on 

upper surface. Inflorescence a short cyme, 1-2-times branched, stalk 11-14 mm long, 

finely greyish pubescent, 1-3-flowered. Flowers 5-merous, subsessile to shortly stalked, 

stalks 2-3 mm long; calyx funnel-shaped, 3-4 mm long, 3 mm across, finely stellatepubescent, 

teeth 5, triangular, to 1 mm high; petals linear-lanceolate, c. 28 mm long, c. 5 

mm wide at base, abaxial side covered with minute stetlate hairs; stamens as many as 

petals, linear, 10-11 mm long; ovary l-celled, style hairy, 22-26 mm long. Fruit and seed 

unknown. 

7


Distribution. Endemic to Sarawak, recorded from Bukit Siol, Sempadi Forest Reserve and 

Gunung Pueh in the 1st Division. Apparently a species of kerangas forest. 

2. Alangium griffithii (Clarke) Harms 

(W. Griffith, 1810-1845, doctor and botanist in India and Malacca) 

in Engl. & Prantl., Pfl. Fam. 3, 8 (1898) 262; Bloembergen I.e. (1935) 266, I.e. (1939) 194; 

Masamune I.e. 517; Kochummen I.e. 58; Cockbum I.e. 16; Anderson I.e. 134; Ashton I.e. 5; 

Whitmore, Tantra & Sutisna I.e. 11; Berhaman I.e. 32. Basionym: Marlea grifJithii C1arke in Hooker 

J, Fl. Brit. Ind. 2 (1879) 742. Type: Griffith 3387, Malacca (lectotype K; isolectotypes B, BM). 

Synonym: Marlea densiflora Koord. & Va1eton, Bijdr. Booms. Java 5 (1899) 84. 

Medium-sized tree to 20 m tall, 30 cm diameter, sometimes with spreading buttresses to 50 

cm wide. Bark smooth, grey to dark brown; inner bark pale yellow. Sapwood white to pale 

yellow. Twigs dark brown, smooth to hairy. Leaves chartaceous, elliptic to narrowly ovate, 

5.5-18.5 x 2-8 cm, length at least twice the width, hairy on the lower surface, mainly on 

the main veins, grey to dark brown when dry; base cuneate, asymmetric, 3-5-veined, apex 

acuminate; midrib flat on upper side, hairy on the lower surface; lateral veins 4-5(-6) 

pairs, usually glabrous between the veins on upper surface; stalk 5-15 mm long, sparse to 

densely hairy. Inflorescence hairy, 3-4-times branched, 3-6.5 cm long, many-flowered, 

stalk 1-2.2 cm long. Flowers 5-merous, white to cream, fragrant; calyx-tube densely hairy, 

with teeth 0.25-0.5 mm long: limb spreading; petals linear, swollen at base, glabrous to 

sparsely hairy outside, cream, 8.5-15 mm long; stamens as many as petals, 8-14 mm long; 

ovary l-celled, style glabrous, cream, 8-11 mm long. Fruit ovoid, 14-21 x 8-10 mm, dark 

brown when dry, faintly grooved, crowned by the persistent calyx-limb, dark blue when ripe. 

Vernacular names. Sabah-gadong hutan (Brunei Malay). Brunei--mayam kampong 

(Malay). 

Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. In 

Sabah, quite common but in Sarawak only found in the Tinjar Forest Reserve. Also in 

Brunei and Kalimantan. 

Ecology. Primary and secondary forest, 60-400 m. 

3. Alangium havilandii Bloemb. 

(lD. Haviland, 1857-190 I, first Sarawak Medical Officer and plant and insect collector) 

I.e. (1935) 277, I.e. (1939) 213; Masamune I.e. 518; Anderson I.e. 134; Ashton I.e. 8; Whitmore, 

Tantra & Sutisna I.e. 1l. Type: Omar 54, Sarawak, 1st Division, Gunung Sedi1u Forest Reserve 

(ho1otype SING). 

Small to medium-sized tree to 25 m tall, 30 cm diameter, with tall prominent thin flying 

buttresses. Bark smooth, greyish; inner bark yellow. Sapwood pale whitish to yellowish. 

Leaves ovate to elliptic, 5-15 x 3.5-6 cm, chartaceous or thinly coriaceous; base typically 

asymmetric, rarely symmetric, pinnately veined, apex long acuminate to acute; midrib flat 

8


on tht.e upper surface; lateral veins, 6-9 pairs, raised on the lower surface; stalk 5-12 mm 

long, slender. Inflorescence a short cyme, 1-2-times brariched, 1-6-flowered, stalk 1-1. 5 

cm long. Flowers 4-5-merous; calyx-tube tomentose, c. 2 mm long, limb with distinct 

triangular teeth; petals 15-20 mm long, prominently dilated at base; stamens as many as 

petals., 14-l7 mm long, filaments broadened and thickly woolly hairy at base; ovary 1- 

celled,style thickly yellow hairy, 11-13 mm long. Fruit ellipsoid-ovoid, finely hairy, 10.5- 

18 x 6-9 mm, flattened when dried, crowned by the persistent calyx-limb, ripening pink. 

Vernacular names. Sarawak---dadam or jadam paya (Melanau), jenangan (Melanau, 

Bintulu), sisit (Malay). 

Distri bution. Sarawak, Brunei and Kalimantan, not yet recorded in Sabah. 

Ecolo-gy. Frequent and locally common in mixed peat swamp forest at low altitude. 

4. Atangium javanicum (Blume) Wangerin 

(ofJava) 

Fig.!. 

in Engl & Prantl., Pfl. Fam. 4, 220b (1910) 14; Bloembergen l.c. (1935) 281, I.c.(l939) 218; 

Merrill I.c. (1921) 458, I.c. (1929) 232, in syn.; Masamune I.c. 518; Cockbum I.c. 16, in syn.; 

KochUIl1men l.c. 57, in syn.; Ashton l.c. 10; Anderson I.c. 133, in syn.; Whitmore, Tantra & Sutisna 

I.c. 12; Berhaman I.c. 33. Basionym: Styraxjavanicum Blume, Bijdr. 13 (1825) 671. Type: Blume, 

s.n., Ja va (ho1otype BO; isotype NY). 

Medium-sized tree to 30 m tall and 35 cm diameter, often with flying buttresses. Bark 

smooth, yellowish to pale brown; inner bark yellowish to reddish. Sapwood yellowish. 

Leaves ovate-elliptic to obovate, (8-)15(-35) x (2.5-)5.5(-15.5) cm, chartaceous to coriaceous, 

lower surface glabrous to short-hairy on the midrib, or sometimes short velvety all over, 

drying pale olive-brown to purplish brown; base cuneate to rounded, symmetric (rarely 

asymmetric), pinnately veined, apex acute to acuminate; midrib flat to raised, rarely sunken 

on the upper surface; lateral veins 6-9(-19) pairs; stalk 0.5-3.8 cm long, slender, 

sometimes grooved on the upper surface and densely hairy. Inflorescence 1-3-times 

branched, with up to 35 flowers, stalk 2-8 mm long. Flowers (4-)6(-7)-merous, 2-2.5 cm 

long; calyx-tube campanulate, limb sub truncate, glabrous to densely covered by silky long 

hairs; corolla usually swollen at base, densely covered by long silky hairs or densely or 

sparsely stellate-hairy; petals 8.5-17 mm long, stamens as many as petals, filaments 

uniformly thick throughout, subglabrous to sparsely short-hairy at base; ovary 1-celled, 

style 4-10 mm long, sparsely white-hairy. Fruit ellipsoid-ovoid, variable in size, smooth to 

strongly ridged, crowned by the persistent calyx, 1-5 mm high and 2-10 mm wide. 

Vernacular names. Sabah-satu inchi (colloquial Malay). Sarawak-jadam (Malay). 

Key to varieties 

1. Corolla densely covered by silky long hairs (velutinous). Lower side of leaf short-hairy 

on midrib only .............................................................................................................. . 

9


var. javanicum 

Synonyms: A. bogoriense Wangerin, Fedde. Repert. 4 (1907) 338; A. bomeense Merr., J. 

Str. Br. R. As. Soc. 86 (1922) 10; A. javanieum "form E" of Ashton I.e. 11; A. ebenaeeum 

"var. COl of Cock bum I.e. 18. 

Sumatra, Java and Borneo. Common in all districts in Sabah and Sarawak. 

Primary mixed dipterocarp and secondary forest. 

Corolla sparsely to densely covered with stellate hairs, not long-hairy. Lower side of 

leaf glabrous, or sometimes short-velvety all over.. ...................................................... 2 

2. Leaves drying pale olive-brown, typically smaller, 8-12 cm long, but exceptionally 

reaching 27 cm long. Calyx-limb of mature fruit smaller, 1-2 mm high and 2-5 mm 

wide .................................................................................. . 

var. meyeri (Merr.) Berhaman 

I.e. 34. Basionym: A. meyeri Merr., Publ. Govern. Labor. 35 (1906) 54. Type: Meyer F.E. 

2284, Phillipines, Luzon, Cagayan Province (lectotype BO; isolectotypes B, K, NY, S). 

Synonyms: A. tutela Ridl., J. Str. Br. R. As. Soc. 61 (1912) 10; A. ebenaeeum var. tutela 

(Ridl.) Kochummen, Fed. Mus. J. 13 (1970) 133; A. javanieum "form COl (pro parte) & 

"form D" of Ashton I.e. 11; A. ebenaeeum "var. E" (pro parte), "var. D", "var. E", and "var. 

G" of Cock bum I.e. 18 (pro parte). 

Peninsular Malaysia, Java, Borneo, and the Philippines. Very common in all districts 

in Sabah and Sarawak, in primary and secondary mixed dipterocarp forest. 

Leaves drying purplish brown, (at least markedly so on the lower side), typically longer 

or larger (24-39 cm long). Calyx-limb of mature fruit larger, 2-5 mm high and 3-10 

mm wide ......................................................... 

var. ebenaceum (Clarke) Berhaman 

I.e. 33. Basionym: Marlea ebenaeea Clarke in Hooker /, Fl. Brit. Ind. 2 (1879) 742. Type: 

Griffith 3383, Malacca (lectotype K). Synonyms: A. ebenaeeum (Clarke) Harms in Engl. & 

Prantl I.e. (1898) 262; A. ridleyi sensu Ashton I.e. 14, non King (1902); A. mezianum 

Wangerin, I.e. 338; A. sessiliflorum Merr. I.e. (1929) 232; A. javanieum "form COl of 

Ashton, I.e. 11 (pro parte); A. ebenaeeum "var. E" (pro parte) and "var. G" of Cock bum I.e. 

18 (pro parte). 

Peninsular Malaysia, Java, Borneo, and the Philippines. Common in all districts in 

Sabah and Sarawak. All these forms appear to be confined to clay-rich soils in 

mixed dipterocarp and associated secondary forest. 

5. Alangium kurzii Craib 

(W.S. Kurz, 1834-1878, German soldier and naturalist, Bogor and Calcutta) 

Kew Bull. (1911) 60; Bloembergen I.e. (1935) 262, I.e. (1939) 183; Kochummen I.e. 58; Berhaman 

I.e. 3l. Type: Kerr 1172, Thailand, Chiangmai, Doi Sutep (holotype K). Synonyms: Alangium 

ehinensis var. tomentosum Merr., Philip. J. Sci., 2l, 5 (1922) 505; Alangium begoniaefolium Ridl., 

FMP 1 (1922) 894. 

Small tree to 15 m tall, 20 cm diameter, with short buttresses. Bark smooth, dark grey, 

lenticellate; inner bark orange and cream mottled. Sapwood soft, pale brown. Leaves thin 

coriaceous to chartaceous, lower surface velvety (seldom sparsely) hairy, ovate to broadly 

ovate, 8-20 x 5-10.5 cm; base markedly asymmetric, broadly rounded to cordate, 3-5- 

veined, apex acuminate-acute; midrib and major veins on upper surface densely hairy; 

10


r(I~, 

\. /}~ fl'J 

. 

1& 

r /) 

·~.0'-[ ~r.¥ 

,j 6 .. ··. 

l~1 

A 

t\ I,> () .. A.~ 

]2cm 

'q\r 

B \1/ 

., fl " 

\./ 

Fig. 1. Alangium javanicum. A-B, var. javanicum; C-E, var. ebenaceum; F-G, var meyeri. A, 

flowering leafy twig; B & D, fruits; C & F, fruiting leafy twigs; E & G, inflorescences. CA from SAN 

110170, B from SAN 39285, C & D from S. 34940, E from SAN 26087, F from SAN 82215, G from 

SAN 75771.) 

11


lateral veins 5-7 pairs; stalk 2.5-3 cm long, glabrous to sparsely hairy. Inflorescence 

hairy, 2-4-times branched with 3-15 flowers per branch, stalk 11-15 mm long. Flowers 

pale greenish to yellow creamy; calyx-tube densely hairy, l.25-2.5 mm long, teeth hairy, 

0.25-0.5 mm long; petals (5-)9(-10), 5.5-10 mm long, swollen at base for 2-3 mm, 

covered by straight hairs; stamens as many as petals, 5-10 mm long, densely covered with 

long straight hairs; ovary (l-)2-celled, style glabrous, 5-8 mm long. Fruit ellipsoid 1-l.5 x 

0.5-0.7 cm, crowned with the persistent calyx-limb. 

Vernacular name. Sabah--marapangi (Dusun Tambunan). 

Distribution. Myammar (Burma), China, lndo-China, Thailand, Sumatra, Peninsular 

Malaysia, Java, and Borneo. In Sabah, recorded only from Kota Marudu (SAN 99531), 

Ranau (SAN 62010) and Tambunan (SAN 11355). Not recorded for Sarawak . 

.Ecology. Lowland to submontane forest to 1300 m. Flowering from January to July; 

fruiting from May to August. 

6. Alangium longiflorum Merr. 

(Latin, longus = long,jlorum = flowers) 

Philip. J. Sci. Bot. 7 (1912) 319; B10embergen I.e. (1935) 253, I.e. (1939) 159; Cockburn I.e. 16; 

Anderson I.e. 134; Ashton I.e. 11; Whitmore, Tantra & Sutisna I.e. 12; Berhaman I.e. 32. Type: 

Darling FE 14773, Philippines, Ma1ueg, Cagayan Province (ho1otype L; isotype BO). Synonyms: 

Alangium salvifolium subsp. hexapetalum sensu MeIT. I.e. (1912) 321; A. lamarekii Thwaites, Enum. 

PI. ZeyI. (1850) 133;A. hirsutum B1oemb. I.e. (1939) 16l. 

Medium-sized tree to 20 m tall and 25 cm diameter. Bark smooth, dark brown; inner bark 

yellowish. Sapwood white. Leaves markedly obovate with narrowed base and 4-7 pairs of 

lateral veins or if elliptic and base rounded then with 6-7 pairs of lateral veins, velvety 

hairy on lower surface, 8-12 x 2.5-8 cm, chartaceous; base 3-veined, apex long-acuminate; 

midrib raised and sparsely hairy on the upper surface; stalk slender, hairy, 8-13 mm long. 

Inflorescence a short cyme with 1-5 flowers, densely hairy, almost sessile or with stalk to 

1.5 mm long. Flowers white, 30-50 mm long; calyx-tube campanulate, densely hairy, teeth 

0.25-0.5 mm long, limb 1-l.5 mm long; petals 5, white, 35-50 mm long, densely hairy 

outside, swollen at the base; stamens 30-50 mm long, 2-6-times as many as petals; ovary 

l-celled, style glabrous, 30-45 mm long. Fruit ellipsoid to ovoid in dry state, rounded at 

both ends, 20-25 cm long, crowned by the persistent calyx-limb, ripening pinkish. 

Distribution. Borneo and Philippines. Very uncommon in Sabah (e.g., SAN 30390, SAN 

83891) and Sarawak (e.g., S. 21704, S. 43342), also known in Kalimantan. 

Ecology. Primary mixed dipterocarp and associated secondary forests to 1200 m, on high 

nutrient clay soils. Flowering in March to July; fruiting in August to December. 

One specimen, SAN 14167 (Sabah, Ranau) bears spines, but the leaf characters are very 

close to that for A. longijlorum. 

12


7. Alangium nobile (Clarke) Harms 

(Latin, nobilis = distinguished, noble, probably referring to the growth habit) 

in Engl. & Prantll.e. (1898) 262; Bloembergen I.e. (1935) 275, I.e. (1939) 211; Masamune I.e. 518; 

Kochummen I.c. 58; Anderson I.e. 134; Ashton I.e. 12; \Vhitmore, Tantra & Sutisna I.e. 12; 

Berhaman I.e. 32. Basionym: Marlea nobilis C1arke in Hooker f I.e. 743; Rid1ey, Agr. Bull. Str. 

Settl. & Fed. Mal. St. 1 (1902) 181. Type: GrifJith 3385, Malacca (lectotype K; isolectotypes B, 

BM). 

Medium-sized tree to 20 m tall, 30 cm diameter. Bark smooth, brown; inner bark thin, 

yellow to pale brownish. Sapwood medium-hard, whitish. Leaves rounded to broadly 

ob ovate, 9-23 x 5-18 cm, chartaceous to coriaceous, upper surface hairy to glabrous, lower 

surface hairy to densely hairy; base rounded to cordate, 5-veined, apex rounded to acute, 

rarely acute to acuminate; midrib flattened to raised on the upper surface, densely hairy on 

the lower surface; lateral veins 9-12 pairs, raised and hairy below, flattened on the upper 

surface; stalk densely hairy, 2.S-4.S cm long and 3-S mm thick. Inflorescence a short 

cyme, stalk S-7 mm long, 1-2-branched with l-S flowers. Flowers 4-S-merous, almost 

sessile; calyx-tube'campanulate, hairy, lobes 5-7; petals swollen at base, 1O-1S mm long, 

densely hairy outside; stamens as many as petals, 9-14 mm long, with filaments broadened 

at base; ovary 1-celled, style hairy, 7-12 mm long. Fruit ellipsoid-ovoid, 25-30 x lS-20 

mm, grooved, velvety hairy, crowned by the persistent calyx-limb. 

Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak, uncommon 

in mixed dipterocarp forest (e.g., SAN 73205 and S. 38479). 

Ecology. Lowland to submontane forest to lS00 m. 

8. Alangium rotundifolium (Hassk.) Bloemb. 

(Latin, rotundus = rounded in outline,jolium = leaves) 

I.e. (1935) 258, I.e. (1939) 179; Masamune I.e. 518; Kochummen I.e. 60; Cockburn I.e. 16; 

\Vhitmore, Tantra & Sutisna I.e. 12; Berhaman I.e. 38. Basionym: Diaeaeearpium rotundifolium 

Hassk., Bonplandia 7 (1859) 172. Type: Sine coil., s.n., Java, probably from Tjibodas, Cult. Bogor 

Botanic Garden (ho10type BO; isotype L). Synonyms: Marlea rotundifolia (Hassk.) Teijsm. & Binn., 

Cat. PI. Hort. Bot. Bogor. (1866) 238; Alangium begoniifolium Harms in Engl. & Prantll.e (1898) 

261; Marlea begonifolia (Harms) Ridl., J. Fed. Malay. St. Mus., 8, 4 (1917) 44; Alangium 

rotundatum Ridl. ex Burkill & Henders., Gard. Bull. Str. Settl. 3 (1925) 380. 

Tree to 28 m tall, 40 cm diameter. Bark smooth to slightly scaly, white to pale grey; inner 

bark yellowish. Sapwood white to pale yellow. Twig dark brown to pale brown. Leaves 

glabrous to sparsely hairy on the lower surface, never velvety, rounded to broadly ovate or 

triangular-ovate, 9.5-22 x 7.S-14 cm, chartaceous; base rounded to cordate, sometimes 

cuneate, 3-5-veined, apex acuminate to acute; midrib and lateral veins on upper surface 

glabrous; lateral veins 5-7 pairs; stalk 2.5-3.5 cm long, glabrous to sparsely hairy. 

Inflorescence a cyme, 3-4-times branched with 4-1S flowers, covered by short hairs, stalk 

12-20 mm long. Flowers fragrant, white or cream to yellow; calyx-tube sparsely hairy, 

1. 5-3 mm long, limb flared with teeth 1-1. 5 mm long; corolla swollen at the base; petals 

13


white or cream, 6-14(-18) mm long; stamens as many as petals, 6-18 mm long; ovary 

usually l-celled, rarely 2-celled, style glabrous, 5-15 mm long. Fruit glabrous or thinly 

hairy, ovate-ellipsoid, cuneate or rounded at base, crowned by the flared persistent calyxlimb 

1-1.5 mm long, reddish when ripe. 

Vernacular name. Sabah-marapangi (Dusun Ranau). 

Distribution. Sumatra, Peninsular Malaysia, Java, and Borneo. In Sabah only recorded 

from Mt. Kinabalu (e.g., SAN 46772, SAN 48064) and Crocker Ranges (SAN 83987). Not 

recorded for Sarawak. 

Ecology. Primary and secondary forest on ultramafic soils, at 400-1600 m. 

Excluded species 

A. kinabaluense w.w. Srn., Not. Bot. Gard. Edinb. 8 (1915) 315; Merilll.c. (1921) 459, is 

based on a specimen collected from Mt. Kinabalu, Native Coli. 49 (E, K), which is 

Polyosma hookeri Stapf. 

14


ANISOPHYLLEACEAE 

K.M. Wong & L. Madani 




Merrill, EB (1921) 420 (as part of Rhizophoraceae); Ridley, FMP 1 (1922) 692 (as Anisophylleae at 

family rank); Masamune, EPB (1942) 515 (as part of Rhizophoraceae); Ding Hou, FM 1, 5 (1958) 

429 (as part of Rhizophoraceae); Burgess, TBS (1966) 431 (as part of Rhizophoraceae); Corner, 

WS1M 1 (1988) 131 (as Anisophylleaceae); Ashton, IvfNDTS 2 (1988) 342 (as part of Rhizophoraceae); 

Juncosa & Tomlinson, Ann. Missouri Bot. Gard. 75 (1988) 1278 (as Anisophylleaceae); Kochummen, 

TFM 4 (1989) 302 (as part of Rhizophoraceae); Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 295 

(as part of Rhizophoraceae). 

Trees or shrubs, very rarely (in Combretocarpus) with small stilt-roots or pneumatophore 

("breathing") roots. Bark invaginating finely and regularly into wood in most species. 

Branches highly differentiated architecturally from the trunk, twigs solid, branch nodes not 

swollen; supernumerary axillary buds present (inAnisophyllea) or not. Leaves alternate (2- 

ranked in Combretocarpus, basically 4-ranked in Anisophyllea), simple, often (in Anisophyllea) 

dimorphic, 3-7-veined from the base or (in Combretocarpus) pinnately veined; stipules 

none. Inflorescence fundamentally paniculate or racemose, axillary, multiflorous, bisexual 

or (rarely) unisexual. Flower unisexual (Anisophyllea,but plants monoecious) or bisexual 

(Combretocarpus), radially symmetrical, 3-5-merous; calyx valvate; petals free, lobed or 

laciniate, rarely entire; stamens free or (in Anisophyllea) epipetalous and episepalous, 2- 

times the number of petals, anthers 4-celled, dorsifixed, splitting lengthwise; ovary inferior, 

3-5-locular, ovules 1-2 per carpel, style single or (mostly) several and free; disc annular, 

lobed. Fruit a drupe or (in Combretocarpus) a dry, 3-winged structure. Seed one; without 

endosperm; embryo a solid structure with reduced or no cotyledons; germination hypogeal. 

Distribution. 4 genera and c. 35 species, tropical Africa to S America and Malesia. In 

Sabah and Sarawak, 2 genera, Anisophyllea and Combretocarpus, with 10 species. 

Ecology. Combretocarpus is found mainly in peat-swamp forest, and Anisophyllea species 

occur mainly in primary moist lowland forest and sometimes in secondary lowland forest. 

Uses. Several Anisophyllea species and Combretocarpus grow to timber size and are exploited 

for logs. 

Taxonomy. The members of the Anisophylleaceae have traditionally been treated as part of 

the Rhizophoraceae. Ridley (l. c.) appears to have been the first to consider the group (only 

Anisophyllea as was known then for Malaya) at family rank, equivalent to Bentham & 

Hooker's designation of the tribe Anisophylleae within the Rhizophoraceae in 1865 (in 

15


their Genera Plantarum 1, 2). Although Melchior (1964) (Engler's Syllabus der Pflanzenfamilien, 

12th Edition, Vol. 2) and Cronquist (1981) (An Integrated System of Classification of 

Flowering Plants) both adopted the Anisophylleaceae, this has not been widely adopted until 

multi disciplinary studies demonstrated the distinctness of the Anisophylleaceae as a family 

(Juncosa & Tomlinson I.c. and Dahlgren, Ann. Missouri Bot. Gard. 75 (1988) 1259). Basically, 

the Anisophylleaceae is distinct from the Rhizophoraceae by the following characteristics: 

alternate leaves, absence of stipules, simple (not scalariform) vessel perforatiop., mostly 

paniculate or racemose (not cymose) inflorescences, inferior ovary with separate styles (the 

Rhizophoraceae have a superior to inferior ovary and a single style), relatively thin pollen 

wall tapetum, lack of subepidermal floral laticifers (in the ovary and calyx), and lack of 

endosperm in the seed. 

Key to genera 

Leaves basically 4-ranked on lateral branches (although in many species the reduced leaves 

of the two upper ranks are hard to notice or fall away early), with 3-7 veins from the leaf 

base. Leafaxils each with several axillary buds in a row. Fruit a drupe ........ 1. AnisophylIea 

Leaves 2-ranked on lateral branches, pinnately veined. Leafaxils each with only one 

distinct axillary bud. Fruit a 3-winged structure ..................................... 2. Combretocarpus 

1. ANISOPHYLLEA R. Br. ex Sabine 

(Greek, anisos = unequal, phullon = leaf; the dimorphic leaves) 

menengang (Murut, Sarawak), mertama (!ban, Sarawak), 

mopu (Bidayuh, Sarawak), sireh sireh (Malay, Sabah) 

Trans. Hart. Soc. 5 (1824) 466; Merrilll.c. 422; Ridley l.c. 701; Masamune I.c. 515; Ding Hou I.c. 

474; Corner I.c. 131; Anderson, CLTS (1980) 290; Ashton l.c. 344; Kochummen I.c. 304; Whitmore, 

Tantra & Sutisna I.c. 295; Madani, Sandakania 3 (1993) 49. Synonyms: Anisophyllum G. Don ex 

Hook., Niger Fl. (1849) 342, 575; Tetracrypta Gardn. & Champ. in Hooker, J. Bot. Kew Misc. 1 

(1849) 314. 

Trees and shrubs, trunk not buttressed but sometimes fluted at base. Bark smooth or 

minutely cracked or flaky; inner bark invaginating finely and regularly into the wood. 

Sapwood pale to reddish brown, with conspicuous pale radiating lines. Branches 

developing on the young stem in widely separated tiers; axillary buds several in a series in 

each leaf axil. Leaves basically 4-ranked, often dimorphic (but reduced leaves of the two 

upper ranks hard to notice or fall away early in most species), 3-7-veined from the base. 

Flower unisexual (but plants monoecious), 3-5-merous; petals entire, lobed or laciniate; 

stamens epipetalous and episepalous, usually unequal in length; ovary 3-5-celled, ovule 1 

per cell, styles 3-4, free. Fruit a drupe, ellipsoid or pear-shaped, smooth or ridged, fruitwall 

usually hard and fibrous (fleshy inA. disticha). Seed solitary, globose to ellipsoid. 

16

ANISOPHYLLEACEAE (WONG & MADANI) 

Distribution. c. 30 species, tropical Africa, Sri Lanka, India, South East Asia, and West 

Malesia (Sumatra, Peninsular Malaysia and Borneo). 9 species in Sabah and Sarawak. 

Ecology. Primary or old secondary lowland and hill forests to c. 1000 m. 

Key to Anisophyllea species 

1. Leaves along branches of two markedly different sizes, arranged in four rows, the 

larger ones on the two lower rows trapezoid ................................................................. 2 

Leaves of only one type easily discernible, basically elliptic or ovate, arranged in two 

rows (reduced leaves scantily developed and falling early, and usually not noticed) ...... 3 

2. Larger leaves not longer than 3-3.5 cm. Fruiting racemes 0.5-2.7 cm long, typically 

bearing a solitary fruit... ............................................................................. .4. A. disticha 

Larger leaves typically 5-10 cm long. Fruiting racemes 4-10 cm long, typically bearing 

several fruits ........................................................................................ 9. A. rhomboidea 

3. Lateral veins all arising from the very base of the leaf-blade ........................................ .4 

One or both pairs of lateral veins arising from the midrib away from the base of the 

blade ............................................................................................................................ 6 

4. Intercostal veins on the lower leaf surface conspicuously raised, forming a fine tessellated 

pattern ................................................................................................................. 3. A. corneri 

Intercostal veins on the lower leaf surface not conspicuously raised, forming a loose 

reticulation and not distinctly tessellated ........ . 

5. Leaves coriaceous, broadly ovate, upper surface drying glossy, the lateral veins raised. 

Inflorescence axes persistently densely covered with velvety purple-brown hairs ........... . 

................................................................................................................... . 8. A. nitida 

Leaves chartaceous, elliptic, upper surface drying dull or matt, the lateral veins sunken 

or flat. Inflorescence axes subglabrous or only minutely pale-brown hairy ..................... . 

............................... 2. A. chartacea 

6. Leaves rusty hairy on lower sUliace, the remains of the hairs always visible with IOX 

magnification, even on old leaves; intercostal veins on lower surface finely tessellate 

A. ferruginea 

Leaves glabrous, intercostal veins on lower surface indistinct or forming only a loose 

network. ........................................................................................................................... 7 

7. All veins, including the intercostal veins, on upper leaf surface sunken. Mature fruit 

globose ............................................................................................ 7. A. impressinervia 

Veins flat or raised on upper leaf surface. Mature fruit ellipsoid or globose ................. 8 

8. Leaves thinly coriaceous; apex rather abruptly caudate; midrib conspicuously thicker 

than lateral veins. Mature fruit ellipsoid ................................................ 1. A. beccariana 

Leaves chartaceous; apex acute; midrib similar to lateral veins in thickness. Mature 

fruit globose .............................................................................................. 6. A. globosa 

17


1. Anisophyllea beccariana Baill. 

(Odoardo Beccari, Italian explorer and botanist, 1843-1920) 

Adansonia II (1875) 311; Mcrrilll.c. 422; Masamune I.c. 515; Ding Hou I.c. 474; Ashton I.c. 345; 

Whitmorc, Tantra & Sutisna l.c. 295. Type: Beccari PB 1001, Sarawak (lectotype K, here chosen; 

iso1ectotypes G, P). 

Small to medium-sized tree to 25 m tall. Bark scrolled to scaly; inner bark brownish. 

Sapwood yellowish. Leaves of one form mainly, in two distinct ranks along the branches, 

elliptic or ovate, 7.5-10.5 x 3-5 cm, thinly coriaceous, drying matt or slightly glossy on 

upper surface, lower side with scattered tiny but conspicuous dot-glands; base cuneate, apex 

abruptly caudate; midrib conspicuous~y thicker than lateral veins, flat to slightly depressed 

on the upper side; lateral veins (one or both pairs) arising from the midrib away from the 

base of the blade, flat or elevated on upper side, elevated on lower side; intercostal veins 

forming only a loose network or somewhat obscure; glabrous; stalk 4-8 mm long. Inflorescence 

a raceme or basally branched panicle to 10 cm long, axes minutely pale-brown pubescent. 

Flowers unisexual; calyx pubescent outside, lobes 4; petals 0.6-0.7 mm long; stamens in 

male 0.5-0.7 mm long, replaced by staminodes (stamen rudiments) in the female; pistil in 

female 0.5-0.7 mm long, with 4 styles, replaced by pistillode (pistil rudiments) in the male. 

Fruit ellipsoid, c. 3 x 2.5 cm, smooth on the surface when dry. 

Vernacular name. Sarawak-pei (Kayan). 

Distribution. CE Sumatra and Borneo. In Sabah, recorded from Lahad Datu and Sandakan 

districts; in Sarawak from the Kuching and Bintulu districts. Also in Brunei and Kalimantan. 

Ecology. Mixed dipterocarp forest on leached, often shallow humult soils, and kerangas 

(heath) forest in the lowlands, to c." 570 m. 

2. Anisophyllea chartacea Madani 

(Latin, chartaceus = papery; the leaves) 

Sandakania 3 (1993) 51. Type: Othman S. 29055, Sarawak, 3rd Division, Kapit, Ulu Balleh, Ng. 

Mengiong (ho1otype KEP; isotypes A, BO, K, L, l'v1EL, SAR, SING). 

Medium-sized tree to 33 m tall, 80 cm diameter. Leaves of one form mainly, in two distinct 

ranks along the branches, elliptic, 5.5-7 x 2.5-3.5 cm, chartaceous, drying matt on upper 

surface, lower side without conspicuous dot-glands; base cuneate, apex acuminate; midrib 

not conspicuously thicker than lateral veins, sunken on upper side; lateral veins arising 

from the very base of the leaf-blade, sunken on upper side, elevated on lower side; intercostal 

veins forming only a loose network; glabrous; stalk 5-7 mm long. Inflorescence a 

raceme 1.5-5 cm long, axis minutely pale-brown pubescent. Flowers unisexual; calyx 

pubescent outside, lobes 4; petals 0.6-0.7 mm long; stamens in male 0.5-0.7 mm long; 

female unknown. Fruit unknown. 

Distribution. Known only from the type collection from Ulu Balleh in the Kapit district in 


18

ANISOPHYLLEACEAE (WONG & ivlA])ANI) 

Ecolo gy. Lowland mixed dipterocarp forest. 

3. Anisophyllea corneri Ding HOll 

(E.J.H. Corner, 20th Century eminent tropical botanist) 

I.c. 47'B; Corner I.e. 132; Ashton I.c. 347; Kochummen I.e. 305; Whitmore, Tantra & Sutisna I.c. 295. 

Type: Corner SFN 25927, Malay Peninsula, Trengganu (holotype SING; isotypes BO, KEP). 

Small to medium-sized tree to 30 m tall, 55 cm diameter. Bark smooth; inner bark dark 

brown. Sapwood reddish brown. Leaves of one form mainly, in two distinct ranks along 

the branches, elliptic or oblong, 7.5-l3.5 x 2.5-5.5 cm, thinly coriaceous, drying matt on 

upper surface, lower side without conspicuous dot-glands; base cuneate, apex acute; midrib 

conspicuously thicker than lateral veins, raised on the upper side; lateral veins arising from 

the very base of the leaf-blade, elevated on both sides; intercostal veins forming a dense 

and tessellated network of raised fine veins; glabrous; stalk 5-10 mm long. Inflorescence 

a raceme-like panicle with reduced side-branches, 4-9 cm long, axes minutely pale-brown 

pubescent. Flowers unisexual; calyx pubescent outside, lobes 4-5; petals 0.1-0.2 mm long; 

stamens in male 0.3-0.5 mm long, replaced by staminodes in the female; pistil in female 

0.5 mm long, with 4 styles, replaced by pistil rudiments in the male. Fruit broadly 

ellipsoid, 6-l3. 5 x 4-6 cm, smooth on the surface when dry. 

Vernacular names. Sarawak--mertama (Iban), sireh sireh (Malay). 

Distribution. Peninsular Malaysia, Borneo. In Sabah and Sarawak in all districts. Also in 


Ecology. Locally common in mixed dipterocarp forest in the lowlands on leached yellow 

humult soils, and to submontane forest at 1200 m. 

4. Anisophyllea disticha (Jack) Baill. 

(Latin, distichus = 2-ranked; the leaf arrangement on branches) 

l.c. (1875) 311; Merrilll.c. 422; Masamune I.e. 515; Ding Hou l.c. 479 (exc!. A. rhomboidea in 

syn.); Corner I.e. 132; Ashton l.c. 347 (exc!. A. rhomboidea in syn.); Kochummcn I.e. 307; 

Whitmore, Tantra & Sutisna I.e. 295. Basionym: Haloragis distieha Jack, Ma!. Misc. 2 (1822) 19. 

Type: Jack, s.n., Sumatra (lectotype L, here chosen; isolectotype FI). Synonyms: Anisophyllum 

trapezoidale Bail!., Adansonia 3 (1862) 24, 36. 

Shrub or small tree to 7 m tall, 5 cm diameter. Bark smooth; inner bark dark brown. 

Sapwood pale brown. Leaves of two forms, in four distinct ranks along the branches, those 

of the lower 2 ranks oblong-rhomboid, those of the upper 2 ranks tiny and subulate to 

fa1cate, the larger leafform 1.5-3.5 x 0.5-1.5 cm, the smaller leaf form 0.5-0.7 x 0.1-0.3 

cm, chartaceous to thinly coriaceous, drying matt on upper surface, lower side without 

conspicuous dot-glands; base cuneate, apex acute; midrib slightly thicker than lateral veins, 

flat on the upper side; lateral veins one pair arising from the very base of the leaf-blade and 

a third (unpaired) lateral vein arising from the midrib away from the base of the leaf-blade 

19


] >om 

A 

~w \ VV' 

Fig. 1. Anisophyllea globosa. A, leafy twig; B, fruit; C, fruit, cut transversely. (From SAN 76131.) 

20


on the acroscopic side of the midrib, flat on upper side, elevated on lower side; intercostal 

veins scalariform or forming a dense and tessellated network of flat to slightly raised pale 

fine veins; glabrous; stalk none or very short, 0-1 mm long. Inflorescence a raceme, 0.5- 

2.7 cm long, in the axils of the large leaves, axes pale-brown fine-hairy. Flowers unisexual; 

calyx pubescent outside, lobes (3-)4(-5); petals 1.2-1.3 mm long; stamens in male 0.3-0.5 

mm long, replaced by staminodes in the female; pistil in female 0.5-0.8 mm long, with 1 

style, replaced by pistil rudiments in the male. Fruit usually one per raceme, ellipsoid, 

1.5-2.5 x 0.5-1 cm, ripening briglit red, smooth on the surface when dry. 

Vernacular names. Sarawak--ambun ambun (Kedayan), kayu runap (Punan Tutoh), 

mertama (Iban). 

Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah and Sarawak in all districts. 

Also in Brunei and Kalimantan. 

Ecology. Understorey of mixed dipterocarp forest and old secondary forest, lowlands to 

ridges to 1000 m, recorded on alluvial, sandstone, granite and ultramafic soils. 

5. Anisophyllea ferruginea Ding HOll 

(Lahn,ferrugineus = red-brown; the hairs on the leaves) 

I.e. 477; Ashton I.e. 349; Whitmore, Tantra & Sutisna I.e. 295. Type: bb. 25118, E Borneo (holotype 

BO; isotype L). 

Medium-sized tree to 30 m tall, 40 cm diameter. Bark smooth to cracking or slightly 

fissured; inner bark dark brown. Sapwood pale brown. Leaves of one form mainly, in two 

distinct ranks along the branches, ovate to elliptic, 12-16 x 4-6 cm, coriaceous, drying 

matt on upper surface, lower side without conspicuous dot -glands; base cuneate to rounded, 

apex acute to shortly cuspidate; midrib slightly thicker than lateral veins, raised on the 

upper side; lateral veins (one or both pairs) arising from the midrib away from the base of 

the leaf-blade, elevated on both sides; intercostal veins forming a dense and tessellated 

network of raised fine veins; glabrous on upper side, rusty hairy on lower side (the remains 

of the hairs always visible even on old leaves under IOX magnification); stalk 10-17 mm 

long. Inflorescence a panicle, 4-9 cm long, axes minutely pale brown pubescent. Flowers 

unisexual; calyx pubescent outside, lobes (3-)4(-5); petals 0.1-0.2 mm long; stamens in 

male c. 0.3 mm long, replaced by staminodes in the female; pistil in female 0.5 mm long, 

with 4 styles, replaced by pistil rudiments in the male. Fruit ellipsoid, to 8 x 5.5 cm, 

smooth on the surface when dry. 

Vernacular names. Sarawak--belian landak, dajak, kepajang, kepajang landak, sangkuak, 

tengoda (Malay). 

Distribution. Wand SE Borneo. In Sarawak recorded from the Semengoh (Kuching) and 

Balingian, Bintulu (Nyabau FR) and Miri (Lambir NP) areas. Not recorded for Sabah. Also 

in Brunei and Kalimantan. 

21


Ecology. A characteristic species of deep sandy humult ultisol soils in mixed dipterocarp 

forest to 600 m; uncommon but locally frequent. 

6. Anisopbyllea globosa Madani 

(Latin, globosus = round; the fruits) 

Fig. 1. 

I.c. 53. Typc: Shea & Minjulu SAN 76131, Sabah, Kudat, Bengkoka Peninsula (holotype SAN; 

isotypes K, L). 

Small tree to 10 m tall, 10 cm diameter. Bark smooth; inner bark red-brown. Sapwood 

white. Leaves of one form mainly, in two distinct ranks along the branches, narrowly 

elliptic, 5-7 x 1.5-2.5 cm, chartaceous, drying matt on upper surface, lower side with 

scattered tiny but conspicuous dot-glands; base cuneate, apex acute to acuminate; midrib 

similar to lateral veins in thickness, flat or slightly sunken on the upper side; lateral veins 

(one or both pairs) arising from the midrib away from the base of the leaf-blade, flat on the 

upper side, elevated on the lower side; intercostal veins forming a loose network of fine 

veins; glabrous; stalk 4-7 mm long. Inflorescence and flower not known. Fruit globose, 5- 

5.5 cm across, irregularly rough-lumpy on the surface when dry. 

Distribution. Probably endemic to Sabah, known only from the type collection from 

Bengkoka in Kudat district. 

Ecology. Open secondary vegetation at edge of steep gully, lowlands at 50 m. 

7. Anisophyllea impressinervia Madani 

(Latin, impressus- = sunken, nervus = veins; the impressed leaf veins) 

l.c. 53. Typc: Shea & Minjulu SAN 76094, Sabah, Kudat, Bengkoka Peninsula (holotype SAN; 

isotypes A, K, L, SAR, SING). 

Medium-sized tree to 26 m tall, 30 cm diameter. Bark irregularly rough-cracking, finely 

flaky at the base; inner bark red grading to yellow-orange. Sapwood pale yellow. Leaves of 

one form mainly, in two distinct ranks along the branches, elliptic or ovate, 4-4.5 x 2-3 

cm, chartaceous to thinly coriaceous, drying matt on upper surface, lower side with 

scattered tiny but conspicuous dot-glands; base cuneate to rounded, apex acute; midrib 

similar to lateral veins in thickness, sunken on the upper side; lateral veins (one or both 

pairs) arising ji-om the midrib away from the base of the leaf-blade, sunken on the upper 

side, elevated on the lower side; intercostal veins forming a loose network of fine veins; 

glabrous; stalk 3-6 mm long. Inflorescence and flower not known. Fruit globose, c. 4.5-5 

cm across, irregularly rough-lumpy on the surface when dry. 

Distribution. Possibly endemic to Sabah, known only from the Lype collection from Bengkoka in 

Kudat district. 

Ecology. Secondary forest, lowlands at c. 30 m. 

22


8. Anisophyllea nitida Madani 

(Latin, nitidus = polished; the shiny dry upper leaf surface) 

I.e. 53. Type: Ampuria SAN 40249, Sabah, Kuala Penyu (holotype SAN). 

Medium-sized tree to 23 m tall, 25 cm diameter. Bark smooth to scaly. Sapwood 

brownish. Leaves of one form mainly, in two distinct ranks along the branches, broadly 

elliptic-ovate to almost round, 5.5-10 x 4-5.6 cm, thickly coriaceous, drying glossy on 

upper surface, lower side without conspicuous dot-glands; base obtuse-rounded, apex acute 

to slightly cuspidate; midrib slightly to conspicuously thicker than lateral veins, raised on 

the upper side; lateral veins arising from the very base of the leaf-blade, raised on both 

sides; intercostal veins forming a loose network of fine veins; glabrous; stalk 10-15 mm 

long. Inflorescence and flower not known. lnfructescence axis covered in persistent velvety 

purple-brown hairs. Fruit broadly ellipsoid, to 5 x 2.5 cm, smooth on the surface when dry. 

Vernacular name. Sabah---engkop engkop (Kedayan Bundu). 

Distribution. Possibly endemic to northwest Borneo and highly localised. Known only 

from Kuala Penyu in SW Sabah and the Niah area in NE Sarawak. 

Ecology. Lowland secondary and disturbed mixed dipterocarp forest on sandy soils. 

9. Anisophyllea rhomboidea Baill. 

(Latin, rhomboideus = rhombic; the leaf shape) 

I.e. (1875) 310; Merrilll.e. 422; Masamune I.e. 515. Type: Eeeeari PE 1514, Sarawak (lectotype 

here chosen: G). Synonyms: A. distieha sensu Ding Hou I.e. 479, pro parte, sensu Ashton I.e. 347, 

pro parte, non (Jack) Bail!. 

Shrub or small tree to 4 m tall. Leaves of two forms, in four distinct ranks along the 

branches, those of the lower 2 ranks oblong-rhomboid, those of the upper 2 ranks much 

smaller and falcate-elliptic to falcate-ovate, the larger leafform 3.5--,-10 x 1.2-3.5 cm, the 

smaller leaf form 0.6-1.4 x 0.3-0.8 cm, chartaceous to thinly coriaceous, drying matt on 

upper surface, lower side without conspicuous dot-glands; base cuneate to right-angled, 

apex acute; midrib slightly or not thicker than lateral veins, flat on the upper side; lateral 

veins one pair arising from the very base of the leaf-blade and a third (unpaired) lateral 

vein arising from the midrib away from the base of the leaf-blade on the acroscopic side of 

the midrib, flat on upper side, elevated on lower side; intercostal veins scalariform or 

forming a loosely tessellated network of flat to slightly raised pale fine-veins or obscure; 

glabrous to slightly hairy on the main veins on the lower side; stalk none or very short, 0-3 

mm long. Inflorescence a raceme or panicle with short side-branches, 1.5-8(-10) cm long, 

in the axils of the large leaves, axes rusty to pale brown fine-hairy. Flowers unisexual; 

calyx pubescent outside, lobes 4; petals 1.5-2 mm long; stamens in male 1-1.5 mm long, 

replaced by staminodes in the female; pistil in female 1-1.5 mm long, with 1 style, 

replaced by pistil rudiments in the male. Fruit usually several per raceme, ellipsoid, 1.5- 

2.5 x 0.5-1 cm, smooth on the surface when dry, ripening pinkish. 

23


veins forming a loose network of veins, often obscure; glabrous; stalk 7-14 mm long. 

Inflorescence of one or more panicles to 9 cm long arising from the leaf-axil, the base 

becoming tuberculate with age. Flowers yellow; calyx-cup conical with 3 sides, 2.5-4 x 2 

mm, glabrous, lobes ovate, 2.5-3 x l.5-2 mm, outside glabrous, inside scantily pale longhairy; 

petals c. 2 mm long, linear, entire or laciniate (the divisions also linear); stamens 

with filaments 2-2.5 mm long, anthers ovoid, c. 0.3 mm long; styles 1-l.5 mm long. Fruit 

3(-4)-winged, 2-3 cm long, l.5-2 cm wide; wings semicircular, membranous. Seed solitary in 

each fruit, elongate to spindle-shaped. 

Vernacular names. Sabah-perepat paya (Brunei Malay), perepat perepat (Bajau). 

Sarawak--kayu tom (Kayan, Kenyah), keruntum (lban), mutun (Pusa Malay), perepat paya 

(Kayan, Kenyah), sabutun (Melanau Oya). Brunei--balak bekatan (lban) , keruntum 

(Dusun, Iban), perepat hutan (Brunei Malay, Belait, Kedayan). 

Distribution. As for genus. In Sabah and Sarawak found in all types of peat swamp forest. 


Ecology. It is often found in alan (Shorea albida) swamp forest in Sarawak and Brunei and 

mixed peat swamp forest in general. In the more open conditions on the domed centres of 

the largest swamps, and on kerangas (heath forest) sites on groundwater podsols it occurs 

as small trees, whereas in the wetter swampy sites truly big trees occur. In the periodically 

drier sites in the centre of the big peat swamp complexes, gregarious stands of small 

Combretocarpus trees have been called "padang keruntum". Keruntum trees coppice easily. 

Medium-sized or large trees of it are often stag-headed (with sparse, large-limbed crown 

portions) and hollow (Browne I.c.). 

Uses. The timber is suitable for railway sleepers and heavy construction if impregnated 

with preservative. When well-seasoned, it can be used for flooring and panelling. The wood 

is moderately hard and moderately heavy (keruntum logs sink in water). The texture is 

coarse and uneven, due to broad rays which produce an oak-like figure (Burgess I.c.; Wong 

(1982) DMT). 

26 

-------


ARAUCARLL\CEAE 

P.c. Yii 

Sarawak Forestry Department, 

Kuching, Malaysia 

HookerJ, Fl. Br. Ind. 5 (1890) 650; Merrill, EB (1921) 32; Ridley, FMP 5 (1925) 277; Masamune, 

EPB (1940) 2; Backer & BakhuizenJ, FJ 1 (1963) 87; Keng, OFMSP (1969) 12, TFM 1 (1972) 39; 

de Laubenfels, Philip. J. BioI. 7,2 (1978) 143, FM 1, 10 (1988) 419; Kochummen, Mal. For. Rec. 17 

(1979) 54; Cockburn, TS 2 (1980) 2; Anderson, CLTS (1980) 347; Corner, WSTM 2.(1988) 762; 

Whitmore, Tantra & Sutisna, CLK 1 (1989) 27. 

Monoecious trees or rarely shrubs. Exudate resinous, colourless or slightly whitish. Leaves 

simple, spiral, opposite, sub-opposite or in whorls, scale- or needle-like or broad leathery 

blades with many faint closed longitudinal veins. Male and ftmale cones on separate branches. 

Male cones solitary, lateral or rarely terminal, cylindrical, consisting of numerous spirally 

arranged microsporophylls, each bearing a few to numerous microsporangia at the abaxial 

or lower side of an enlarged shield-like apex. Female cones solitary, terminal on robust shoots 

or pedunculate with bracts at the base, ovoid or round, consisting of an axis with numerous 

spirally arranged seed-scales (megasporop~ylls), each bearing a solitary seed (ovule) on the 

adaxial (upper) surface; mature female cones large, woody; cone-scales wedge-shaped. Seed 

winged or not; cotyledons either in 2 fused pairs or of 4 independent units. 

Distribution. 2 genera (Agathis and Araucaria) and 40 species, chiefly in countries bordering 

the South Pacific Ocean (Indo-China, Malesia, New Caledonia, Australia, New Zealand, 

Melanesia, and South America). In Sabah and Sarawak represented by a single genus (Agathis) 

with 5 species. 

AGATHIS Salisb. 

(Greek, agathis = a clew; the shape ofthe cone) 

Trans Linn. Soc. London 8 (1807) 311; Hooker J I.c. 650; Merrilll.c. 32; Ridley I.c. 277; Meijer 

Drees, Bull. Jard. Bot. Btzg. 3, 16 (1940) 455; Backer & BakhuizenJ l.c. 87; Keng I.c. (1969) 12, 

I.c. (1972) 39; Cockburn I.c. 2; Anderson I.c. 347; Whitmore, PI. Syst. Evol. 41 (1980) 46, New 

Phytol. 84 (1980) 407; de Laubenfels, Blumea 24 (1978) 499, Blumea 25 (1979) 531, I.c. (1988) 

419; Whitmore, Tantra & Sutisna l.c. 27. Synonym: Dammara Link, Enum. Hort. Berol. Alt. 2 

(1822) 411. 

Usually large trees. Bole straight and cylindrical, often swollen at the base or with large 

superficial roots. Bark light-grey to greyish brown, usually smooth and lenticellate when 

young, gradually peeling off into thin irregular flakes, leaving behind irregular pock-marks 

on larger trees; inner bark granular, reddish brown. Twigs glabrous, greenish. Terminal 

27


buds globular, usually covered with several pairs of overlapping scales. Petioles usually 

very short, hardly distinguishable from the blades, slightly channelled above. Leaves 

entire, spirally arranged on the older branches and decussate on the young twigs, size and 

shape extremely variable, lanceolate, ovate, to elliptic, without distinct midrib; juvenile 

leaves distinctly larger than the adult leaves and often have more acuminate apices. Male 

cones subtended by several pairs of bracts, sessile or shortly pedunculate. Female cones 

with numerous closely appressed more or less triangular seed-scales. Seed flattened ovoid, 

with 2 unequal lateral wings; cotyledon 2, opposite and leaf-like. 

Distribution. About 21 species; Indo-China, Malesia, Australia, New Zealand and Melanesia 

(except Solomon Islands). 5 species occur in Sabah and Sarawak. 

Ecology. Large and emergent trees with 2 species, Agathis borneensis and A. endertii, 

widely distributed from lowland peat swamp, kerangas (heath), mixed dipterocarp and 

montane forest to about 2400 m. Young trees often produce female cones several years 

ahead of males. 

Uses. Agathis produces a beautiful softwood which fetches a very good price. Its pleasing 

yellow timber has a natural sheen which is very suitable' for furniture and cabinet making, 

wall and ceiling panelling. It is not durable for outdoor use. The white crystalline resin is 

also collected for sale as copal. For more information, see Whitrnore, Econ. Bot. 34 (1980) 1. 

Taxonomy. The species of Agathis occurring in Sabah and Sarawak are characterised by 

the decussate leaves with close faint longitudinal veins without prominent midrib, and by 

the male and female cones which are borne on separate branches. For a detailed account of 

the systematy of the genus see papers by de Laubenfels, Blumea 24 (1979) 531 and 

Whitmore, New Phytol. 84 (1980) 407 and PI. Syst. Evol. 41 (1980) 41. 

Key to Agathis species 

1. Leaf lower surface glaucous ......................... . ..... 2 

Leaf lower surface not glaucous ................................. ................... .4 

2. Adult leaves distinctly convex-lens-shaped, asymmetric; apex and base sharply acute 

............................................................................................................... 4. A. lenticula 

Adult leaves not so ................................................................................................................ 3 

3. Adult leaves ovate or orbicular, apex obtuse or rounded, usually less than 4 cm. Male 

cone usually less than 6 mm in diameter, shortly pedunculate. Seed-scales without 

distinct lip ............................................................................................... 5. A. orbicula 

Adult leaves elliptic, apex acute or rarely blunt, usually longer than 5 cm. Male cones 

about 10 mm in diameter, sessile. Seed-scales with distinct protruding lips at the apical 

margin ..................................................................................................... 2. A. endertii 

4. Adult leaves elliptic, usually longer than 5 cm, apex acute. Male cones oblong, pedunculate, 

usually larger than 20 mm in diameter.. ............................................... 1. A. borneensis 

28

ARAUCARIACEAE (YII) 

Adult leaves ovate, usually less than 5 cm long, apex usually acuminate or round and 

blunt with a distinct tip. Male cones about 10 mm in diameter.. ...... 3. A. kinabaluensis 

1. Agathis borneensis Warb. 

(of Borneo) 

Fig. 1. 

Monsunia 1 (1900) 184; Merrilll.e. 32; Masamune I.e. 2; Meijer Drees I.e. 459; de Laubenfels I.e. 

(1979) 532, I.e. (1988) 433; Whitmore I.e. (1980) 54. Type: Beeeari 491 (B) & 596 (B, K), Sarawak 

(syntypes). Synonyms: Pinus dammara Lamb., Descr. Pinus 1 (1803) 61; A. dammara (Lamb.) 

Richard, Comm. Bot. Conif & Cycad. (1826) 83;A. loranthiifolia Salisb.l.e. 312; Dammara loranthiifolia 

(Salisb.) Link I.e. 411; A. beeearii Warb. I.e. 184; A. maerostaehys Warb. I.e. 183; A. rhomboidalis 

Warb. I.e. 184; A. alba Foxw., Philip. J. Se. 4 (1909) 442; A. latifolia Meijer Drees I.e. 459. 

Emergent tree to 55 m tall and 100 cm diameter. Bark warty lenticellate, papery scaly or 

flaky, maurish grey; inner bark granular, pale brown. Leaves when juvenile ovate to lanceolate, 

up to 14 x 4 cm; when adult elliptic to ovate, 6-12 x 2-3.5 cm, leathery, glabrous; base 

cuneate rounded, apex acute. Male cones oblong, 4-7 cm long, 20-25 mm across, on short 

peduncles, 2-10 mm long; microsporophylls 6-7 x 4.5-5 mm, with broad, rounded apex. 

Female cones ovoid, 6-8.5 x 5.5--6.5 cm; seed-scales 30-40 x 25-28 mm, roughly triangular 

in shape with well rounded apical margin. Seeds flattened ovoid, c. 12 x 9 mm, with one 

well-developed wing, c. 20 x 16 mm. 

Vernacular names. Sabah-manggilan (Dusun), salang (Kedayan). Sarawak-bindang 

(Malay), bulu (lban), kayu jadi (Malay, Iban). Brunei-tulong (Brunei, Malay, Dusun). 

Distribution. Sumatra, Peninsular Malaysia and throughout Borneo. In Sabah and Sarawak 

widespread; also in Brunei and Kalimantan. 

Ecology. Infrequent in the lowlands, although found in sandy margins of some peat 

swamps and kerangas, and lower montane forests to 1200 ill. Generally prefers more acidic 

soils; often forming dense, nearly pure stands on certain areas of low-lying wet kerangas 

forest and dry kerangas forest on sandstone formations at higher altitude, elsewhere 

uncommon and scattered. 

2. Agathis endertii Meijer Drees 

(F.R. Endert, 1891-1953, forester at the Forest Research Institute, Bogor, Indonesia) 

I.e. 470; Masamune I.e. 3; de Laubenfels I.e. (1979) 534, I.e. (1988) 439. Type: bb. 21694, Southern 

Borneo (holotype BO). 

Emergent tree to 60 m and 100 cm diameter. Bark smooth with shallow pock-marks and 

scales, grey-purplish brown; inner bark granular, straw-brown. Exudate whitish, opaque. 

Leaves when juvenile lanceolate, with blunt or acute apex; when adult elliptic, 5-9 x 1.8- 

3.5 cm, leathery and slightly glaucous on undersurface; base cuneate rounded, apex 

broadly acute or rarely blunt. Male cones cylindrical, 2.6-3.8 cm long, 7-10 mm across, 

sessile; microsporophylls spoon-shaped, c. 25 x 20 mm, with spreading and slightly angled 

29


apical margin. Female cones shortly m'oid, c. 4.5 x 7 cm; seed-scales c. 32 x 42 mm, more 

or less triangular with well rounded apical margin, and a distinct protruding lip (c. 8 x 3 

mm) at the apical margin. Seed flattGned ovoid, c. 11 x 8 mm; wing one, c. 18 x 14 mm. 

Vernacular names. Sarawak--bulok (Iban). 

Distribution. Endemic to Borneo. In Sabah and Sarawak scattered. 

Ecology. Scattered throughout lowland rain forests from sea-level to sandstone kerangas 

forest at 1600 m. 

3. Agathis kinabaluensis de Laub. 

(of Mt. Kinabalu) 

I.c. (1979) 535, I.c. (1988) 439. Type: de Laubenfels P625, Sabah, Mt. Kinabalu (holotype L; 

isotypes A, K, SAN). 

Tree up to 36 m tall, becoming relatively stunted on exposed ridges and summits. Bark 

dark brown, with numerous lenticels, peeling off into irregular flakes; inner bark granular, 

reddish brown. Exudate white. Leaves when juvenile ovate, up to 9 x 4.4 cm, apex strongly 

acuminate; when adult ovate, 3.5-7 x 1.8-3.2 cm; base cuneate rounded, apex slightly 

acuminate or more or less round and blunt with distinct tips on smaller leaves. Male cones 

cylindrical, 1.8-3 cm long, 8-10 mm across, nearly sessile or on very short peduncles; 

microsporophylls spoon-shaped, c. 1.7 x l.4 mm, with very slightly angled apical margin. 

Female cones ovoid, c. 8 x 11 cm; seed-scales 28-32 x 40-45 mm, with distinct narrow 

ridges along apical margin and broadly rounded at the upper end. Seed c. 11 x 7 mm, 

distinctly acute at one end and with a broad wing c. 20 x 12 mm at the other. 

Vernacular name. Sarawak-tumu (Kelabit, Murut). 

Distribution. Only known from Mt. Kinabalu, Sabah, and Mt. Murut in Sarawak. 

Ecology. Upper mossy montane forest at 1500-2400 m. 

4. Agathis lenticula de Laub. 

(Latin, lenticularis = a double convex lens; the leaf shape) 

I.c. (1979) 537, I.c. (1988) 436. Type: de Laubenfels P619, Sabah, Mt. Kinabalu, Park HQ (holotype 

L; isotypes A, K, SAN). 

Tree to 45 m tall. Bark greyish brown, with numerous small lenticels; young bark 

distinctly covered with thin exfoliating flakes, gradually breaking into irregular plates; 

inner bark granular, reddish brown. Exudate whitish, gradually turning yellow. Leaves 

when juvenile lenticular, up to 11 x 4.7 cm, tapering toward both ends; when adult lenticular, 

5-7 x 1.7-2.4 cm, leathery, glaucous on the undersurface, apex and base more or 

less acute. Male cones cylindrical, 3-4 cm long, 9-10 mm in diameter, on short peduncles, 

30

ARAUCARIACEAE (YII) 

c 

D 

~ "~""i 

~.' , 

\~\. 

2cm 

B 

50mm 

'om [ 

;(Po.o 

A 

Fig. 1. Agathis bomeensis. A, leafy branch with lateral male cone and terminal female cone; B, 

detail ofleafvenation; C, seed-scale; D, winged seed. (All from S. 9633.) 

31


2-6 mm long; microsporophylls spoon-shaped, 2-2.5 x 1.5-2 mm, with blunt spreading 

apical margin. Female cones spherical, c. 7 x 6 cm; seed-scales c. 40 x 27 mm. Seeds c. 11 

x 7 mm, with a broadly rounded wing, c. 8 x 14 mm at one end. 

Vernacular names. Sabah-tangilan, tungilan (Dusun). 

Distribution. So far only known from the Crocker Range, including Mt. Kinabalu in Sabah. 

Ecology. Emergent trees in mossy montane rain forest, at 1140-1680 m. 

5. Agathis orbicula de Laub. 

(Latin, orbicularis = circular; the leaf shape) 

I.e. (1979) 540, I.e. (1988) 437. Type: de Laubenfels P614, Sarawak, Lawas (holotype L; isotypes A, 

K, SAN, SAR). 

Tree to 40 m tall. Bark dark brown, peeling off into irregular flakes, exposing scattered 

lenticels; inner bark granular, reddish brown. Exudate resinous, light yellow and produced 

in abundance. Leaves when juvenile ovate, up to 6.5 x 2.8 cm; apex bluntly acute; when 

adult ovate to orbicular, 2.4-4 x 1.2-2.4 cm, broadly rounded to slightly angled at the 

apex, tapering toward the base. Male cones minute, cylindrical, 0.8-1.4 cm long, 4-6 mm 

through; microsporophylls helmet-shaped, 1.2-1.5 x 1-1.2 mm, with blunt apex. Female 

cones ovoid, c. 7 x 4.5 cm; seed-scales ovate, c. 20 x 33 mm. Seeds similar to that of A. 

lenticula. 

Vernacular names. Sarawak-bulok (Iban), tubu (Kenyah & Kayan), tumuh (Mumt) .. 

Distribution. Endemic to Borneo; known from southern parts of Sabah to Central Sarawak. 

Ecology. Scattered in sandstone kerangas forest at 450-1050 m. 

32


BIGNONIACEAE 

A. Berhaman 




Merrill, EB (1921) 525; van Steenis, Rec. Trav. Bot. Neerl. 24 (1927) 787, Bull. Jard. Bot. Btzg. 3, 

10 (192S) 173, FM 1, 8 (1977) 114; Masamune, EPB (1942) 652; Kochummen, TFM 3 (1978) 36; 

Anderson, CLTS (1980) 152; Cockbum, TS 2 (1980) 19; Santisuk, Flora of Thailand 5 (1987) 32; 

Corner, WSTM 1 (1988) 172; Whitmore, Tantra & Sutisna, CLK 1 (1989) 28. 

Trees, shrubs or woody climbers; twigs often lenticellate; stipules absent. Leaves generally 

opposite and J-2-4-times pinnate, or simple and whorled, often with glands on the lower 

surface of the leaflets; domatia sometimes present. Inflorescences generally terminal, 

sometimes borne on the trunks and branches. Flowers bisexual, usually very showy; calyx 

tubular, subtruncate or with 5 lobes or teeth, closed in bud and later splitting into lobes, 

often glandular outside; corolla in most genera bilaterally symmetrical, in some genera 

somewhat radially symmetrical, basally tubular, upper part expanded, generally campanulate 

or salver-shaped, with 5 lobes, 2 on the upper side, 3 on the lower; stamens 5, almost equal, 

or (mostly) 4 and didynamous (2 long and 2 short), attached to the corolla (epipetalous); ovary 

superior, 2-cellecl, style long, stigma 2-lobed, nectary usually present and ring-like; ovules 

several to many per cell, arranged longitudinally in 1-2 rows on the septum. Fruit a pod 

divided by a longitudinal partition into 2 compartments, splitting open when ripe. Seeds 

many, flat; embryo near the centre; peripheral wing very thin and transparent or thickened 

and opaque; germination epigeal. 

Distribution. About 120 genera and 650 species in the tropics and subtropics. In Sabah and 

Sarawak, 10 species in 5 genera including 5 apparently unnamed species (4 woody climbers 

and 1 tree). A number of exotic species have been introduced as ornamental plants: 

Arrabidaea magnifica, Jacaranda rhombifolia (jambul merah, formerly J jilicifolia), lvIansoa 

hymensala, lvIillingtonia hortensis, Spathodea campanulata (African tulip tree), Tabebuia rosea, 

and Tecoma stans (YeHow bells). 

Ecology. Mostly evergreen but Oroxylilm indicum and Dolichandrone spathacea have been 

noted as leafless for many months during the dry season of seasonal parts of Asia. 

Dolichandrone spathacea is confined to the mangroves, Deplanchea bancana is found in 

kerangas (heath) and peat swamp forest, some woody climber species appear restricted to 

limestones, and some Radermachera species may be restricted to ultramafic soils. In many 

species, the flowers open at night, and some have fragrant white flowers with long tubes 

which are pollinated by moths. O. indicum has flowers with lurid colour, musky odour, 

fleshy funnel-shaped corollas, which are pollinated by nectarivorous bats. The winged seeds 

are dispersed by wind. 

33


1. DEPLANCHEA Vieill. 

(E.F. Deplanche 1824-1874, French physician & naturalist) 

Bull. Soc. Linn. Norrnand. 7 (1862) 96; van Steenis I.e. (1927) 906, I.e. (1977) 135; Kochummen I.e. 

37; Anderson I.e. 152; Cockburn I.e. 19; Corner I.e. 175; Whitmore, Tantra & Sutisna I.e. 28. 

Small to medium-sized trees. Bark often lenticellate. Sapwood soft. Leaves simple, in whorls, 

quite large, elliptic to ob ovate, chartaceous to coriaceous; base with two large saucershaped 

glands on upper surface. Inflorescence terminal, a thyrse with a somewhat flattopped 

structure, the branches many (more than 20) and held horizontally at almost right 

angles to the main inflorescence axis, large and conspicuous. Flowers large; calyx 5-lobed; 

corolla irregular (zygomorphic), funnel-shaped, tube with a short constricted basal part and 

a flared upper part, lobes 5, yellow; stamens strongly exserted, 4 (rarely 5), didynamous; 

disc annular to lobed; ovary 2-celled; style long exserted; stigma 2-lobed. Fruit a 2-valved 

capsule. Seeds many, thinly winged. 

Distribution. 5 species, Malesia to N Australia. 1 species in Borneo (Sabah, Sarawak, Brunei, 

and Kalimantan). 

Ecology. Being light-demanding, species of the genus are mainly found in secondary rain 

forests (including kerangas forest), woodland savannahs, and grasslands, from sea-level to 

1000 m. 

DepJanchea bancana (Scheff) Steenis 

(of Banka Island) 

Fig. 1. 

I.e. (1927) 921, I.e. (1928) 221, I.e. (1977) 137; Kochummen I.e. 37; Anderson I.e. 152; Cockburn 

I.e. 19; Corner I.e. 175; Whitmore, Tantra & Sutisna I.e. 28. Basionym: Diplanthera baneana 

Scheff., Nat. Tijd. Ned. Ind. 31 (1870) 334. Type: Teijsmann, H.B. 9666, Banka (BO). Synonym: 

Deplanehea eoriaeea Steenis I.e. 224. 

Small to large trees 10-45 m tall, 25-70 cm diameter; bole fluted at base, buttresses small 

or large. Bark white to brown, lenticellate and slightly fissured; inner bark yellow to pale 

brown. Sapwood white to pale yellow, soft. Young twigs lenticellate, glabrous to pale hairy, 

pale brown to yellow. Leaves ob ovate to elliptic, 9-34 x 5.5-20 cm, chartaceous to coriaceous; 

base cordate to cuneate, apex rounded, rarely acuminate-acute; midrib glabrous to yellowhairy 

on lower surface; lateral veins 7-8 pairs, raised and glabrous to yellow-hairy on the 

lower surface; stalk 3-6 cm long, glabrous to yellow-hairy. Inflorescences showy, stalk 5- 

20 cm long. Flowers showy; calyx 12-16 mm long, with scattered glands, teeth 5; corolla 

c. 3.5 x 2.5 cm, yellow, 5-lobed, the upper 2 lobes recurved; stamens 4,jilaments erect and 

curved over the upper corolla-lobes, base glandular hairy; ovary sub sessile, each locule 

with two placentas; style greenish yellow, exserted and curved like the stamens; stigma 

bilobed. Fruits oblong, 10-14 x 3.5 cm. Seeds flat, c. 3 x 2 cm; wing broad, very thinly 

transparent, irregularly lobed. 

Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak uncommon. 

36

BIGNONIACEAE (BERHAMAN) 

Ecology. In Sabah and Sarawak, it occurs in heath forests on white sands or in peat 

swamps, in primary and secondary forests, from sea-level to 1000 m. 

2. DOLICHANDRONE Seemen 

(Greek, dolichos = long, andron = male; the long stamens) 

Ann. Mag. Nat. Hist. Ser. 3, 10 (1862); van Steenis I.e. (1927) 928, I.e. (1928) 227, I.e. (1977) 141; 

Kochummen I.e. 39; Anderson I.e. 152; Cockburn I.e. 19; Corner I.e. 176; Whitmore, Tantra & 

Sutisna I.e. 28. 

Medium-sized trees. Leaves once pinnate, leaflets elliptic, entire. Inflorescence a terminal 

raceme. Flowers fragrant, nocturnal; calyx spathaceous, somewhat curved; corolla funnelshaped, 

equally 5-10bed; stamens 4, didynamous, included; disc annular; ovary with many 

ovules in many rows; style slender. Fruit a subcylindric capsule. Seeds hyaline-winged. 

Distribution. 9 species, from East Africa through tropical Asia to Australia; 4 species in 

tropical SE Asia; 1 species in Sabah and Sarawak. 

Dolichandrone spathacea (L. f) K. Schum. 

(Latin, spathaceus = resembling a spathe; the calyx form) 

Fig. 2. 

Fl. Kais. Wi1h. Land. (1889) 123; Merrill, I.e (1921) 525; van Steenis I.e. (1927) 937, I.e. (1928) 

227, I.e. (1977) 142; Masamune I.e. 652; Kochummen I.e. 39; Anderson I.e. 152; Cockburn I.e. 19; 

Santisuk I.e. 56; Corner I.e. 176; Whitmore, Tantra & Sutisna I.e. 28. Basionym: Bignonia spathaeea 

L. j, Suppl. (1781) 283. Type: Konig, Herb. Linn. No. 776. 8 (LlNN). Synonyms: Doliehandrone 

rheedii Seemen, J. Bot. 8 (1870) 380; Doliehandrone longissima K. Schum. in Engl. & Prantl, Pfl. 

Fam. 4, 3b (1894) 240. 

Tree to 25 m tall, 10-40 cm diameter. Bark pale brown, smooth to fissured; inner bark 

laminated, reddish pink. Sapwood white, soft. Young twigs lenticellate. Leaves once pinnate, 

opposite; rachis smooth, brown; leaflets usually 3-4 pairs with one terminal leaflet, ovateoblong 

to lanceolate, 5.5-15 x 2.5-6 cm, drying dark to brown-black; base oblique, apex 

acuminate pointed; lateral veins 5-8 pairs, lower surface with hairy domatia; stalks short in 

lateral leaflets but 2-4 cm in terminal leaflet. Inflorescences 2-8-flowered, with one 

flower opening at a time. Flowers fragrant; calyx conical, closed and with hooked apex in 

bud, with scattered glands, splitting along one side; corolla white, with a tube to 14.5 cm 

long, bell-shaped near the mouth, 3-4 cm wide, with many scattered glands outside, lobes 

5 with frilled margin; stamens 4, didynamous with a fifth rudiment, included; style long, 

exserted, persistent in young fruit; anthers glabrous, bilocular. Fruits 20-33.5 x 1.5-3 cm, 

generally curved, hanging in clusters from the ends of the branches. Seeds many, dark 

grey, in many rows, c. 15 x 9 mm including the thick corky wings. 

Vernacular names. Sabah-kelaju, tui (Dusun). Sarawak-tuih (Iban). 

Distribution. From the coasts of Malabar to tropical SE Asia and the whole of Malesia. 

37


Ecology. Restricted to the back mangroves and along banks of tidal rivers and estuaries. In 

Sabah, common in the mangrove forests of the west coast; in Sarawak occasional on clay 

soils at inland limits of mangroves. 

Uses. The wood is soft and light with density of about 480 kg/m 3 , easy to saw and work 

with and seasons well although it must be protected against sapstains. So far, there is no 

record of its use in Sabah and Sarawak but in various parts of its geographical range, the 

wood is used for wooden clogs, net-floats, scabbards and household utensils. Due to its 

anatomical and physical characteristics the wood may also be suitable for pattern making 

and matches. 

3. OROXYLUM Vent. 

(Greek, oros = mountain, xylon = wood or tree; the natural habitat of the tree) 

Dec. Gen. Nov. (1808) 8; van Steenis, I.e. (1927) 816, I.e. (1928) 181, I.e. (1977) 128; Masamune 

I.e. 652; Kochu11ll11en I.e. 40; Anderson I.e. 152; Cockburn I.e. 19; Corner I.e. 178; Whitmore, Tantra 

& Sutisna I. e. 28. 

Small to medium-sized trees. Leaves opposite, 2-4-times pinnate; leaflets with 4-5 pairs 

of lateral veins, glands on the lower surface present only in young and some mature leaves, 

and then only at the basal part of leaflets. Inflorescence a terminal raceme, erect, long. 

Flowers fragrant, showy; calyx closed in bud, with an apical pore opening as a somewhat 

bell-shaped structure; corolla purplish, with 5 subequallobes; stamens 5; ovules in many 

rows in each cell. Fruit a long, flattened woody capsule. Seeds quite large, with thin wings 

all around, in many rows. 

Distribution. Monotypic genus distributed in India, Sri Lanka, and throughout SE Asia. 

Oroxylum indicum (L.) Vent. 

(ofIndia) 

Fig. 3. 

I.e. 8; van Steenis I.e. (1927) 816, I.e. (1928) 181, I.e. (1977) 128; Masamune I.e. 652; Kochu11ll11en 

I.e. 40; Anderson I.e. 152; Cockburn I.e. 19; Santisuk I.e. 36; Corner I.e. 166; Whitmore, Tantra & 

Sutisna I.e. 28. Basionym: Bignonia indiea L., Sp. PI. (1753) 625. Type: Herb. Hermann No. 236; 

vol. 5: Icon. 28, 29, 72 (BM). Synonyms: Bignonia pentandra Lour., Fl. Coch. 2 (1970) 379; 

Bignonia tripinnata Noronha, Verh. Bat. Gen. 5 (1790) 8; Calosanthes indiea Blume, Bijdr. (1826) 

760; Hippoxylum indieum Rafin., Tellur. (1838) 78. 

Medium-sized tree, partly deciduous, 20-40 m tall, 10-40 cm diameter. Bark smooth, 

whitish; inner bark pale yellowish. Sapwood pale whitish, soft. Leaves 3-4-times pinnate, 

up to 2 m long, tufted at the branch ends; petioles long, rachis swollen at points of 

articulation; leaflets ovate to oblong, 6-9.5 x 2.5-5 cm, terminal one always bigger; base 

cuneate or mostly oblique, apex acute to acuminate; lateral veins 4-5 pairs; intercostal veins 

reticulate; glands scattered on the flower surface. Inflorescence 25-100 cm long. Flowers 

nocturnal, c. 9 cm long; calyx coriaceous, 2.5-3.5 cm long, 1.5-2 cm across, containing 

38


c 

A 

2 cm [ 

,; ~~ 

1 

'~~~ 

,~ ,;:1 

'~ ~ 

~'" " 

[~ 

f:~ 

1 cm 2:_·_,.'-( 

;:~~~~ 

D 

Fig. 2. Dolichandrone spathacea. A, leafy twig; B, f1ower; C, fruit; D, seed, CA from SAN 31498, B 

fromFRI12490, C & D from SAN A 1771.) 

39


8 cm 

D 

Fig. 3. Oroxylum indicum. A, part of compound leaf; B, detail of leaflet; C, part of inflorescence; D, 

pod. (From SAN 89947.) 

40


water in bud, campanulate, dirty brown-violet, becoming almost woody in the fruit; corolla 

c. 10 cm long, reddish outside, yellowish to pinkish inside, fleshy, funnel-shaped, the 5 

unequal lobes with wrinkled margin; stamens all fertile, hairy at the base, included. Fruit 

25-100 x 5-10 cm, hanging conspicuously resembling the big flat blade of a slash-knife. 

Seeds numerous, 4-8 x 2.5-4 cm including the membranous and transparent wing. 

Vernacular names. Sabah-parang pamol (colloquial Malay). Sarawak--binkuli (!ban), 

gimurai, murai (padawan Bidayuh), parang nyabor (!ban). 

Distribution. As for genus. 

Ecology. Secondary forest, mostly below 1000 m. 

4. RADERMACHERA Zoll. & Moritzi 

(lC.M. Radermache, l741-1783, patron of science in the Netherlands Indies) 

in Zollinger, Syst. Verz. Pfl. Ind. Arch. 3 (1855) 53; Merrilll.e. (1921) 525; van Steenis I.e. (1927) 

953, I.e. (1928) 238, I.e. (1977) 149, Blumea 23 (1976) 121; Masamune I.e. 652; Kochummen I.e. 

42; Anderson I.e. 152; Cockburn I.e. 20; Corner I.e. 180; Whitmore, Tantra & Sutisna I.e. 29. 

Trees or shrubs. Twigs lenticellate. Leaves J-3-times pinnate, opposite; leaflets often with 

gland-fields on the lower surface. Inflorescence a raceme or thyrse, terminal on branches 

or borne on the trunk and branches (plant cauliflorous or ramiflorous). Flowers white, pink 

or greenish yellow; calyx 2-5-lobed, often glandular; corolla 5-lobed, slightly two-lipped; 

stamens 4 or 5; ovary elongate, glabrous or covered with tiny scales or tubercles. Fruit a 

straight or twisted capsule with a corky central placenta. Seeds many, tiny and narrow, the 

ends winged. 

Distribution. 15 species; tropical Asia, from India and south China to Malesia; 3 species in 

Sabah and Sarawak. 

Key to Radermachera species 

1. Leaves typically once pinnate, sometimes 1-2-times pinnate. Inflorescences always 

borne on leafy shoots ................................................................................ 1. R pinnata 

Leaves 2-3-times pinnate. Inflorescences borne directly on main branches or on trunk 

and branches (plant cauliflorous or ramiflorous) .......................................................... 2 

2. Leaves 3-times pinnate, the secondary rachises never winged; leaflet tips caudate 

....................................................... 2. R ramiflora 

Leaves 2-3-times pinnate, the secondary rachises with 1-2-rnm-wide wings; leaflet tips 

acuminate to cuspidate ............................................................ 3. Radermachera sp. A. 

41

TREE FLORA OF SABAl-l AND SARAWAK VOL. 1 (1995) 

1. Radermachera pinnata (Blanco) Seemen 

(Latin, pinnatus = feathery; the pinnate leaves) 

Fig. 4. 

1. Bot. 8 (1870) 147; Merrill, Sp. Blanc. (1918) 350; van Steenis l.c. (1927) 973, l.c. (1928) 248, I.c. 

(1976) 129, I.c. (1977) 153; Kochummen I.c. 42; Corner I.c. 181; Whitmore, Tantra & Sutisna I.c. 

29. Basionym: Millingtonia pinnata Blanco, Fl. Filip. (1827) 501. Type: Merrill Sp. Blanc. 834, 

Philippines, Luzon, Rizal Province (neotype A; isoneotypes BO, K, L). Synonym: R. whitfordii 

Merr., Philip. 1. Sc. 75 (1912) 352. 

subsp. acuminata (Steenis) Steenis 

I.c (1976) 129, l.c. (1977) 154; Whitmore, Tantra & Sutisna l.c. 29. Basionym: R. lobbii (Teijsm. & 

Binn.) Miq. subsp. acuminata Steenis I.c. (1928) 247. Type: Endert E 1309, S Sumatra, Lampong, 

Tandjong Karang (holotype BO; isotype L). Synonyms: R. lobbii Teijsm. & Binn., Nat. Tijd. Ned. 

Ind. 25 (1863) 413; R. corymbosa Steenis I.c. (1928) 249. 

Tree 8-40 m tall, 30 cm diameter; buttresses to 2 m high. Bark scaly to flaky, light brown 

to grey; inner bark pale yellow. Sapwood white. Leaves J-2-times pinnate, petioles 8-10 

cm long; leaflets obovate to obovate-elliptic, 7-9 x 2.5-4 cm, coriaceous to chartaceous; 

base cuneate, apex acute to rounded; lateral veins 7-9 pairs. Inflorescence a terminal 

thyrse. Flowers mildly fragrant; calyx purple, c. 1.5 cm long, c. 1-1.5 cm across, with a few 

scattered glands outside, lobes 3; corolla about 1.5 cm long, c. 1-1.2 cm across, basal part 

narrow tubular and white, upper inflated portion pale lilac, throat and medium inner 

surface of the lobes orange, lobes 5 with pale, minute hairs on the margin; stamens 4, 

didynamous; ovary glabrous, style glabrous, 1.2-2 cm long. Fruit a capsule, hanging in 

clusters at the end of branches, 16-27.5 x 0.5-1 cm; persistent calyx c. 1.8 cm long. Seeds 

c. 2.5 x 0.5 cm, winged. 

Vernacular names. SarawaklKalimantan--binutan, kujuk langit (Kapuas Dayak). 

Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Borneo, and the Philippines. 

In Sabah and Sarawak, generally uncommon but localised populations may be encountered 

in uItramafic and limestone areas in the northern, eastern and southern parts of Sabah, and 

around the Niah Caves and Kuching areas in Sarawak. 

Ecology. Primary and secondary forest, to 800 m, associated with ultramafic soils (Sabah) 

or limestones (Sarawak). Flowering in May, July-March; fruiting in May-November. 

Taxonomy. This species consists of two subspecies, according to van Steenis l.c. (1977) 

153, I.c. (1976) 129. The other subspecies, R. pinnata subsp. pinnata, is restricted to 

Celebes, the Moluccas and the Philippines. The main difference is that in subsp. pinnata 

the intercostal veins on the lower leaflet surface form a dense and prominent network, 

whereas in subsp. acuminata the veins are not conspicuous or prominent. Subsp. acuminata 

is slightly variable in leaflet shape and texture. 

2. Radermachera ramiflora Steenis 

(Latin, ramiflorus = flowering on the branches) 

J. Bot. 74 (1934) 5, l.c. (1976) 130, I.c. (1977) 152; Masamune I.c. 653; Whitmore, Tantra & Sutisna 

l.c. 29. Type: Clemens 28672, British North Borneo, Mount Kinabalu (holotype BO; isotypes K, L). 

42


1 'om 

c 

"I ~ I1 

!fI 

.J 

I" 

ill, 

III 

','!!1 

'·i,1 

, 

E 

F 

III 

B 

3cm 


Medium-sized tree, 30-45 m tall, 30-50 cm diameter; trunk fluted at base. Bark greyish, 

fissured; inner bark whitish, turning yellowish on exposure. Sapwood yellowish white. 

Twigs with corky bark. Leaves 3-times pinnate, crowded at the branch-tips; leaflets 4-6 

pairs, elliptic-lanceolate, coriaceous, 3.5-8.5 x 1.5-3.5 cm; apex caudate; lateral veins 5-7 

pairs and raised below, lower surface with scattered small glands near the base. Inflorescence a 

raceme, borne on the trunk and branches (plants cauliflorous or ramiflorous), 11-14 cm 

long. Flowers yellow, erect on curved pedicels of c. 20 cm long; calyx dull reddish green to 

dark brown when dry, in the bud pear-shaped with many scattered glands outside especially 

near the base, 18-21 x 6-8 mm, lobes 3; corolla deep ochre-yellow, the limb red, narrowly 

salver-shaped with scattered 2-4 glands outside, slightly curved, the narrow basal part 3-5 

cm long, lobes 5; stamens 4, exserted, didynamous, hairy at the base; ovary ribbed, style to 

3-3.5 cm long. Fruit straight or twisted, 35-70 cm long. Seeds 4-5 x 2.4 mm, the wing 6- 

7 mm. 

Vernacular name. Sabah-tuik-tuik hufan (Malay). 

Distribution. Endemic to Sabah, known from Mt. Kinabalu, Ranau, Kota Marudu, and Mt. 

Rara. 

Ecology. Rainforest, also in disturbed forest on hill sides, to 1500 m, associated with 

ultramafic soils. Flowering in January to March and around August; fruiting recorded in 

April and December. 

van Steenis I.c. (1977) noted that the calyx is eglandular, but the specimen Chew & Corner 

RSNB 4290, examined by him, has a few scattered glands outside the calyx. 

3. Radermachera sp. A. 

van Steenis I.c. (1977) 152 included the only collection of this undescribed species 

(Nooteboom & Aban 1603, Sabah, Kinabalu, at Kampong Kiau) in R. ramiflora, a conclusion 

we disagree with. The species is poorly known; the young flower buds are not sufficient for 

describing it properly, but the winged secondary rachis and cuspidate leaflet tips clearly 

distinguish it from R. ramiflora. 

44


BURSERACEAE 

K.M. Kochummen 



A.w. Bennett in Hooker [, Fl. Brit. Ind. 1 (1875) 527; King, J. As. Soc. Beng. 62, 2 (1893) 235; 

Merrill, EB (1921) 316, PEB (1929) 116; Ridley, FMP 1 (1922) 368; H:J. Lam, Bull. Jard. Bot. 

Btzg. 3, 12 (1932) 281; Masamune, EPB (1942) 362; Browne, FTSB (1955) 70; Leenhouts, FM 1, 5 

(1956) 208, FM 1, 6 (1972) 917, FM 1, 7 (1976) 820; Backer & Bakhuizen[, FJ 2 (1965) 112; 

Smythies, CST (1965) 30; Burgess, TBS (1966) 60; Kochurnmen, TFM 1 (1972) 121; Co(;kburn, TS 

1 (1976) 34; Anderson, CLTS (1980) 155; Wong, DMT (1982) 30; Corner, WSTM 2 (1988) 199; 

Whitmore, Tantra & Sutisna, CLK 1 (1989) 34; Ng, Mal. For. Rec. 34, 1 (1991) 35. 

Medium-sized to large buttressed trees, rarely shrubs; dioecious or occasionally monoecious; 

crushed living parts with strong resinous smell. Bark pale grey to brown, smooth or scaly, 

often lenticellate, surface sometimes with black or white resinous gum; inner bark pinkish, 

or creamy, laminated or mottled, with droplets of clear or white resinous gum, with strong 

resinous smell in most species (especially those of Canarium and Triomma). Sapwood 

often whitish, shiny, sometimes pinkish or yellowish brown, vessels usually visible to the 

unaided eye. Pith of twigs, petioles and petiolules often with vascular strands and resin 

ducts. Leaves spiral, usually crowded at the tips of twigs, pinnately compound with opposite 

leaflets and a terminal leaflet, rarely trifoliolate; petiolules of leaflets with distinct swellings 

at both ends in most species (leaflets subsessile in Garuga); margin entire except in some 

Canarium and Garuga species. Stipules present in all species of Garuga and in most 

species of Canarium. Inflorescences usually axillary or terminal panicles, sometimes 

spikes, racemes or thyrses. Flowers unisexual (except in Garuga), male and female on 

different trees, 3- or 5-merous, usually greenish to cream, the remains of the other sex 

persisting; sepals valvate, mostly united; petals valvate, free; stamens usually twice as many 

as petals, filaments free or united, sometimes fused to the disc, anthers dehiscing inwards; 

disc intrastaminal (except in Triomma), nectariferous, brightly coloured; ovary superior, 

with 3-5 cells, each cell with 2 ovules; style simple, stigma globular, often slightly lobed. 

Fruit a drupe with a fleshy or leathery resinous rind and a more-or-Iess woody stone 

(pyrene) or in Triomma a woody capsule; stone 3-celled or less. Seed one; cotyledons fleshy, 

variously folded, rolled and convoluted; endosperm absent; germination epigeal or 

hypogeal, mostly rapid; cotyledons divided into 3, 5 or more lobes (entire in Scutinanthe). 

Seedling with first 2 leaves opposite or alternate, subsequent leaves alternate, spiral, simple 

for several nodes, then pinnate (in some Santiria the leaves are pinnate from start); sapling 

leaflets thinner, with long drawn-out tips, longer petioles and more distantly spaced when 

compared to the adult leaves. 

Distribution. 16 genera and about 550 species; tropics and subtropics. In Sabah and Sarawak, 

8 genera and 59 species are known. 

45


Ecology. Burseraceae are common constituents of the main storey of mixed dipterocarp and 

kerangas forests; they also occur in submontane forests to 1800 m. The drupaceous fruits 

are dispersed by animals. In Triomma the winged seeds are dispersed by wind. 

Timber. The standard Malaysian name for the timber of Burseraceae is kedondong (Malay). 

It is a very homogeneous group except for slight colour differences, and a light hardwood, 

moderately hard and moderately heavy; in strength Class 'C', strong; working quality 

variable, some easy, others difficult and blunting tools excessively. Not durable in tropical 

conditions. Moderately susceptible to powder post-beetles. Difficult to treat with preservatives. 

Seasoning without serious degrade, with low shrinkage. Sapwood pale, not sharply defined; 

heartwood yellow-brown, pink-brown or red-brown, only slightly darkening on exposure, 

surface mostly glossy, without figure; grain interlocked. Texture moderately fine and even. 

Most species of Dacryodes, Santiria and Scutinanthe have siliceous timbers while Canarium 

(except c. apertum) and Triomma have no silica in the timber. 

Uses. Timber suitable for general building construction and carpentry work, plywood and 

weather boarding. The family abounds in fragrant balsams and resins (e.g., balsam from 

Commiphora opobalsamum; frankincense from Boswellia species especially B. sacra; 

myrrh from Commiphora myrra). The resins are used in traditional medicine in Peninsular 

Malaysia. The resin from Canarium luzonicum, the "Manila-elemi" is used in pharmacy in 

ointments and plasters. The oil from the kernel is used locally. Commercial quantities of 

the oil are now being produced in the Solomon Islands from Canarium species for 

industrial use in the manufacture of skin- and hair-care products under the trade name 

"Solomon Nut Oil". The seeds of Canarium album, Canarium indicum (ngali-nut; Solomon 

Islands), C. ovatum and C. vulgare (kenari, Indonesian) are eaten. Certain species, e.g., C. 

album, C. ovatum and C. vulgare, are planted as avenue trees, or for wind-breaks and also 

as shade-trees for nutmeg plantations. 

Taxonomy. Although the Burseraceae is taxonomically closely related to the Meliaceae and 

Simaroubaceae it is liable to be confused with the pinnate-leaved species of the Anacardiaceae. 

The sole distinguishing character is the number of ovules per cell which is two in every cell 

in the Burseraceae and one in the Anacardiaceae. Vegetatively, the swollen petiolule is a 

good diagnostic feature for most members of the Burseraceae. The family is divided into 3 

tribes. Except for Triomma which belongs to tribe Bursereae with capsular fruit, all the 

genera in Borneo belong to the tribe Canarieae with a drupaceous fruit. Triomma retains a 

capsular fruit which must be the primitive fruit of the family and which is perhaps to be 

regarded as an ancient relic character. 

Key to genera 

1. Flowers 4-5-merous. Fruit with 1 or more pyrenes .. . 

Flowers 3-merous. Fruit with 1 pyrene ................................ . 

... 2 

.5 

2. Flowers bisexual. Stipellae present. Leaflet margin toothed ........................... 3. Garuga 

Flowers unisexual. Stipellae absent. Leaflet margin entire or occasionally toothed ....... 3 

3. Stamens 5; disc extrastaminal. Fruit a 3-winged capsule. Leaflets withering yellow, 

with pink veins .................................................................................................... 8. Triomma 

46

BURSERACEAE (KOCHUMMEN) 

Stamens 8 or 10; disc intrastaminal. Fruit a globular or ellipsoid drupe. Leaflets not 

so ................................................................................................................................. 4 

4. Receptacle of flower cup-shaped; ovary 3-celled ..................................... 7. Scutinanthc 

Receptacle flat; ovary 4-5-celled .................................................................. 5. Protium 

5. Fruit thick-walled, bony, seated on enlarged calyx. Leaves often stipulate, leaflet 

margin often toothed ................................................................................. 1. Canarium 

Fruit thin-walled, calyx not enlarged in fruit. Leaves without stipules, leaflet margin 

entire ........................................................................................................................... 6 

6. Remains of stigma on the fruit always distinctly off-centre. Fruit surface smooth when 

dry ................................................... , ............................................................ 6. Santiria 

Remains of stigma on the fruit not off-centre. Fruit smooth or coarsely wrinkled when 

7. Dry fruits smooth. Cotyledons entire. Inflorescences axillary, to 10 cm 10ng .. .4. Haplolobus 

Dry fruits coarsely wrinkled. Cotyledons divided. Inflorescences axillary and terminal, 

much longer.. ........................................................................ , ............................ 2. Dacryodes ~ 

1. CANARIUM Stickman 

(from the Moluccan name kenari) 

kedondong (Malay) 

Herb. Amb. (1754) 10 (erroneously Cenarium); King l.c. 236; Rid1ey I.c. (1922) 369; H.J. Lam, Bull. 

Jard. Bot. Btzg. 3, 12 (1932) 422; Burkill, EPMP 1 (1935) 424; Browne I.c. 70; Leenhouts I.c. 

(1956) 249, I.c. (1972) 92, Blumea9 (1959) 275; Backer & BakhuizenJ l.c. 114; Kochummen I.c. 

(1972) 126, Sandakania 5 (1994) 73; Cockburn I.c 37; Anderson l.c. 155; Corner I.c. 200; Wong I.c. 

30; Whitmore, Tantra & Sutisna l.c. 34; Ng l.c. 37. 

Medium-siz,ed to large buttressed trees, rarely shrubs. Bark grey-fawn or light yellowbrown, 

smooth, scaly or dippled, with many small lenticels; inner bark pinkish or reddish, 

laminated, soft, aromatic, with clear sticky exudate, rarely with non-sticky exudate, which 

becomes dark brown on exposure. Sapwood whitish, darkening inwards. Twigs usually 

round, the pith nearly always with vascular strands. Leaves often with stipules of various 

shapes and sizes, at base of rachis or on it, soon falling off; leaflets generally with pointed 

tips, the margin entire, dentate, or serrate, basal pair of leaflets usually smaller; petiole 

rounded, flattened or channelled especially towards base, often swollen at base. Inflorescences 

axillary or terminal panicles, spikes, racemes, or thyrses, female ones often reduced and 

smaller. Flowers unisexual, male and female on separate trees, 3-merous; receptacle flat or 

concave; calyx cup-shaped, lobes deltoid, outside glabrous or hairy, inside always densely 

silky-hairy; petals 3, creamy, free, usually overlapping in bud, nearly always ovoid-oblong, 

the tip injlexed, jleshy and thick with thin margins, outside hairy in the centre, inside 

usually glabrous; stamens 6 in one whorl, rarely only 3, free to entirely connate, sometimes 

adnate to the disc, in female flowers sterile and often less-developed, filaments flattened, 

47


anthers opening by a longitudinal slit; disc inside to the stamens (intrastaminal), 6-lobed, 

strongly developed in male flowers; ovary in female flowers stalked if receptacle is concave, 

ovoid to ellipsoid, style cylindrical, stigma globular, slightly 3-lobed; in the male much 

reduced, often fused to disc. Fruit a drupe seated on persistent enlarged calyx, with apical 

stigma, blue black when ripe (rarely ivory white or red), hairy especially near base and 

apex, or glabrous; pericarp fleshy or fibrous, wrinkled when dry; stone hard, woody, round 

or triangular in cross-section, with 3 cells, often reduced to 2 or 1, containing woody 

intrusions of the placenta, penetrating between cells as wings, often visible as surface ribs. 

Seed I per cell; testa brown; without endosperm; cotyledons oily, 3-lobed in most species 

but 5-lobed in C. megalanthum. 

Distribution. c. 100 species; tropical Wand E Afri~a, Madagascar, Mauritius, Sri Lanka, 

SE Asia from S Deccan to S China and Hainan, Malesia, NE Australia and Melanesia; 23 

species in Sabah and Sarawak. 

Ecology. Common in the lowland mixed dipterocarp forests, rare in submontane forests to 

1800 m. 

Uses. Canarium species produce general utility timber. Burkill (l.c.) documents some ofthe 

uses. C. pimela is cultivated in SE China as an ornamental and as a fruit tree. The fruits are 

highly esteemed by the Chinese. C. indicum, C. ovatum and C. vulgare are planted for their 

nuts. They are an important constituent of the diet in the Solomon Islands. In Malaysia and 

China they are a valued titbit. In Sarawak C. odontophyllum is widely cultivated for the 

edible fruits. C. pseudodecumanum has edible fruits; the oil pressed out of the seeds is 

locally used. The resin is used in pharmacy in ointments and plasters, mainly from C. 

luzonicum, the "Manila-elemi". It is also a constituent of cellulose lacquers. The shell of the 

nut is used as fuel substitute in the Solomon Islands. Very little is known about the 

economic value of the other species. 

Taxonomy. The genus can be subdivided into 3 subgenera: subgenus Canarium consisting 

of sections Canarium and Pimela is centred in Malesia, subgenus Ajricanarium in W 

Africa, and subgenus Canariellum restricted to E Queensland and New Caledonia. In 

Sabah and Sarwak, section Canarium can be recognised by its foliaceous or pectinate 

(comb-like) stipules, its leaflets which dry brown, its stamens that are free or adnate to the 

disc, the usually glabrous disc and pistil, and the larger fruit (5-7 cm long) with frequently 

tomentose calyx. In contrast, section Pimela has lanceolate or narrow stipules, leaflets 

which dry greyish green, stamens that are often partly or entirely connate, a disc and pistil 

that are mostly pilose, and smaller fruit (often 2.5-3.5 cm long) with usually glabrous 

calyx. 

Key to Canarium species 

1. Leaflets glabrous ............................................... ......... 2 

Leaflets hairy .......................... .. ................................................. 16 

2. Leaflet margin toothed ............................................................................................... 3 

Leaflet margin entire ........................................................................................................ 6 

48


3. Leaflets distinctly whitish below ............................................................. 13. C. littorale 

Leaflets not so, rarely faintly whitish below ....................................................................... .4 

4. Stipules absent. Leaflet base rounded, margin faintly toothed ..... 20. C. pseudodecumanum 

Stipules present. Leaflet base not rounded, margin distinctly toothed ........................... 5 

5. Stipules deeply irregularly lobed. Rachis yellowish .......................... 5. C. denticulatum 

Stipules suborbicular, not lobed. Rachis brownish ........................ l0. C. kinabaluensis 

6. Leaflets sessile, lateral veins perpendicular to midrib. Trees with stilt-roots 

.............................................................................. 4. C. decumanum 

Leaflets stalked, lateral veins not so. Trees without stilt-roots ..................................... 7 

7. Plants without stipules .. . ......... 8 

Plants with stipules ............................. ............................................. 10 

8. Lateral veins 5-7 pairs. Leaflet apex with 1-2-cm-long tip ........... 22. C. pseudopimela 

Lateral veins 7-14 pairs. Leaflet apex without such long tips ...................................... 9 

9. Twigs with prominent leaf-scars. Petals clawed ............... 21. C. pseudopatentincrvium 

Twigs without prominent leaf-scars. Petals not clawed ........................... 19. C. pilosum 

10. Stipules needle-like ... .................... 11 

Stipules broad and flat... ...................... ............... 13 

1l. Rachis black. Flowers reddish when fresh ......................................... 6. C. dichotomum 

Rachis not black. Flowers not reddish when fresh ...................................................... 12 

12. Leaflet stalk about 5 mm long; lateral veins 15-18 pairs ................ 11. C. kostermansii 

Leaflet stalk longer, to 20 mm long; lateral veins 6-11 pairs ................... 2. C. asperum 

13. Leaves faintly glaucous below ............................ ..... 14. C. littorale 

Leaves not glaucous below ......................... ..................... 14 

14. Lateral veins and reticulations invisible .......... .......... 18. C. patentinervium 

Lateral veins and reticulations visible ... ..... 15 

15. Lateral veins prominently raised above; leaflet base usually rounded ..... 7. C. divergens 

Lateral veins not raised; leaflet base cuneate ......................................... 3. C. caudatum 

16. Leaflet margin toothed .......................................... ............... 17 

Leaflet margin entire ............... 

......................................... M 

17. Leaflets sessile, margin minutely toothed ............................. 20. C. pseudodecumanum 

Leaflets stalked, margin coarsely and distinctly toothed ............................................. 18 

18. Stipules absent... ............................. . 

Stipules present.. .......... . 

. .. 19 

. .... 21 

49


19. Apex ofleaflet blunt, rounded or notched ................................................ 1. C. apertum 

Apex of leaflet pointed ............................................................................................... 20 

20. Leaflet margin curled inwards ......................................................... 9. C. grandifolium 

Leaflet margin not so ................................................................... 23. C. sarawakanum 

21. Stipules linear. Twigs woolly. ............................ 19. C. pilosum 

Stipules broad and flat. Twigs not woolly ... ............................................ 22 

22. Stipules deeply incised ............. ......................... 17. C. odontophyllum 

Stipules not deeply incised .......... . ........................................................ 23 

23. Stipules kidney-shaped with wavy margin .............................................. 14. C. Iittorale 

Stipules rounded to ovate, margin not wavy .................................. 13. C. latistipulatum 

24. Stipules 3-4-lobed. Leaflet base rounded ...................................... 15. C. megalanthum 

Stipules linear, not lobed. Leaflet base not rounded .................................................... 25 

25. Leaflet margin curled inwards .............................. . ....... s. C. fusco-calycinum 

Leaflet margin not curled inwards ............. . ..................... 26 

26. Ultimate leafy twigs stout, more than 1 cm thick. Leaflets (except basal ones) subsessile, 

base rounded or subcordate ............................. : ........................ 10. C. hirsutum 

Ultimate leafy twigs slender, less than 1 cm thick. Leaflets distinctly stalked, base 

cuneate .................................................................................................. 16. C. merrillii 

1. Canarium apertum H.J. Lam 

(Latin, apertus = open or unshielded; the basal gaps between petals) 

Ann. Jard. Bot. Btzg. 42 (1932) 214, Bull. Jard. Bot. Btzg. 3, 12 (1932)491; Masamune I.c. 362; 

Leenhouts I.c. (1956) 275, I.c. (1959) 386; Burgess l.c. 60; Kochummen l.c. (1972) 128; Anderson 

l.c. 155; Whitmore, Tantra & Sutisna I.c. 34. Type: Beccari PB 1630, Sarawak, Matang (holotype FI; 

isotypes K, L). Synonym: Santiria serrnlata Engl. in DC., Mon. Phan. 4 (1883) 160; non C. 

serrnlatum Miq., Fl. Ind. Bat. 1,2 (1859) 646. 

Very large tree to 40 m tall, 80 cm diameter; buttresses to 2 ill high. Bark grey-brown, 

lenticellate, scaly; inner bark yellow-brown. Twigs angled, powdery reddish brown hairy 

when young. Stipules absent. Leaves with 3-4 pairs of leaflets; rachis and petiole powdery 

brown-hairy; blade oblong, elliptic or ovate, 4.5-10.5 x 3-5.5 cm, sparsely rough-hairy 

below and on midrib above; base rounded to subcordate, margin finely toothed towards 

apex, apex blunt or rounded; midrib flattened above; lateral veins 8-15 pairs, prominently 

raised below, giving a bullate appearance to the blade, flat above; intercostal veins reticulate, 

raised below, faint above; petiolules 2-3 mm long, powdery hairy. Flowers (female) 

yellow-brown, in terminal broadly thyrsoid panicles; petals clawed. Fruits ovoid, pointed, 

hairy, circular in cross-section, 4-5 x 2-2.5 cm. 

Distribution. Sumatra, Peninsular Malaysia and Borneo. Uncommon in Sabah, recorded from 

Beaufort, Keningau, Sandakan, Tawau and Tenom. In Sarawak, reported from Belaga, 

Lundu, Miri and the Sabal Forest Reserve. Also occurs in Kalimantan. 

50


Ecology. In lowand mixed dipterocarp forests to 500 m, on clay-rich and yellow sandy soils 

and alluvium. Flowers collected in January and April and fruits in May and October. 

2. Canarium asperum Benth. 

(Latin, asper = rough, uneven; possibly the rough-hairy inflorescence) 

in Hooker, Lond. J. Bot. 2 (1843) 215; H.J. Lam, Bull. lard. Bot. Btzg. 3, 12 (1932) 461; Masamune 

I.e. 362; Leenhouts I.e. (1956) 293, I.e. (1959) 439; Backer & Bakhuizen f I.e. 115; Whitrnore, 

Tantra & Sutisna I.e. 34. Type: Hinds, s.n., New Guinea (K). 

subsp. asperum var. asperum 

Synonyms: Canariopsis aspera (Benth.) Miq., FI. Ind. Bat. 1, 2 (1859) 653; Canarium 

villosum Benth. & Hook.f ex F.-ViiI., Nov. App. (1880) 40; C. molle EngI. in DC. I.c. 109 

(for a complete list of synonyms, cf Leenhouts l.c. (1956). 

Small to medium-sized tree to 30 m tall, 50 cm diameter. Bark pale brown, scaly with 

large flakes; inner bark dull brown with white sticky exudate. Sapwood cream. Stipules 

needle-like, inserted near the base of petiole. Leaves with up to 6 pairs of leaflets; rachis 

glabrous, rarely hairy; petiolules of lateral leaflets 0.5-2 cm long, swollen towards apex 

(only 2 mm long on collections from Pulau Gaya), rarely hairy; blade elliptic to lanceolate, 

7.5-16 x 4-6.5 cm; base rounded to broadly cuneate, slightly unequal, margin entire, apex 

pointed; midrib raised above; lateral veins 6-11 pairs, raised below, faintly raised above; 

intercostal veins scalariform-reticulate, raised below, faint above. Flowers in axillary 

spikes or racemes. Fruits ovoid to subglobose, 9-14 x 4-11 mm. 

Distribution. E Java, Borneo, Lesser Sunda Islands, Philippines, Celebes, Moluccas, New 

Guinea and the Solomon Islands. In Sabah uncommon, recorded only from Pulau Selipol, 

P. Gaya, P. Banggi, Lahad Datu and Kudat. Not yet recorded from Sarawak. Also known 

from Kalimantan. 

Ecology. Rocky coasts and inland lowland mixed dipterocarp forests. 

Leenhouts I.c. (1956 & 1959) recognised two subspecies, viz. subsp. asperum andpapuanum, of 

which only subsp. asperum occurs in Sabah and Sarawak. Of subsp. asperum, he distinguished 

two varieties, namely var. asperum and var. clementis with the latter endemic to the 

Philippines. 

3. Canarium caudatum King 

(Latin, caudatus = tailed; the gradually tapering leaflet apex) 

I.e. 240; Ridley I.e. (1922) 370, Kew Bull. (1930) 81; H.J. Lam, Bull. lard. Bot. Btzg. 3, 12 (1932) 

443; Masamune I.e. 362; Leenhouts I.e. (1956) 259, I.e. (1959) 343; Kochummen I.e. (1972) 129. 

Type: King's collector 10227, Perak (holotype CAL; isotypes A, BM, G, L). 

Medium-sized to large tree to 36 m tall, 40 cm diameter. Bark grey, cracking and scaly; 

inner bark with strong resinous smell. Stipule usually falling off early, kidney-shaped. 

51


Leaves with 2-4 pairs of leaflets; petiolules of lateral leaflets c. 1 cm long, swollen at both 

ends; blade elliptic or ovate, 5.5-16.5 x 3-8 cm; base rounded or broadly cuneate, margin 

entire, apex pointed, tip c. 1 cm long; midrib raised above; lateral veins 8-11 pairs, distinct 

on both surfaces; intercostal veins reticulate, visible on both sides. Flowers in terminal 

thyrsoid inflorescences; stamens 3 in male flowers and 6 in female flowers. Fruits spindleshaped, 

5.5-8 x 2-3.5 cm, with saucer-shaped persistent calyx. 

Key to forms 

Stipules small, kidney-shaped, inserted partly on the twig ..... 

forma caudatum 

Synonym: C. pauciflarum Rid!. l.c. (1930) 80. 

Sumatra, Peninsular Malaysia and Borneo. Of scattered distribution in Sabah and Sarawak. 

Stipules auricle-shaped, inserted on the petiole ..................... . 

forma auriculiferum Leenh. 

Blumea 8 (1955) 181. Type: Haviland 2877, Sarawak (holotype SING; isotype BM, K, L, SAR). 

Sumatra, Peninsular Malaysia, and Borneo. In Borneo uncommon, known by a few 

collections from Sabah (SAN 33828), Sarawak (e.g., S. 5840, S. 12032 and S. 46003), and 

Kalimantan. 

Ecology. Lowland mixed dipterocarp and kerangas forests to 230 m. 

4. Canarium decumanum Gaertn. 

(Latin, decumanus = the largest or greatest; the size of the fruit and tree) 

Fruct. 2 (1791) 99; El Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 435; Leenhouts I.c. (1956) 276, I.c. 

(1959) 389; Burgess l.c. 60; Cockbum I.e. 39; Whitmore, Tantra & Sutisna l.c. 34. Type: Hart. Bot. 

Bog. VI E 5 (= P!. Bog. Exsicc. 116), Moluccas (neotype L; isoneotypes B, BO, BRSL, G, K, L, NY). 

Very large tree to 54 m tall, 150 cm diameter; buttresses to 5 m tall, stilt-roots present. 

Bark grey, smooth to scaly or dippled; inner bark orange-red, granular, with strong mango 

smell, exudate brownish and sticky. Sapwood white. Twigs pale white with prominent leafscars. 

Stipules inconspicuous. Leaves closely spirally arranged, with 4-5 pairs of almost 

sessile leaflets; rachis powdery yellowish hairy; blade oblong, elliptic or ovate, 6-15.5 x 3- 

7 cm; base rounded or subcordate, margin entire, apex pointed; midrib flattened above; 

lateral veins 12-23 pairs, almost perpendicular to midrib, forking and forming 

reticulations afew millimeters away from margin, visible on both surfaces; intercostal veins 

scalariform-reticulate, very faint; petiole flattened above with sharp edges. Flowers hairy, 

in axillary thyrsoid inflorescences. Fruits ellipsoid, 7-8.5 x 4.5-6 cm, rough hairy. 

Vernacular name. Sabah-pamatudon (Malay, Dusun). 

Distribution. Borneo, Moluccas, New Guinea. In Sabah uncommon, recorded from Lahad 

Datu and Sandakan; not yet reported from Sarawak; also known in Kalimantan. 

Ecology. Lowland forests. 

52

BURSERACEAE (KO CHUM MEN) 

5. Canarium denticulatum Blume 

(Latin, denticulatus = with very small teeth; the leaflet margin) 

Bijdr. (1826) 1162; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 521; Mfisamune I.e. 362; Leenhouts 

I.e. (1956) 272, I.e. (1959) 367; Backer & BakhuizenJ I.e. 115; Burgess I.e. 60; Kochummen I.e. 

(1972) 129; Cockburn I.e. 41; Whitmore, Tantra & Sutisna I.e. 34. Type: Blume 743, Java CL, 

lectotype ). 

Medium-sized tree to 30 m tall, 50 cm diameter; buttresses short. Bark grey-white, smooth; 

middle bark green; inner bark pinkish. Sapwood white. Twig pale whitish. Stipules persistent, 

yellowish, inserted on petiole, deeply irregularly lobed. Leaves with 2-6 pairs of leaflets; 

rachis pale yellow; petiolules of lateral leaflets 8-15 mm long, yellowish (rachis hairy in 

saplings); blade elliptic or oblong, 6-23.5 x 3-9 cm; base unequal, rounded or cuneate, 

margin faintly toothed towards apex, apex pointed; midrib raised above; lateral veins 9-13 

pairs, curving and joining near margin, visible on both surfaces; intercostal veins reticulate, 

faintly visible on both surfaces; glabrous or reddish brown-hairy on the lower side. Flowers 

white, hairy, in an axillary thyrsoid inflorescence. Fruits ellipsoid, 2.5-3 x 1.5 cm. 

Key to subspecies 

Leaflets reddish brown-hairy below, margin toothed .................... . 

subsp. kostermansii Leenh. 

Blumea 8 (1955) 18!. Type: Kostermans 5226, E Borneo, Sg. Menubar region (holotype L; 

isotypes BM, BO, K). 

Endemic to Borneo. Uncommon, in Sabah only known from several collections, all 

from the east coast districts; also in Kalimantan. 

Leaflets glabrous, margin entire or faintly toothed. 

subsp. denticulatum 

Synonyms: C.fissistipulum Miq., Fl.lnd. Bat. Suppl. (1861) 521; C. kunstleri King I.e. 184; C. 

laeiniatum Elmer, Leafl. Philip. Bot. 3 (1911) 1084. 

Andarnans, Burma, Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sabah 

reported from Ranau, Kalabakan, Kinabatangan, Lahad Datu, Sandakan, Tawau. In 

Sarawak known from Batu Laga, Bt. Mersing, Ulu Tinjar, Bt. Raya, Ulu Rajang, 

Segam FR, Lambir NP and Marudi; also known in Kalimantan. Common in the 

lowlands, rarely to 750 m, in mixed dipterocarp forest on friable fertile soils, particularly on 

basic volcanic rocks. 

6. Canarium dichotomum (Blume) Miq. 

(Latin, dichotomus = having divisions always in pairs; branching ofthe inflorescence) 

I.e. (1859) 648; Merrilll.e. (1929) 116; H.J. Lam, Bull Jard. Bot. Btzg. 3, 12 (1932) 447; Masamune 

I.e. 363; Leenhouts I.e. (1956) 283, I.e. (1959) 423; Burgess I.e. 60; Anderson I.e. 155; Whitmore, 

Tantra & Sutisna I.e 34. Basionym: Pimela diehotoma Blume, Mus. Bot. Lugd. Bat. 1 (1850) 222. 

Type: Korthals 957, Sumatra (holotype L; isotype U). Synonym: C. endertii H.J. Lam, Ann. Jard. 

Bot. Btzg. 42 (1932) 210. 

53


Medium-sized tree to 27 m tall, 25 cm diameter. Bark brownish to reddish, smooth to 

scaly; inner bark reddish brown. Sapwood pale white. Twigs 0.5-2(-2.5) cm thick, grey or 

dark brown to blackish, powdery brown-hairy when young. Stipules linear, hairy, on the 

base of rachis, persistent. Leaves with 1-4 (rarely 6-8) pairs of leaflets; rachis black, 

powdery brown-hairy; petiolules of lateral leaflets 0.5-2 cm long, hairy; blade elliptic to 

lanceolate or oblong, 8-20 x 4-9.5 cm, sparsely hairy below or glabrous; base broadly 

cuneate, unequal, margin entire, apex pointed, tip c. I cm long; midrib raised above; lateral 

veins 9-15 pairs, raised below, distinct above; intercostal veins reticulate, faintly visible on 

both surfaces. ·Flowers in terminal and axillary thyrsoid inflorescences, main branches in 

male inflorescences repeatedly dichotomously branched, axis reddish. Fruits oblong, 3-4 x 

1.2 -1.5 cm, triangular; calyx funnel-shaped. 

Distribution. Sumatra, Borneo. Of scattered distribution in Sabah and Sarawak. Also known in 

Kalimantan. 

Ecology. Lowland forest, rarely in submontane forest to 1200 m. 

7. Canarium divergens Engl. 

(Latin, divergium = going different ways; the laxly branched inflorescence) 

in DC. I.e. 143; Merrilll.e. (1921) 316; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 503; Masamune 

I.e. 363; Leenhouts I.e. (1956) 260, I.e. (1959) 346; Anderson I.e. 155. Type: Beeeari PB 2237, 

Sarawak, Matang (holotype P; isotypes Fr, K). 

Medium-sized tree to 30 m tall, 45 cm diameter; buttresses short. Bark grey-white, 

lenticellate, scaly. Twigs brownish, glabrous. StipUles deciduous. Leaves with 2-4 pairs of 

leaflets; petiolules of lateral leaflets 7-15 mm long; blade elliptic, ovate or obovate, 7-18 x 

3.5-9.5 cm, glabrous; base cuneate to rounded, margin entire to faintly toothed, apex 

pointed, tip c. 1 cm long; midrib raised above; lateral veins 7-13 pairs, raised below, faint 

above, curving and joining near margin; intercostal veins scalariform-reticulate, raised 

below, faint above. Inflorescences thyrsoid, terminal and axillary; male to 37 cm long, 

hairy. Fruits yellowish green when fresh, ellipsoid, 6-8.5 x 2.5-3.5 cm; calyx funnelshaped 

with wavy margin. 

Distribution. Endemic to Borneo. Uncommon in Sabah, known by a single collection SAN 

50076 from Beaufort; more common in Sarawak, (Semengoh, G. Buri, Matang, Lambir 

National Park, Bintulu and Ulu Anap); also recorded from Brunei. 

Ecology. Mixed dipterocarp forests on deep sandy humult ultisols, to 450 ill. 

8. Canarium fusco-calycinum Stapf ex Ridl. 

(Latin,fuscus = dark or dark brown, calycinus = calyx; the colour of the calyx) 

I.e. (1930) 82; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 499; Masamune I.e. 363; Leenhouts I.e. 

(1956) 283, I.e. (1959) 424; Anderson I.e. 155. Type: Haviland 1981, Sarawak, Kuching (holotype 

K; isotypes BM, SAR). 

54


Medium-sized tree to 20 m tall, 50 cm diameter. Twigs rusty brown-hairy. Stipules linear, 

falling off early. Leaves with 2 pairs of leaflets; rachis hairy; petiolules of lateral leaflets 2- 

3 mm long, hairy; blade hairy below, ovate to oblong, 4.5-15 x 3-9 cm; base broadly 

cuneate, margin entire, incurled, apex pointed, tip c. 1 cm long; midrib raised above, hairy; 

lateral veins 12-17 pairs, raised below, visible above, curving and joining near margin; 

intercostal veins scalariform-reticulate, raised below, faint above. Flowers (male) in terminal 

thyrsoid inflorescences, hairy; stamens united. Fruits ellipsoid, 3-3.5 x 1.2-1.5 cm, triangular 

in cross-section; calyx red-brown hairy, with distinct lobes. 

Distribution. Endemic to Sarawak. Uncommon, reported from the 1st, 4th and 7th Div. 

Ecology. Lowland mixed dipterocarp forest. 

9. Canarium grandifolium (Rid!.) H.J. Lam 

(Latin, grandis = large;Jolium = leaf) 

Ann. Jard. Bot. Btzg. 42 (1932) 215, Bull. Jard. Bot. Btzg. 3, 12 (1932) 527; Leenhouts I.c. (1956) 

275, I.c. (1959) 386; Kochununen I.c. (1972) 129. Basionym: Trigonochlamys grandifolia Ridl., J. 

Str. Br. R. As. Soc. 54 (1916) 31, I.c. (1922) 38l. Type: Cantley's collector, s.n., Singapore 

(lectotype SING). 

Medium-sized to large tree to 40 m tall, 60 cm diameter; buttresses tall. Bark greyish, 

dippled and scaly. Twigs densely fulvous tomentose. Stipules absent. Leaves with 5-7 pairs 

of leaflets, leaflet stalks glabrous to hairy; blade elliptic to ob ovate, 10-17 x 5-8 cm, 

densely hairy beneath and on midrib above; base cuneate to rounded, margin re curved, 

minutely toothed to entire, apex rounded with short tip; midrib sunken above; lateral veins 

9-14 pairs, prominently raised below; intercostal veins equally prominent, sunken above. 

Inflorescences terminal, rarely lateral, densely red-tomentose; male thyrsoid, female racemose 

to spicate. Flowers pubescent; petals clawed; stamens free. Fruits ellipsoid, c. 5 x 3.5 cm, 

slightly hairy at apex. 

Distribution. Peninsular Malaysia and Borneo. In Sabah and Sarawak very uncommon, 

known only from two collections (SAN 44553 and S. 36641); also known from Brunei. 


10. Canarium hirsutum Willd. 

(Latin, hirsutus = covered with rough hairs; the fruit) 

Sp. PI. 4 (1805) 760; Ridley I.c. (1922) 374; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 466; 

Masamune I.c. 363; Leenhouts I.c. (1956) 287, l.c. (1959) 424; Backer & BakhuizenJ l.c. 115; 

Burgess I.c. 60; Kochununen I.c. (1972) 130; Cockburn I.c. 42; Anderson I.c. 155; Whitmore, Tantra 

& Sutisna I.c. 35. Type: bb. 33772, Moluccas (neotype L; isoneotypes A, BO, SING). Synonyms: C. 

hispidum Blume, Cat. (1823) 109; C. subcordatum Ridl. l.c. (1922) 374. 

55


Medium-sized tree to 25 m tall, 60 cm diameter. Twigs very stout, 1.5-3.5 cm thick, reddish 

brown-hairy near tip, pith large with many small vascular strands. Stipules present (except 

in two varieties), caducous, inserted on the petiole, subulate, 4-12 mm long. Leaves to 2 m 

long, with 4-13 pairs of leaflets; blade subsessile except basal ones, glabrous or hairy, 

ovate to lanceolate, 5-45 x 2.5-15 cm; base rounded to cordate, margin entire, apex pointed; 

midrib raised above; lateral veins 12-30 pairs, visible on both surfaces; intercostal veins 

finely reticulate, visible; petiole to 2 cm thick near base with sharp edges. Inflorescences 

axillary, male thyrsoid, female subracemose. Flowers c. 1 cm long, shortly stalked. Fruits 

with irritant reddish brown hairs, oblong to ovoid, 2.8-3.5 x 2-2.5 cm. 

Distribution. Throughout Malesia (except Lesser Sunda Islands), Carolines and Solomon 

Islands. In Sabah rather common, but in Sarawak very uncommon and represented by a 

single collection (s. 25192). Also known in Kalimantan. 

Ecology. Mixed dipterocarp forest, rarely to 1800 m. 

Two subspecies and a few varieties are recognised for this species. In Sabah and Sarawak 

only subspecies hirsutum is present with its variety hirsutum and forma scab rum (Blume) 

Leenh. 

11. Canarium kinabaluensis Leenh. 

(ofMt. Kinabalu) 

I.e. (1955) 182, I.e. (1956) 260, I.e. (1959) 346; Burgess I.e. 60; Whitmore, Tantra & Sutisna I.e. 35. 

Type: Ramos 1698, North Borneo, Sandakan (holotype L; isotypes A, BO, BM, K, P). 

Medium-sized tree to 25 m tall, 50 cm diameter; buttresses steep, to 1 m high. Bark grey, 

smooth; inner bark orange with strong resinous smell. Sapwood white. Twigs blackish. 

Stipules suborbicular, inserted on the base of the petiole. Leaves with 2-4 pairs of leaflets; 

rachis black, cracking; petiolule black on drying; blade drying to reddish brown, thickly 

leathery, elliptic-ovate or lanceolate, 8-20 x 3.5-10 cm; base broadly cuneate, margin very 

faintly toothed, apex pointed; midrib raised above, sharply keeled below; lateral veins 11- 

13 pairs, raised below; intercostal veins scalariform. Inflorescences and flowers unknown. 

Fruits spindle-shaped, green drying brown, strongly wrinkled, 6-8 x 2-4 cm. Seeds one. 

Distribution. Endemic to Borneo. Uncommon, collected from Papar, Sandakan and Mt. 

Kinabalu in Sabah; also known from Kalimantan. 

Ecology. Lowland to submontane forests at 300-1500 m. 

12. Canarium kostermansii Leenh. 

(A.J.G.R. Kostermans, 1907-1994, botanist at the Forest Research Institute Bogor and the 

Herbarium Bogoriense, Indonesia) 

I.e. (1955) 191, I.e. (1956) 281, I.e. (1959) 398; Burgess I.e. 61; Whitmore, Tantra & Sutisna I.e. 35. 

Type: Kostermans 5315, E Borneo (holotype L; isotypes BM, K). 

56


Medium-sized to large tree reaching 35 m tall, 60 cm diameter. Bark brown, smooth. 

Twigs slender, to 1 cm thick, pith with 2 concentric cylinders of vascular strands. Stipules 

subulate, caducous. Leaves with 4-7 pairs of leaflets; leaflet stalk about 0.5 cm long; blade 

lanceolate, 10-22 x 3-7 cm; base unequal, broadly cuneate, margin entire, apex pointed; 

midrib raised above; lateral veins 15-18 pairs, faintly curving towards the margin, raised 

above; intercostal veins reticulate, distinct on both surfaces. Inflorescences (female) axillary, 

racemose. Fruits spindle-shaped, 3 x 1-1.5 cm, glabrous; calyx funnel-shaped. 

Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak; also found in 

Kalimantan. 

Ecology. Lowlands and hill forests to 900 m. 

13. Canarium latistipulatum Rid!. 

(Latin, latus = broad, wide, stipula = stipule) 

I.e. (1930) 81; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 508; Masamune I.e. 364; Leenhouts I.e. 

(1956) 258, I.e. (1959) 341; Whitmore, Tantra & Sutisna I.e. 35. Type: Haviland & Hose 2245, 

Sarawak, Kuching (holotype K; isotype SAR). 

Small to medium-sized tree to 27 m tall. Young twigs powdery brown-hairy. Stipules subpersistent, 

rounded to ovate, 1-2 xl-I. 7 cm, inserted on the base of petiole. Leaves with 3 

pairs of leaflets; rachis powdery brown-hairy; petiolule of lateral leaflets 5-10 mm long, 

hairy; blade sparsely hairy below, drying reddish brown, oblong or elliptic, 6-15 x 3.5-6 

cm; base broadly cuneate, margin toothed, apex pointed, tip c. 1.5 cm long; midrib raised 

above; lateral veins 7-10 pairs, raised below, curving and joining near margin; intercostal 

veins scalariform-reticulate, distinct below, faint above. Flowers (male) hairy, in terminal 

thyrsoid inflorescences. Fruits oblong, 6.2-7.2 x 2.2-3 cm, with tapered base; calyx 

funnel-shaped, powdery brown-hairy. 

Distribution. Endemic to Borneo. Common in Sabah but uncommon in Sarawak and 

known only by 2 other collections (s. 3458 & s. 52998) beside the type. 


14. Canarium Iittorale Blume 

(Latin, littoralis = of the seashore; its main habitat) 

I.e. (1826) 1164; Merrill I.e. (1929) 116; H.J. Lam, Bull. Jard. Bot. Btzg. 3,12 (1932) 498; 

Masamune I.e. 364; Leenhouts I.e. (1956) 256, I.e. (1959) 337; Backer & BakhuizenJ I.e. 115; 

Burgess I.e. 61; Kochummen I.e. (1972) 130; Cockbum I.e. 42; Anderson I.e. 155; Whitmore, Tantra 

& Sutisna I.e. 35. Type: Blume 1736, Java, Nusa Kambangan (holotype L; isotypes BO, MEL, U). 

Medium-sized to large tree to 40 m tall, 60 cm diameter. Bark grey, smooth to scaly; inner 

bark orange-red or brownish. Sapwood pale white. Twigs glabrous or hairy. Stipules 

dropping off early or semi-persistent, kidney-shaped with wavy to deeply lobed margins. 

57


Leaves with 2-6 pairs of leaflets; rachis glabrous or softly hairy; petiolules of lateral 

leaflets 1-2 cm long, glabrous or hairy, often swollen at both ends; blade ovate to oblong or 

lanceolate, 3-25 x 2-9 cm, glabrous or sparsely to densely hairy below, sometimes white 

waxy below; base cuneate to subcordate, margin entire to faintly or prominently toothed, 

apex pointed; midrib raised above; lateral veins 9-20 pairs, sometimes curving and joining 

near margin, raised below, faint or sunken above; intercostal veins scalariform-reticulate or 

reticulate, distinct below, faint above, rarely sunken above. Flowers usually in terminal 

thyrsoid (male) or subracemose (female) inflorescences. Fruits ellipsoid or ovoid, 4.5-7 x 

1.5-3 cm, sparsely hairy or glabrous with rugose and wrinkled surface when dried. 

An extremely variable species in vegetative and floral characters. Leenhouts (l.c. 1959) 

recognises 5 forms (littorale, pruinosum, purpurascens, rujum, and tomentosum), of which 

3 occur in Sabah and Sarawak. 

Key to forms 

1. Leaflets whitish beneath ......................................... . 

forma pruinosum (Engl.) Leenh. 

I.c. (1956) 258. Basionym: C. pruinosum Engl. in DC. I.c. 106, Merrilll.c. (1921) 317, 

Masamune l. c. 365. Type: Beceari PB 1970, Sarawak (holotype K; isotype FI). 

Confined to Sabah and Sarawak, common in secondary forest, especially on 

periodically flooded sandy alluvium. 

Leaflets not whitish beneath .. 

2. Leaflets densely hairy below; margin prominently toothed; lateral veins sunken above .......... . 

forma rufum (A.W. Benn.) Leenh. 

l.c. (1959) 339. Basionym: C. ru/um AW. Benn. in Hooker f I.c. 533, Masamune I.c. 365. 

Type: Maingay 1434, Malacca (holotype K; isotypes CAL, L). 

Widely distributed in Sabah, less common in Sarawak. Also in Indo-China, Sumatra, 

Peninsular Malaysia, and Java. 

Leaflets glabrous below; margin faintly toothed; lateral veins not sunken above ........... . 

forma littorale 

Synonyms: C. glaucum Blume I.c. (1850) 219; C. serricuspe Miq. I.c. (1859) 649; c. 

serrulatum Miq. I.c. (1859) 646; C. acutum Engl. in DC. I.c. 113; c. giganteum Engl. in 

DC.I.c. 106; c.jlavum Ridl.l.c. (1930) 81. 

Sumatra, Peninsular Malaysia, Java and Borneo. In Sabah and Sarawak common 

in the lowland mixed dipterocarp to submontane forests to 1100 m. Also found in 

Kalimantan. 

15. Canarium megalanthum Merr. 

(Greek, mega = large, anthos = flower) 

Philip. J. Sc. 30 (1926) 81; HJ. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 446; Masamune I.c. 324; 

Leenhouts I.c. (1956) 274, I.e. (1959) 370; Burgess I.c. 61; Kochummen I.c. (1972) 131; Anderson 

I.c. 155; Whitmore, Tantra & Sutisna l.c. 35. Type: Wood 1213, British North Borneo (holotype VC; 

isotypes A, K, L). 

58


Emergent tree to 35 m tall, 50 cm diameter; buttresses present. Bark grey, smooth to 

dippled and scaly; inner bark reddish brown. Sapwood yellowish white. Stipules subpersistent, 

inserted on the petiole near base, 3-4-lobed. Leaves with 4-5 pairs of leaflets; 

rachis powdery yellowish hairy; petiolules of lateral leaflets 1-1.5 cm long, powdery hairy; 

blade powdery hairy, oblong to lanceolate, 8-17 x 4-7 cm; base rounded, margin entire, 

apex pointed; midrib flattened above; lateral veins 10-12 pairs, raised on both surfaces; 

intercostal veins scalariform-reticulate, visible below. Flowers hairy, in terminal thyrsoid 

inflorescences. Fruits ellipsoid, sparsely hairy, 5-7.5 x 2.8-4.5 cm. 

Distribution. Sumatra, Peninsular Malaysia, Borneo; uncommon in Sabah and Sarawak. 

Also known in Kalimantan. 

Ecology. Ridges in mixed dipterocarp forest to 360 m. Flowering in May and fruiting in 

May and September. 

Uses. In Brunei this species is cultivated for its edible fruits which are among the largest in 

the genus. 

16. Canarium merrilIii H.I. Lam 

(E.D. Merrill, 1876-1954, American botanist) 

in Merrilll.c. (1929) 117; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 489; Masamune I.c. 364; 

Leenhouts I.c. (1956) 282, I.c. (1959) 402; Burgess l.c. 61; Anderson l.c. 156; Whitmore, Tantra & 

Sutisna I.c. 35. Type: Elmer 20326, British North Borneo, Sandakan (ho1otype PNH; isotypes A, 

BM, G, K, L, NY, P, SING, U, Z). 

Small to medium-sized tree to 25 m tall, 30 cm diameter. Bark grey, scaly; inner bark 

brownish. Sapwood white. Twigs pale grey, rough hairy. Stipules linear, to 5 mm long, 

inserted at base of petiole, semi-persistent. Leaves with 4-5 pairs of leaflets; rachis hairy; 

petiolules of lateral leaflets 2-5 mm long, hairy; blade rough hairy below (like sandpaper 

to the touch), elliptic, oblong or ob ovate, 5.5-16 x 3-5.5 cm; base cuneate, unequal, 

margin entire, apex pointed, tip c. 1 cm long; midrib faintly sunken above; lateral veins 6- 

14 pairs, curving and joining near margin, raised below, visible above; intercostsal veins 

reticulate, distinct below, faint above. Flowers yellow, in terminal or axillary thyrsoid 

inflorescences. Fruits ellipsoid, pointed, .3-4.2 x 1.2-1.5 cm. 

Distribution. Endemic to Borneo. Of scattered distribution in Sabah, Sarawak and 

Kalimantan. 

Ecology. Mixed dipterocarp to submontane forests to 1400 m. 

17. Canarium odontophyllum Miq. Fig. 1. 

(Greek, odontos = toothed, phyllon = leaf; the toothed leaflets) 

I.c. (1861) 521;.Merrilll.c. (1929) 118; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 519; Masamune 

l.c. 364; Leenhouts I.c. (1956) 271, I.c. (1959) 365; Burgess I.c. 60; Cockburn l.c. 43; Anderson I.c. 

156; Whitmore, Tantra & Sutisna I.c. 35. Type: Teijsmann HB 692, Sumatra (ho1otype U; isotype L). 

Synonyms: C. beccarii Engl. in DC. I.c. 107; C. palawanense E1mer, Leafl. Phi lip. Bot. 5 (1913) 

1754; C. multifidum H.J. Lam, Ann. Jard. Bot. Btzg. 42 (1932) 215. 

59

TREE FLORA OF SABAI-I AND SARAWAK VOL. \ (\995) 

3cm 

'cm 

~ I~.,~" ,~ 

"-- ----.:': ~ 

~" 

" 

C/;)I-, 

/' 

,cm' 

f-----I 

2cm 

?"512"1.Ao.W ~~~~,_ 

Fig. 1. Canarium odontophyllum. A, mature leaf; B, details of leaflet venation and margin; C, young 

leaflet; D, young shoot with stipules and inflorescences; E, flower bud; F, female flower; G, 

infructescence. CA, B from SAN 106079; C, D, E & F from SAN 37543; G from SAN 106079.) 

60


Tree to 30 m tall, 30 cm diameter. Bark grey-brown; inner bark brownish. Sapwood pale. 

Twigs stout, whitish, 1.5-2 cm thick, rusty hairy. Stipules persistent, irregularly toothed or 

lobed, c. 6 x 3 cm. Leaves with 3-8 pairs of leaflets; rachis yellowish hairy; petiolules of 

lateral leaflets 3-10 mm long, hairy; blade densely velvety hairy below, oblong to 

lanceolate, 9.5-28 x 4-11 cm; base broadly cuneate to rounded, margin toothed, apex 

pointed; midrib raised above, hairy; lateral veins 12-23 pairs, raised below, faint to sunken 

above; intercostal veins scalariform-reticulate, raised below, faint to sunken above. Flowers 

hairy, in axillary or terminal thyrsoid inflorescences. Fruits ovoid to ellipsoid, 3-4 x 2.5-3 

Cill. 

Distribution. Sumatra, Borneo, and the Philippines. Widely distributed in Sabah, uncommon 

in Sarawak. Also known in Kalimantan. 

Ecology. Mixed dipterocarp forests on fertile clay soils to 500 m. Flowering in May, June 

and November and fruiting in March and August. 

Uses. Widely cultivated in Sarawak and Brunei for its fruits. 

18. Canarium patentinervium Miq. 

(Latin, patens = spreading, nervus = nerves; the leaflet venation) 

I.e. (1861) 526; H.J. Lam, Bull. Jard. Bot. Btzg. 3,12 (1932) 506; Masamune I.e. 365; Leenhouts I.e. 

(1956) 258, I.e. (1959) 342; Burgess I.e. 61; Kochurnmen I.e. (1972) 134; Whitmore, Tantra & 

Sutisna I.e. 36. Type: Teijsmann HE 3736, Sumatra, Palembang (holotype V; isotype L). Synonyms: 

C. nitidum AW. Berm. in Hooker! I.e. 533; c. parvifolium AW. Berm. in Hooker! I.e. 536. 

Tree to 24 m tall, 25 cm diameter. Bark grey, smooth. Twigs brownish, lenticellate. Stipules 

falling off early, kidney-shaped. Leaves with 2-4 pairs of leaflets; petiolules of lateral 

leaflets 1-1. 5 cm long, cracking; blade thickly leathery, glabrous, drying to dark brown, 

ovate to oblong, 7-12.5 x 2.5-6 cm; base cuneate, margin entire, apex pointed; midrib 

flattened above, rarely sunken; lateral veins 7-10 pairs, raised below, faint above, curving 

and joining near margin; intercostal veins reticulate, faintly visible on both surfaces, 

sometimes inconspicuous. Flowers hairy, usually in terminal thyrsoid inflorescences. Fruits 

ellipsoid, 6-6.5 x 2.2-3 cm, apex pointed. 

Distribution. Sumatra, Peninsular Malaysia, and Borneo. Widespread, collected from Tenom, 

Tawau and Tuaran in Sabah, and from Lundu, Semengoh to Bukit Lambir in Sarawak. Also 

found in Kalimantan. 

Ecology. Mixed dipterocarp forest on yellow sandy and leached clay soils, and kerangas 

forest, to 450 m. 

19. Canarium pilosum AW. Benn. 

(Latin, pilosus = having soft and distinct hairs; the twigs and leaves) 

in Hooker! I.e. 533; King I.e. 243; Ridley I.e. (1922) 372; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 

(1932) 472; Masamune I.e. 365; Leenhouts l.c. (1956) 281, I.e. (1959) 398; Burgess I.e. 61; 

61


Kochummen I.e. (1972) 134; Anderson I.c 156; Whitmore, Tantra & Sutisna I.c. 36. Type: Maingay 

3103, Malacca (holotype K; isotype CAL). 

Small to medium-sized tree to 27 m tall, 25 cm diameter. Bark pale brown, smooth; inner 

bark pink. Sapwood white. Twigs densely woolly hairy. Stipules persistent, linear, inserted 

at the base of petiole or on the petiole, rarely absent. Leaves with 2-3 pairs of leaflets; 

rachis densely hairy, rarely glabrous; petiolules of lateral leaflets 2-3 mm long, densely 

hairy, rarely glabrous; blade densely hairy below, sparsely hairy above, rarely completely 

glabrous, elliptic, oblong or obovate, 7-19 x 3.5-9 cm; base cuneate or rounded, margin 

faintly to distinctly toothed, rarely entire, apex pointed, tip c. 1 cm long; midrib raised 

above; lateral veins 7-14 pairs, raised on both surfaces, curving and joining near margin; 

intercostal veins scalariform-reticulate, raised below, faint above. Flowers hairy, in axillary 

or terminal panicles. Fruits oblong, 2-3 x 1-1.5 cm. 


Stipules present. Leaflets hairy below, margin toothed ................ . 

subsp. piIosum 

Synonyms: C. grandifolium AW. Berm. in Hooker! I.e. 533; C. hirtellum AW. Berm. in 

Hooker! I.e. 533; C. pilosum var. hirtellum (AW. Berm.) RidI. I.e. (1922) 372; C. motleyanum 

EngI. in DC. I.c. 133. 

Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak uncommon, recorded 

from Beaufort, Beluran, Papar, and Sandakan (Sabah), and from Kapit, Marudi, Miri 

(Sarawak). Also known in Kalimantan. Lowland forest. 

Stipules absent. Leaflets glabrous below, margin entire ..................................... . 

subsp. borneensis Leenh. 

I.e. (1955) 193. Type: Clemens 40163, British North Borneo, Mt. Kinabalu (holotype L; isotypes 

A, BM, BO, G, NY). Synonym: Daeryodes seandens Husson, Blumea 7 (1952) 164. 

Uncommon, known from Mt. Kinabalu, Lahad Datu and Sandakan in Sabah, and G. 

Lambir in Sarawak. Also found in Kalimantan. 

20. Canarium pseudodecumanum Hochr. 

(Latin, pseudo = false; resembling C. decumanum) 

PI. Bog. Exs. (1904) 61; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 438; Leenhouts l.c. (1956) 

275, I.e. (1959) 388; Burgess I.e. 61; Kochummen l.c. (1972) 135; Whitmore, Tantra & Sutisna I.e. 

36. Type: Hart. Bot. Bog. VI E 6a, Sumatra, Lampong (holotype L; isotypes B, BO, BRSL, G, K, L, 

WY, P). Synonym: Canarium deeumanum (non Gaertn.) EngI. in Merrilll.e. (1929) 116. 

Tree to 40 m tall and 120 cm diameter; buttresses to 6 m high. Bark grey-white, smooth to 

scaly; inner bark pale. Sapwood yellowish white. Twigs stout, 1-1.5 cm thick, with large 

leaf-scars. Terminal bud 4-5 cm long, densely brown tomentose. Stipules absent. Leaves 

with 4-6 pairs of leaflets; petiole stout, light brown pubescent, swollen at base and flattened 

above; blade almost sessile, lanceolate or oblong, 17-23 x 7-11 cm, glabrous above except 

the midrib and lateral veins, densely minutely woolly tomentose below; base rounded or 

62


sub cordate, margin minutely toothed, apex pointed; midrib flattened to raised above; lateral 

veins 17-23 pairs, distinctly arching near margin, prominent below; intercostal veins 

prominent giving pitted appearance to the lower surface, veins and reticulations raised 

above. Inflorescences (male) axillary thyrses. Flowers pubescent. Fruits ellipsoid, 7-8 x 

4-6 cm, densely tomentose when young. 

Distribution. Sumatra, Peninsular Malaysia, and Borneo. In Sabah recorded from the Ranau, 

Sandakan and Tawau districts; not recorded from Sarawak. Also known in Kalimantan. 

Ecology. Lowland forest to 280 m. 

Uses. The resin is used for caulking boats. The fruit is edible. An edible oil is extracted 

from the seeds. 

Closely related to C. decumanum but differing in the absence of stipules and by the toothed 

and densely woolly tomentose lower surface of the leaflets. 

21. Canarium pseudopatentinervium H.J. Lam 

(resembling c. patentinervium) 

Ann. Jard. Bot. Btzg. 42 (1932) 214, Bull. Jard. Bot. Btzg. 3, 12 (1932) 490; Masamune I.c. 365; 

Leenhouts I.c. (1956) 274, l.c. (1959) 385; Burgess I.c. 61; Anderson I.c. 155; Whitmore, Tantra & 

Sutisna I.c. 36. Type: bb. 153, T. 3P. 336, Sumatra, Palembang, Lamatang Ilir near G. Megang 

(lectotype BO). 

Medium-sized to large tree to 45 m tall, 100 cm diameter; buttresses to 5 m tall. Bark pale 

yellow-brown, irregularly scaly to dippled. Twigs whitish, with prominent leaf-scars, pith 

with many peripherally arranged vascular strands and often with a central cavity. Stipules 

absent. Leaves with 1-4 pairs of leaflets; blade drying greenish, ovate to elliptic, 5-12 x 

2-7 cm; base asymmetric, rounded to broadly cuneate, margin entire, apex blunt to pointed; 

rnidrib flattened above; lateral veins 7-14 pairs, curving near margin; intercostal veins 

reticulate, prominent on both surfaces; petiolules of lateral leaflets 5-9 mm long, not 

swollen at both ends. Inflorescences (female) in terminal panicles. Flowers (female) subsessile, 

densely tomentose, petals distinctly clawed, apex inflexed, stamens free. Fruits 

ellipsoid, 5-7 x 2-3 cm; calyx triangular, hairy inside. 

Distribution. Sumatra, Borneo. In Sarawak uncommon, known only by a single collection 

S. J 82 J 5 from Belaga; not yet found in Sabah. Also occurs in Kalimantan. 

Ecology. Lowland forest at c. 100 ill. 

') 

22. Canarium pseudopimela Kochummen 

(resembling c. pimela) 

l.c. (1994) 73. Type: Ding Hou 489, Sarawak, G. Raya (holotype SAR; isotype L). 

Small tree to 12 m tall, 15 cm diameter. Twigs brown, c. 3 mm thick, rounded. Stipules 

absent. Leaves with 2-3 pairs ofleaflets; blade thinly leathery, elliptic to narrowly obovate, 

63


6-13 x 2.5-5 cm, slightly falcate; base cuneate, margin entire, apex cuspidate, tip 1. 5-2 

cm long; midrib raised above; lateral veins 5-7 pairs, arching and joining near margin, 

visible below, faint above; intercostal veins finely reticulate, distinctly visible below, faint 

above; petiolules of lateral leaflets 4-5 mm long, petiolules of terminal leaflet 2-3.5 cm 

long. Inflorescences (female) in racemes, terminal, glabrous, c. 23 cm long, with l3-14 

cm long branches in few-flowered racemes, buds obovate. Flowers (female) c. 9 mm long, 

stalk c. 5 mm long; calyx funnel-shaped, with obtuse lobes, fleshy, with few irregular lines 

towards base; petals fleshy, irregularly wrinkled on drying; rudimentary stamens 6, 

connected into a cup-shaped staminal ring; disc undulate, c. 1 mm high; ovary hairy, 

faintly ridged, stigma capitate. Infructescences c. l3 cm long. Fruits ellipsoid, 3 x 1-1. 5 

cm, triangular, apex narrowed to sharp point; calyx almost flat; stalk c. 1.5 cm long. 

Distribution. Endemic to Borneo. Uncommon, known by two collections only from G. 

Lambir (S. 16613) and G. Raya (Ding Hou 489) in Sarawak. 


Very close to C. pimela but differing in the terminal inflorescences, hairy ovary and in the 

long-pointed leaflets. 

23. Canarium sarawakanum Kochummen 

l.c. (1994) 75. Type: Au S. 24802, Sarawak, Kapit (holotype SAR; isotypes A, BO, K, KEP, L, SAN, 

SING). 

Small tree to 8 m tall, 10 cm diameter. Twigs reddish brown hairy when young. Stipules 

absent. Leaves with 1-2 pairs of leaflets; rachis finely reddish brown hairy; blade thinly 

leathery, drying to reddish brown, glabrous above, sparsely reddish brown hairy below, 

elliptic to· narrowly obovate, 8-20 x 3.5-8.5 cm; base cuneate, margin sub-entire to faintly 

toothed, apex acuminate, tip 1-2 cm long; midrib raised above; lateral veins 6-8 pairs, 

distinct below, faintly raised above, curving and joining near margin; intercostal veins 

reticulate, visible below, faint above; petiolules of lateral leaflets swollen near the apex, 1- 

1.5 cm long, that of terminal leaflet to 5 cm long. Inflorescences (female) axillary or 

terminal, thyrsoid, to 22 cm long, with few side branches; rachis brownish hairy. Flowers 

(male) with the calyx hairy outside, margins wavy to lobed; petals oblong, c. 4.5 mm long, 

hairy outside; stamens 3, joined at the base, filaments to 2.5 mm long; rudimentary ovary 

and style present. Fruits (immature) ellipsoid, 4.5-5.5 x 1.7-2 cm. 

Distribution. Endemic to Borneo. Uncommon, known from a few collections from Kapit, 

Lubok Antu, G. Penrissen and Simanggang in Sarawak. 

Ecology. Lowland to low submontane forests to 900 m. 

Close to C. latistipulatum but that species has semi-persistent stipules. 

64


2. DACRYODES Vahl 

(Greek, dakruon = a tear; the resin droplets on the bark surface) 

kedondong (Sabah and Sarawak Malay), kimayau (Bidayuh in Sarawak) 

kembayau (Sabah Dusun), seladah (Sarawak Iban) 

Skrift. Dansk. Nat. Hist. Selsk. 4 (1810) 116; H.J. Lam, Bull. Jard. Bot. Btzg. 3,. 12 (1932) 334; 

Kalkman, Blumea 7 (1954) 500; Leenhouts I.e. (1956) 219, I.e. (1972) 917, I.e. (1976) 820; Backer 

& BakhuizenJ I.e. 114; Kochummen I.e. (1972) 136; Cockbum i.e. 43; Anderson i.e. 156; Wong I.e. 

42; Whitmore, Tantra & Sutisna I.e. 36; Ng I.e. 37. Synonyms: Paehylobus G. Don, Gen. Syst. 2 

(1832) 89; Canarium section Tenuipyrena Engl. in DC. I.e. 104; Curtisiana Ridl., J. Str. Br. R. As. 

Soc. 82 (1920) 180; Hemisantiria H.J. Lam in Merrilll.e. (1929) 118. 

Small to medium-sized, dioecious trees; buttresses small, short; sometimes with stilt-roots. 

Bark smooth to scaly; inner bark orange, yellowish white or pink, exudate often clear, 

rarely white. Sapwood pale. Stipules absent (in D. laxa the first pair of leaflets are smaller 

and stipule-like). Leaflets entire, the base often strongly unequal; stalks (petiolules) strongly 

swollen at both ends. Inflorescence an axillary or terminal panicle. Flowers unisexual, 3- 

merous; sepals free or united; petals usually with thickened injIexed apex; stamens 6, 

filaments free, bases united with the disc; disc intrastaminal, glabrous; ovary 3(-2)-celled, 

usually moderately reduced in male flowers; stigma sessile. Fruit a drupe, oblong or 

ellipsoid, I-seeded; stigma usually apical; pericarp fleshy and thick, coarsely wrinkled 

when dry, glabrous; stone containing one fertile and two reduced cells; calyx persistent or 

caducous. Seeds round in cross-section; cotyledons 9-11-1obed in D. rostrata, and 5-10bed 

in D. costata, D. laxa and D. rugosa. 

Distribution. About 40 species; tropical America, Africa and Asia. In Malesia mainly 

centred in Sumatra, Peninsular Malaysia and Borneo; 11 species in Sabah and Sarawak. 

Ecology. Lowland (including swamp) to submontane forests to 1500 m. 

Uses. The timbers of Dacryodes are essentially similar to that of Canarium in anatomical 

structure but they are on the whole heavier and the rays contain silica which makes them 

difficult to saw. Fruits of Dacryodes rostrata f. cuspidata, known in Sarawak as keramoh 

(Malay) and kembayau (Murut and Iban), are eaten. 

Key to Dacryodes species 

l. Rachis and young twigs hairy .......................................................... . 

Rachis and young twigs glabrous ...................................... . 

2. Rachis and young twigs covered with long stiff hairs ....................... . 

Rachis and young twigs powdery,hairy ......................................... . 

.. ..... 2 

.. ..... 7 

.5. D. laxa 

.. ..... 3 

3. Ultimate leafy twigs c. 1 cm thick. Terminal bud c. l.5 cm long ................. 2. D. elmeri 

Ultimate leafy twigs to 0.5 cm thick. Terminal bud to 1 cm long .................................. 4 

65

TREE FLORA OF SABAH AND SARAW AI< VOL. 1 (1995) 

4. Intercostal veins prominently raised below. Hairs not yellowish ................................... 5 

Intercostal veins not raised, but visible below. Hairs yellowish ........... l0. D. rubiginosa 

5. Intercostal veins scalariform. Leaflets densely hairy below ....................... 8. D. nervosa 

Intercostal veins reticulate. Leaflets sparsely hairy below ............................................. 6 

6. Petiolules strongly swollen at both ends; rachis glabrous .......................... 11. D. rugosa 

Petiolules not strongly swollen at both ends; rachis hairy ........................... 1. D. costata 

7. Adaxial side of petiole flattened with sharp edges, petiolules of lateral leaflets 2.5-3.5 

cm long .................................................................................................... 3. D. expansa 

Petiole and petiolules not so ......................................................................................... 8 

8. Midrib below sharply keeled; petiole strongly grooved or flattened above ...................... . 

... 6. D. longifolia 

Midrib below not keeled; petiole rounded or flattened above ........................................ 9 

9. Reticulations fine and more distinct on the upper than on the lower leaflet surface. 

Inner bark reddish; cut bark and twigs with white sap .............................. 9. D. rostrata 

Reticulations more distinct on the lower surface. Inner bark yellowish; cut bark and 

twigs without white saD .............................................................................................. 10 

10. Leaflets usually of 2 pairs only. Trees with stilt-roots ........................ 7. D. macrocarpa 

Leaflets more than 2 pairs. Trees without stilt-roots ................................................... 11 

11. Lateral veins sunken above; intercostal veins raised below. Petiolule strongly swollen 

at both ends .............................................................................................. 11. D. rugosa 

Lateral veins not sunken above; intercostal veins faintly visible below. Petiolule not 

prominently swollen .............................................................................. 4. D. incurvata 

1. Dacryodes costata (AW. Benn.) HJ. Lam 

(Latin, costatus = ribbed; probably the prominent veins on the lower leaflet surface) 

Ann. Jard. Bot. Btzg. 42 (1932) 204, Bull. Jard. Bot. Btzg. 3, 12 (1932) 359; Masamune l.c. 366; 

Kalkman I.c. (1954) 508; Leenhouts I.c. (1956) 222; Burgess l.c. 61; Kochummen I.c. (1972) 140; 

Anderson l.c. 156; Whitmore, Tantra & Sutisna l.c. 36. Basionym: Santiria costata AW. Benn. in 

Hooker f I.c. 537. Type: Maingay 313/3295, Malacca (holotype K; isotype CAL). Synonym: 

Canarium costatum (AW. Benn.) Ridl.l.c. (1930) 82. 

Medium-sized to large tree to 45 m tall, 45 cm diameter; buttresses short. Bark greybrown, 

smooth to flaky; inner bark yellow-brown, with droplets of white sap. Sapwood 

whitish. Ultimate leafy twigs to 0.5 cm thick, rusty-brown velvety hairy when young. 

Leaves with 1-3 pairs of leaflets; rachis powdery rusty hairy; petiole hairy, flattened 

above; petiolules of lateral leaflets 5-15 mm long, powdery hairy to glabrous, slightly 

swollen at both ends; blade sparseZy hairy on midrib and veins below, elliptic to oblong, 6- 

15 x 2.5-6 cm; base cuneate, often unequal, apex pointed, tip 1-2 cm long; midrib raised 

above; lateral veins 10-14 pairs, prominently raised below, faint above, looping and joining 

66


near margin; intercostal veins reticulate, distinct below, faint above. Inflorescences 

terminal or from upper leafaxils, axes densely hairy. Flowers pubescent; petals whitish, 

glabrous; stamen filaments free from the disc; disc annular to 6-lobed. Fruits ellipsoid or 

ovoid, l.7-2.2 x 1-l.2 cm; stalkc. 1 cm long. 

Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sabah 

reported from Ranau, Sandakan and Tawau; in Sarawak, known only from the Lambir 

National Park and Semengoh FR. Also known in Kalimantan. 

Ecology. Widely distributed from lowland to hill forests to 540 m, mainly on ridges and 

hillsides. Flowering in May and September, and fruiting in July, October and November. 

This species is very similar to D. rugosa from which it can be distinguished by the hairy 

petioles, and the less swollen leaflet stalks. The flowers of both species are quite distinct, in 

D. costata the disc is annular to 6-lobed and the stamen filaments free from disc, while in 

D. rugosa the disc is cup-shaped and stamen filaments adnate to disc. 

2. Dacryodes elmeri H.J. Lam 

(A.D.E. Elmer, 1870-1942; plant collector with the Bureau of Science, Manila, the 

Philippines) 

Ann. Jard. Bot. Btzg~ 42 (1932) 203, Bull. Jard. Bot. Btzg. 3, 12 (1932) 344; Kalkman I.c. 

(1954) 521; Leenhouts l.c. (1956) 225. Synonym: Hemisantria ?n. sp. H.J. Lam in Merrill 

I.c. (1929) 199. Type: Elmer 21573, British North Borneo, Tawau (holotype L; isotypes A, 

BKH,BM). 

Tree, to 75 cm in diameter. Ultimate leafY twigs c. 1 cm thick, densely minutely villous; 

pith with many scattered vascular strands. Terminal bud c. l.5 cm long, densely pubescent. 

Leaves with 3-4 pairs of leaflets; rachis strongly flattened at base, hairy at first becoming 

glabrous; blade oblong, 12-22 x 5-8.5 cm, chartaceous; base oblique-cuneate, on one half 

rounded, apex pointed; lateral veins 13-18 pairs, prominent beneath. Inflorescence and 

flower unknown. Fruits ovoid, 4-4.7 x 2.25 cm, with rounded apex and base. 

Distribution. Endemic to Borneo. Very uncommon, known from the type collection (Sabah) 

and from Brunei. 

Ecology. Lowland forest. 

3. Dacryodes expansa (Ridl.) H.J. Lam 

(Latin, expansus = spread out, diffuse; the thin, spread out petals) 

Ann. Jard. Bot. Btzg. 42 (1932) 204, Bull. Jard. Bot. Btzg. 3, 12 (1932) 366; Masamune I.c. 366; 

Kalkman I.c. (1954) 510; Leenhouts I.c. (1956) 228, l.c. (1972) 919, l.c. (1976) 821. Basionym: 

Canarium expansum Ridl.l.c. (1930) 83. Type: Haviland 2271, Sarawak, Kuching (K). 

67


Small tree. Buds reddish brown, hairy. Leaves with 4 pairs of leaflets; adaxial side of 

petiole flattened with sharp edges; petiolules of lateral leaflets 2.5-3.5 cm long, strongly 

swollen at both ends; blade oblong to oblong-lanceolate, 17-23 x 6-7.5 cm, brownish when 

dried; base cuneate, apex pointed; midrib raised above; lateral veins 10-12 pairs, prominent 

below; intercostal veins faint. Inflorescences (male) probably lateral on axillary shoots, to 

24 cm long, glabrous; pedicels 3-7 mm long, articulated. Flowers (male) glabrous; calyxlobes 

deltoid; petals very thin, spreading; stamens free from disc; disc annular, thick; ovary 

in male flowers reduced. Infructescence and fruit unknown. 

Distribution. Endemic to Borneo. Uncommon, known from the type collection only and 

from Brunei. 


4. Dacryodes incurvata (Engl.) H.J. Lam 

(Latin, incurvatus = bending inward; the leaflet margin) 

Ann. Jard. Bot. Btzg. 42 (1932) 204, Bull. Jard. Bot. Btzg. 3, 12 (1932) 362; Ka1kman l.c. (1954) 

506; Leenhouts l.c. (1956) 224; Burgess I.c. 61; Koehurnrnen l.c. (1972) 140; Anderson I.c. 156; 

Whitmore, Tantra & Sutisna l.c. 36. Basionym: Canarium incurvatum Engl. in DC. I.c. 138. Type: 

Eeccari PE 2937, Sarawak (holotype P; isotype FI). Synonyms: Santiria nitida Merr., Publ. Govt. 

Lab. Philip. 35 (1906) 29; Canarium nitens Merr., Philip. J. Se. 10 (1915) Bot. 24; Hemisantiria 

nitida H.J. Lam in Merrill I.c. (1929) 118; Canarium angulatum Ridl., Kew Bull. (1931) 493; 

Dacryodes angulata (Ridl.) HJ. Lam, Ann. Jard. Bot. Btzg. 42 (1932) 204. 

Medium-sized to large tree to 35 m tall, 30 cm diameter; buttresses steep to 2 m high. Bark 

grey-brown, smooth, with horizontal rings; inner bark pink with droplets of white sap. 

Sapwood pale white. Twigs dark brown. Leaves with 1-4 pairs of leaflets; petiole strongly 

flattened at base; petiolules of lateral leaflets 1-2 cm long, swollen at both ends; blade 

thinly to thickly leathery, shiny above, elliptic, oblong or lanceolate, 6-16 x 2-9 cm; base 

rounded, subcordate or cuneate, apex pointed; midrib raised or flattened above; lateral 

veins 10-16 pairs, raised on both surfaces; intercostal veins reticulate, faintly visible on 

both surfaces. Flowers hairy, usually in terminal panicles. Fruits ovoid or ellipsoid, 2.5-3 

x 1.7-2 cm, ripening to yellow and then purplish. 

Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. Common in Sabah, 

recorded from Beaufort, Sipitang and Tawau. In Sarawak recorded from Bintulu, Lambir 

National Park, Serian and Kuching. Also found in Brunei and Kalimantan. 

Ecology. Mixed dipterocarp forests to 860 m; uncommon in mixed peat swamp forest. 

Flowering in April, June and September, and fruiting in March and June-November. 

5. Dacryodes laxa (AW. Benn.) HJ. Lam 

(Latin, laxus = loose or distant; the inflorescence) 

Ann. Jard. Bot. Btzg. 42 (1932) 204, Bull. Jard. Bot. Btzg. 3, 12 (1932) 355; Kalkman I.c. (1954) 

503; Leenhouts I.c. (1956) 224; Koehurnrnen I.c. (1972) 141; Coekburn I.c. 45; Anderson I.c. 156; 

Whitmore, Tantra & Sutisna I.c. 37. Basionym: Canarium laxum A.W. Benn. in Hooker f I.c. 535. 

68


Type: Maingay 366, Malacca (holotype K; isotypes CAL, L). Synonyms: Canariumfragile Eng!. in 

DC. I.c. 138; Santiria laxa (AW. Benn.) King, I.c. 254. 

Small to medium-sized tree to 30 m tall, 30 cm diameter. Bark grey, smooth to scaly; inner 

bark pale yellow. Sapwood pale white. Twigs with long rough hairs when young. Leaves 

with up to 5 pairs of leaflets; rachis rough-hairy; petiolules of lateral leaflets 3-10 mm 

long, hairy; blade drying gretnish, oblong, lanceolate or oblanceolate, 8.5-34 x 4.5-9 cm; 

base cuneate, apex pointed, tip to 1.5 cm long; midrib raised above; lateral veins 9-30 

pairs, raised on both surfaces, curving and joining near margin; intercostal veins reticulate, 

raised below, sunken above; petiole strongly swollen at base. Flowers in branched, long (c. 

80 cm) terminal panicles; petals glabrous. Fruits pink ripening to blue, oblong or ovoid, 

2.5-4.5 x 1-2 cm, apex pointed; stalk 2-2.5 cm long, slender. 

Distribution. Sumatra, Peninsular Malaysia, and Borneo. Widely distributed in Sabah. In 

Sarawak reported only from Bako National Park, Semengoh Arboretum and Lundu. Also 

known in Brunei. 

Ecology. Common in mixed dipterocarp forests on yellow sandy clay soils, to 700 m. 

Flowering from April to June, and fruiting from June to November. 

6. Dacryodes longifolia (King) H.J. Lam 

(Latin, longus = long,jolium = leaf) 

Ann. lard. Bot. Btzg. 42 (1932) 202, Bul!. lard. Bot. Btzg. 3, 12 (1932) 340; Kalkman I.c. (1954) 

509; Leenhouts I.c. (1956) 228, I.c. (1972) 919; Kochummen I.c. (1972) 141. Basionym: Santiria 

longifolia King I.c. 258. Type: King's collectors 6838, Perak (holotype K; isotype L). Synonyms: 

Curlisiana penangensis Rid!. I.c. (1920) 180; Dacryodes longifolia var. penangensis (Rid!.) H.J. 

Lam, Ann. lard. Bot. Btzg. 42 (1932) 202. 

Medium-sized to large tree to 40 m tall. Bark dark grey, smooth; inner bark orange-yellow. 

Twigs brown, with scattered white lenticels. Leaves with 2-6 pairs of leaflets; petiole 

strongly grooved or flattened above; petiolules of lateral leaflets 1-2 cm long, strongly 

swollen at both ends; blade oblong to lanceolate, 6.5-18 x 2.5-4.5 cm; base usually 

unequal, broadly cuneate to rounded, apex pointed; midrib raised above, sharply keeled 

below; lateral veins 7-12 pairs, distinct below, faint above; intercostal veins reticulate, 

faintly visible on both surfaces. Flowers (male) in axillary panicles, glabrous. Fruits ellipsoid, 

2.5-3.5 x 1.5-2.5 cm, much wrinkled on drying; stalk c. 7 mm long. 

Distribution. Sumatra, Peninsular Malaysia, Borneo and the Philippines. Common and 

widely distributed in Sabah; less common in Sarawak. 

Ecology. Lowland to submontane forests to 1500 m. 

7. Dacryodes macrocarpa (King) H.I. Lam 

(Greek, makros = large, karpos = fruit) 

Ann . .lard. Bot. Btzg. 42 (1932) 203, Bul!. lard. Bot. Btzg. 3, 12 (1932) 342; Masamune I.c. 366; 

Kalkman I. c. (1954) 514; Leenhouts I. c. (1956) 228, I. c. (1972) 919, I. c. (1976) 820; Burgess l. c. 61; 

69


] 2 cm 

2cm [ 

B 

Fig. 2. Dacryodes rostrata. A, fruiting leafy twig; B, part of inflorescence. CA from SAN 39742, B 

from SAN 30412.) 

70

I3URSERACEAE (KOCHUMMEN) 

Kochummen I.e. (1972) 142; Anderson I.e. 156; Whitmore, Tantra & Sutisna I.e. 37. Basionym: 

Santiria maeroearpa King I.e. 256. Type: King's collectors 7298, Perak (holotype K; isotypes CAL, 

SING). 

Medium-sized to tall tree; stilt-roots and buttresses present. Bark reddish brown, cracking. 

Twigs dark brown to blackish when dry. Leaves with 1-2 pairs of leaflets; petiolules of 

lateral leaflets 0.7-2 cm long, swollen at both ends; blade elliptic, obovate or ovate, 6-12 x 

3-6 cm; base often unequal, rounded to broadly cuneate, apex blunt, rounded or shortly 

pointed; midrib flattened above; lateral veins 6-9 pairs, visible on both surfaces; intercostal 

veins scalariform-reticulate, faintly visible on both surfaces. Flowers glabrous, in axillary 

panicles. Fruits ovoid or ellipsoid, 2.5-4 x 2.5 cm. 


1. Leaflets widest around the middle, equal-sided at base; lateral veins at right angle to 

midrib ................................................................................................. . 

var. patentinervia Leenh. 

I.e (1976) 821. Type: Sinclair & Kadim 10492, Brunei, Bt. Labi FR (holotype L; isotypes 

K, SAR, SING). / 

Endemic to Borneo. In Sabah uncommon, known only from Bt. Hampuan, Ranau 

district (SAN 25331). In Sarawak scattered throught 3rd, 4th and 5th Div. in the 

lowland to submontane forest to 1500m. Also in Brunei. Vernacular names: 

Sarawak-seladah (Iban). Brunei-sabal (Iban), sibut (Tutong, Dusun). 

Leaflets widest in the lower half, oblique at base; lateral veins at acute angle with 

midrib .......................................................................................................................... 2 

2. Twigs blackish when dry. Midrib and veins not prominent beneath ... 

var. macrocarpa 

Synonym: D. maeroearpa var. genuina H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 343. 

Sumatra, Peninsular Malaysia, and Borneo. Uncommon in Sabah, more common 

in Sarawak. Also in Brunei and Kalimantan. Vernacular names: Sabah---asamasam 

(Dusun). Sarawak-icerawas burung, keruas (Malay). Brunei---kedondong, 

pasoh-pasoh (Malay). 

Twigs and rachis scaly to powdery hairy and brown when dry. Midrib and veins 

prominent below ... 

var. kostermansii (Kalkman) Kalkman 

in Leenhouts I.e. (1956) 228. Synonym: D. kostermansii Kalkman I.e. (1954) 515. Type: 

Kostermans 6725, SE Borneo, Loa Djanan, W Samarinda (holotypc L; isotypes DO). 

Endemic to Borneo. In Sabah known by a single collection (SAN 25572) from the 

Silam FR, Lahad Datu district; not recorded in Sarawak. Also found in Kalimantan. 

Ecology. Coastal and mixed swamp forests. Flowering in September and fruiting in 

March-April and November. 

71


8. Dacryodes nervosa (H.J. Lam) Leenh. 

(Latin, nervosus = prominently nerved; the leaves) 

Blumea 12 (1964) 19, l.c. (1972) 919. Basionym: Santiria nervosa H.J. Lam, Ann Jard. Bot. Btzg. 

42 (1932) 206, Bull. Jard. Bot. Btzg. 12,3 (1932) 387. Type: . GrashojJ 960, Sumatra (L). 

Medium-sized tree to 30 m tall, 35 cm diameter; buttresses to 1.5 m tall. Bark smooth to 

scaly, dark brown. Ultimate leafy twigs to 0.5 cm thick. Terminal bud 0.5-1 cm long. 

Leaves with 1-4 pairs ofleaflets; blade oblong to ovate, 5.5-17 x 3-7.5 cm, upper surface 

greenish when dried, lower surface pubescent to glabrous; base broadly cuneate, asymmetric, 

apex pointed; midrib raised above; lateral veins 10-15 pairs, prominent below; 

intercostal veins scalariform, raised below; petiolule 8-11 mm long, hairy. Inflorescences 

axillary. Flowers sessile or shortly stalked. Fruits pink when fresh, ellipsoid, 1.6-1.7 x 1.1 

cm. 

Distribution. Sumatra, Peninsular Malaysia, and Borneo. Uncommon, in Sarawak so far 

recorded only from Bako National Park, Bintulu and Sadong (S 13389, S16228, S18192, 

S 37488); not yet found in Sabah. Also occurs in Kalimantan. 

Ecology. Lowland forest to 350 ill. 

9. Dacryodes rostrata (Blume) HJ. Lam Fig. 2. 

(Latin, rostratus = :with a beak or narrowed tip; the leaves) 

Ann. Jard. Bot. Btzg. 42 (1932) 203, Bull. Jard. Bot. Btzg. 3, 12 (1932) 349; Masamune l.c. 366; 

Ka1kman l.c. (1954) 519; Leenhouts I.c. (1956) 225; Burgess l.c. 161; Kochummen I.c. (1972) 143; 

Cockburn I.c. 45; Anderson I.c. 156; Whitmore, Tantra & Sutisna l.c. 37. Basionym: Santiria 

rostrata Blume l.c. (1850) 213. Type: Korthals, s.n. (= Leiden no. 898.321-228), Borneo (L). 

Synonym: Hemisantiria rostrata (Blume) H.J. Lam in Merrilll.c. (1929) 119. 

Small to very large tree to 45 m tall, 200 cm diameter. Bark dark grey, smooth to scaly or 

dippled; inner bark reddish or pinkish with droplets of white sap. Sapwood whitish. Twigs 

thin to very thick (0.4~2.5 cm), dark brown, lenticellate. Leaves with 2-8 pairs of leaflets; 

petiolules of lateral leaflets 1-2 cm long, swollen at both ends; blade thinly to thickly 

leathery, glabrous or (sometimes) with tiny inconspicuous hairs below, ovate to oblong, 8- 

30 x 3.5-11 cm; base strongly unequal, apex pointed, tip c. 2 cm long; midrib raised above; 

lateral veins 5-15 pairs, raised on both surfaces; intercostal veins reticulate, faint to distinct 

on both surfaces or sometimes more distinct above than below; petioles strongly flattened at 

base in those with thick twigs. Flowers densely hairy, in axillary panicles 5-35 cm long. 

Fruits ovoid to oblong, 2-3.5 x 1-2 cm, often strongly wrinkled on drying, ripening to 

blue, with white sap. 

Key to forms 

Ultimate leafy twigs 1.5-2.5 cm thick. Leaflets at least 10 cm long, drying greenish grey; 

reticulations more distinct on the upper than on lower surface ............................ . 

72


forma cuspidata (Blume) H.J. Lam 

Bull. Jard. Bot. Btzg. 3, 12 (1932) 351. Basionym: Draeontomelon euspidatum Blume I.e. 

(1850). Type: Korthals, s.n. (= Leiden no. 897. 363-270), Borneo (L). Synonym: Canarium 

minahassae Koord. I.e. 96; Santria samarensis Merr. I. e. (1915) 315; C. erassifolium Merr. I. e. 

(1915) 274; C. euspidatum (Blume) Merr. I.e. (1921) 316; C. retieulatum Ridl. I.e. (1930) 83. 

lndo-China, Sumatra, Borneo, Philippines, and N Celebes. In Sabah, known from a 

few collections from Labuan FR (FDBNB 44236), Bt. Batangan, Sipitang district (SAN 

16612), and Nabutan, Ranau district (SAN 100230). In Sarawak recorded from Bt. 

Rawan (S 45575) and Semengoh Arboretum (S 32406) in 1st Div., Ulu Balleh, Kapit, 

3rd Div. (S 29100); and Ulu Dapoi, Marudi (S 23056 & S 23474), Kebulu Protected 

Forest (S 49126), and Dulit Range (S 46672) in the 4th Div. Vernacular names: 

Sabah-kedondong, salong banggi (Malay). Sarawak-seladah (Iban). 

Ultimate leafy twigs to 0'.5 cm thick. Leaflets smaller, drying reddish brown; reticulations 

not so ....................................................................... . 

forma rostrata 

Synonyms: D. rostrata f. genuina H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 351; Santiria 

montana Blume I.e. (1850) 212; C. montanum (Blume) Korth. ex Miq. I.e. (1859) 649: C. 

kedondon A.W. Benn. in Hooker! I.e. 535. 

Sumatra, Peninsular Malaysia, and Borneo. Common and widely distributed in Sabah 

and Sarawak. Vernacular names: Sabah-kedondong, kambayau (Dusun). 

Ecology. Very common in mixed dipterocarp forest to 800 m. Flowering in February 

September, and fruiting in April and June-December. 

10. Dacryodes rubiginosa CAW. Benn.) H.J. Lam 

(Latin, rubiginosus = of rusty colour; the tomentum) 

Ann. Jard. Bot. Btzg. 42 (1932) 204, Bull. Jard. Bot. Btzg. 3, 12 (1932) 361; Masamune I.e. 367; 

Kalkman I.e. (1954) 521; Leenhouts I.e. (1956) 225; Kochummen I.e. (1972) 143; Whitmore, Tantra 

& Sutisna I.e. 37. Basionym: Canarium rubiginosum A.W. Berm. in Hooker! I.e. 535, non C. 

rubiginosum Miq. I.e. (1859) 651 (= Santiria rubiginosa Blume). Type: Maingay 309, Malacca 

(holotype K; isotypes CAL, L). 

Medium-sized tree to 20 m tall, 20 cm diameter. Ultimate leafY twigs to 0.5 cm thick, 

yellowish brown velvety hairy when young. Leaves with 2-3 pairs of leaflets; rachis 

appressed hairy; petiolules of lateral leaflets 0.5-1 cm long, swollen at both ends; blade 

sparsely hairy below, elliptic or oblong, 9.5-19.5 x 2.5-6.5 cm; base cuneate, unequal, 

apex pointed, tip c. 1.5 cm long; midrib raised above; lateral veins 8-12 pairs, prominently 

raised below, faintly raised above; intercostal veins scalariform-reticulate, visible below. 

Flowers (male) tomentose, in terminal inflorescences, axes densely yellowish brown hairy. 

Fruits ellipsoid, 2-2.5 x 1 cm. 

Distribution. Peninsular Malaysia and Borneo. In Sabah scattered. In Sarawak uncommon, 

known only from two collections (S 43194 and S 43377) from Samunsam Wildlife Sanctuary 

and UIu Sg. Semawat respectively. Also found in Kalimantan. 


73


11. Dacryodes rugosa (Blume) H.J. Lam 

(Latin, rugosus = wrinkled; the bullate leaflets) 

Arm. Jard. Bot. Btzg. 42 (1932) 203, Bull. Jard. Bot. Btzg. 3, 12 (1932) 345; Masamune I.e. 367; 

Kalkman I.e. (1954) 505; Leenhouts I.e. (1956) 221; Backer & BakhuizenJ I.e. (1972) 114; Burgess 

I.e. 62; Kochummen I.e. 144; Cockburn I.e. 47; Anderson I.e. 157; Whitmore, Tantra & Sutisna I.e. 

37. Basionym: Santiria rugosa Blume I.e. 212. Type: Korthals, S.n. (= Leiden no. 898.321-232), 

Java (L). Synonyms: Canarium rugosum (Blume) Miq. I.e. (1859) 649 (including var. sumatranum); 

Hemisantiria rugosa (Blume) H.J. Lam in Merrilll.e. (1929) 119. 

Small to medium-sized tree, rarely to 30 m tall and 40 cm diameter. Bark grey-white, 

smooth to scaly; inner bark yellowish. Sapwood pale. Ultimate leafy twigs 0.3-0.5 cm 

thick. Leaves with 1--4 pairs ofleaflets; rachis flattened above; petiole glabrous; petiolules 

of leaflets strongly swollen at both ends; blade glabrous or appressed pubescent below or 

with erect hairs on midrib and veins below (var. virgata), elliptic, ovate to oblonglanceolate, 

6-22 x 2.5-11 cm, sometimes bullate; base cuneate, apex pointed with long tip; 

midrib raised above; lateral veins 7-12 pairs, raised below, flattened above (sunken in var. 

rugosa), often curving and joining near margin; intercostal veins reticulate, raised below, 

faint or sunken above. Flowers in axillary panicles, glabrous (var. virgata) or hairy; stamen 

filaments adnate to the disc; disc cup-shaped. Fruits ovoid, slightly oblique, 1.5-2.5 x 1- 

1.5 cm. 


Lateral veins sunken on upper leaflet surface. Leaflets glabrous or appressed-hairy on lower 

surface. Flowers densely hairy ............................................................................................. . 

var. rugosa 

Synonyms: Daeryodes rugosa var. genuina H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 347; 

Santiriafascieulata AW. Berm. in HookerJ I.e. 539. 

Sumatra, Peninsular Malaysia, Java, and Borneo. Widespread in Sabah and Sarawak in 

mixed dipterocarp forest on yellow sandy clay soils. Also in Kalimantan. 

Lateral veins not sunken on upper leaflet surface. Leaflets with scattered erect hairs on 

midrib and veins on lower side. Flowers glabrous ................................................ . 

var. virgata (Blume) H.J. Lam 

Bull. Jard. Bot. Btzg. 3, 12 (1932) 348. Basionym: Santiria virgata Blume I.e. (1850) 213. 

Synonym: Canarium virgatum (Blume) Miq. I.e. (1959) 650. Type: Korthals, s.n., W Borneo, G. 

Pamaton (holotype L; isotype BO). 

Widely distributed in Sabah and Sarawak; also in Kalimantan. 

Ecology. Lowland to hill mixed dipterocarp forests to 900 m. Flowering in April-November, 

and fruiting in March-December. 

3. GARUGA Roxb. 

(an lndo-Malayan plant name, origin uncertain) 

PI. Corom. 3 (1811) 5; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 325; Kalkman I.e. (1954) 459; 

Leenhouts I.e. (1956) 215; Kochurnnien I.e. (1972) 144; Whitmore, Tantra & Sutisna I.e. 37. 

74


Deciduous trees. Pith of branches and petioles without vascular strands. Leaves with 

stipules, stipellae (stipule-like outgrowths at the base of leaflets) often present; margin of 

leaflets toothed. Inflorescence an axillary panicle. Flowers bisexual, 5-merous; receptacle 

concave, globose or cup-like; sepals free; stamens 10; disc adnate to the receptacle, glabrous 

with 10 lobes between the stamens; ovary 5-celled. Fruit drupaceous; pericarp fleshy; 

pyrenes 1-5, furrowed, bony. Cotyledons twisted and folded. 

Distribution. 4 species; continental SE Asia, with a variety of one species in Malesia, 

Melanesia, N Australia, and the Solomon Islands. 


Gartiga floribunda Decne. var. floribunda 

(Latin,floribundus = with abundant flowers) 

Fig. 3. 

Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 477; H.J. Lam, Bull. lard. Bot. Btzg. 3, 12 (1932) 326; 

Kalkman I.e. (1954) 463; Leenhouts I.c. (1956) 215; Kochummen l.c. (1972) 144, Whitmore, Tantra 

& Sutisna I.c. 37. Type: sine coli., S.n., Timor (holotype G; isotype NY). Synonyms: Guaiacum 

abilo Blanco, Fl. Filip. (1837) 364; Garuga abilo (Blanco) Merr., Publ. Govt. Lab. Philip. 35 (1905) 

73; Garuga littoralis Merr.lc. (1915) 27; Garuga clarkei Merr.l.c. (1915) 29. 

Small to medium-sized tree to 12 m tall, 10 cm diameter. Bark grey, smooth. Twigs 

covered with many leaf-scars, minutely tomentose at tips. Stipules inserted at base of 

petioles, caducous. Leaves crowded at ends of twigs, leaflets 4-10(-15) pairs; blade 

subsessile, elliptic to oblong or lanceolate, 5-19 x 2-5.5 cm; base oblique, cordate or 

rounded, margin toothed, ape)\. pointed; midrib raised above; lateral veins 10-20 pairs, 

distinct on both surfaces; intercostal veins reticulate, visible on both surfaces. Flowers with 

cup-shaped receptacle; petals tomentose inside. Fruits obliquely pear-shaped, 5-9 x 5-12 

mm. 

Distribution. Peninsular Malaysia, Java, Borneo, Philippines, Celebes, Lesser Sunda Islands, 

Moluccas, New Guinea, Melanesia, and W Australia. In Borneo uncommon, known only from 

some islands (Gaya and Sipadan) in Sabah. 

Ecology. Coastal forests at low altitude. 

A second variety, var. gamblei, is known from E India, Sikhim, Bangladesh, W China and 

Hainan. 

4. HAPLOLOBUS H.J. Lam 

(Greek, haplos = single, lobus = lobe; the single-lobed seed) 

Ann. lard. Bot. Btzg. 42 (1932) 25, Bull. lard. Bot. Btzg. 3, 12 (1932) 404, Blumea 9 (1958) 237; 

Husson & H.J. Lam, Blumea 7 (1953) 413; Leenhouts I.c. (1956) 239, Blumea 20 (1972) 283 & 311; 

Whitmore, Tantra & Sutisnal.c. 37; Kochummen, Sandakania 5 (1994) 75. 

75


5cm 

A 

B 

] 'mm 

Fig. 4. Haplolobus leenhoutsii CA) and H. kapitensis CB-D). A, fruiting leafy twig;. B, flower bud; C, 

male flower with sepals and petals removed; D, stamen. CA from S. 41304, B-D from S. 23966.) 

78


Tree. Twigs stout to 2.5 cm diameter, lenticellate. Leaves with 6 pairs of leaflets; pith of 

petioles with 15-20 vascular strands; rachis to 23 cm long; blade lanceolate-oblong, 21,"""36 

x 7.5-10 cm, densely hairy on the under surface; base cuneate, apex pointed; lateral veins 

25-30 pairs, arching near margin; petiole flattened on the upper side, and distinctly striate 

on the lower side. Inflorescences and flowers unknown. Infructescences borne on leafless 

twigs or stems. Fruits oblong, with pointed tip, 13-15 x 12 mm. 

Distribution. Endemic to Sarawak. Uncommon, known only from the type specimen from 

Mt. Matang. 


2. Haplolobus bintuluensis Kochummen 

(of Bintulu in Sarawak) 

I.c. (1994) 75. Type: Sibat S. 24562, Sarawak (holotype SAR; isotypes A, K, L, SAN, SING). 

Treelet to 3 m tall, 5 cm diameter. Twigs c. 7 mm thick, greyish, with white lenticels. 

Terminal bud oblong, c. 10 x 3 mm, rusty hairy. Leaves: rachis c. 23 cm long, slightly 

swollen at base, dark brown-hairy with 2-3 pairs of leaflets; petiolules of lateral leaflets 8- 

15 mm long, dark brown hairy, swollen at both ends, petiolules of terminal leaflets 3.5-4.5 

cm long; blade thinly leathery, glabrous above except midrib, dark brown hairy on the 

midrib a.-zd veins below, 10.5-17 x 5.5-7 cm; base cuneate, apex long-pointed, tip narrow, 

1-2 cm long; midrib flattened above; lateral veins 7-9 pairs, distinct below, faint above, 

curving and joining near margin; intercostal veins reticulate, faintly visible below. 

Inflorescences and flowers unknown. Infructescence c. 2 cm long. Fruits oblong, reddish 

when fresh, yellowish brown on drying, 16-20 x 11-12 mm, with pointed tip. 

Distribution. Endemic to Sarawak. Uncommon, known from the type collection only, from 

the Nyabau Catchment area, Bintulu. 

Ecology. Mixed dipterocarp forest on yellow-red sandy humult ultisol, at about 100 m. 

3. Haplolobus inaequifoIius Kochummen 

(Latin, inaequalis = unequal,folium = leaf; the asymmetric leaflets) 

I.c. (1994) 78. Type: Yii S. 40756, Sarawak (holotype KEP; isotypes K, L, MO, SAN, SAR). 

Small tree to 15 m tall, 10 cm diameter. Twigs brownish, c. 5 mm thick with whitish lenticels. 

Terminal bud oblong, c. 12 mm long, rusty hairy. Leaves glabrous, with 2 pairs of leaflets; 

rachis grey, not swollen at base; petiolules of lateral leaflets 8-15 mm long, of terminal 

leaflet c. 4.5 cm long, the ends black on drying; blade greenish brown on drying, leathery, 

with 1-mm-wide circular galls, elliptic to ovate or oblong, 7.5-15.5 x 2.5-6 cm, strongly 

asymmetric; base broadly cuneate, apex pointed; midrib flattened above, sharply keeled 

below; lateral veins 10 pairs, faint on both surfaces; intercostal veins finely reticulate, 

visible on both surfaces. Inflorescence (male) glabrous, axillary, 4-8 cm long, often 

79


terminating in a vegetative shoot. Flowers (male): calyx-lobes triangular with pointed tip; 

petals oblong; stamens 6 with short stout filaments; disc with wavy margin; rudimentary 

ovary with 3 distinct stigma. Fruits unknown. 

Distribution. Endemic to Borneo. Uncommon, known only by the type specimen from the 

Sabal Forest Reserve in Sarawak. 

Ecology. Hill forest at about 360 m. 

4. Haplolobus kapitensis Kochummen 

(ofKapit, Sarawak) 

Fig.4B-D. 

I.c. (1994) 78. Type: Wright S. 23966, Sarawak, Kapit (holotype KEP; isotypes A, BO, K, L, SAN, 

SAR, SING). 

Emergent tree to 40 m tall, 80 cm diameter. Twigs greyish white or grey-brown, c. 1 cm 

thick, lenticellate, youngest strongly angled. Leaves glabrous, with 2-3 pairs of leaflets; 

rachis c. 26 cm long, channelled above near base; petiolules of lateral leaflets 1.5-2 cm 

long, of terminal leaflets c. 4.5 cm long, strongly swollen at both ends; blade thickly 

leathery, drying to reddish brown below and greenish brown above, oblong to lanceolate, 

11.5-20 x 4.5-9 cm; base asymmetric, almost rounded to broadly cuneate, apex pointed; 

midrib raised above; lateral veins 9-12 pairs, distinctly looping near margin, distinct on 

both surfaces; intercostal veins reticulate, more distinct above than below. Inflorescence 

(male) axillary, to 21 cm long, on leafless vegetative shoot. Flowers (male): calyx cupshaped 

with truncate apex; petals yellow, oblong; stamens 3 or 6, with short filaments; disc 

cup-shaped with wavy margin; pistil rudimentary. Fruits unknown. 

Distribution. Endemic to Borneo. Uncommon, known by four collections, S. 23966 and S. 

29162 from Kapit, Sarawak, and SAN 22391 and SAN 62657 from Ranau in Sabah. 

Ecology. Lowland and hill forests to 600 m. 

5. Haplolobus leenhoutsii Kochummen Fig. 4A. 

(P.W. Leenhouts, botanist at Rijksherbarium, Leiden, the Netherlands) 

I.c. (1994) 81. Type: Othman et a!. S. 41304, Sarawak, Kapit (holotype SAR; isotypes K, KEP, L, 

MO, SAN). 

Small tree to 20 m tall, 20 cm diameter. Bark brownish green, scaly. Twigs brown, c. 1 cm 

thick. Leaves with 3 pairs of leaflets; rachis c. 23 cm long; petioles channelled with incurled 

edges above, with many shallow grooves on the remaining part; petiolules of lateral and 

terminal leaflets c. 2 cm long, strongly swollen at both ends and with horizontal cracks; 

blade drying to greenish brown, glabrous, leathery, lanceolate to elliptic, 14-23 x 4.5-7.5 

cm, with many circular (3-mm-diameter) domatia holes; base rounded or broadly cuneate, 

apex pointed; midrib raised above, channelled below; lateral veins 14-17 pairs, distinct 

below, faint above; intercostal veins reticulate, visible on both surfaces. Inflorescences and 

80


flowers unknown. Infructescences axillary, c. 15.5 cm long, with many branches. Fruits 

(immature) red when fresh, black on drying, oblong, c. 1 cm long, with prominent apical 

stigma, surface slightly rugose, with remains of floral parts at base, stalk c. 1 cm long. 

Distribution. Endemic to Borneo. Uncommon, known only from the type specimen from 

Ulu Balleh, Kapit, Sarawak. 

Ecology. Mixed dipterocarp forest at about 500 m, on ridge. 

6. Haplolobus sarawakanus Kochummen 

(of Sarawak) 

I.c. (1994) 81. Type: Othman et al. S. 41339, Sarawak (holotype SAR; isotypes K, KEP, L, SAN, 

MO). 

Small tree to 11 m tall, 15 cm diameter. Bark greyish brown, scaly. Twigs c. 5 mm thick, 

pale brown, shallowly grooved, with pale lenticels. Leaves glabrous, with 3-4 pairs of 

leaflets; rachis c. 25 cm long, slightly swollen at base; petiolules of lateral leaflets c. 1 cm 

long, of terminal leaflet c. 3 cm long, the ends swollen and whitish or blackish; blade thinly 

leathery, drying to yellowish brown, elliptic or oblong, 12-15 x 3.5-5 cm; base cuneate, 

slightly asymmetric, apex pointed; midrib flattened above; lateral veins 8-9 pairs, distinct 

below, faint above, curving and joining near margin; intercostal veins reticulate, very faint 

on both surfaces. Inflorescences and flowers unknown. Infructescences c. 4 cm long. 

Fruits (immature) c. 8 x 5 mm, calyx funnel-shaped. . 

Distribution. Endemic to Borneo. Uncommon, known only by the type from Ulu Balleh, 

Kapit, Sarawak. A collection from Sabah, SAN 97191, probably belongs to this species but 

better material is needed to confirm this. 

Ecology. Mixed dipterocarp forest at about 500 m, on ridge. 

5. PROTIUM Bunn.J, nom. cons. 

(Greek, proteion = first-class; probably 

referring to the hardness and strength of the wood) 

Fl. Ind. (1768) 88; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 318; Leenhouts I.c. (1952) 154, l.c. 

(1956) 213; Backer & BakhuizenJ l.c. 113. 

Trees or shrubs. Pith of branchlets and petioles withouts vascular strands. Leaves without 

stipules, tips of the leaflets usually distinctly mucronulate. Inflorescences paniculate, axillary, 

rarely pseudo-terminal. Flowers 4-5-merous, structurally not always completely unisexual; 

sepals united; petals valvate with inflexed margins, glabrous; stamens double the number of 

petals, free; disc intrastaminal, annular, truncate to undulate, glabrous; ovary 3-5-celled, 

stigma subsessile. Fruits drupaceous, pericarp fleshy, calyx persistent, not enlarged, with 

reflexed lobes. Cotyledons plicate, lobed to palmatifid. 

81


4cm r 

~ 

~~ 

Fig. 5. Protium connarifolium. Fruiting leafy twig. (From SAN 89964.) 

82


Distribution. About 85 species; mainly in the American tropics, Madagascar, Mascarenes, 

India and Malesia. Only one species has been recorded in Sabah; not yet found in Sarawak. 


Protium connarifolium (Perkins) Merr. 

(with the leaflets resembling those of Connarus) 

Fig. 5. 

I.e. (1915) 30; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 320; Leenhouts I.e. (1952) 155, I.e. 

(1956) 215. Basionym: Canarium eonnarifolium Perkins, Fragm. Fl. Philip. 2 (1904) 92. Type: 

Merrill 787, Philippines, Palawan (GH, K, L). Synonym: P. philippinensis Elmer, Leafl. Philip. Bot. 

7 (1915) 2571. . 

Small tree. Leaves with up to 3 pairs of leaflets; petiolules of lateral leaflets to 2 cm long, 

swollen at both ends; blade ovate to elliptic, 3-11 x 2-5 cm; base cuneate, margin entire, 

apex pOinted with mucronate tip; midrib slightly sunken above; lateral veins 5-7 pairs, 

looping, visible below, faint above; intercostal veins reticulate, very faintly visible on both 

surfaces. Inflorescences axillary, or female inflorescences sometimes pseudo-terminal. 

Flowers tomentose. Fruits 7-8 x 5-13 mm, sparsely pubescent to glabrous, with 1-4 

pyrenes. 

Distribution. Borneo, Philippines (Palawan). Very uncommon, known by a single 

collection SAN 89964 from Semporna in Sabah. According to Leenhouts I.c. (1956), this 

species is rather isolated taxonomically from the rest. 


6. SANTIRIA Blume 

(Santir, Blume's native guide to G. Salak, Java) 

kerantai (Sabah), seladah (Sarawak) 

I.e. (1850) 209; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 367; Ridley I.e. (1922) 376; Kalkman 

I.e. (1954) 522; Leenhouts I.e. (1956) 229; Kochummen I.e. (1972) 145, Sandakania 5 (1994) 83; 

Cockburn I.e. 47; Anderson I.e. 157; Wong I.e. 212; Whitmore, Tantra & Sutisna I.e. 38; Ng I.e. 38. 

Synonyms: Trigonoehlamys Hook. f, Trans. Liml. Soc. 23, 1 (1860) 170; Ieieaster Ridl., J. Str. Br. 

R. As. Soc. 75 (1917) 15. 

Medium-sized to large dioecious trees. Bark brown, fawn or grey, smooth to fissured, scaly 

or flaky, with many small or large lenticels; inner bark either pink, soft, and laminated or 

yellowish white, hard mottled and granular (s. grifJithii, s. ruMginosa), with white to 

colourless exudate darkening on drying. Pith o/twigs and petioles with or without vascular 

strands. Leaves without stipules, imparipinnate; leaflets entire, petiolules only slightly 

swollen at both ends; petiole usually flat or channelled on the upper surface. Inflorescences 

usually axillary, rarely terminal panicles. Flowers unisexual, 3-merous; sepals free or 

united; petals usually with thickened in flexed apex; stamens 6 or 3, anthers basifixed or 

83


dorsifixed; disc intrastaminal, glabrous; ovary 3-celled, glabrous, in male flowers reduced; 

stigma (sub)-sessile. Fruits brightly coloured drupes, irregularly globose or ellipsoid, more 

or less oblique, in big bunches, usually seated on persistent calyx; stigma off-centre, 

sometimes nearZv basal; rind thin and firm, almos.t smooth when dry; stone thinly woody 

containing one fertile and two sterile cells. Seeds almost round, not angled; germination in 

S. laevigata and S. oblongifolia epigeal, cotyledons 5-lobed, fleshy, first two leaves 

opposite, subsequent leaves alternate, leaves in young seedling simple; in S. grifjithii and S. 

rubiginosa germination hypogeal, first two leaves opposite, subsequent leaves alternate and 

then spiral, leaves in young seedlings pinnate. 

Distribution. About 22 species, with 6 species restricted to W Mrica and the rest in 

Malesia (mainly Sumatra, Peninsular Malaysia, Borneo). 15 species in Sabah and Sarawak. 

Ecology. Lowland forest, including swamp forest, to submontane forest to 1650 m. 

Uses. The timber is commonly grouped together with other members of the family and sold 

as kedondong. The wood is light hardwood. It is essentially similar in anatomical structure 

to the timber of Canarium and Dacryodes but is more variable in colour, weight and 

hardness. Silica is present in all species. S. laevigata and S. tomentosa are important sources of 

kedondong, timber in Peninsular Malaysia. The buttresses of S. tomentosa are interlocked 

and frequently used for parang sheaths. 

Taxonomy. The genus is sub-divided into two sections: section Santiria and section 

Jcicopsis. Apart from the differences in floral characters by which these sections were 

established, evidence from wood anatomy and germination supports the distinction of these 

sections. In Sabah and Sarawak, section Jcicopsis is represented by S. grifjithii and S. 

rubiginosa. 

Key to Santiria species 

1. Leaflets hairy on the lower side, at least on the midrib .............................................. 2 

Leaflets glabrous .......................................................................................................... 6 

2. Leaflets densely velvety or woolly hairy on the lower side ............................................ 3 

Leaflets sparsely hairy on the lower side ...................................................................... 5 

3. Intercostal veins scalariform ................................................................ 15. S. tomentosa 

Intercostal veins reticulate .............. . 

4. Leaflets concave, hairs dark brown; petiolule not swollen at ends. Twigs covered with 

lenticels. Stigma on fruit more than 90 0 excentric ...................................... l0. S. mollis 

Leaflets not concave, hairs yellowish brown; petiolule swollen at ends. Twigs not so. 

Stigma prominently excentric, near pediceL .......................................... 1. S. apiculata 

5. Twigs stout, c. 1.5 (fm thick. Terminal bud c. 2 cm long. Petiole strongly channelled 

above with deep groove ...................................................................... .4. S. grandiflora 

Twigs slender, c. 6 mm thick. Terminal bud up to 0.5 cm long. Petiole not channelled 

or grooved ....................................................................................... 14. S. sarawakana 

84


6. Lateral veins sunken above .......................................................... 6. S. impressinervis 

Lateral veins not sunken above .................................................................................. 7 

7. Twigs stout, 1.5-3 cm thick. ........................................................................................ 8 

Twigs slender, to 1 cm thick. ....................................................................................... 9 

8. Terminal bud c. 6 cm long with curved tip. Leaflets 7-9 cm wide; petiolules of lateral 

leaflets to 2.5 cm long. Stigma of fruit more than 90 0 excentric ........ 7. S. kalkmaniana 

Terminal bud c. 1.5 cm long, tip not curved. Leaflets 13-22 cm wide; petiolules of 

lateral leaflets 4-5 cm long. Stigma of fruit near pedicel... .................. 9. S. megaphylla 

9. Stigma offruit more than 90 0 excentric, near pedicel... .................................................. lO 

Stigma offruit less than 90 0 excentric ....................................................................... 12 

10. Terminal bud c. 2 cm long. Petiole strongly channelled at base with sharp edges. 

Midrib raised above .................................................................................. 2. S. conferta 

Terminal bud up to 1 cm long. Petiole not so. Midrib flattened to slightly sunken 

above ......................................................................................................................... 11 

11. Midrib below channelled. Leaflet margin not curled inwards; intercostal veins distinct 

below ...................................................................................................... l. S. apiculata 

Midrib below not channelled. Leaflet margin curled inwards; intercostal veins invisible 

below ......................................................................................................... 3. S. dacryodifolia 

12. Terminal bud 0.5-2.5 cm long, often resin-coated. Leaflets drying reddish brown or 

dark brown .............................................................................................. 8. S. laevigata 

Not this combination of characters .......................................................................... 13 

13. Rachis blackish, oily, shiny. Lateral veins of leaflets very faint. Stigma on fruit only 

slightly excentric .................................................................................. 11. S. nigricans 

Rachis not so. Lateral veins ofleaflets distinct. Stigma in fruit 90 0 excentric ............. 14 

14. Lateral veins distinctly looping near margin; intercostal veins parallel to lateral veins. 

Fruits to 1.3 cm long .................................................................................................. 15 

Lateral veins faintly looping near margin; intercostal veins not parallel to lateral veins. 

Fruit to 3 cm long ............................................................................. 12. S. oblongifolia 

15. Leaflets not asymmetric at base, petiolules of lateral leaflets to 5 mm long, not swollen 

at ends ..................................................................................................... 5. S. griffithii 

Leaflets asymmetric at base, petiolules of lateral leaflets to 2 cm long, swollen at both 

ends .................................................................................................... 13. S. rubiginosa 

1. Santiria apiculata AW. Benn. 

(Latin, apiculatus = furnished with a little point; the pOInted leaflets) 

in Hooker f I.c. 537; Ridley I.c. (1922) 378; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 375; 

Masamune I.c. 367; Kalkman I.c. (1954) 538; Leenhouts I.c. (1956) 234; Burgess I.c. 62; 

85

TREE FLORA OF SABAH AND SARA W AK VOL. 1 (1995) 

Kochummen I.e. (1972) 146; Cockbum I.e. 48; Anderson I.e. 157; Whitmore, Tantra & Sutisna I.e. 

38. Type: Maingay 303, Malacca (holotype K; isotype L). 

Small to medium-sized tree rarely more than 30 m tall. Bark grey or brown, smooth to 

scaly; inner bark pinkish with droplets of white sap and with strong resinous smell. 

Sapwood pale yellow. Twigs pale or whitish, glabrous or densely brownish hairy (var. 

pilosa). Leaves trifoliolate or with 2-S pairs of leaflets; blade glabrous, drying greenish or 

hairy below and drying brownish (var. pilosa), elliptic or lanceolate, 3.S-16 x l.S-7 cm; 

base broadly cuneate, apex pointed with long tip; midrib flattened to slightly sunken above, 

distinctly grooved below; lateral veins S-14 pairs, distinct below, faint above, curving and 

joining a few miIlimeters away from margin; intercostal veins reticulate, faintly visible 

below, faint to invisible above; with a strong resinous smell when freshly crushed; 

petiolules usually whitish, glabrous or hairy, 0.S-3 cm long. Flowers yellow or reddish, 

usually in short axillary inflorescences, glabrous or hairy (var. pilosa). Fruits globose or 

ellipsoid, 1-2 x 1 cm, stigma usually more than 90 0 excentric, sometimes near the pedicel. 


Twigs, leaflets and flowers densely hairy ...... . 

var. pilosa (Engl.) Kalkman 

in Leenhouts I.e. (1956) 236. Basionym: Santiria pilosa Engl. in DC. I.e. 159. 

Endemic to Borneo. In Sabah and Sarawak widely distributed from 

submontane forest but less common compared to var. apiculata. 

lowland to 

Twigs, leaflets and flowers glabrous ....................................... . 

var. apiculata 

Synonyms: S. beeearii Engl. in DC. I.e. 159; S. glabra MeIT. I.e. (1915) 30; S. minutiflora Ridl. 

I.e. (1922) 377; S. braehystaehys Ridl. I.e. (1925) 79; Canarium paueiflorum Ridl. I.e. (1930) 

80; Haplolobus bomeensis H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 418. 

Sumatra, Peninsular Malaysia, Borneo, Philippines, Celebes, Moluccas. Common and 

widely distributed throughout Sabah and Sarawak. Mixed dipterocarp to submontane 

forests to 1300 m. 

2. Santiria conferta AW. Benn. 

(Latin, confertus = crowded; the flowers) 

in I-looker! I.e. 537; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 381; Kalkman I.e. (1954) 528; 

Leenhouts I.e. (1956) 233; Burgess I.e. 62; Kochummen I.e. (1972) 146; Whitmore, Tantra & Sutisna 

I.e. 38. Type: Griffith 1150, Malacca (holotype K; isotype P). Synonyms: S. wrayi King I.e. (1893) 

259; S. eonferta var. wrayi (King) HJ. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 381. 

Small to medium-sized tree to 30 m tall, SO cm diameter. Bark grey-brown; inner bark 

yellow. Terminal bud c. 2 cm long. Twigs greyish yellow, c. 1 cm thick. Leaves: petiole 

strongly channelled with sharp edges; leaflets thickly leathery, oblong, ovate or elliptic, 

sometimes glaucous below, 11-17 x S.S-8.5 cm; base rounded to broadly cuneate, apex 

86


pointed; midrib raised above; lateral veins 11-14 pairs, arching near margin, raised on both 

surfaces; intercostal veins reticulate, faintly visible on both surfaces. Flowers in axillary 

inflorescences. Fruits reddish when fresh, subglobose, 1.5-2 x 1.2-1.5 cm, stigma strongly 

excentric, near pedicel; surface warty. 

Distribution. Sumatra, Peninsular Malaysia, Borneo. In Borneo uncommon, known from a 

few collections from Sabah and Sarawak. 

Ecology. Submontane forest at 1200-1650 m. Fruiting in November and December. 

3. Santiria dacryodifolia Koch4mmen 

(with leaves resembling those of Dacryodes) 

I.c. (1994) 83. Type: Tong & lugah S. 32944, Sarawak (holotype KEP; isotypes K, L, SAN, SAR, 

SING). 

Small tree to 12 m tall, 25 cm diameter; bole fluted. Bark light brown, flaky. Twigs 

brownish, finely lenticellate, c. 7 mm thick. Leaves with 2 pairs of leaflets; rachis pustular, 

rounded, not swollen at base; blade coriaceous, glabrous, drying to brownish, elliptic to 

rhomboid, 8.5-10.5 x 3.5-5.5 cm; base cuneate, margin curled inwards, apex pointed; 

midrib sunken above; lateral veins 5-7 pairs, faintly visible on both surfaces; intercostal 

veins reticulate, visible above, invisible below; petiolules of lateral leaflets 1-1.5 cm long, 

of terminal leaflet c. 4.5 cm long, strongly swollen at both ends. Inflorescences and flowers 

unknown. Infructescence axillary and terminal, c. 14 cm long. Fruits yellowish to deep 

red when fresh, drying to dark brown, obliquely oblong, 7-8 x 5-6 mm, surface slightly 

rugose, stigma more than 90 0 excentric, near the pedicel. 

Distribution. Endemic to Sarawak. Uncommon, known from the type only from Ulu Sg. 

Maria, Lawas. 

Ecology. Ridge tops at about 1400 m. 

Appears close to S. apiculata differing in the stout twigs, thick leaflets with recurved 

margins and in the petiolules with strongly swollen ends. 

4. Santiria grandiflora Kalkman 

(Latin, grandis = large,florus = flower) 

I.c. (1954) 525; Leenhouts I.c. (1956) 232; Anderson I.c. 157; Whitmore, Tantra & Sutisna I.c. 38. 

Type: Richards 1278, Sarawak (holotype K; isotype SING). 

Medium-sized tree to 30 m tall. Bark grey-brown, scaly; inner bark pink, with droplets of 

white sap. Sapwood pale. Twig stout, c. 1.5 cm thick, lenticellate, dark brown powdery 

hairy towards the tip. Terminal bud c. 2 cm long. Leaves with 4-6 pairs of leaflets; 

87


petiolules of lateral leaflets 1-2 cm long, powdery hairy; blade glabrous above except the 

midrib, powdery hairy below, oblong-elliptic, 16-29 x 5-9 cm; base rounded or broadly 

cuneate, apex pointed; midrib raised below, flat and faint above; lateral veins 15-24 pairs; 

intercostal veins scalariform-reticulate, distinct below, faint above; petiole strongly 

channelled above with deep groove, powdery brown-hairy. Flowers (male) in axillary 

inflorescences. Infructescences 9-18 cm long. Fruits obliquely ellipsoid, yellowish green, 

ripening pink, with white waxy bloom on drying, 1.5-l. 9 x 1.1-1.3 cm, with faintly 

wrinkled surface; stigma slightly excentric; persistent calyx reflexed. 

Distribution. Endemic to Borneo. Uncommon, recorded from Mt. Dulit, Mulu National 

Park, and Lambir Hills in Sarawak. Also known from Brunei. 

Ecology. Mixed dipterocarp forest to 300 m, on ridges or yellow sandy clay soils. Fruiting 

in March and April. 

This species is somewhat similar to S. tomentosa but the strongly channelled petiole will 

distinguish it. 

5. Santiria griffithii (Hook. f) Engl. 

(W. Griffith, 1810-45, surgeon at Malacca) 

Bot. Jahrb. 1(1881) 43; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 394; Masamune I.e. 368; 

Ka1kman I.e. (1954) 545; Leenhouts I.e. (1956) 236; Kochummen I.e. (1972) 147; Burgess I.e. 62; 

Anderson I.e. 157; Whitmore, Tantra & Sutisna I.e. 38. Basionym: Trigonoehlamys grifJithii Hook. 

f I.e. (1860) 170. Type: GrifJith 1148, Malacca (ho1otype K; isotype L). Synonym: Santiria 

bomeensis Engl. I.e. 43. 

Medium-sized tree to 30 m tall, 55 cm diameter; buttresses to 2 m tall. Bark grey-brown, 

smooth to scaly; inner bark brownish. Sapwood pale. Twigs reddish brown-hairy when 

young. Leaves with 5-11 pairs of opposite or subopposite leaflets; rachis minutely hairy; 

petiolules of leaflets c. 5 mm long, hairy, not swollen; blade glabrous or sparsely hairy 

below, lanceolate or elliptic, 5-10.5 x l.5-3 cm; base broadly cuneate to rounded, slightly 

unequal, apex pointed with long tip; midrib flattened above; lateral veins 15-18 pairs, 

faintly visible below, faint to inconspicuous above, distinctly curving and joining near 

margin; intercostal veins reticulate, visible below, faint above. Inflorescences axillary, 

pubescent. Flowers tomentose; calyx deeply divided; petals pubescent; stamens 6, filaments 

adnate to disc, pistil in male flowers reduced. Fruits obliquely globose, 8-13 x 8-16 mm; 

stigma almost 90 0 excentric. 

Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah uncommon, recorded from 

Keningau and Tawau only. In Sarawak more common and widely distributed. Also occurs 


Ecology. Mixed dipterocarp forest on yellow sandy soils to 300 m. Flowering in March 

November and fruiting in September-October. 

88


6. Santiria impressinervis Kochummen 

(Latin, impressus = pressed in, nervus = nerve; the sunken veins on the upper leaflet 

surface) 

I.c. (1994) 83. Type: Nooteboom & Chai 2113, Sarawak (holotype KEP; isotype SAR). 

Small tree to 15 m tall, 15 cm diameter. Bark yellowish, smooth. Twigs c. 4 mm thick, 

brownish, with small lenticels. Leaves with 2-3 pairs of leaflets; rachis slender, slightly 

flattened above towards the base; petiolules of lateral leaflets 0.8-1.5 cm long, that of 

terminal leaflet c. 2 cm long, strongly swollen at both ends; blade glabrous, coriaceous, 

drying to yellowish brown, ovate to lanceolate, 6-9 x 2-4 cm; base broadly cuneate, slightly 

asymmetric, apex pointed, tip 5-10 mm long; midrib faintly sunken above; lateral veins 7- 

9 pairs, sunken above, curving and joining near margin; intercostal veins invisible. 

Inflorescences and flowers unknown. Infructescences axillary, to 11 cm long. Fruits 

glaucous green when fresh, drying to brownish, smooth, ovoid, c. 10 x 7 mm, stigma 

prominently off-centre, near pedicel. 

Distribution. Endemic to Sarawak. Uncommon. known only from the type from the 

Pamerario river at Bario in the Kelabit Highlands. 

Ecology. Old secondary forest at 1000 m. 

Appears close to S. apiculata but differing in the thickly coriaceous leaflets with invisible 

intercostal veins and in the sunken lateral veins. 

7. Santiria kalkmaniana Kochummen Fig. 6. 

(C. Kalkman, former director of the Rijksherbarium, Leiden, Netherlands) 

I.c. (1994) 86. Type: Saikeh SAN 72293, Sabah (holotype SAN; isotypes A, K, SAR, SING). 

Medium-sized tree to 30 m tall, 45 cm diameter; buttresses to 3 m tall. Bark greyish, 

smooth, lenticellate; inner bark yellowish. Sapwood pale yellow. Twigs very stout, 1.5-2 

cm thick, grey-brown, lenticellate. Terminal bud lanceolate, rusty brown, c. 6 cm long with 

curved tip. Leaves with 5 pairs of leaflets, glabrous; rachis c. 54 cm long, petiole 

channelled above; petiolules of lateral leaflets 2-2.5 cm long, slightly swollen at both ends, 

petiolules of terminal leaflets 5-8.5 cm long; blade thinly leathery, oblong to lanceolate, 

18-35 x 7-9 cm; base rounded or broadly cuneate, asymmetric, apex pointed; midrib raised 

above; lateral veins 16-19 pairs, raised below, faint above; intercostal veins reticulate, more 

distinct on the upper surfaces. Inflorescences (male) axillary, glabrous, panicles to 27 cm 

long, axes sharply angled. Flowers (male) glabrous; calyx red, cup-shaped; petals deep 

scarlet, oblong; stamens 6, inserted on rim of disc; disc annular; ovary rudimentary. 

Infructescences c. 6 cm long, with short branches. Fruits light yellow when fresh drying 

black, subglobose or hemispherical, 1.7-2 x 1.0-1.8 cm, flattened above; stigma more than 

90 0 excentric. 

89


Distribution. Endemic to Borneo. Uncommon, known from few collections from Sabah 

(SAN 72293, Sipitang; SAN 49754, Ranau), and Sarawak (s. 14727, Kapit; S. 39782, G. 

Mulu, and S. 4030 from Baram). 

Ecology. Lowland and hill forests to 450 m. 

This species appears close to S. laevigata but differs in the long rusty brown curved 

terminal bud and in the fruit stigmas that are more than 90 0 off-centre. 

8. Santiria laevigata Blume 

(Latin, laevigatus = smooth and polished; the leaflets) 

l.c. (1850) 211; King I.c. 257; Rid1ey l.c. (1922) 378; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 

382; Masamune I.c. 368; Ka1kman I.c. (1954) 535; Leenhouts I.c. (1956) 232; Burgess l.c. 62; 

Kochummen l.c. (1972) 147; Cockbum l.c. 48; Anderson l.c. 157; Whitmore, Tantra & Sutisna l.c. 

38. Type: Korthals. S.n. (= Leiden no. 898.321-201), Sumatra (L). Synonym: Canarium laevigatum 

(Blume) Miq. I.c. (1859) 648. 

Medium-sized tree to 30 m tall, 80 cm diameter; buttresses to 4 m tall. Bark grey-brown, 

fissured and scaly, lenticellate; inner bark pinkish, laminated, with droplets of white sap. 

Sapwood pale white. Terminal bud 0.5-2.5 cm long, often resin-coated. Twigs c. 1 cm 

thick, lenticellate, dark brown. Leaves with 1-4 pairs of leaflets; petiole flattened above 

with sharp edges (forma laevigata) or not flattened (forma glabrifolia); petiolules of lateral 

leaflets 1-1.5 cm long, slightly swollen at both ends, sometimes the swollen parts drying 

black; blade drying reddish or dark brown, elliptic, ovate or oblong, 7-12 x 2.5-6.5 cm; 

base rounded to broadly cuneate, sometimes unequal, apex pointed; midrib raised above; 

lateral veins 7-15 pairs, fairly raised on both surfaces, curving and joining near margin; 

intercostal veins reticulate, very faintly visible on both surfaces. Flowers glabrous, in 

axillary inflorescences. Fruits rounded to oblong, 1-2 cm across; stigma less than 90 0 

excentric. 

Key to forms 

Petiole flattened above, at base with sharp edges .................................................................. . 

forma laevigata 

Synonyms: Santiria laevigata forma typica H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 384; s. 

violacea HJ. Lam, Ann. Jard. Bot. Btzg. 42 (1932) 205. 

Sumatra, Peninsular Malaysia, Borneo, and Celebes. In Sarawak uncommon, known 

from two collections (s. 32817 and S. 41457). Not yet recorded from Sabah. 

Petiole rounded at base, without sharp edges ................................................. . 

forma glabrifolia (Engl.) HJ. Lam 

Ann. Jard. Bot. Btzg. 42 (1932) 205. Basionym: Santiria glabrifolia Engl. in DC. l.c. 164. Type: 

Eeccari PE 3756, Sarawak (FI). 

Sumatra, Peninsular Malaysia, and Borneo. Common and widely distributed in Sabah 

and Sarawak. 

Ecology. Mixed dipterocarp, mixed peat swamp and kerangas forests on low-nutrient soils 

to submontane forests to 1200 m. 

90

I3URSERACEAE (KOCHUMMEN) 

~;t~~~ B 

2cm 

}cm 

Fig. 6. Sanliria kalkmaniana. A, Leafy shoot with infruetescences; B, inflorescence with flower buds; 

C, flower bud; D, flower bud with sepals and petals removed. CA from SAN 72293, B from S. 39782, 

C &]) from S. 4030.) 

91


9. Santiria megaphylla Kalkman 

(Greek, mega = large, phyllon = leaf) 

I.e. (1954) 533; Leenhouts I.e. (1956) 236; Burgess l.c. 62; Anderson l.c. 157; Whitmore, Tantra & 

Sutisna I.e. 38. Type: Beccari PB 3059, Sarawak (FI). 

Medium-sized tree to 25 m tall. Twigs very stout, c. 3 cm thick, grey with circular lenticels 

and with stipule-like undeveloped leaves. Terminal bud c. 1.5 cm long. Leaves with 3-5 

pairs of leaflets; petiole stout, strongly channelled at base; petiolules of lateral leaflets 4-5 

cm long, petiolules of terminal leaflets c. 11 cm long, strongly swollen at both ends; blade 

thickly leathery, very large, oblong, 25-61 x 13-22 cm; base broadly cuneate, apex 

pointed; midrib raised above; lateral veins 8-14 pairs, raised on both surfaces, curving and 

joining near margin; intercostal veins reticulate, faintly visible on both surfaces. Flowers in 

axillary inflorescences. Fruits pink when fresh, asymmetric, ellipsoid, 1.5-1. 7 x 1-1.25 

cm; stigma excentric, near pedicel. 

Distribution. Endemic to Borneo. Uncommon, known from few collections from Sabah 

(SAN A 1746, Beaufort; SAN 15152, Sipitang) and Sarawak (s. 38747 and S. 43958 from 

Semengoh Arboretum and Mt. Dulit), and ecological plots at Bako and Lamoir NP, and 

Segam FR. Also known from Brunei. 

Ecology. Mixed dipterocarp forests on deep sandy humult ultisols. 

The stout twigs and very large leaflets are good diagnostic features for the species. 

10. Santiria mollis Engl. 

(Latin, mollis = soft-hairy; the indumentum of young twigs, leaves and flowers) 

in DC. l.c. 156; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 388; Masamune l.c. 368; Kalkman I.c. 

(1954) 530; Leenhouts I.e. (1956) 231; Burgess I.e. 62; Anderson I.e. 157; Whitmore, Tantra & 

Sutisna I.e. 38. Type: Beccari PB 3497, Sarawak (FI). Synonym: Canarium hirtipetalum Rid!. I.e. 

(1930) 84. 

Medium-sized tree to 24 m tall, 70 cm diameter; buttresses to 3 m high. Bark brown, 

smooth; inner bark reddish brown, with white latex. Sapwood pale yellow. Twigs densely 

reddish brown-hairy with abundant smalllenticels when young. Terminal bud stout, c. 1 x 

0.5 cm, hairy as the twig. Leaves with 2-4 pairs of leaflets; rachis densely reddish brownhairy; 

petiolules of lateral leaflets to 1 cm long, not swollen at both ends, hairy; blade 

densely velvety hairy below and on midrib above, obovate, oblong or elliptic, often 

concave, 7.5-15.5 x 3-7 cm; base cuneate or rounded, often unequal, margin curled 

inwards, apex pointed with long tip; midrib raised above; lateral veins 10-13 pairs, raised 

below, faint to sunken above; intercostal veins reticulate, raised below, faint to invisible 

above. Flowers (male) in hairy axillary inflorescences; sepals almost free, sparsely hairy 

outside; petals densely hairy on both sides. Fruits sub globose, 1. 5-1. 8 cm across; stigma 

more than 90 0 excentric. 

Distribution. Endemic to Borneo. Uncommon, in Sabah recorded from Beaufort and 

Tawau. In Sarawak collected from Semengoh Arboretum and Lambir National Park. Also 

known from Brunei and Kalimantan. 

92


Ecology. Mixed dipterocarp forest to 200 m, often by streams. Flowering in March and 

November and fruiting in February and March. 

Very atypical of Santiria in vegetative features. 

11. Santiria nigricans Kochummen 

(Latin, nigricans = becoming black; the leaf rachis) 

I.e. (1994) 89. Type: Ilias & Yeo S. 38319, Sarawak (holotype KEP; isotypes K, L, MO, SAN, SAR). 

Large tree to 30 m tall, 100 cm diameter. Bark brownish, scaly; inner bark reddish brown. 

Sapwood whitish. Twigs brown or greyish brown, 3-4 mm thick. Leaves with up to 3 pairs 

of leaflets, glabrous; rachis slender, dark brown, surface oily, slightly swollen at the very 

base; petiolules of lateral leaflets slender, l-1.S cm long, of the terminal leaflets to 4 cm 

long, slightly swollen at both ends, drying to dark brown; blade thinly leathery, drying to 

grey-brown, elliptic to lanceolate, 6.S-l1.S x 2.8-4.7 cm; base cuneate, strongly asymmetric, 

apex pointed; midrib raised above; lateral veins 7-11 pairs, very faint on both 

surfaces; intercostal veins reticulate, faintly visible on both surfaces. Inflorescences (male) 

axillary and terminal, glabrous. Flowers (male) with green calyx and white petals. Fruits 

green, ripening deep purple to blackish, drying to pale brownish, subglobose, 11-14 mm 

across, surface with shallow reticulate veins; stigma only slightly off-centre. 

Distribution. Endemic to Sarawak; of scattered distribution in Lambir National Park (S 

38319 and S 46599), Mulu National Park (S 42406), Balingian (S 23687), Selampit (S 

24939) and Sg. Jelalong (S 48804). 

Ecology. Mixed dipterocarp forest from lowlands to 1600 m. 

Somewhat close to S oblongifolia but the dark brown rachis, very faint lateral veins and 

the sub globose fruits distinguish this species. 

12. Santiria oblongifolia Blume 

(Latin, oblongus = rather long,folium = leaf; the leaflet shape) 

I.e. (1850) 211; H.I. Lam,Bul!. Jard. Bot. Btzg. 3, 12 (1932) 372; Masamune I.e. 369; King I.e. 257; 

Ridley I.e. (1922) 379; Kalkman, I.e. (1954) 537; Leenhouts I.e. (1956) 233; Burgess I.e. 62; 

Kochummen I.e. (1972) 150; Anderson I.e. 157; Whitmore, Tantra & Sutisna I.e. 39. Type: 

Praetorius, s.n. (= Leiden no. 898.321-221), Sumatra (L). Synonyms: S. eaesia Eng!. in DC. I.e. 

166; S. latifolia Stapf. ex Rid!. I.e. (1930) 86. 

Medium-sized tree 30 m tall, very rarely to SO m tall, 80 cm diameter; buttresses to 1.S m 

high. Bark grey-brown, smooth, lenticellate, rarely scaly or flaky; inner bark pink, laminated. 

Sapwood pale white. Leaves with 2-4 pairs of leaflets; petiolules of lateral leaflets l-3.S 

cm long, swollen at both ends; blade thinly to thickly leathery, oblong to lanceolate, 8':"'21.S 

x 3-7.S cm; base broadly cuneate, apex pointed; midrib raised above; lateral veins 8-10 

pairs, visible on both surfaces, curving and joining near margin; intercostal veins reticulate, 

93


faint on both surfaces. Flowers in axillary or terminal inflorescences, glabrous or sparsely 

hairy. Fruits oblong-ellipsoid, l.2-3 x 1.1-2.2 cm; stigma slightly excentric. 

Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely distributed in Sabah. In 

Sarawak collected mainly from the Balm National Park and Semengoh Forest Reserve area. 

Also found in Kalimantan. 

Ecology. Mixed dipterocarp forest on yellow sandy and clay soils, heath forest and 

submontane forest to 1000 m. Flowering in March, May and September and fruiting in 

January, March-May and September. 

13. Santiria rubiginosa Blume 

(Latin, rubiginosus = rusty coloured; the indumentum) 

I.e. (1850) 213; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 399; Masamune I.e. 369; Kalkman I.e. 

(1954) 542; Leenhouts I.e. (1956) 237; Burgess I.e. 62; Kochumrnen I.e. (1972) 151; Anderson I.e . 

. 158; Whitmore, Tantra & Sutisna I.e. 39. Type: Praetorius, s.n. (=Leiden no. 898.321-229), 

Sumatra (L). Synonym: Canarium rubiginosum (Blume) Miq. I.e. (1859) 651. 

Medium-sized tree to 28 m tall, 40 cm diameter; buttresses to 1.5 m high. Bark grey, 

smooth, lenticellate. Twigs brownish, glabrous, except the tip. Leaves with 1-4 pairs of 

leaflets, sometimes mixed with simple leaves; petiolules of lateral leaflets 0.5-2 cm long, 

either rounded or strongly flattened, swollen at both ends; blade glabrous or very sparsely 

powdery hairy below, elliptic to lanceolate, 4.5-13.5 x 1.5-6 cm; base cuneate, often 

unequal, apex pointed with long tip; midrib raised above; lateral veins 9-13 pairs, visible 

below, faint above, arching and joining near margin; intercostal veins reticulate, faintly 

visible on both surfaces. Flowers in axillary inflorescences, glabrous; sepals almost free; 

stamens 3. Fruits globular or ellipsoid, 10-14 x 8-10 mm; stigma less than 90 0 excentric. 

S. rubiginosa is very similar to S. griffithii in vegetative characters but the asymmetric 

leaflets of S. rubiginosa are distinctive. 


1. Pedicels 1-3 mm long, shorter or as long as the flowers ............. , ................................. 2 

Pedicel 2-8 mm long, longer than the flowers .............. . 

var. pedicellata (Ridl.) Kalkman 

I.e. (1954) 544. Basionym: Santiria pedieellata Ridl. I.e. (1930) 86. Synonym: S. 

minimiflora Ridl. I.e. (1930) 87. Type: Haviland 1866, Sarawak, Sibu, Rejang (SING). 

Endemic to Borneo. Common in Sarawak but not yet reported from Sabah. Freshwater, 

heath and peat swamp forest. 

2. Leaflet apex caudate-acuminate, more than 1 cm long; petiolule flattened, the ends 

blacken on drying .................................................................................... . 

var. latipetiolata Kochummen 

I.e. (1994) 89. Type: George S. 40254, Sarawak (holotype KEP; isotypes E, K, L, MO, 

SAN, SAR). 

94


Endemic to Sarawak, known from two collections from the Lambir National Park 

(S 40254 and S 47190). Mixed dipterocarp forest on ridges at 250 m. 

Leaf apex not so, petioiule not flattened ............. . 

var. rubiginosa 

Synonyms: Santiria planehonii A.W. Benn. in Hooker f I.e. 536; C. planehonii CAW. 

Benn.) King I.e. 240; lcieaster planehonii CAW. Benn.) Ridl. I.e. (1917) 15; S. havilandii 

Ridl. I.e. (1930) 85. 

Sumatra, Peninsular Malaysia, Borneo, and New Guinea. Common and widely 

distributed in Sarawak in mixed dipterocarp forest but uncommon in 3abah and 

known only by a single collection, SAN 16678, from Sipitang at 750 m. 

14. Santiria sarawakana Kochummen 


I.e (1994) 9l. Type: Tong et aI., S. 34287, Sarawak (holotype SAR; isotypes K, KEP, L, MO, SAN). 

Medium-sized tree to 26 ill tall, 30 cm diameter. Bark dark brown, flaky. Twigs whitish, 

powdery brown-hairy. Terminal bud ovate, c. 5 x 3 mm. Leaves with 3 pairs of leaflets; 

rachis hairy as the twig, rounded; petiolules of lateral leaflets 7-15 mm long, of terminal 

leaflets 1.5-3 cm long, hairy and swollen at both ends; blade leathery, glabrous above 

except the midrib, densely hairy on midrib and lateral veins below, ovate to oblong or 

elliptic, 6-14 x 3-5 cm; base broadly cuneate, asymmetric, margin curled inwards, apex 

pointed; midrib raised above; lateral veins 9-12 pairs, curving and joining near margin, 

raised below, visible above; intercostal veins scalariform-reticulate, visible on both surfaces. 

Inflorescences and flowers unknown. Infructescences axillary, to 11.5 m long, axes hairy. 

Fruits glaucous black when fresh, black on drying, oblong, 11-13 x 7-10 mm; stigma less 

than 90 0 excentric. 

Distribution. Endemic to Sarawak. Uncommon, known only from Miri and the Sabal 

Forest Reserve (S 24119 and S 34287). 


Close to Santiria tomentosa but the sparsely hairy leaflets and short terminal buds are quite 

distinct. 

15. Santiria tomentosa Blume 

(Latin, tomentosus = densely pubescent with matted, woolly or short hairs; the young twigs, 

buds and leaflets) 

I.e. (1850) 211; H.J. Lam, Bull. lard. Bot. Btzg. 3, 12 (1932) 391; Masamune I.e. 369; Kalkman I.e. 

(1954) 529; Leenhouts I.e. (1956) 231; Burgess I.e. 62; Kochummen I.e. (1972) 151; Cockbum I.e. 

49; Anderson I.e. 158; Whitmore, Tantra & Sutisna I.e. 39. Type: Korthals, S.n. C= Leiden no. 

95


". [ 

o 

c 

E 

, 

3mm 

Fig. 7. Scutinanthe brunnea. A, fruiting leafy twig; B, inflorescence; C, flower bud; D, open flower; 

E, longitudinal section of flower. CA from S. 43904, B & C from S. 21666, D & E after FM 1, 5 

(1956) 247, fig. 18.) 

96


898.321-245), Sumatra (L). Synonyms: Canarium korthalsii Miq. I.e. (1859) 645; Santiria 

multiflora AW. Beilll. in Hooker f I.e. (1875) 538; C. mierantherum Stapf ex Ridl. I.e. (1930) 82; S. 

mollissima Ridl. I.e. (1930) 85. 

Medium-sized tree to 30 m tall, 50 cm diameter; buttresses to 2 m high. Bark grey-brown, 

scaly; inner bark pinkish with droplets of white sap. Sapwood pale yellow. Twigs 5-15 mm 

thick, slightly angled, woolly hairy. Terminal bud to 2 cm long, hairy. Leaves with 1-4 

pairs of leaflets; rachis woolly hairy; petiolules of lateral leaflets 0.5-2 cm long, hairy; 

blade woolly hairy below and on midrib above, ovate to oblong, 11-27 x 4-12 cm; base 

rounded or cuneate, sometimes unequal, apex pointed; midrib raised above; lateral veins 

10-20 pairs, raised on both surfaces, curving and joining near margin; intercostal veins 

scalariform-reticulate, distinct below, faint above. Flowers hairy, in axillary inflorescences. 

Fruits globose or oblong, 2-2.7 x 1.5-2 cm; stigma less than 90 0 excentric. 

Distribution. Sumatra, Peninsular Malaysia, Borneo. Common and widely distributed in 

Sabah and Sarawak. Also in Brunei and Kalimantan. 

Ecology. Mixed dipterocarp forests, rarely in swamps, to 300 ill. Flowering in May-October 

and fruiting in March, April, June, October and November. 

There is great variation in the size of leaflets and in the degree of pubescence. 

7. SCUTINANTHE Thwaites 

(Latin, scutum = shield, anthus = flower; the shape of the flower) 

in Hooker f, J. Bot. Kew Misc. 8 (1856) 266; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 420; 

Leenhouts I.e. (1952) 160, I.e. (1956) 246; Kochummen I.e. (1972) 152; Anderson I.e. 158; Wong I.e. 

215; Ng I.e. 38; Whitmore, Tantra & Sutisna I.e. 39. 

Dioecious trees; bole smooth with horizontal ring or occasionally scaly; inner bark 

raspberry-red with large beads of creamy to colourless exudate. Pith of branchlets and 

petioles without vascular strands. Leaves without stipules; leaflets entire. Inflorescences 

axillary, paniculate. Flowers unisexual, 5-merous, receptacle cup-shaped; sepals free; 

petals free with slightly thickened apex; stamens 10, confluent at base, episepalous ones 

longer, in female flowers slightly reduced in size, sterile; disc intrastaminal, adnate to 

receptacle; ovary hairy, 3-celled, only slightly reduced in male flowers. Fruits drupaceous, 

stigmatic scar nearly apical, pericarp fleshy; pyrene hard and bony, usually 2 cells strongly 

reduced; persistent calyx not enlarged. Seeds one; germination in S. brunnea epigeal; 

cotyledons leafY, entire; leaves of young seedling simple, alternate to spiral. 

Distribution. Two species in Sri Lanka, S Burma, W Malesia and Celebes. One species in 


Ecology. Lowland forests to 100 m. 

Uses. The light hardwood timber is commonly grouped together with other members of the 

family and sold as kedondong. 

97


c 

E 

2mm [ 

~ 

-........ ( 

G 1 2cm [ 

A 

Fig. 8. Triomma malaccensis. A, fruiting leafy twig; B, dehisced fruit; C, seed; D, part of 

inflorescence; E, flower bud; F, female flower; G, section through male flower. CA from SAN 37050, 

B-C from FRI 21209, D-E from SAN 40617, F-G after FM 1, 5 (1956) 219, fig. 8.) 

98


Scutinanthe brunnea Thwaites Fig. 7. 

(Latin, brunneus = brown; the indumentum on the inflorescence) 

in Hooker f I.e. (1856) 267; H.J. Lam, Bull. lard. Bot. Btzg. 3, 12 (1932) 420; Leenhouts I.e. (1952) 

162, I.e. (1956) 247; Burgess I.e. 62; Kochummen I.e. (1972) 152; Anderson I.e. 158; Whitmore, 

Tantra & Sutisna I.e. 39. Type: Thwaites 1149, Ceylon (K). Synonyms: Garuga brunnea (Thwaites) 

Marchand, Adansonia 8 (1867) 34, 66; Canarium brunneum (Thwaites) Beddome, Fl. Sylv. 1 (1868) 

t. 127. 

Medium-sized tree to 30 m tall, 30 cm diameter. Sapwood pale. Twigs brownish, 

lenticellate. Leaves with 3-6 pairs of leaflets; petiolules of lateral leaflets 1-2.5 cm long, 

swollen at both ends; blade elliptic to ob ovate, 8-21 x 4-8 cm; base broadly cuneate, 

unequal, apex pointed; midrib sunken above; lateral veins 8-10 pairs, raised below, faint 

above, curving and joining near margin; intercostal veins reticulate, visible on both 

surfaces. Inflorescences rusty red-pubescent. Flowers densely pubescent with greenish 

white inner face. Fruits ellipsoid, 4-6.5 x 2-3 cm, green, ripening yellow, with persistent 

calyx. 

Distribution. Sri Lanka, Sumatra, Peninsular Malaysia, and Borneo. In Sabah and Sarawak 

uncommon, known only by a few collections. Also in Brunei and Kalimantan. 


8. TRIOMMA Hook.! 

(Greek, tri = 3, omma = eyes or openings; 

the triangular seed showing 3 openings in cross-section) 

Trans. Linn. Soc. 23 (1860) 171; Ridley I.e. (1922) 369; H.J. Lam, Bull. lard. Bot. Btzg. 3, 12 (1932) 

331; Kalkman I.e. (1954) 499; Leenhouts I.e. (1956) 218; Burgess I.e. 62; Kochummen I.e. (1972) 

154; Cockbum I.e. 49; Anderson I.e. 158; Wong I.e. 248; Ng I.e. 38; Whitmore, Tantra & Sutisna I.e. 

39. 

Dioecious trees. Pith of branchlets without vascular strands. Stipules absent. Leaflets with 

entire margin, strongly asymmetrical at base; stalk not swollen. Inflorescences axillary 

panicles. Flowers unisexual, 5-merous; sepals and petals free; stamens 5, episepalous, base 

of filaments adnate to disc; disc extrastaminal, 5-lobed, lobes emarginate; ovary triangular, 

3-celled, in male flowers entirely reduced. Fruits 3-winged, dry, woody capsule, splitting 

into 3 valves. Seeds 3, broadly winged; germination epigeal; cotyledons shallowly 5-lobed, 

leafy; leaves of young seedling alternate to spiral, simple for the first few nodes, later with 

3 and 5 leaflets, margin initially toothed becoming entire later. 

Distribution. One species confined to W Malesia: 


99


Uses. The timber is commonly grouped with other members of the family and sold as 

kedondong. The wood is light hardwood, and used for indoor construction. The resin is 

sometimes used for torches. 

Triomma malaccensis Hook. f 

(of Malacca) 

Fig. 8. 

I.e. (1860) 171; King I.e. 236; Rid1ey I.e. (1922) 369; Merrilll.e. (1929) 119; H.J. Lam, Bull. Jard. 

Bot. Btzg. 3, 12 (1932) 332; Kalkman I.e. (1954) 499; Leenhouts I.e. (1956) 218; Burgess I.e. 62; 

Kochummen I.e. (1972) 154; Cockbum I.e. 49; Anderson I.e. 158; Whitmore, Tantra & Sutisna I.e. 

39. Type: Maingay 299, Malacca (ho1otype K; isotype L). 

Tree to 60 m tall, 100 cm diameter; buttresses to 5 m high, spreading. Bark smooth to 

scaly, grey-brown; inner bark pink or reddish brown, with a strong mango smell. Sapwood 

pale yellow. Twigs dark brown. Leaves with 2-5 pairs of leaflets; petiolules of leaflets not 

swollen; blade withering yellow with pink midrib and veins, oblong or ovate, 4-12 x 2-8 

cm; base strongly unequal with one half rounded, apex pointed; midrib raised above; 

lateral veins 6-11 pairs, raised on both surfaces; intercostal veins reticulate, distinct on 

both surfaces. Inflorescences many-branched, hairy. Flowers: sepals and petals densely 

tomentose. Fruits 5-7 x 2-2.5 cm. Seeds 3 with broad membranous wing. Saplings with 

densely hairy rachis and leaflets. 

Vernacular name. Sabah-kedondong asam (Malay). 

Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah a common emergent tree of 

the lowland forests, especially by streams. Widespread but not common in Sarawak, known 

from only two collections from Lambir National Park (s. 46465) and Limbang district (s. 

42833). Also occurs in Kalimantan. 

Ecology. Mixed dipterocarp forest on yellow sandy clay soils. Flowering in April-September 

and fruiting in May-December. 

100


CAPPARACEAE 

Deborah Kennard 

Department of Botany, 

University of Florida, U.S.A. 

Merrill, EB (1921) 280; Masamune, EPB (1940) 318; Jacobs, FM 1, 6 (1960) 61, FM 1, 7 (1976) 

822; Whitmore, TFM 2 (1973) 25; Anderson, CLTS (1980) 158; Ashton, MNDTS 2 (1988) 83; 

Corner, WSTM 1 (1988) 201; Whitmore, Tantra & Sutisna, CLK 1 (1989) 40. 

Trees, shrubs, woody climbers, or herbs. Leaves simple and entire, or trifoliolate or deeply 

5-7-lobed, alternate or spirally arranged, petioled; stipules thorny, minute or absent. 

Inflorescences racemose or paniculate, terminal or lateral, sometimes individual flowers 

axillary or serial; bracts, if present, small and caducous, rarely with stipular bracteoles. 

Flowers bisexual, regular or slightly irregular, sometimes big and showy, mostly in bud 

until anthesis but in Crateva opening at a very early stage; sepals 4, either equal or in two 

whorls of 2, free; petals 4 or (in Stixis) absent, equal or sometimes 2 of the petals slightly 

asymmetric and joined at the base, often free, usually stalked; receptacle more or less 

conical; stamens (4-)6 to many, all fertile, either free or their bases connate with the stalk 

of the ovary (gynophore) and forming an androgynophore; anthers dorsifixed; ovary superior, 

1-6-loculed, generally on a long gynophore, sometimes sessile, ovoid to cylindrical, with a 

small, simple sessile stigma. Fruit a leathery berry or a capsule. Seeds many, rarely 1 

(Stixis), mostly kidney-shaped; endosperm scanty. 

Distribution. Approximately 45 genera and 700 species in tropical and subtropical regions. 

In Sabah and Sarawak represented by 4 genera; Crateva with 2 species of trees, the other 3 

genera are mainly shrubs, woody climbers, and herbs. 

Ecology. Mostly in open places in the lowlands, a few ascend to 1000 m. Although species 

of the Capparaceae are usually found in dry regions, both Bornean species of Crateva are 

riverine trees. The flower of this family is characterised by long-radiating stamens, 

suggesting pollination by butterflies and moths. Several species are known to be nightflowering. 

Birds, bats, and ants have all been cited as dispersal agents. Trees of Crateva, 

however, have water-dispersed fruits. 

Uses. The familiar capers are from Capparis spinosa originating in the Mediterranean 

region. Several species are noted for their medicinal properties (c! Perry, MPESA (1980) 

68-70). Some tree species of Crateva are deemed to have religious significance and are 

planted for ornamental purposes as welL 

Taxonomy. The link between Capparaceae and Brassicaceae (the cabbage family) has long 

been recognised. The distinct mustard-like taste common in Brassica~eae is also present in 

101


many Cappparaceae, e.g. in the capers. Recent cladistic studies (Judd et aI., Harvard Papers 

in Botany 5 (1994) 1-51) based on morphological and genetic data strongly suggest that 

the Capparaceae is paraphyletic, and the.subfam. Cleomoideae is more closely related to the 

Brassicaceae than to the Capparaceae. 

Key to genera 

1. Trees. Leaves trifoliolate ................................................................................... Cratcva 

Shrubs, woody climbers, or herbs. Leaves simple or deeply 3-7-lobed ........................... 2 

2. Herbs. Leaves palmately, deeply 3-7-lobed. Fruit a long capsule ................................... . 

Clcomc L. 

Gen. PI. ed. 5 (1754) 302; Jacobs I.c. (1960) 99; Whitmore l.c. 25. 

A pantropical and subtropical genus with over 150 species, well represented in 

America; 4 species in Sabah and Sarawak. 

Annual or perennial herbs. Flowers in leafy, terminal racemes or panicles; sepals 

4, free; petals clawed, normally 4; stamens 6 to many. Capsule linear and 

cylindrical. Seeds many. 

Shrubs or woody climbers. Leaves simple. Fruit a leathery berry ................................. 3 

3. Plants with stipular thorns. Flowers on long stalks. Fruit with one to many seeds ......... . 

Capparis Tourn. ex L. 

Gen. PI. ed. 5 (1753) 222; Jacobs l.c. (1960) 70, B1umea 12, 3 (1965) 405; Whitmore I.c. 

25. 

About 250 species in the tropics and subtropics, half of which are in the New 

World; 7 species in Sabah and Sarawak. 

Shrubs, woody climbers, or rarely small trees. Leaves simple, spiral, with a pair of 

stipular thorns which are occasionally wanting. Flowers single or in small, sessile 

groups just above the leafaxils, feathery with long stamens; sepals biseriate, 

mostly imbricate; petals 4; stamens many, rarely (7-)8(-12), free, radiating, 

longer than the petals; ovary I-locular. Berry globular to ellipsoid with a corky or 

leathery pericarp. Seeds 1 to many, kidney-shaped. 

Plants without thorns. Flowers on short stalks. Fruit with 1 seed ............ . 

Stixis Lour. 

Fl. eoch. (1790) 295, ed. Willdenow (1793) 361; Jacobs l.c. (1960) 95, Blumea 12 (1964) 

5; Whitmore I.e. 25. 

7 species distributed throughout India, Burma, Cambodia, Hainan, and W Malesia. 

One species endemic to Borneo, S. ovata, occurs in Sabah and Sarawak. 

Rather small unarmed, woody climbers. Branches lenticellate. Leaves simple, 

rather large. Inflorescences racemose or paniculate. Flowers many, axillary or 

terminal; petals absent; sepals mostly 6, in two whorls of 3; stamens 20-50(-100), 

about as long as sepals; ovary subglobular. Fruits few, on a thick woody stalk, 

ellipsoid, 2.5-5 cm long. Seed one, large. 

102

CAPPARACEAE(KENNARD) 

CRATEVAL. 

(Kratevas, 132-63 RC., an ancient Greek medicine man or root gatherer) 

I.e. (1754) 203; Jacobs I.e. (1960) 63, I.e. (1964) 186; \Yhitmore I.e. 26; Anderson I.e. 158; Corner 

I.e. 203; Whitmore, Tantra & Sutisna I.e. 40. 

Small to medium-sized trees. Bark grey or brown, smooth. Twigs round, glabrous, with 

distinct leaf-scars, stipules small, not persistent; axillary buds small. Leaves spiral, 

palmately lobed or trifoliolate, sessile· to shortly stalked, base of stalks bearing small 

appendages to 1 mm. Racemes terminal, corymbiform, either with arrested growth or 

growing through and developing into a leafy twig with lateral flowers; pedicels leaving 

distinct scars on the rachis. Flowers large and showy, to 5 cm across, opening while green 

and young, subtended by bracts; sepals 4, clawed, green; petals 4, clawed, first white then 

cream-coloured; stamens (8-)12-30, filaments at the very base connate with the gynophore, 

5-7 cm, thread-like, pinkish violet; gynophore approximately as long as the stamens; 

receptacle dish-shaped, floral parts not persistent. Fruits large, round or oblong berries, 

with tough, sometimes papillate skin, hanging on a thickened woody pedicel and 

androgynophore, together to 14 cm long; torus leaving a distinct scar. Seeds numerous, 

heart- or horseshoe-shaped, embedded in pulp. 

Distribution. About 6 species distributed throughout the tropics, except in Australia and 

New Caledonia. Two species in Sabah and Sarawak. 

Ecology. Found mostly in periodically flooded lowland forest near rivers, usually below 

700 m. Fruits are dispersed by water, possibly also by fish. 

Key to Crateva species 

Leaflets with 10-15 pairs oflateral veins, paler beneath than above .................. 1. C. magna 

Leaflets with 5-11 pairs oflateral veins, same colour beneath as above .......... 2. C. religiosa 

1. Crateva magna (Lour.) DC. Fig. lA-C. 

(Latin, magnus = great, large; referring to the stature) 

Prod. 1 (1824) 243; Merrill, Comm. Lour. (1935) 172; Masamune I.e. 318; Jacobs I.e. (1964) 206, 

I.e. (1976) 822; Whitmore I.e. 26; Corner I.e. 203; Whitmore, Tantra & Sutisna I.e. 40. Basionym: 

Capparis magna Lour., Fl. Cochinch. 1 (1790) 331. Type: Loureiro, s.n., "Indochina" (BM). 

Synonyms: Crateva nurvala Buch.-Ham., Trans. Linn. Soc. 15 (1827) 121; C. lophosperma Kurz, 1. 

Bot. 12 (1874) 1,95, t. 147, f. 4-6. 

Tree to 20 m tall, 40 cm diameter. Twigs terete, lenticellate. Petiole 3-11 cm. Leaflets 

firmly herbaceous to slightly coriaceous, paler beneath, mostly lanceolate; apex gradually 

acuminate with an acute tip; lateral leaflets symmetrical, 8-17 x 2-6 cm, usually subsessile, 

sometimes with stalk 3-6 mm long; lateral veins 10-15 pairs, prominent beneath. 

Inflorescences terminal on leafy twigs to 12 cm, bearing 20 to 100 flowers; pedicels 4-7 

103


cm long; petals 4, ovate-spathulate with a 5-12 mm stalk, lobes 15-30 x 15-22 mm; sepals 

4, ovate, 2-3.5 x 1-1.5 mm; stamens 15-25, 3.5-4.5 cm long, filaments purple; ovary 

oblong to cylindrical, gynophore 3.4-4.5 cm. Fruits ellipsoid, sometimes globose, c. 5 x 4 

cm; pericarp covered with a yellow-greyish crust which eventually breaks off. Seeds 

horseshoe-shaped, 6-9 mm across, dull dark-brown. 

Vernacular names. Sabah-mempulak (Murut), pangos (Dusun Kinabatangan). Sarawakserang 

(Than, in confusion with Me/icope). Brunei-kebuan (Than). Kalimantan-sasgah, 

sebelu, tigarun (Than). 

Distribution. India, Burma, China, Hainan, Indo-China, Thailand, Sumatra, Peninsular 

Malaysia, Java and Borneo. In Sabah, recorded from the Kalabakan, Lahad Datu, Lamas, 

Sandakan and Telupid districts. In Sarawak from the 3rd, 4th and 7th Div., in all cases 

near limestone or basalt exposures. Also in Brunei and Kalimantan. 

Ecology. Found along streams and rivers or the edge of swamps; rarely found on ridges 

although it can occur up to 1000 m. Flowering and fruiting irregular. 

Uses. In Sabah and Sarawak, poultices made from the root, leaf, or bark are used externally 

for saki! angin ("wind"), poultices made from the bark alone are used externally to reduce 

high fever. The wood is sometimes used as timber. Elsewhere, the juice from the bark has 

been used to increase the appetite and as a laxative. The fruits are used in Thailand as fish 

bait. 

2. Crateva religiosa G. Forst. Fig. ID. 

(alluding to its common cultivation around temples in Tahiti) 

PI. Escul. Ins. Oc. Austral. (1786) 45; Masamune I.e. 318; Jacobs I.e. (1960) 65, I.e. (1964) 191; 

Whitmore I.e. 26; Anderson I.e. 158; Corner I.e. 203; Ashton I.e. 84; Whitmore, Tantra & Sutisna I.e. 

40. Type: G. Forster, s.n., Pacific Society Isl. (K). Synonyms: C. membranifolia Miq., Fl. Ind. Bat., 

Suppl. (1861) 387; C. brownii Korth. ex Miq., Illustr. (1870) 21; C. macroearpa Kurz I.e. 195, t. 

148, f. 8-10. 

Shrub or tree, to 30 m tall and 30 cm diameter. Twigs with many large lentice1s. Petiole 

(3-)6':"'10 cm. Leaflets thin herbaceous, oblong to ovate, top rather abruptly acuminate, (5- 

)8-16(-27) x (3-)4-10 cm, mostly subsessile, sometimes stalked to 5 mm, concolorous 

when dried; lateral veins 7-11 pairs. Flowers few to 25, toward the end of leafy twigs 

which do not grow through; pedicels 2-

CAPPARACEAE(KENNARD) 

Fig. 1. Crateva magna CA-C) and C. religiosa CD). A, flowering leafy twig; E, fruit on thickened 

pedicel and androgynophore; C, seed; D, single leaf. (Drawn from fresh material from Danum Valley 

Conservation Area, Sabah.) 

105


Ecology. Common in periodically inundated forest along rivers; rarely in secondary or 

primary dryland forest. Mostly found below 100 m but occasionally also at about 700 m. 

Flowering and fruiting irregular. 

Uses. Occult power has been ascribed to C. religiosa in India and Polynesia where it is 

planted around temples. In the Solomon Islands the leaves are used for ear-ache and the 

fruit is used to relieve constipation. In Borneo, C. religiosa is occasionally planted for 

ornamental purposes, and the fruits are also used for fish bait. 

106


CELASTRACEAE 

K.M. Kochummen 



Merrill, EB (1921) 354; Rid1ey, FMP 1 (1922) 443; Masamune, EPB (1942) 417; Browne, FTSB 

(1955) 76; Ding Hou, FM 1,6 (1962) 227, FM 1, 6 (1964) 389, FM 1, 6 (1972) 930, Blumea 17 

(1969) 97; Backer & BakhuizenJ, FJ 2 (1965) 53; Smythies, CST (1965) 38; Burgess, TBS (1966) 

73; Kochummen & Whitmore, TFM 1 (1972) 156; Cockburn, TS 1 (1973) 51; Anderson, CLTS 

(1980) 159; Wong, DMT (1982) 33; Corner, WSTM 1 (1988) 212; Ashton, MNDT 2 (1988) 87; 

Whitmore, Tantra & Sutisna, CLK 1 (1989) 42; Ng, MFR 34 (1991) 41. 

Trees, erect or scandent shrubs or woody climbers; bole sometimes buttressed, rarely with 

pneumatophores. Leaves simple, decussate, opposite, sub-opposite, spiral or rarely alternate, 

sometimes black-dotted especially beneath, margin faintly toothed, wavy or entire. Stipules 

small, falling off early or absent. Incipient bracts absent except in Microfropis. Inflorescences 

axillary and/or terminal, sometimes extra-axillary or borne on the branches, thyrsoid, 

paniculate, racemose, cymose or fasciculate, usually with bracts. Flowers regular, bisexual 

or unisexual (plants dioecious, rarely polygamous); calyx 4-5-lobed, lobes imbricate, rarely 

valvate, persistent; petals 4 or 5, imbricate, contorted or rarely valvate, falling off early, 

rarely persistent, free or united at base, sometimes united with the staminal ring, always 

larger than calyx-lobes except in Perrottetia; stamens 2-5, alternate with petals, filaments 

inserted on or on the inner side of the disc or on a basal ring (Microtropis), anthers 2- 

celled, longitudinally or laterally dehiscent, introrse or extrorse; disc often conspicuous, 

fleshy or membranous, cup-shaped, saucer-shaped or flat, entire, toothed, angled or lobed, 

extrastaminal or intrastaminal, rarely absent (Microtropis); ovary superior or rarely semiinferior, 

partly or fully immersed in the disc, usually glabrous, sometimes with a tuft of 

hairs at top (Bhesa), rarely with papilla-like or fleshy subulate processes at the base 

(Euonymus), (l-)2-5-celled, rarely many-celled (Siphonodon), very rarely hollow at the top 

(Siphonodon), style distinct, or obscure or lacking, simple or divided to the base (Bhesa), 

stigmas simple or lobed; ovules usually 2 in each cell, sometimes 1 or 3-18, anatropous. 

Fruits drupes, capsules, or berries, surface smooth or spiny. Seeds erect or pendulous, 

sometimes winged, aril present or absent, when present usually partly or entirely enveloping 

the seed or forming a cushion-like structure situated at the base of seed or with long, 

jilamentous appendages (Sarawakodendron), often orange or orange-red, rarely white; 

endosperm present or absent; cotyledons flat, foliaceous. 

Distribution. About 90 genera and over 1000 species distributed in both hemispheres, 

predominant in the tropics and subtropics. In Malesia, 18 genera and c. 115 species, of 

which 15 genera and 70 species are recorded in Sabah and Sarawak. 

Ecology. From lowland including mangrove and peat swamp, to montane forests to about 

3200 m. Most members of the family have nectariferous disc in the flowers which are 

107


visited by various kinds of insect -pollinators. Seeds of species having brightly coloured arils 

are dispersed by birds; the winged seeds of Kokoona, Lophopetalum and other taxa are 

dispersed by wind. The fruits of the mangrove tree, Cassine viburnifolia, have a corky 

mesocarp that facilitates dispersal by sea water. 

Uses. Timbers of the genera Kokoona, Lophopetalum, and Siphonodon have commercial 

value. Some species contain alkaloids. Aborigines in Peninsular Malaysia make use of the 

bark of Lophopetalum pallidum in the preparation of dart poison. The ochre yellow middle 

bark of some Lophopetalum and Kokoona species burn readily and is useful tinder in the 

forest. 

Key to genera 

1. Woody climbers or scandent shrubs ................................................................................. 2 

Trees, or erect shrubs ............................................................................................................. 6 

2. Leaves spirally arranged ........................................................................................................ 3 

Leaves opposite, decussate, or rarely spiraL ...................................................................... 4 

3. Woody climber. Ovary free from the disc ...................................................................... . 

Celastrus L. 

Gen. PI. ed. 5 (1754) 91; Ridley I.c. 450; Ding Hou I.c. (1963) 233. 

About 30 species, tropics and subtropics; 2 species in Sabah and Sarawak. 

Leaves spirally arranged, margin toothed. Inflorescences cymose or paniculate. 

Flowers unisexual, 5-merous, pedicel articulated; calyx bell-shaped; disc membranous 

or fleshy, cup-shaped; ovary superior, 3-celled; style columnar, stigma 3-lobed; 

ovules 2 in each cell. Fruit a 3-valved capsule, tipped by persistent style. Seeds 1- 

6, enveloped by crimson aril. 

Scandent shrubs. Ovary partially immersed in the disc ............. . 

May tenus Mollina 

Saggio Chile (1782) 177; Ding Hou I.c. (1962) 238. 

About 225 species; tropics and subtropics of both the Old and New World. In 

Malesia 5 species, one of which, M emarginatus (Willd.) Ding Hou, is present in 

Sabah; uncommon, known only by a single collection (Sugau 34) from Balambangan 

island. 

Scandent shrubs with spines on leafy shoots. Leaves spirally arranged, margin 

toothed. Flowers in fascicles, unisexual, 4-5-merous; calyx-lobes deltoid; petals 

obovate or oblong, entire; disc rounded; stamens inserted on the margin of the 

disc; ovary partly immersed in the disc, 3-celled, style cylindric, stigma 3. Fruits a 

capsule. Seeds ellipsoid, red, subtended by white fleshy aril at the base. 

4. Flowers with distinct 

Flowers without or with inconspicuous disc .......................... . 

. Rcissantia Halle 

Mus. Hist. Nat. Paris 30 (1958) 466; Ding Hou l.c. (1964) 400. 

7 species, Old World tropics; 4 species in Sabah and Sarawak. 

108

CELASTRACEAE (KOCHUMMEN) 

Leaves decussate, rarely subopposite. Inflorescences axillary. Flowers bisexual; calyxlobes 

5, imbricate; petals 5, imbricate; disc inconspicuous; stamens 3; ovary semisuperior, 

3-celled; style short; stigma obscure; ovules 2 (rarely 4-8) in each cell. 

Fruits capsular consisting of 3 divergent separate follicles which split into 2 valves. 

Seeds with basal more or less transparent membranous wing. 

5. Fruits capsular with 3 follicles. Seeds winged. Petals with inflexed tip .......................... . 

LocscncricIla A. C. Smith 

Am. J. Bot. 28 (1941) 438; Ding Hau I.e. (1964) 397. 

26 species, Tropical Africa, Asia, Malesia, New Hebrides. In Malesia, 4 species of 

which 2 are present in Sabah and Sarawak. 

Leaves decussate. Inflorescences axillary, cymose. Flowers bisexual; calyx deeply 

5-lobed; petals 5, thick with injlexed tip; disc fleshy; stamens 3; ovary superior or 

semi-inferior. Fruits capsular, with 3 divergent follicles, each dehiscing into 2 

valves. Seeds usually with basal membranous wing. 

Fruits drupaceous. Seeds embedded in pulp. Petals without inflexed tip ........................ . 

Salacia L. 

Mant. (1767) 159; Ridleyl.e. (1922)456; Ding Hau I.e. (1964)404. 

About 150 species, pantropical; 33 species in Malesia of which l7 are found in 

Sabah and Sarawak. Leaves decussate, sub opposite or rarely spiral. Flowers bisexual, 

axillary, in clusters or organised in cymes or panicles; calyx deeply 5-lobed 

or 3-5-lobed in apical part, and circumscissile at the base or lengthwise splitting or 

not lobed; petals usually 5, rarely 4 or 7; disc fleshy, ring-like, truncate-conical or 

flattened, occasionally cup-like; stamens 3, rarely 2; ovary semi-inferior, 3-celled, 

rarely 2-celled, ovules 2-8 in each cell; style distinct or obscure; stigma obscure. 

Fruits drupaceous, 1-3-celled. Seeds 1 to several, embedded in mucilaginous pulp. 

6. Leaves alternate or spirally arranged ............................................................................ 7 

Leaves decussate, opposite, subopposite, or very rarely alternate or spiral.. .......... 10 

7. Leaf margin toothed ..................................................................................................... 8 

Leaf margin 

8. Petals usually similar in size to calyx-lobes. Ovary not hollow in apical part ... S. Pcrrottetia 

Petals always larger than calyx-lobes. Ovary hollow in apical part ......... l0. Siphonodon 

9. Leaves and twigs with pustules; petiole to 1.5 cm long, not swollen towards apex; 

stipules not prominent, leaving no stipular scars on twigs ............. 9. Sarawakodcndron 

Leaves and twigs without pustules; petioles longer than 1.5 cm, swollen at the apex or 

both ends; stipules prominent, leaving scars on twigs ....................................... 1. Bhcsa 

10. Incipient bracts prominent. Disc absent.. ................................................ 7. Microtropis 

Incipient bracts absent. Disc present. .............................................................................. 11 

11. Trees confined to tidal rivers and mangroves. Ovary 2-celled ..... .2. Cassinc 

Trees or shrubs of inland forests. Ovary 3 or 4-5-celled ............ .. .. ........... 12 

109


E 

, 

(D.!ilI 1 L 

D:t, 

.. I 

l' .. 

B 

A 

Fig. 1. Bhesa paniculata. A, flowering leafY twig; E, flower; C, flower with pistil removed; D, pistil; 

E, part of infructescence. CA from SAN 79200, E-D after FM 1, 6 (1962) 281, fig. 16, E from SAN 

82521.) 

110


12. Ovary 4-5-celled. Seeds not winged ........................................................................... 13 

Ovary 3 -celled. Seeds winged .................................................................................. 14 

13 . Ovules 2 in each cell... ............. . ................................................ 3. Euonymus 

Ovules 1 in each 

Glyptopetalum 

14. Petals twisted and overlapping. Wing attached to the apex of seed ......... 5. Kokoona 

Petals imbricate. Wing surrounding seed .............................................. 6. Lophopetalum 

1. BHESA Buch.-Ham. ex Am. 

(origin unknown, probably an English plant name) 

Edin. NewPhil. J. (1834) 315; Ding Hou, Blumea Suppl. 4 (1958) 149, I.c. (1962) 280; Kochummen 

& Whitmore I.c. 158; Cockburn I.c. 54; Anderson I.c. 159; Wong l.c. 22; Corner l.c. 213; Ashton l.c. 

89; Whitmore, Tantra & Sutisna l.c. 42; Ng I.e. 43. Synonym: Kurrimia Wall. ex Am., Nov. Act. Ac. 

Caes. Leop.-Car. 18 (1836) 328. 

Medium-sized to tall ~rees; bole with buttresses, often fluted at base. Bark grey-brown, 

smooth to cracking and scaly; inner bark mottled cream and orange. Sapwood pale yellow. 

Twigs smooth, with distinct stipular scars. Stipules prominent, lanceolate, soon falling off 

and leaving scars on the twigs. Leaves spiral, entire, with silky sheen; intercostal veins 

fine, scalariform; petioles slender, longer than 1.5 cm, swollen at both ends or at the apex 

only. Inflorescences axillary, solitary or paired, paniculate or racemose; bracts falling off 

early; pedicels jointed. Flowers bisexual, 5-merous (rarely 4-merous); calyx deeply lobed, 

lobes imbricate or valvate; petals twisted; disc fleshy, entire or lobed; stamens 4-5, seated 

on or just below the disc; ovary superior, usually with terminal tuft of hairs, 2-celled, with 

2 ovules in each cell, styles 2. Fruit a capsule, entire or 2-lobed, splitting into 2 valves or 

on one side only. Seeds 1-2 per cell, with bright orange-red or pink basal aril; endosperm 

copious; germination epigeaJ. 

Distribution. 5 species; Sri Lanka and Malesia; 2 species in Sabah and Sarawak. 

Ecology. Widely distributed from lowland to submontane forests to 1500 m. 

Key to Bhesa species 

Stipules to 3 cm long. Petioles strongly swollen at both ends. Intercostal veins distinct. 

Inflorescences usually paniculate. Disc deeply 5-lobed. Fruits two-Iobed ..... 1. B. paniculata 

Stipules to 1 cm long. Petioles only slightly swollen at apex. Intercostal veins faint to 

invisible. Inflorescence racemose. Disc entire or obscurely notched. Fruits not lobed, with 

pointed tip ........................................................................................................ 2. B. robusta 

III


1. Bhesa paniculata Am. Fig. 1. 

(Latin, paniculatus = having a loose, branched flower-cluster or inflorescence) 

l.c. (1834) 315; Ding Hou I.c. (1958) 151, I.c. (1962) 282; Burgess I.c. 73; Koehummen & Whitmore 

I.c. 160; Coekburn I.c. 55; Anderson I.c. 159; Corner I.c: 213; Ashton I.c. 92; Whitmore, Tantra & 

Sutisna I.c. 42. Type: Wallich no. 4336, Penang (BO, K). Synonyms: Kurrimia paniculata Wall. ex 

Am. I.c. 328; K. luzonica Vidal, Rev. PI. Vase. Filip. (1886) 88; K. minor Ridl., Kew Bull. (1938) 

235. 

Medium-sized tree to 35 m tall, 50 cm diameter; bole often fluted at base. Bark greybrown, 

smooth to cracking; inner bark orange-yellow. Sapwood pale white. Twigs pale 

brown to dark brown, 5-10 mm thick, with prominent stipular and leaf scars. Stipules 

lanceolate, c. 3 cm long. Leaves elliptic to oblong, rarely obovate, 5.5-27 x 2.2-13 cm; 

base cuneate or rounded, apex pointed or blunt; midrib flattened or raised above; lateral 

veins 5-20 pairs, prominently raised below, visible above; intercostal veins scalariform, 

very close, faint above, distinct beneath; petioles 1-9.5 cm long, strongly swollen at both 

ends. Inflorescences panicles, 10-37 cm long or racemes to 20 cm long, from axils of 

upper leaves. Flowers greenish yellow or dark purplish red, pedicels 2-3 mm long; calyxlobes 

hairy outside; disc deeply 5-lobed; petals oblong or ovate, hairy inside; stamens l.5-2 

mm long, anthers triangular; ovary ellipsoid, styles free, about half as long as ovary. Fruits 

2-lobed, red when fresh, drying dark brown, 1-2 cm long, with the biggest lobe c. 0.5 cm 

wide. Seeds 2-4, more than half covered with pink aril; cotyledons leafy. 

Vernacular names. Sabah-biku-biku (Malay). Sarawak-simun (lban). Brunei-serunai 

(Malay). 

Distribution. S India, S Thailand, Sumatra, Peninsular Malaysia, Borneo and the 

Philippines. Common and widely distributed in Sabah and Sarawak. Also in Brunei and 

Kalimantan. 

Ecology. Lowland, including mixed dipterocarp, heath, and peat swamp, io submontane 

forests to 1500 m. Submontane samples have comparatively smaller and thicker leaves. 

Flowering in March-May and August-October, fruiting in April-December. 

2. Bhesa robusta (Roxb.) Ding HOll 

(Latin, robustus = strong-growing, robust; the habit) 

I.c. (1958) 152,I.c. (1962) 283; Koehummen & Whitmore l.c. 161; Anderson I.c. 159; Corner I.c. 

214; Ashton l.c. 93; Whitmore, Tantra & Sutisna I.c. 42. Basionym: Celastrus robustus Roxb., Fl. 

Ind. 2 (1824) 395. Type: Roxburgh,Icones no. 2185 (K). Synonyms: Kurrimia pulcherrima Wall. ex 

Laws. in Hookerf, Fl. Br. Ind. 1 (1875) 622; Kurrimia maingayi Laws. in Hookerf I.c. 622. 

Medium-sized tree to 18 m tall and 40 cm diameter. Bark grey-brown; inner bark yellow. 

Sapwood pale. Stipules lanceolate, 5-10 mm long. Leaves elliptic or oblong, 7-9 x 3-4 

cm; base cuneate, apex blunt or pointed; midrib raised above; lateral veins 11-15 pairs; 

intercostal veins very faint to invisible; petioles 1-3 cm long, slightly swollen at apex. 

Flowers in racemes, subsessile; calyx lobed, lobes broadly ovate to rounded; petals oblong 

112


to elliptic; disc cup-shaped, sub entire or obscurely notched; stamens c. 2 mm long, attached 

beneath the outer margin of disc, anthers deltoid; ovary subglobose with tuft of hairs at 

apex, styles free, longer than ovary. Fruits ovoid, not lobed, c. 2 x 1 cm, apex pointed. 

Seeds 1, enveloped by aril, sometimes only the lower half; cotyledons fleshy. . 

Distribution. NE India, Bhutan, Chittagong, Burma, Andaman Islands, Thailand, Indo-China, 

Sumatra, Peninsular Malaysia, and Borneo. In Sabah and Sarawak, uncommon; in Sabah, 

known from 3 collections (SAN A 6613, SAN 50057, and SAN 76690) from Beaufort and 

Sandakan districts, and in Sarawak by one collection (Beccari 2624). 

Ecology. Mixed dipterocarp forests at 300-600 m, on clay-rich soils. 

Uses. Though of no commercial importance in Sabah and Sarawak, the timber is used in 

house-building in other countries where it occurs. 

2. CASSINE L. 

(origin and meaning unknown) 

Gen. PI. (1737) 338; Ding Hou I.c. (1962) 284; Backer & BakhuizenJ l.c. 55; Kochummen & 

Whitmore I.c. 161; Anderson I.c. 159; Kostermans, Gard. Bull. Sing. 39 (1986) 177; Ashton l.c. 93. 

Synonym: Elaeodendron Jacq.J ex Jacq., lc. PI. Rar. 12 (1782) t. 48. 

Shrubs or small trees. Leaves decussate. Inflorescences axillary or extra-axillary cymes, 

with distinct peduncles. Flowers bisexual, 4-5-merous; calyx-lobes imbricate; petals free, 

imbricate; stamens 4-5, inserted on the disc or on its outer margin, filaments subulate, 

anthers introrse; disc prominent, fleshy, flat, orbicular or lobed; ovary semi-inferior, 2- 

celled, conical or flask-like, the base slightly united with the disc or partly immersed in it; 

style very short or obscure, stigma obscure or 2-lobed; ovules 2 in each cell, erect, attached 

at the base. Fruit a drupe, 1-2-celled. Seeds 1-2, not winged, without aril, with endosperm. 

Distribution. About 80 species throughout the tropics, mainly in Africa; 2 species in 

Malesia of which only one is present in Sabah and Sarawak. 

Ecology. In Sabah and Sarawak, confined to mangroves and banks of tidal rivers. 

Cas sine viburnifolia (Juss.) Ding Hou 

(with leaves resembling those of Viburnum) 

Fig. 2. 

l.c. (1962) 286; Kochummen & Whitmore I.c. 161; Cockbum I.c. 54; Anderson I.c. 159; Ashton l.c. 

93. Basionym: Aegiphila viburnifolia Juss., Ann. Mus. Hist. Nat. Paris 7 (1806) 76. Type: Sine 

call., s.n., Philippines (P). Synonyms: Euonymus viburnifolius (Juss.) Merr., Philip. 1. Sc. 9 (1914) 

Bot. 312; Elaeodendron subrotundum King, J. As. Soc. Beng. 65,2 (1896) 356. 

Shrub or small tree to 10 m tall, 20 cm diameter. Bark yellowish grey, smooth. Twigs 

black. Leaves broadly ob ovate, 4-10 x 2-6 cm; base cuneate, margin curled inwards, 

113


remotely minutely toothed, apex blunt to rounded; lateral veins 4-6 pairs, faint; intercostal 

veins reticulate, equally prominent as the lateral ones; petioles 8-13 mm long. Inflorescences 

cymose, axillary. Flowers white, 4-merous; calyx-lobes broadly ovate to rounded, 

almost free; petals oblong to ovate-oblong. Fruits obovoid, often rhomboid in cross-section; 

mesocarp thick and corky. Seed 1, c. 6 x 3 mm, obovate to oblong. 

Vernacular name. Sarawak~barat-barat (Malay). 

Distribution. Andaman Islands, Thailand, Sumatra, Peninsular Malaysia, Borneo, Philippines, 

and Celebes. In Sabah known from Kudat, Sandakan and Tawau districts, and in Sarawak 

from the Rejang delta in 3rd. Div. Also in Brunei. 

Ecology. Common on the banks of tidal rivers and mangrove channels near the inland 

limits of salinity. The fruits are dispersed by water. 

3. EUONYMUS Tourn. ex L. 

(Greek, eu = good, onuma = name; a plant name) 

Gen. PI. ed. 5 (1754) 91; Ridley I.c. (1922) 445; Masamune l.c. 417; Blake1ock, Kew Bull. (1951) 

232; Ding Hou l.c. (1962) 245; Backer & Bakhuizenj l.c. 53; Kochununen & Whitmorc i.c. 162; 

Cockbum I.c. 55; Andcrson I.c. 159; Ashton I.c. 94; Whitmore, Tantra & Sutisna l.c. 42. 

Shrubs or small trees. Leaves opposite-decussate sometimes with dark spots on the under 

surface; stipules lanceolate, falling off early. Inflorescences axillary, cymose; rarely flowers in 

fascicles (E. javanicus); pedicels jointed. Flowers bisexual, 5- or 4-merous; calyx deeply 

lobed, lobes imbricate, entire or minutely toothed; petals free, imbricate, spreading or 

reflexed; disc prominent, fleshy or thin, flat, 5- or 4-angled or 5- or 4-lobed or rounded, 

smooth or covered with fleshy papilla-like or subulate processes; stamens 4-5, inserted on 

the disc, anthers 2-ceIIed, lateral or introrse, dehiscence apical, filaments obscure or 

distinct; ovary semi-inferior 4-5-celled, partly or wholly immersed in the disc, stigma 

obscure or discoid, ovules mostly 2 in each cell. Fruits capsules, usually (3-)4-5-angular 

or lobed, smooth or armed with rigid prickles, apex obtuse, acute, truncate or concave. 

Seeds usually black, not winged, partly or completely covered by orange ariI. 

Distribution. About 180 species, mainly in the tropics and subtropics. In MaIcsia, 12 

species of which 5 are recorded in Sabah and Sarawak. 

Ecology. In primary and secondary forests from lowlands to mountains to 3200 m. 

Key to Euonymus species 

(based on inflorescences and flowers) 

1. Flowers 5-merous. Leaves without black gland-dots below ............................................ 2 

Flowers 4-merous. Leaves with black gland-dots below ............................................... .4 

114

CELASTRACEAE (KO CHUM MEN) 

@]o.5mm 

c 

o 

"m f 

\ 

Fig. 2. Cassine viburnifolia. A, flowering leafy twig; B, half-flower; C, transverse section through 

ovary; D, infructescence. (A from SAN 61255, B & C after FM I, 6 (1962) 285, fig. 18, D from SAN 

38943.) 

115


D ~ ]o.5mm 

~ 

c 

I1mm 

4cm 

B 

sd 

E 

A 

Fig. 3. Euonymus castanaefolius. A, flowering leafy twig; B, flower with petals and stamens 

removed; C, petal; D, young stamen; E, part of infructescence. CA from SAN 93113, B-D after FM 1, 

6 (1962) 246, fig. 4, E from S. 47567.) 

116


2. Petals fimbriate ................................................................................................................ 3 

Petals entire ........... 

...... 1. E. acuminifolius 

3. Peduncles distinct... .......................... E. cochinchinensis 

Peduncles obscure ............. 

........................ 5. E. javanicus 

4. Lateral veins of the leaves sunken above .......................................... 2. E. castaneifolius 

Lateral veins of the leaves not sunken above ...................................... .4. E. glandulosus 

Key to Euonymus species 

(based on leaves and twigs) 

1. Leaf margin toothed ..................................................................................................... 2 

Leaf margin entire (sometimes faintly toothed in E. cochinchinensis) ............................. 3 

2. Leaf lower surface with scattered black gland-dots. Lateral veins sunken above ............ . 

E. castaneifolius 

Leaf lower surface without black gland-dots. Lateral veins not sunken above ................ . 

....................................................................................................... . 1. E. acuminifolius 

3. Twigs usually 4-angled. Leaf lower surface with scattered black gland-dots; margin 

strongly recurved ............................................................................... .4. E. glandulosus 

Twigs not 4-angled. Leaf lower surface without scattered black gland-dots; margin not 

strongly recurved ........................................................................................................ 4 

4. Leaf margin entire. Peduncle obscure ....................................................... 5. E. javanicus 

Leaf margin faintly toothed. Peduncle distinct... ........................... 3. E. cochinchinensis 

1. Euonymus acuminifoIius Blakelock 

(Latin, acuminatus = long-pointed,folius = leaves) 

I.e. 253; Ding Hou I.e. (1962) 251; Whitmore, Tantra & Sutisna I.e. 43. Type: Bunnemeijer 581, 

Sumatra, Ophir district, NW slopes ofMt. Ta1amau (Ba). Synonym: E. aeuminifolius Blakelock var. 

borneensis Blakelock I.e. 253. 

Shrub to 4 m tall. Twigs 4-angled. Leaves membranous to chartaceous, ovate-oblong to 

lanceolate or elliptic to oblong, 7-10.5 x 2-4 cm, lower surface without black gland-dots; 

base cuneate, margin toothed, apex pointed, tip c. 2 cm long; lateral veins 4-6 pairs, 

arching near margin, not sunken above; petioles 2-6 mm long. Flowers in cymes, purplish 

red, 5-merous; calyx-lobes rounded, inner two usually larger; petals almost rounded, entire; 

disc obscurely 5-angular; stamens with very short filaments, anthers c. 0.5 mm long; ovary 

faintly 5-angled, stigma discoid, obscurely 5-angled. Fruits obcordate, distinctly 5-lobed, 

1.5-2 x 1-1.7 cm. Seeds with aril at base. 

Distribution. Sumatra, Borneo and Celebes. In Sabah uncommon, known only from Mt. 

Kinabalu (Clemens 30350, type of E. acuminifulius var. borneensis, and Clemens 34478). 

Not recorded from Sarawak. 

Ecology. Montane forest at about 2700 m. 

117


~ ",] 

"."" 

F;


Distribution. Nicobar Is., Burma, Thailand, Indo-China, Java, Borneo, Natuna Is., Philippines, 

Celebes, Lesser Sunda Isl., Moluccas, and New Guinea. In Sabah, known from Lahad Datu, 

Ranau, Tawau and Tenom districts. In Sarawak, uncommon, recorded from the Bukit 

Numpang and Tai Ton forests. 

Ecology. Lowland (including limestone) to montane forests to 2400 m. 

4. GLYPTOPETALUM Thwaites 

(Greek, glypto = incised, petalum = petals) 

in Hooker, J. Bot. Kew Mise. 8 (1856) 267; Ridley I.c. (1922) 446; Ding Hou l.c. (1962) 254; 

Koehummen & Whitmore I.c. 162; Anderson I.c. 160; Ashton I.c. 95. 

Shrubs or small trees. Leaves opposite. Inflorescences cymose, axillary or extra-axillary. 

Flowers bisexual, 4-merous; calyx spreading, inner pair larger; petals free, fleshy with 

small appendage or depression inside; disc conspicuous, fleshy; stamens 4, inserted on the 

disc, near base of ovary, anthers opening apica//y; ovary semi-inferior, immersed in the 

disc, 4-ceIIed, style obscure, stigma obscure, ovule one in each ceII. Fruits capsules, when 

splitting leaving a persistent central columeIIa. Seeds with fleshy aril at lower half. 

Distribution. c. 20 species; India, Sri Lanka, Burma to Hainan, and Malesia. 8 species are 

present in Malesia; 2 in Sabah and Sarawak. 


Taxonomy. The genus is closely allied to Euonymus from which it differs in having only 

one ovule per ovary-cell. 

Key to Glyptopetalum species 

Leaf apex blunt or rounded. Twigs rounded ............................................ 1. G. palawanense 

Leaf apex pointed. Twigs sharply 4-angled .......................................... 2. G. quadrangulare 

1. Glyptopetalum palawanense Merr. 

(of Palawan) 

Philip. 1. Se. 26 (1925) 466; Ding Hou l.c. (1962) 257. Type: Forestry Bureau 29181, Philippines, 

Palawan (BO, K). 

Small tree to 5 m tall. Twigs grey-green, rounded. Leaves leathery, drying to greenish 

yellow, obovate or elliptic, 10-14 x 5-8 cm; base cuneate, margin distantly toothed towards 

the upper half, apex blunt or rounded; midrib raised above; lateral veins 6-8 pairs, almost 

invisible below, faintly raised above; intercostal veins invisible; petioles 8-12 mm long. 

Flowers unknown. Fruits globose to depressed globose, c. 15 x 8 mm, 1-4-celled. 

121


Distribution. Borneo and Philippines (Palawan). Uncommon in Borneo, known only from 

a sterile collection (Wong & Payne, s.n., 27 April 1993) from Balambangan Island, off 

Kudat Peninsula, Sabah. 

Ecology. Coastal limestone ridge. 

2. Glyptopetalum quadrangulare Prain ex King 

(Latin, quadrangularis = 4-angled; the twigs) 

Fig. 4. 

I.c. 345; Rid1ey I.c. (1922) 446; Ding Hou l.c. (1962) 257; Kochummen & Whitmore l.c. 162; 

Anderson l.c. 160; Ashton I.c. 95. Type: King's collector 7106, Perak (lectotype K; iso1ectotype BO). 

Shrub or small tree to 5 m tall, 5 cm diameter. Twigs strongly 4-winged. Leaves ellipticlanceolate, 

9-30 x 3-14 cm; base cuneate to rounded, margin toothed from base to apex, 

apex pointed; lateral veins 8-12 pairs, raised below, sunken above; intercostal veins 

reticulate; petioles 5-10 mm long. Flowers greenish yellow, in 12 cm long cymes; calyx 

almost divided to base, lobes reniform; petals suborbicular, fleshy, margin thinner, wavy; 

disc flat; stamens inserted near the base of ovary; ovary pyramidal, style and stigma 

obscure. Fruits c. 2 x 1-1.3 cm, depressed globose, sulcate, 3-4-celled. 

Distribution. Burma, Sumatra, Peninsular Malaysia, and Borneo. In Sarawak uncommon, 

known only by two collections (s. 16363 from Serian, and S. 12549 from Bekup); not yet 

recorded from Sabah. Also known from Kalimantan. 

Ecology. Lowland forests on limestone. 

5. KOKOONA Thwaites 

(kokoon = a Ceylonese plant name) 

mata ulat (Malay), bajan (!ban) 

in Hooker, J. Bot. Kew Misc. 5 (1853) 379; King I.c. 346; Merrilll.c. (1921) 354; Masamune l.c. 

417; van Steenis, Sarawak Mus. J. 8 (1958) 437; Ding Hou I.c. (1962) 258, I.c. (1969) 105; Balan 

Menon, MF 17 (1964) 18; Backer & BakhuizenJ I.c. 54; Smythies l.c. 38; Kochummen & Whitinore 

I.c. 163; Jansen et al., Blumea 21 (1973) 153; Cockburn I.c. 56; Anderson l.c. 160; Wong I.c. 109; 

Ashton I.c. 97; Whitmore, Tantra & Sutisna I.c. 43; Ng I.c. 43; Kochummen, Sandakania 5 (1994) 51. 

Small to very large trees; bole often with short buttresses. Bark grey to chocolate-brown, 

often with horizontal rings, smooth, cracking or fissured, with large lenticels; middle bark 

ochre or orange; inner bark pinkish or yellowish, fibrous. Sapwood white to yellow-brown 

with prominent pale bands. Twigs flattened at nodes, drying black. Stipules small. Leaves 

decussate, occasionally sub opposite or alternate, margin entire, wavy or toothed. Flowers 

bisexual, in axillary panicles or racemes, pedicels jointed; sepals 5; petals 5, free, overlapping 

and twisted (contorted); disc conspicuous, fleshy, cup-shaped, corrugated or 5- 

lobed; stamens 5, inserted on the inner edge of the disc, filaments abruptly narrowed 

122


towards the apex and transparent at the upper end, anthers usually with prominent 

connective; ovary superior or semi-inferior, 3-celled, style obscure, stigma capitate, ovules 

6-16 in each cell, in two rows down the central axis. Fruits capsules, 3-angled, 3-valved, 

splitting loculicidally. Seeds overlapping, flat, with conspicuous membranous wing at the 

apical end; endosperm absen; germination duriari-type; cotyledons fleshy; seedlings with 

opposite leaves, without or with fine hair-like, deciduous stipules. 

Distribution. 10 species; Southern India, Sri Lanka, Burma, Malesia. 8 species in Malesia 

of which 7 are present in Sabah and Sarawak. 

Ecology. Found scattered in a wide range of inland forests including swamps to 1500 m. 

Uses. The timber is a medium hardwood. It is easy to saw and cross-cut; suitable for heavy 

construction if treated and can be used as posts, beams, joints and railway sleepers. Also 

suitable for heavy-duty furniture, parquet-flooring, veneers, window and door-frames. 

Taxonomy. The genera Kokoona and Lophopetalum are difficult to distinguish on vegetative 

characters alone. However, some species of Kokoona have wavy to toothed leaf margins 

and they dry with greenish tinge, while in Lophopetalum the leaves have entire margins 

and they usually dry to a dark brown colour. The important diagnostic character is in the 

seed which is surrounded by a membranous wing in Lophopetalum, while in Kokoona the 

wing is apical. There is a sharp difference in pollen grains between the two genera; pollen 

grains of Kokoona are single while that of Lophopetalum are in tetrads or polyads. 

Key to Kokoona species 

(based on flowers and fruits) 

1. Fruit surface with abundant pustules. Twigs whitish ............................ 2. K leucoclada 

Not this combination of characters .............................................................. . 

2. Anthers with distinctly prolonged connective ...................... . 

Anthers without or with obscure connective .................................................................... 5 

3. Connective longer than anther-cells ............................................................ .4. K ochracea 

Connective shorter or as long as anther-cells ..................................................................... .4 

4. Flower buds ovoid to globose; apex of calyx-lobes rounded or truncate .... 3. K littoralis 

Flower buds broad-ellipsoid; apex of calyx-lobes acute ............................... 1. K coriacea 

5. Stamen filaments not broadened at base. Leaf margin strongly wavy and toothed 

toward apex ..................................................................................................... 6. K reflexa 

Stamen filaments with broad thickened base. Leaf margin recurved, entire or only 

slightly wavy .................................................................................................................. 6 

6. Inflorescences many-branched. Stigmas not papillose. Leaf margin rec\lrved; intercostal 

veins equally prominent as the lateral ones ......................... 5. K ovatolanceolata 

123


Inflorescences with few short branches only. Stigmas papillose. Leaf margin entire or 

slightly wavy; intercostal veins very faint... ................................................ 7. K sabahana 

Key to Kokoona species 


1. Leaf-margin entire to faintly 

Leaf-margin toothed .......................................................................................................... 5 

2. Twigs whitish. Leaves 6.5-10 cm wide; lateral veins 10-12 pairs ........ 2. K leucoclada 

Twigs blackish or dark-brown. Leaves to 6 cm wide; lateral veins 5-9 pairs .................. 3 

3. Leaf-base almost rounded; margin strongly recurved. Common trees of peat and 

freshwater swamp forests ............................................................. 5. K ovatolanceolata 

Leaf-base not rounded; margin not recurved. Except for K. sabahana, usually not 

found in swamp forests ......................................................................................................... .4 

4. Small pole-sized tree. Petiole to 7 mm long, channelled above ............... 7. K sabahana 

Medium-sized tree. Petiole to 2 cm long, wrinkled on drying .................. ,3. K littoralis 

5. Twigs strongly 4-angled. Leaves drying to dark brown above and purplish below ......... . 

. . .............. . 1. K coriacea 

Twigs not angled. Leaves drying pale yellow on both surfaces or pale greyish below .... 6 

6. Leaves when dry pale yellow on both surfaces; margin strongly wavy and distinctly 

toothed toward the apex ............................................................................. 6. K reflex a 

Leaves when dry pale greyish below; margin shallowly toothed from the base to apex 

.................... .4. K ochracca 

1. Kokoona coriacea King 

(Latin, coriaceus = leathery; the leaves) 

I.e 347; Ding Hou I.e. (1962) 261; Kochummen & Whitmore I.e. 164. Type: Kunstler 4226, Perak 

(ho1otype K; isotype BO). Synonym: Lophopetalum eoriaeea (King) Ridl. I.e. (1922) 450. 

Medium-sized tree to 25 m taU, 35 cm diameter. Bark grey-white. Twigs 4-angled (somewhat 

like that of Glyptopetalum quadrangulare). Leaves leathery, drying to dark brown 

above, and purplish below, oblong, 11-13 x 5-7.5 cm; base cuneate, margin toothed, 

slightly recurved, apex pointed; midrib sharply keeled below; lateral veins 5-7 pairs, faint; 

intercostal veins invisible below; petioles c. 1 cm long. Flowers 5-merous, greenish yellow, 

fragrant; buds broad ellipsoid; calyx-lobes triangular, apex acute; petals with pale margin; 

connective of anthers prolonged, almost as long as anthers; free part of the ovary ovoid, 

gradually narrowed into an obscure style, stigmas globose. Fruits unknown. 

Distribution. Peninsular Malaysia, Borneo. In Sarawak uncommon, known by a single 

collection (S. 40130) from Niah Cave National Park. Not yet recorded from Sabah. 

124


Ecology. Lowland forest below 100 m. 

This is the second collection of this species and a new record for Borneo. This species was 

until now known only from the type collected from Perak .in Peninsular Malaysia. 

2. Kokoona leucoclada Kochummen 

(Greek, leuco = white, cladus = branch) 

I.e. (1994) 51. Type: Amin & Francis SAN 129399, Sabah, Ranau (holotype SAN; isotypes K, L). 

Medium-sized to large tree to 35 m tall, 20 cm diameter. Bark grey, smooth; inner bark 

pale brown. Sapwood ochre. Twigs whitish, youngest flattened, grooved in the centre. 

Leaves leathery, drying to grey-brown, elliptic to oblong, 16.5-20 x 6.5-10 cm; base 

cuneate, margin faintly wavy, slightly recurved, apex pointed; midrib raised above, keeled 

below; lateral veins 10-12 pairs, visible on both surfaces, distinctly looping near margin; 

intercostal veins reticulate, very faint; petioles 1-1.5 cm long. Inflorescences axillary, to 

14 cm long, rachis rectangular, greyish. Flowers (immature) sessile, subtended by 5 bracteoles. 

Fruits c. 16 x 6 cm, with large pustules outside, on 20 cm long stalk. Seeds c. 12 x 2.5 cm. 

Distribution. Endemic to Sabah. Uncommon, and known only from two collections, SAN 

129399 (the type) from Ranau and SAN 61073 from Sandakan. 


The flowers are too young to give details of anther and ovary characters. The pustulate 

fruits and the white twigs distinguish this species from the others. 

3. Kokoona littoralis Laws. 

(Latin, littoralis = ofthe sea-shore) 

in Hooker! I.e. 617; Ding Hou I.e. (1962) 261; Burgess I.e. 73; Kochummen & Whitmore I.e. 164; 

Cockbum I.e. 57; Anderson I.e. 160; Ashton I.e. 100; Whitmore, Tantra & Sutisna I.e. 43. Type: 

Maingay 396/2, Malacca (holotype K; isotype BO). 

Medium-sized tree to 27 m tall, 30 cm diameter, or rarely pole-sized treelet to 6 m tall and 

2 cm diameter; bole slightly fluted with irregular hoops. Bark greyish or grey-brown with 

yellowish tinge, smooth to scaly. Twigs blackish on drying, slightly swollen at the nodes, 

youngest ones rectangular. Leaves thickly leathery, drying to dark green with purplish 

blotches above and pale yellowish below, elliptic-lanceolate to oblong or narrowly obovate, 

5-20 x 1.5-6.5 cm; base broadly cuneate to rounded, apex pointed; midrib faintly raised 

above; lateral veins 6-9 pairs, very faint below, faint to inconspicuous above, looping and 

joining near margin; intercostal veins visible or invisible; petioles 0.5-2 cm long, wrinkled 

on drying. Flowers with yellow petals, pedicels 1-2.5 mm long; flower buds ovoidsubglobose, 

calyx-lobes semi-orbicular, apex rounded or truncate; petals broadly ovate, 

elliptic or suborbicular; anthers with protruding connective to 1 mm long, slightly shorter 

125


4 cm 

'-----' 

lmm 

Fig. 5. Kokoona ovatolanceolata. A, leafy twig; B, longitudinal section through fruit; C, seed; D, 

inflorescence; E, longitudinal section through flower. (A from SAN A 1756. B & C from Anderson 

9291 (22 November 1957). D & E after FM 1, 6 (1962) 259, fig. 8.) 

126


than the anthers; ovary triangular, style obscure, stigma capitate. Fruits 13-18 x 3-5.5 cm, 

base tapered. Seeds 7.5-12.5 x 2.5 cm (including wing). 


1. Leaves c. 20 x 6.5 cm, margin pinkish ................. . 

var. longifolia Kochummen 

I.c. (1994) 53. Type: Wright & Ismawi S. 32289, Sarawak, Limbang (holotype SAR; isotype 

K). 

Small tree to 18 m tall, 20 cm diameter. Bark greyish, smooth to scaly. Leaves 

thick-leaiherly, elliptic to lanceolate, c. 20 x 6.5 cm; petiole c. 2 cm long, channelled 

above; base cuneate, margin pinkish, apex pointed; midrib raised above; lateral 

veins 7-9 pairs, faint on both surfaces; intercostal veins laxly reticulate, very faint. 

Uncommon, known by a single collection (s. 32289) from Limbang, Sarawak. 

Lowland forest. 

Leaves to 11 x 6 cm, margin not pinkish ........... . . ......... 2 

2. Leaf apex blunt or rounded; intercostal veins invisible .................................................. . 

var. bakoensis Kochummen 

l.c. (1994) 53. Type: Ding Hou 534, Sarawak, Kuching (holotype SAR; isotype L). 

Small pole-sized tree to 6 m tall, 2 cm diameter. Leaves coriacous, oblong with 

rounded or blunt apex, 5-11 x 1.5-4 cm; lateral and intercostal veins invisible. 

Locally frequent, known only from Bako National Park, Sarawak. Heath forest. 

Leaf apex pointed; intercostal veins visible ..... . 

var. littoraIis 

Synonyms: Lophopetalum dubium Laws. in Hooker! l.c. 616; L. maingayi Ridl. I.c. (1922) 

450; L. littoralis (Laws.) Ridl. l.c. (1922) 450; K. scortechinii King I.c. 347; K. lanceolata 

Ridl., Kew Bull. (1938) 237, Masamune I.c. 417; Solenospermum littorale (Laws.) Loes., 

Notizbl. Berl. Dahl. l3 (1936) 223. 

Sumatra, Peninsular Malaysia, Borneo. In Sarawak, known from Semengoh Arboretum, 

Balingian, Belaga and Lundu. One sample was collected from G. Matang 

at 1000 m. Uncommon in Sabah with only 4 collections. 

Ecology. Mixed dipterocarp forest on leached soils and heath forest to submontane forest to 

1000 m, including limestone. Flowering in August-October and fruiting in May-July. 

4. Kokoona ochracea (Elmer) Merr. 

(Latin, ochraceus = yellow or yellowish brown; the colour of the flowers) 

En. Philip. 2 (1923) 484, PEB (1929) 171; Masamune I.c. 417; van Steenis l.c. 438; Ding Hou I.c. 

(1962) 260; Burgess l.c. 74; Cockburn I.c. 57; Anderson I.e. 160; Ashton I.e. 101; Whitmore, Tantra 

& Sutisna I.e. 43. Basionym: Ardisia oehraeea Elmer, Leafl. Philip. Bot. 5(19l3) 1819. Type: 

Elmer 12881, Philippines, Palawan (holotype K; isotype BO). 

Medium-sized tree to 30 m tall, 50 cm diameter. Twigs black, terete. Leaves elliptic to ovate or 

lanceolate, 7-13.5 x 3.5-6 cm, drying pale greyish below; base broadly cuneate, margin 

127


shaIlowly toothed, ape)\ pointed; midrib raised or flattened above; lateral veins 5-8 pairs, 

slightly raised on both surfaces; petioles c. 1 cm long. Flowers almost sessile, in 12 cm 

long panicles; calyx-lobes semi-orbicular; petals fleshy, ovate or elliptic, margin thin and 

transparent; anthers subglobose, connective prolonged, longer than the anthers; ovary 

conical, style obscure, stigma cylindrical. Fruits unknown. 

Vernacular name. Sabah-perupok kuning (Dusun, Malay). 

Distribution. Peninsular Malaysia, Borneo, and Palawan Is. In Sabah, reported from 

Kinabatangan, Lahad Datu, Tenom and Tawau districts. In Sarawak, collections were made 

from northeastern parts. 

Ecology. Scattered in mixed dipterocarp forests at low altitude. 

5. Kokoona ovatolanceolata Ridl. Fig. 5. 

(Latin, ovatus = egg-shaped, lanceolatus = tapering toward both ends; the leaf shape) 

I.e. (1938) 236; Masamune I.e. 417; Ding Hou I.e. (1962) 261; Smythies I.e. 39; Cockburn I.e. 57; 

Anderson I.e. 160; Ashton I.e. 100; Whitmore, Tantra & Sutisna I.e. 43. Type: Beeeari PB 3471, 

Sarawak (holotype FI; isotypes BO, K). Synonym: K. seorteehinii (non King) Steenis I.e. 438. 

Tree to 36 m tall. Bark smooth to scaly, hoop-marked. Twigs dark brown to blackish. 

Leaves ovate to ovate-lanceolate or elliptic, 7-11.5 x 4-5 cm; base rounded to broadly 

cuneate, margin strongly re curved, apex pointed; midrib flattened above; lateral veins 5-6 

pairs, very faint on both surfaces and hardly distinguishable from the equally prominent 

reticulate intercostal veins; petioles 1-1. 5 cm long, wrinkled and yellowish on drying. 

Inflorescences many-branched panicles, to 14 cm long. Flowers yellowish, fragrant, buds 

globose, c. 2 mm across; calyx-lobes suborbicular or reniform; petals ovate; anthers ovoid, 

connective not prolonged, filaments with broad thickened base; ovary ovoid, style distinct, 

stigma obtuse, not papillose. Fruits narrow-oblong, 10-17 x 3-5 cm. Seeds 7-11 x 2-2.5 

cm (including wing). 

Vernacular name. Sarawak-bajan paya (Iban, Malay). 

Distribution. Endemic to Borneo. In Sabah, collected from Papar district and in Sarawak 

from Baram, Marudi and Sibu. Common in Brunei. 

Ecology. Known from heath, peat and freshwater swamp forests. 

6. Kokoona reflex a (Laws.) Ding Hou 

(Latin, rejlexus = recurved; the leaf tip) 

I.e. (1962) 262; Burgess I.e. 74; Kochummen & Whitmore I.e. 164; Cockbum I.e. 57; Anderson I.e. 

40; Ashton I.e. 101; Whitmore, Tantni & Sutisna I.e. 44. Basionym: Lophopetalum reflexum Laws. 

in Hooker f I.e. 616. Type: Maingay 383/2, Malacca (holotype K; isotype BO). Synonym: 

Hippoeratea maingayi Laws. in Hooker f I.e. 625. 

Emergent tree, to 50 m tall and 130 cm diameter; bole with stout buttresses. Bark dull 

grey-brown, smooth, hoop-marked, becoming scaly. Leaves elliptic to lanceolate, 5.5-ll x 

128


2.5-4 cm; base cuneate, margin strongly wavy and toothed towards the apex; apex pointed; 

midrib raised above; lateral veins 5 pairs, slightly raised on both surfaces; petioles slender, 

7-15 mm long, pale yellow on drying. Flowers pale yellowish green; calyx suborbicular; 

petals ovate, or broadly elliptic; anthers oblong, connective not protruding, filaments not 

broadened at base; ovary ovoid, narrowed to an obscure style, stigma rounded with flat top. 

Fruits c. 12 x 3 cm. Seeds 9-11 x 2.5 cm including the wing. 

Distribution. Sumatra, Peninsular Malaysia, and Borneo. Uncommon in Sabah, known 

from a single collection (SAN 64149) from Ranau. In Sarawak, locally frequent, recorded 

from Lundu, Semengoh, G. Gaharu, G. Matang, and Bako National Park. 

Ecology. Mixed dipterocarp forests. 

7. Kokoona sabahana Kochummen 

(of Sabah) 

I.c. (1994) 55. Type: Asik SAN 129928, Sabah, Nabawan (holotype SAN; isotypes A, BO, K, L, OX, 

SAR, SING). 

Small tree to 12 m tall, 10 cm diameter. Bark pale grey; inner bark brownish. Sapwood 

yellowish. Twigs dark brown to black, swollen at the nodes. Leaves·elliptic to oblong, 7-1 I 

x 4-6 cm, shiny, drying to yellowish brown; base broadly cuneate, margin entire or slightly 

wavy, apex pOinted or blunt; midrib very faintly visible above; lateral veins 6-7 pairs, 

looping near margin, visible below, faint above; intercostal veins reticulate, faintly visible 

below; petioles 5-7 mm long, channelled above. Inflorescences axillary or terminal panicles, to 

6 cm long, with a few branches. Flowers 5-merous; buds subglobose, c. 3 mm across; 

pedicels c. 2 mm long, articulated; sepals triangular; petals slightly obovate with pale 

margin; disc 5-lobed; stamens inserted within the disc, filaments broad and flat below, 

tapering towards the apex, anthers without protruding connective; ovary immersed in the 

disc, style distinct, stigma capitate, with papillae at the top. Fruits unknown. 

Distribution. Endemic to Borneo. Uncommon, known only from 3 collections from Sabah, 

SAN 129928 (the type) from Nabawan, SAN 61465 from Tawau, and SAN 43778 from 

Sandakan. 

Ecology. Lowland (including swamp) to hill forests to 600 m. 

Close to K. ovatolanceolata from which it differs by the few-branched, short panicles, 

larger flowers, papillose stigma, and shorter petiole. 

6. LOPHOPETALUM Wight ex Am. 

(Greek, lophos = crested, petalum = petal) 

perupok (Malay, Iban) 

Ann. Mag. Nat. Hist. 1, 3 (1839) 150; Merrilll.c. (1921) 354; Ridley I.c. (1922) 447; Masamune I.c. 

418; Browne l.c. 77; Ding Hou I.c. (1962) 262, l.c. (1969) 108; Backer & BakhuizenJ I.c. 54; Balan 

Menon I.c. 18; Smythies I.c. 38; Burgess I.c. 76; Kochummen & Whitmore I.c. 165; Jansen et al. l.c. 

129


153; Cockburn I.e. 58; Anderson I.e. 160; Wong I.e. 121; Ashton I.e. 102; Whitmore, Tantra & 

Sutisna I.e. 44; Ng I.e. 44. Synonym: Solenospermum Zoll., Nat. Tijd. Ned. Ind. 14 (1857) 168, 

Masamune I.e. 419. 

Small to very large trees to 45 m tall and 180 cm diameter; bole with or without buttresses, 

occasionally with pneumatophores. Bark similar to Kokoona. Twigs pale whitish, dark 

brown to blackish, often flattened at the nodes. Leaves opposite or sub opposite, margin 

entire. Inflorescences axillary, peduncles distinct or obscure, pedicels jointed. Flowers 

bisexual, 5-merous (except ovary); calyx-lobes spreading, inflexed or reflexed; petals free, 

imbricate, inner surface partly covered with appendages or without; disc conspicuous, 

usually fleshy and flat, surface smooth or denticulate, 5-angular, rounded or 5-10bed; 

stamens 5, inserted on the disc, anthers introrse, pollen grains in clusters of fours (tetrads); 

ovary usually semi-inferior, 3-celled, style short, stigma obscure, ovules 4-18 in each cell, 

. arranged in two series. Fruit a capsule, 3-angled. Seeds flat, surrounded by membranous 

wing; endosperm absent; germination as in Kokoona; cotyledons very thin, seedlings leaves 

minutely bistipulate, the first two leaves opposite, subsequent leaves alternate on the leader 

shoots but opposite on the branches. 

Distribution. About 18 species; India, Burma, Thailand, Indo-China, Malesia, and Australia. 

In Malesia 15 species, 10 of which are in Sabah and Sarawak. 

Ecology. Lowland (including swamp) to submontane forests to 1500 m. 

Uses. The timber is a light hardwood. It is suitable for interior finishing, panelling, partitioning, 

furniture, veneers, plywood, boxes, crates and mathematical instruments. 

Key to Lophopetalum species 


1. Leaves short-petioled or almost sessile ......................................................................... 2 

. Leaves with distinct petioles .............................................................................................. 3 

2. Twigs rounded, not winged. Lateral veins 5-7 pairs, sunken above .......... 2. L. glabrum 

Twigs winged, lateral veins 10-15 pairs, not sunken above ................. 8. L. sessilifolium 

3. Leaves drying to greenish brown above, lower surface with black gland-dots ................ . 

L. bcccarianum 

Not this combination of characters .............................................................................. .4 

4. Intercostal veins invisible below ...................................... . 

Intercostal veins faint to distinctly visible below .............................................................. 7 

5. Blade thinly leathery, apex rounded or notched; lateral veins 4-6 pairs ......................... . 

........... ........... ... ..... .......................................... .................................... 9. L. subovatum 

Blade thickly leathery, apex pointed, blunt or rounded; lateral veins 4-11 pairs ........... 6 

130


6. Petioles 2-2.5 cm long. Blade densely papillose below, margin curled inwards ............. . 

............................................................................ : ......................... 5. L. pachyphyllum 

Petiole 3-8 mm long. Blade not papillose below, margin not curled inwards . 

................................................................................................................ 7. L. rigidum 

7. Intercostal veins prominently raised below ....................................... l0. L. wightianum 

Intercostal veins faint below ......................................................................................... 8 

8. Leaves pale whitish below. Twigs pale whitish ......................................... 6. L. pallidum 

Leaves not pale whitish below. Twigs dark brown to blackish ...................................... 9 

9. Lateral veins 8-10 pairs. Petiole 1-1.5 cm long .................................... 3. L. javanicum 

Lateral veins 10-15 pairs. Petiole 1.5-3 cm 10ng ............................ .4. L. multinervium 

Key to Lophopetalum species 

(based on flowers and leaves) 

1. Flower buds almost flat, or wider than long; petals without appendages on the inner 

surfaces .............................................................................................................................. 2 

Flower buds conical or subglobose, longer than wide; petals with appendages on the 

inner surfaces .................................................................................................................... 5 

2. Twigs distinctly 4-angled to winged. Inflorescences 18-50 cm long; pedicels 7-15 mm 

long ................................................................................................... 8. L. scssilifolium 

Twigs usually rounded. Inflorescences to 15 cm long; pedicels to 9 mm 10ng .............. .3 

3. Disc dish-shaped, 5-lobed ........................................................................................... .4 

Disc suborbicular, flat or obscurely 5-angled ........................................... 7. L. rigidum 

4. Petioles 7-15 mm long. Leaves with black gland-dots below; lateral veins not sunken 

above ..................................................................................................... 1. L. beccarianum 

Petioles to 3 mm long. Leaves without black gland-dots below; lateral veins sunken 

above ....................................................................................................... 2. L. glabrum 

5. Disc 4.5-9.5 mm in diameter at anthesis ................................ .......................... 6 

Disc 1-3 mm in diameter at anthesis ........... 

6. Leaves densely papillose beneath, intercostal veins invisible ............ 5. L. pachyphyllum 

Leaves not papillose beneath, intercostal veins prominent below ....... l0. L. wightianum 

7. Disc with fleshy subulate processes around the base of filaments. Leaves pale whitish 

below ...................................................................................................... 6. L. pallidum 

Disc smooth or minutely papillose. Leaves not pale whitish below ................................ 8 

8. Leaf-apex rounded or notched; lateral veins 4-6 pairs .......................... 9. L. subovatum 

Leaf-apex pointed; lateral veins 8-15 pairs ......................................................................... 9 

131


9. Petals 1-1.5 cm long. Lateral veins 8-10 pairs ......................................... 3. L. javanicum 

Petals 1. 5-3 cm long. Lateral veins 10-15 pairs ............................. .4. L. multincrvium 

1. Lophopetalum beccarianum Pierre Fig.6A-D. 


Fl. For. Coch. 4 (1894) sub t. 307; Merrill I. c. (1921) 354; Masamune I. c. 418; Ding Hou I. c. (1962) 

266; Smythies I.c. 39; Kochummen & Whitmore I.c. 165; Cockburn I.c. 59; Anderson I.c. 160; 

Ashton I.c. 103; Whitmore, Tantra & Sutisna I.c. 44. Type: Eeccari PE 2475, Sarawak (ho1otype FI; 

isotypes BO, K). Synonyms: Lophopetalum scortechinii King I.c. 350; L. havilandii Rid!. I.c. (1931) 

37, Masamune I.c. 418. 

Small to medium-sized tree, rarely to 36 m tall and 60 cm diameter. Bark grey-brown, 

smooth, hoop-marked; inner bark pale brown. Sapwood pale yellow. Twigs dark brown to 

black, rounded. Leaves ovate, elliptic or oblong, drying to greenish brown above, with 

abundant black gland-dots below, 4-30 x 2.5-9 cm; base cuneate, apex pointed or blunt; 

midrib raised above; lateral veins 5-8 pairs, visible below, faint above, often pinkish when 

dried; intercostal veins very faint; petioles 7-15 mm long. Flowers yellowish, in axillary 

panicles to 15 cm long; buds flat; pedicels c. 5 mm long; calyx-lobes triangular or 

suborbicular; petals sub orbicular or broadly ovate, usually without appendages on inner 

side, sometimes with distinct veins; disc dish-shaped, 5-lobed; stamen filaments with 

cushion-like thickening at base; ovary pyramidal, style and stigma obscure. Fruits 5 2-11 x 

3-3.2 cm. Seeds 4.5 x 1.5-1.2 cm (including wing). 

Distribution. Peninsular Malaysia and Borneo. In Sabah and Sarawak frequent. 

Ecology. Mixed dipterocarp forest to submontane forest to 1800 m, mainly on ridges and 

hillsides on clay-rich soils. Flowering in February-November. 

2. Lophopetalum glabrum Ding HOll 

(Latin, glabrus = smooth; without pubescence) 

Fig.6E. 

I.c. (1962) 266; Anderson I.c. 160; Ashton I.c. 105; Whitmore, Tantra & Sutisna I.c. 44. Type: 

Rutten 83, E Borneo (ho1otype U; isotype BO). 

Small tree to 15 m tall, 15 cm diameter. Bark grey-white; inner bark yellowish. Sapwood 

white. Twigs grey-brown, rounded. Leaves elliptic or oblong, 8-16 x 2.5-5.5 cm, without 

black gland-dots below; base cuneate, apex pointed; midrib raised above; lateral veins 5-7 

pairs, raised below, usually sunken above, curving and joining near margin; intercostal 

veins reticulate, faint below, visible above; petioles to 3 mm long, cracking. Flowers 

yellowish, in branched panicles, to 9 cm long; buds flat; pediccls 5-7 mm long; calyx-lobes 

deltoid with pointed tips; petals suborbicular, without appendages inside; disc dish-shaped, 

5-10bed; anthers suborbicular; ovary pyramidal, style and stigma obscure. Fruits reddish 

brown when fresh, obovoid, 10-12.5 x 3 cm, apex rounded, drying brownish, surface 

pustulate. Seeds 4 in each locule, c. 7.5 x 1.5 cm. 

Distribution. Endemic to Borneo. Widespread in Sabah and Sarawak, also found in Brunei 

and Kalimantan. 

132


o.5mm[4 

o 

A 

],mm 

] 'mm 

Fig. 6. Lophopetalum beccarianum CA-D) and L. glabrum CE). A, flowering leafy twig; B, flower, 

top view; C, longitudinal section through flower, with petals removed; D, transverse section through 

ovary; E, fruit. CA from SAN 16022, B-D after FM 1,6 (1962) 263, fig. 10, E from SAN 91886.) 

133


Tree to 45 m tall, 80 cm diameter. Bark grey, scaly; inner bark orange-brown. Sapwood 

whitish. Twigs pale whitish. Leaves thinly leathery, glaucous below, elliptic to narrowly 

obovate, 7-11 x 2.5-6.5 cm; base cuneate, apex cuspidate with short tip; midrib raised 

above, pale whitish below; lateral veins 8-10 pairs with short intermediate veins, very faint 

on both surfaces; intercostal veins very faint; petioles 1.5-2 cm long, pale whitish. Flowers 

in panicles; buds short-conical or subglobose; calyx semi-orbicular or triangular; petals 

with fleshy lobed processes in the central part inside; disc 1-3 mm across at anthesis; 

stamens surrounded by fleshy subulate processes of the disc; ovary triangular, style cylindric. 

Fruits c. 15 cm long. Seeds c. 9 x 2.3 cm. 

Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah uncommon, known by only 

two collection (SAN 74348 and SAN 65428) from Sandakan and Tawau respectively. In 

Sarawak known from Bintulu, Bt. Mersing, Bt. Raya, and Kapit. 

Ecology. Mixed dipterocarp forests on clay soils. 

Uses. In Peninsular Malaysia the bark is used as an ingredient in making dart poison by the 

aborigines. 

7. Lophopetalum rigidum Rid!. 

(Latin, rigidus = stiff; the leaf texture) 

l.c. (1931) 38; Masamune I.c. 418; Ding Hou l.c. (1962) 267; Smythies I.c. 39; Burgess I.c. 77; 

Cockbum I.c. 58; Anderson /.t. 161; Ashton I.c. 106; Whitmore, Tantra &, Sutisna I.c. 44. Type: 

Haviland 2236, Sarawak, Kuching (BO, K). Synonym: L. subsessile Ridl.l.c. (1931) 37, Masamune 

I.c.418. 

Shrub or small tree, very rarely reaching 30 m tall and 40 cm diameter. Bark grey-brown, 

smooth, hoop-marked with vertical line of lenticels; inner bark purplish brown. Sapwood 

yellow-brown. Twigs dark brown, rounded. Leaves thickly leathery, drying to greenish 

brown or reddish brown, not papillose below, ovate-oblong, 4.5-21 x 2-4 cm; base broadly 

cuneate to rounded, margin not curled inwards, apex pointed or rounded; midrib raised 

above; lateral veins 4-11 pairs, visible below, faint to invisible above, looping near margin; 

intercostal veins invisible; petioles stout, 3 (-8) mm long. Flowers yellowish green, in 10 

cm long panicles, pedicels to 5 mm long; buds almost flat, suborbicular or obscurely 5- 

angled; calyx-lobes ovate or triangular, pointed, with small papilla-like processes outside; 

petals triangular or suborbicular, without appendages inside, with papilla-like processes 

outside; disc suborbicular, flat or obscurely 5-angled; stamens with small short-apiculate 

anthers; ovary pyramidal, style cylindrical. Fruits to 11 cm long, surface shortly tuberculate. 

Seeds c. 5.5 ~ 1.5 cm, including wing. 

Distribution. Endemic to Borneo. In Sabah, recorded from Lahad Datu (common), 

Keningau and Ranau. In Sarawak, locally frequent. 

Ecology. Freshwater swamps, peat swamps, kerangas swamps and also in montane forests 

to 2400 m. 

136


8. Lophopetalum sessilifolium Rid!. 

(Latin, sessile = stalkless,folium = leaves) 

I.e. (1931) 37; Masamune I.e. 418; Ding Hou I.e. (1962) 265; Anderson I.e. 161; Ashton I.e. 107; 

Whitmore, Tantra & Sutisna I.e. 44. Type: Haviland 1744, Sarawak, Kuching (BO, K). 

Small tree, rarely reaching 20 m tall, 20 cm diameter. Twigs dark brown, strongly 4-angled 

to winged. Leaves elliptic to oblong or lanceolate, 22-50 x 4.5-12 cm, drying to dark 

brown with brown gland-dots below and grey-brown above; base broadly cuneate to 

rounded, apex pointed, acumen to 2 cm long; midrib raised above; lateral veins 10-15 

pairs, raised on both surfaces, with short intermediate veins; intercostal veins reticulate 

(rarely scalariform-reticulate), distinct below, faint above; petioles very short to almost 

absent, rarely to 1 cm long. Flowers yellow, in 18-50 cm long panicles, pedicels 7-15 mm 

long; buds wider than long; calyx-lobes triangular, pointed; petals suborbicular or deltoid 

with blunt apex, without appendages on the inner side; anthers obtuse; ovary immersed in 

disc without distinct style or stigma. Fruits with tuberculate surface, 7.5-10.5 x 3 cm. 

Seeds 3.8-4.5 x 1-1.3 cm. 

Distribution. Endemic to Borneo. In Sarawak, locally abundant in the W Baram valley; 

also known from Bau, Kapit, Sarimnsam Wild Life Sanctuary, Semengoh Arboretum, Sg. 

Jeong, Sg. Kelawit and Sg. Temulan. Not yet reported from Sabah. 

Ecology. Lowland forests by rivers. Flowering in February, September and October and 

fruiting in August and September. 

9. Lophopetalum subovatum King 

(Latin, sub = somewhat, ova/us = egg-shaped; the leaf shape) 

I.e. 349; Ridley I.e. (1922) 448; Ding Hou I.e. (1962) 271; Burgess I.e. 77; Kochummen & Whitmore 

I.e. 169; Cockbum I.e. 59; Anderson I.e. 161; Ashton I.e. 109; Whitmore, Tantra & Sutisna I.e. 45. 

Type: Curtis 1501, Penang (BO, K). Synonym: Solenospermum apieulatum Rid!. I.e. (1938) 235. 

Medium to large-sized tree to 36 m tall, 75 cm diameter. Bark pale brown, cracking to 

scaly, hoop-marked; inner bark pinkish brown. Sapwood pale yellow. Leaves thinly 

leathery, drying to brownish, not pale whitish beneath, obovate to elliptic, 4.5-13 x 2.5-7 

cm; base tapered, apex rounded to notched; midrib raised above; lateral veins 4-6 pairs, 

curving and joining near margin, faintly visible below, faint to inconspicuous above; 

intercostal veins invisible; petioles 5-15 mm long. Flowers in axillary panicles; buds shortconical 

or subglobose; calyx-lobes spreading, triangular; petals inside with small appendage on 

the upper half; disc 1-3 mm across at anthesis, smooth or minutely papillose; anthers 

deltoid, obtuse; ovary triangular, narrowed towards apex, style cylindrical. Fruits to 7 cm 

long. Seeds c. 4 x 0.6 cm. 

Distribution. Sumatra, Peninsular Malaysia and Borneo. Frequent in Sabah and Sarawak. 

Ecology. In mixed dipterocarp forest~ on leached clay-rich soils, on periodically inundated 

alluvium, undulating lands and ridges to 900 m. 

137


10. Lophopetalum wightianum Am. 

(R. Wight, 19th Century botanist in India) 

I.c. (1839) 151; Ding Hou I.c. (1962) 267; Kochummen & Whitmore l.c. 169; Cockbum I.c. 59; 

Whitmore, Tantra & Sutisna I.c. 45. Type: Wight (Icon. t. 162). S India, Malabar (K). Synonyms: L. 

fimbriatum Wight in Ridley I.c. (1922) 448; L. winkleri Loes. I.c. 22l. 

Emergent tree to 40 m tall, 100 cm diameter. Bark dark grey, fissured; middle bark 

whitish; inner bark purplish brown. Leaves elliptic to ovate, 10-18 x 5-7 cm, not papillose 

beneath: base rounded to wedge-shaped, slightly peltate, apex pointed; midrib raised above; 

lateral veins 6-12 pairs, pale below; intercostal veins prominent below; petioles l.5-2.5 cm 

long. Flowers yellow, in panicles, pedicels 5-9 mm long; buds short-conical or subglobose; 

calyx distinctly 5-IQbed; petals broadly ovate to rounded with wavy margin, appendages 

lamellate to cristate and attached to the lower half of petals; disc 4.5-9.5 mm across at 

anthesis; stamens with oblong apiculate anthers; ovary triangular, narrowed into style. 

Fruits to 15 cm long. Seeds c. 6 x l.5 cm. 

Distribution. India, Indo-China, Sumatra, Peninsular Malaysia, and Borneo. In Sabah, 

uncommon and recorded from Sandakan and Tawau only. Not yet recorded from Sarawak. 


7. MICROTROPIS Wall. ex Meisn., nom. cons. 

(Greek, micro = small, tropis = keel; a small keel on the inner side of the petal) 

PI. Vasc. Gen. TabuI. Diagn. (1837) 68; Ridley I.c. (1922) 443; Merrilll.c. (1921) 354; Merr. & 

Freem., Proc. Am. Acad. Arts. Sc. 73 (1940) 276; Masamune I.c. 418; Ding Hou I.c. (1962) 272; 

Backer & BakhuizenJ I.c. 54; Kochummen & Whitmore I.c. 170; Cockbum I.e. 61; Anderson I.c. 

161; Ashton I.c. 109; Kochummen, Sandakania 5 (1994) 55. Synonyms: Microtropia Reichb., 

Nomencl. (1841) 190; Paraeelastrus Miq., Fl. Ind. Bat. 1,2 (1859) 590. 

Shrubs or small trees. Terminal node with one or two pairs of prominent, subpersistent. 

reduced leaves (incipient bracts). Leaves opposite, glabrous, entire. Inflorescences axillary or 

extra-axillary, dichotomous or paniculate cymes, sometimes in sessile clusters. Flowers 

bisexual, 5- or 4-merous; calyx-lobes almost free, imbricate, persistent, unequal in size, the 

outer 2 or 3 usually smallest, margin often thinner, transparent, entire, gnawed, irregularly 

split or fimbriate; petals free or united, imbricate; disc absent; stamens 4-5, filaments 

united at base into a ring or a short tube, the united part free from the petals or adnate to it, 

sometimes the stamens inserted in the mouth of corolla, anthers dorsifixed, usually introrse; 

ovary superior, cylindrical, conical or flask-shaped, 2-celled, ovules 2 in each cell, style 

very short or cylindrical, stigma obscure, rarely 2-4-lobed. Fruit a capsule, striated 

lengthwise, splitting along one side, apex with short beak, calyx persistent. Seeds usually 

1, erect on a knob-like thickened hilum enveloped by aril, with endosperm. 

Distribution. About 80 species, C America, SE to E Asia and Malesia. 20 species in Malesia, 

of which 14 are in Sabah and Sarawak. 

Ecology. Lowland to montane forests to 2700 m, more common in montane forests. 

138


Key to Microtropis species 

(based mainly on leaves) 

1. Leaves thickly leathery, margin distinctly curled inwards ............................................... 2 

Leaves thickly or thinly leathery, margin not distinctly curled inwards (except in M 

grandifolia and M sarawakensis) ............................................................................... .4 

2. Leaves greyish below; lateral veins and intercostal veins faintly sunken above . 

.............................................................................................................. 1. M. argentea 

Leaves not greyish below; lateral veins and intercostal veins not sunken above ............ 3 

3. Petioles 7-10 mm long. Intercostal veins invisible below .................... 2. M. borneensis 

Petioles absent or to 2 mm long. Intercostal veins visible below .................. 9. M. rigida 

4. Leaves drying to greenish yellow. Incipient bracts greenish .................. ....... 5 

Leaves drying brownish. Incipient bracts brown. 

5. Incipient bracts needle-like, c. 5 x 1 mm ........................................... 10. M. sabahensis 

Incipient bracts elliptic, lanceolate or oblong, at least 4 mm broad .................................. 6 

6. Intercostal veins equally prominent as the faint lateral veins. Incipient bracts oblong or 

lanceolate ....................................................................................... 6. M. kinabaluensis 

Intercostal veins invisible or very faint. Incipient bracts elliptic or lanceolate ................ . 

.................................................................................................... . 11. M. sarawakensis 

7. Leaves to 4 cm long; base usually rounded to subcordate, apex blunt or rounded .......... 8 

Leaves longer than 4 cm; base cuneate, apex usually pointed ......................................... 9 

8. Ovary ovoid-oblong ................. . .7. M. ovata 

Ovary conical... ........... 

................. 14. M. wallichiana 

9. Incipient bracts foliaceous, 1.7-3 cm long ............................................. 4. M. grandifolia 

Incipient bracts stiff, not foliaceous, to 12 mm long .................................................. : .... 10 

10. Leaves uniformly papillose below with scattered pustules .............. 5. M. keningauensis 

Leaves not papillose, without pustules ........................................................................ 11 

11. Inflorescences in sessile clusters ..................... ... 3. M. fascicularis 

Inflorescences distinctly stalked. 12 

12. Petioles 0.7-1 cm long ......... M. sumatrana 

Petioles 1-2 cm long. ................ 13 

13. Ovary cylindrical, slightly constricted in the middle ................................. 13. M. valida 

Ovary conical, without constriction in the middle ................................ 8. M. platyphylla 

139


Key to Microtropis species 

(based mainly on flowers) 

1. Inflorescences or infructescences in sessile clusters or condensed cymes; peduncle if 

present to 1 cm long .......................................................................................................... 2 

Inflorescences and infructescences with peduncle more than 1 cm long ....................... 6 

2. Lower surface ofleafuniformly papillose and with scattered pustules .... S. M. keningauensis 

Lower surface of leaf not papillose, without pustules ....................................................... 3 

3. Ovary cylindrical or ovoid-oblong .................................... .............. 4 

Ovary conical or gradually tapered to apex ....... .................... 5 

4. Ovary cylindrical with slight constriction in the middle. Petioles 1-2 cm long .............. . 

................. 13. M. valida (in part) 

Ovary ovoid-oblong. Petioles absent or very short, to 2 mm long ................. 7. M. ovata 

5. Stigmas 4-lobed; ovary gradually tapered towards apex ..................... 3. M. fascicularis 

Stigmas not lobed; ovary conicaL .................................................. 14. M. wallichiana 

6. Leaves thickly leathery; margin distinctly curled inwards ............................................ 7 

Leaves thinly to thickly leathery; margin not distinctly curled inwards (faintly so in M 

grandifolia and M sarawakensis) .............................................................................. lO 

7. Petioles very short, to only 2 mm long or almost absent... ..... . ....... 9. M. rigida 

Petioles distinct, at least 7 mm long ................................. ..................... 8 

8. Lower surface ofleaf greyish; lateral veins and intercostal veins sunken above ............. . 

..................................... . 1. M. argentea 

Lower surface of leaf greenish brown; lateral veins and intercostal veins invisible 

above ........................................................................................................................... 9 

9. Incipient bracts 1.7-3 x 1.2-2 cm; petioles 2.3-6 cm long ... .4. M. grandifolia (in part) 

Incipient bracts to 5 mm long. Petioles 7-10 mm long ........................ 2. M. borneensis 

10. Petals distinctly united, at least up to lower half... ............................. l0. M. sabahensls 

Petals free or only slightly united at the very base .................................................. ll 

11. Ovary cylindricaL ..................................................................... 13. M. valida (in part) 

Ovary conical, flask-shaped or gradually narrowed towards apex ............. : ................. 12 

12. Ovary flask-shaped .................................................................................................... 13 

Ovary conical or gradually narrowed towards apex .................................................... 14 

13. Incipient bracts ovate, 1.7-3 x 1.2-2 cm. Leaves 20-30 x 8.5-14.5 cm; petiole 2.3-6 

cm long ............................................................................... .4. M. grandifolia (in part) 

140

CELASTRACEAE (KO CHUM MEN) 

Incipient bracts lanceolate or oblong, to 1.5 cm long. Leaves 11-24.5 x 3.5-9 cm; 

petiole 1-2 cm long ........................................................................ 6. M. kinabaluensis 

14. Stigmas 4-lobed; ovary gradually narrowed towards apex ................... 12. M. sumatrana 

Stigmas or apex of ovary not lobed; ovary conicaL ..................................................... 15 

15. Leaves thinly leathery. Incipient bracts green, lanceolate or elliptic ..... 11. M. sarawakensis 

Leaves thickly leathery. Incipient bracts brown, lanceolate ........................ 8. M. platyphylla 

1. Microtropis argentea Kochummen 

(Latin, argenteus = silvery; the lower leaf surface) 

I.c. (1994) 55. Type: Lee S. 39980, Sarawak, Belaga (holotype SAR; isotypes E, K, KEP, L, SAN). 

Small tree to 5 m tall. Incipient bracts lanceolate, stiff, brown, c. 5 mm long. Twigs swollen at 

nodes, grey, rounded. Leaves thickly leathery, drying to pale green above, greyish below, 

elliptic, lO.5-l2 x 5.5 cm; base cuneate, margin prominently re curved, with thick rim, 

apex shortly pointed, incurved; midrib prominently raised above; lateral veins 6-7 pairs, 

invisible below, faintly sunken above; intercostal veins invisible below, reticulate and 

faintly sunken above; petioles 0.7-1 cm long, wrinkled on drying. Inflorescences in cymes 

to 1.5 cm long; peduncles c. 1 cm long; buds globose, c. 2 mm across. Flowers white, 5- 

merous; sepals rounded with gnawed margins, wrinkled outside; petals united to more than 

half the length, lobes oblong; stamens at mouth of corolla tube, filaments very short, 

anthers globose; ovary conical, apical part ridged with no distinct style or stigma. Fruit 

unknown. 

Distribution. Endemic to Borneo. Uncommon, known only from the type specimen from 

Belaga, Sarawak. 

Ecology. Kerangas forest on plateau at 700 m. 

2. Microtropis borneensis Merr. & Freem. 


I.c. 296; Masamune l.c. 418; Kochummen I.c. (1994) 63. Type: Clemens 31742, British North 

Borneo, Mt. Kinabalu (holotype A; isotypes B, BO, GE, N, UC). 

Shrub or small tree. Twigs grey, youngest ones reddish brown and slightly angled. 

InCipient bracts triangular, stiff, c. 5 mm long. Leaves thickly leathery, drying to greenish 

brown on both surfaces, elliptic, 5.5-9 x 2.5-4.5 cm; base cuneate, margin with thick rim, 

wavy and strongly recurved, apex pointed; midrib prominently raised above; lateral veins 

5-7 pairs, very faintly visible below only; intercostal veins invisible; petioles 7-10 mm 

long, wrinkled on drying. Inflorescences and flowers unknown. Infructescences with a 

peduncle c. 2 cm long. Fruits when fresh oblong, c. 2 x 1.5 cm, yellow, drying black. 

141


Distribution. Endemic to Borneo. Locally abundant on Mt. Kinabalu in Sabah. In 

Sarawak, known from a single collection (s. 40412), from Lambir National Park. 

Ecology. Kerangas and montane forests to 2400 m. 

Taxonomy. Ding Hou I.c. 279 considered this a synonym of M platyphylla Merr. After 

studying the types and other recently collected specimens of both species, I have come to 

the conclusion that M borneensis should be reinstated as a distinct species. 

3. Microtropis fascicularis Kochummen 

(Latin,fascicularis = in a small bundle; the flower dusters) 

Fig. 7. 

I.e. (1994) 57. Type: Lee S. 44277, Sarawak, Lingga (holotype KEP; isotypes A, K, L, SAN, SAR). 

Treelet to 2 m tall. Incipient bracts stiff, brown, lanceolate, c.7 x 1 mm, with sharp tip. 

Twigs dark brown, rounded. Leaves leathery, surface not papillose nor pustulate, elliptic 

or oblong, 1O.S-20.S x 4-8.S cm; base cuneate, margin slightly recurved, apex pointed; 

midrib raised above; lateral veins 10-12 pairs, looping near margin, very faint on both 

surfaces; intercostal veins reticulate, very close, visible on both surfaces; petioles 0.S-2.2 

cm long, wrinkled on drying. Flowers in sessile fascicles, 4-merous; sepals rotund, margin 

gnawed, c. 2 x 1.5 mm; petals free, oblong to obovate, c. 1.S x 1.2 mm; stamens free, 

anthers oblong, c. 1 mm long; ovary gradually tapered towards apex, striate, stigma 4- 

lobed. Fruits green when fresh, drying black, oblong or ellipsoid, IS-18 x 4-6 mm, with 

pointed tip. 

Distribution. Endemic to Borneo. Uncommon, known by S collections from Sarawak. 

Ecology. Mixed dipterocarp forest and kerangas forest, to SOO m, usually by streams. 

Flowering in December and fruiting in March-May. 

4. Microtropis grandifolia Kochummen 

(Latin, grandis = large,folius = leaf) 

I.e 1994) 59. Type: Awa & llias S. 47380, Sarawak, Lundu (holotype KEP; isotypes K, SAR). 

Smail tree to S m tall. Twigs brownish, youngest ones flattened. Incipient bracts prominent, 

foliaceous, ovate to elliptic, 1.7-3 x 1.2-2 cm. Leaves thinly to thickly leathery, elliptic, 

drying greenish brown on both surfaces, 20-30 x 8.S-14.S cm; base broadly cuneate, 

margin faintly wavy, faintly curled inwards, apex pointed; midrib flattened to raised above; 

lateral veins ,10-12 pairs, looping near margin, faintly raised on both surfaces with short 

intermediate veins; intercostal veins reticulate, faintly visible above; petioles 2.3-6 cm 

long, wrinkled on drying. Inflorescences paniculate cymes, c. 6-7 cm long; peduncles 2-3 

cm long; bracteoles triangular, transparent. Flowers 4-merous; outer two sepals larger than 

inner 2, rotund; petals slightly united at base; anthers sessile, jointed at base on a staminal 

ring, connective slightly prolonged; ovary flask-shaped, style ridged, stigma 2-4-lobed. 

Fruits unknown. 

142


4cm 

4cm 

: 

:.\. 

c··r 

~ 

;~\;! 

Fig. 7. Microtropisfascicularis. A, flowering leafy twig; B, fascicled inflorescence; C, fruits. CA & B 

fromS. 44277, C fromS. 36677.) 

143



Leaves thinly leathery, margin slightly incurved. Petioles c. 3 cm. Stigmas 4-lobed ............. . 

var. grandifolia 

Endemic to Sarawak. Uncommon, known only by two collections (s. 47380 from Bukit 

Panjo, Lundu, and S. 34033 from Lubok Antu). Lowland and hill forests to 600 m. 

Flowering in May. 

Leaves thickly leathery, margin strongly incurved. Petioles c. 6 cm. Stigma 2-lobed ............ . 

var. longipetiolatus Kochummen 

I.c. (1994) 59. Type: Chai S. 34033, Sarawak, Lanjak Entimau (holotype SAR; 

isotypes K, KEP, L, MO, SAN). Uncommon, known only from the type collection. 

5. Microtropis keningauensis Kochummen 

(ofKeningau, in Sabah) 

l.c. (1994) 6l. Type: Fidilis SAN 118405, Sabah, Keningau (holotype SAN). 

Shrub or small tree. Twigs grey-brown. incipient bracts brown, lanceolate, c. 10 x 2 mm, 

with distinct median ridge. Leaves leathery, drying brownish, uniformly papillose below 

with scattered pustules, elliptic, 11.5-23.5 x 4.5-8.5 cm; base cuneate, margin not curled 

inwards, apex pointed; midrib flattened above; lateral veins 9-12 pairs, looping near 

margin, distinct below, faintly visible above; intercostal veins reticulate, distinct below, 

almost invisible above; petioles 8-15 mm long, drying black. Inflorescences and flowers 

unknown. Infructescences almost sessile to l-cm-stalked. Fruits yellowish green when 

fresh, black on drying, oblong, 2-2 ..-3 x 0.7-0.8 cm, apex with distinct point. 

Distribution. Endemic to Borneo. Uncommon, known by two collections only (SAN 118364 

and SAN 118405), from Lanas, Keningau in Sabah. 

Ecology. Lowland forest by stream and ridge top. Fruiting in October. 

6. Microtropis kinabaluensis Merr. & Freem. 

(of Mt. Kinabalu, Sabah) 

I.c. 304; Masamune I.c. 418; Ding Hou I.c. (1962) 277. Type: Clemens 29518, British North Borneo, 

Mt. Kinabalu (holotype A; isotypes B, BO, GE, K, L, N, UC). Synonyms: M. kinabaluensis var. 

acuminata Merr. & Freem. l.c. 305, Masamune l.c. 418; M. sterrophylla Merr. & Freem l.c. 305, 

Masamune l.c. 419. 

Shrub or small tree to 5 m tall. incipient bracts pale green, to 1.5 cm long, 0.5 cm broad, 

lanceolate or oblong. Young twigs slightly angled. Leaves greenish yellow on drying, 

leathery, elliptic-oblong, 11-24.5 x 3.5-9.5 cm; base cuneate to tapered, margin slightly 

recurved, apex pointed; midrib raised above; lateral veins 6-12 pairs, looping near margin, 

very faint on both surfaces; intercostal veins reticulate, equally prominent as lateral veins; 

144


petioles 1-2 cm long, drying wrinkled and yellowish. Inflorescences cymose panicles, to 

5.5 cm l()ng, peduncles 1.5-2.5 cm long. Flowers white; calyx-lobes with 3-5 longitudinal 

veins; petals free, ovate or broadly elliptic; stamen filaments united at base; ovary flaskshaped. 

Fruits red when ripe, oblong or ovoid to ellipsoid, 1.5-2 x 0.7-1 cm, faintly 

furrowed, crowned by the persistent style. 

Distribu tion. Endemic to Borneo. Widely distributed in Sabah and Sarawak. 


7. Microtropis ovata Merr. & Freem. 

(Latin, o'Vatus = egg-shaped; the leaf shape) 

l.c. 297; Masamune I.c. 419; Ding Hou l.c. (1962) 279. Type: Clemens 40046, British North Borneo, 

Mt. Kinabalu (BM, BO). 

Shrub to 3 m tall. Twigs 4-angled. Leaves thickly leathery, under surface not papillose nor 

pustulate, ovate to ovate-oblong, 4.5-10 x 2-7 cm; base rounded to cordate, margin not 

curled inwards, apex acute; lateral veins 5-9 pairs; petioles very short to 2 mm long or 

leaves sessile. Flowers in condensed cymes, peduncle very short; calyx-lobes suborbicular; 

petals free, ovate or ovate-oblong; stamen filaments united at the lower half, ovary ovoidoblong, 

not constricted at the middle. Fruits unknown. 

Distribution. Endemic to Sabah. Uncommon, known from Mt. Kinabalu only. 

Ecology. Submontane forests at 1200-1500 m. 

8. Microtropis platyphylla Merr. 

(Greek, platus = broad, phyllon = leaf; broad-leaved) 

Philip. J. Se. 10 (1915) Bot. 319; Ding Hou l.c. (1962) 279. Type: Loher 5779, Philippines (BO, L). 

Shrub or small tree to 10 m tall, 10 cm diameter. Incipient bracts brown, 4-12 mm long, 

lanceolate. Leaves thickly leathery, not papillose nor pustulate beneath, elliptic to lanceolate or 

ovate-lanceolate, 4.5-24 x 4.5-11 cm; base cuneate, margin not curled inwards, apex blunt 

to pointed; lateral veins 6-12 pairs; petioles 1-2 cm long. Inflorescences paniculate, 

peduncles 1-1.5 cm long. Flowers (4-)5-merous; calyx-lobes suborbicular or subreniform 

with gnawed margins; petals free, elliptic or broadly ovate, obtuse; stamen filaments united 

at base; ovary conical, not constricted at the middle, apex obtuse or notched. Fruits ovoid 

or ellipsoid, 1-2.5 x 1 cm. 

Distribution. Borneo and the Philippines. Common in Sabah, but uncommon in Sarawak. 

Ecology. Hill and montane forests to 2700 m. 

145


9. Microtropis rigid a Ridl. 

(Latin, rigidus = stiff; the thick leaves) 

I.e. 36; Merr & Freem. I.e. 297; Masamune I.e. 419; Ding Hou I.e. (1962) 278. Type: Beeeari PB 

1702, Sarawak (holotype FI; isotypes BO, K). 

Small tree to 20 m tall, 15 cm diameter. Incipient bracts lanceolate, c. 0.5 cm long, brown, 

stiff. Twigs swollen at nodes. Leaves thickly leathery, elliptic-oblong, 7-12 x 2.5-6 cm, 

pustulate below; base obtuse, rarely cuneate, margin curled inwards, apex pointed; midrib 

raised above; lateral veins 5-7 pairs, very faint; intercostal veins reticulate, visible below, 

invisible above; petioles very short to 2 mm long to almost absent. Inflorescences to 2.5 

cm long, peduncles 1-1.5 cm long. Flowers white; calyx-lobes suborbicular, sometimes 

irregularly split; petals free, oblong, obtuse; stamen filaments subulate, lower half united; 

ovary globose, narrowed towards apex, style very short, stigmas obscurely 4-6-lobed. 

Fruits oblong, 12-15 x 7-9 mm, apex rounded with short persistent style. 

Distribution. Endemic to Sarawak; uncommon and known only from Bako National Park, 

G. Santubong and Mulu National Parks. 

Ecology. Lowland to submontane forests to 1000 m, including kerangas. 

10. Microtropis sabahensis Kochummen 

(ofSabah) 

I.e. (1994) 61. Type: Banang SAN 51915, Sabah (holotype KEP; isotype SAN). 

Treelet to 5 m tall. Youngest twigs reddish brown. Incipient bracts greenish, needle-like, c. 

5 x 1 mm. Leaves thinly leathery, elliptic, 13.5-20.5 x 3.5-7 cm; base cuneate, margin 

slightly wavy, apex pointed; midrib raised above; lateral veins 8-10 pairs, looping and 

joining near margin, faint on both surfaces; intercostal veins reticulate, very faint; petioles 

1-1.5 cm long, drying greenish yellow. Inflorescences axillary, 3-4.5 cm long, cymose 

panicles, peduncles 1-1.5 cm long; bracteoles triangular, transparent. Flowers 4-merous; 

sepals ovate, transparent, wrinkled outside; petals united halfway, lobes oblong with 

transparent margins; stamens seated on the mouth of corolla-tube, filaments flat; ovary 

gradually tapered towards apex, slightly ridged towards apex, style indistinct, stigma distinct, 

4-lobed. Fruits unknown. 

Distribution. Endemic to Sabah. Uncommon, known only from the type collection from 

Kinabatangan. 

Ecology. Lowland and seasonal swamp forests. Flowering in June. 

Very similar to M. kinabaluensis but differing in the united petals, shape of ovary and the 

prominent stigma. 

146


11. Microtropis sarawakensis Kochummen 


I.e. (1994) 63. Type: Ilias S. 36501, Sarawak (holotype KEP; isotypes K, L, MO, SAN, SAR). 

Small tree to 5 m tall, 10 cm diameter. Incipient bracts green, elliptic or lanceolate, c. 10 x 

4 mm. Twigs reddish brown? rounded, youngest ones angular. Leaves thinly leathery, 

drying to greenish yellow, elliptic or oblong, 1O-l7 x 7.5 cm; base cuneate, margin wavy 

and slightly recurved, apex pointed; midrib raised above; lateral veins 7-9 pairs, very faint 

on both surfaces; intercostal veins invisible to very faint; petioles 1-1.5 cm long, 

channelled above, wrinkled and yellowish on drying. Inflorescences cymose panicles, 2-5 

cm long, peduncles 1-1.5 cm long. Flowers 4- or 5-merous; calyx-lobes suborbicular, 

transparent with few fine veins in the middle; petals free, imbricate, oblong, transparent; 

stamens with short filaments which are joined at the bottom by a staminal ring, connective 

slightly prolonged; ovary short conical, without distinct style, with wavy surfaced, and 

unlobed stigma. Fruits (immature) green when fresh, ovoid, 7-10 x 4-5 mm, apex with 

short style and stigma, calyx persistent. 

Distribution. Endemic to Sarawak. Uncommon, known only by the type collection. 

Ecology. Lowland forests by streams. Flowering in April and June, and fruiting in October. 

The leaves are somewhat similar to those of M sabahensis, but the shape and size of 

incipient bracts of M sarawakensis distinguishes it from M sabahensis. Furthermore, in 

M sabahensis the stigma is distinctly lobed, and the corolla is united almost half way. In 

contrast, in M sarawakensis the stigma is unlobed and the petals are free. 

12. Microtropis sumatrana Merr. 

(of Sumatra) 

Pap. Mich. Aead. Se. 19 (1934) 164; MeIT. & Freem. I.e. 304; Ding Hou I.e. (1962) 279. Type: 

Rahmat 214, Sumatra (A, BO). 

Small tree to 20 m tall and 20 cm diameter. Bark grey, smooth; inner bark pale grey. 

Sapwood pale white. Incipient bracts lanceolate, to 12 mm long. Twigs greyish. Leaves 

thinly leathery, pale below, dark brown on drying, not papillose nor pustulate beneath, 

elliptic, 6.5-11 x 3.5-5.5 cm; base cuneate, apex pointed; midrib flattened above; lateral 

veins 5-7 pairs, faint to distinct below, curving and joining near margin; intercostal veins 

laxly reticulate, faint; petioles 0.7-1 cm long. Inflorescences dichotomously branched, 

peduncles 1-3 cm long. Flowers with calyx-lobes irregularly split; petals free, oblong, 

obtuse; stamens united at the lower half; ovary gradually narrowed towards apex, stigma 4- 

lobed. Fruits oblong, 1.5-1.7 x 0.7-1.0 cm, with pointed tip, calyx persistent at base. 

Distribution. Sumatra and Borneo. Uncommon, in Sabah known only from Sipitang, 

Tawau, and Mostyn. 


147


o 

Fig. 8. Perrottetia alpestris subsp. philippinensis. A, flowering leafy twig; B, flower; C, infructescence; D, 

fruit. CA from S. 34060, B after FM 1, 6 (1962) 289, fig. 21, C & D from S. 43345.) 

148


13. Microtropis valid a Ridl. 

(Latin, validus = robust-growing) 

J. Str. Br. R. As. Soc. 75 (1917) 19, I.e. (1922) 445; MeIT & Freem. I.e. 304; Ding Hou I.e. (1962) 

276; Kochummen & Whitmore l.c. 170. Type: Curtis 1331, Perak (holotype K; isotype BO). 

Synonyms: M. bieolor MeIT. & Freem. I.e. 298; M. paueiflora Boerl. ex MeIT. & Freem. I.e. 303. 

Shrub or small tree to 5 m tall. incipient bracts brown, stiff, 3-12 mm long. Leaves 

leathery, under surface not papillose nor pustulate, ovate to lanceolate or elliptic, 11-27 x 

4.5-12 cm; base rounded to cuneate, margin not curled inwards, apex acute or acuminate; 

midrib raised above; lateral veins 6-14 pairs; intercostal veins reticulate, distinct on both 

surfaces; petioles 1-2 cm long. Inflorescences usually paniculate, rarely a simple cyme or 

fascicle, peduncles J-4 cm long. Flowers 4(-5)-merous; calyx-lobes suborbicular, slightly 

wrinkled outside, margin transparent; petals free, elliptic or oblong, obtuse; stamen filaments 

united at base; ovary cylindric, slightly constricted at the middle, truncate or discoid at the 

top. Fruits ellipsoid, c. 2 x 1 cm, apex pointed and furfuraceous with persistent style. 

Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak uncommon, 

known by only two collections (SAN 75680 and SAN 44667) in Sabah, and three (including 

the type of M bicolor by Beccari) collections in Sarawak. 


14. Microtropis wallichiana Wight ex Thwaites 

(N. Wallich, 1786-1854, Superintendent, Calcutta Botanic Gardens) 

En. PI. Zeyl. (1858) 71; Ding Hou I.e. (1962) 279. Type: Wight 528, Ceylon (BO, K). Synonym: M. 

suborbieulata MeIT. & Freem. I.c. 282, Masamune I.e. 419. 

Shrub or small tree. incipient bracts brown, 3-8 mm long. Twigs rounded to slightly 

angled. Leaves thickly leathery, elliptic, lanceolate or rounded, 3-18 x 1-6 cm, lower 

surface not papillose nor pustulate; base cuneate to rounded, margin not curled inwards, 

apex pointed, blunt or rounded and notched; midrib raised above; lateral veins 4-9 pairs, 

invisible above; intercostal veins reticulate, faint to invisible; petioles to 1.5 cm long, rarely 

very short. Inflorescences condensed cymes, less than 1 cm long, peduncles very short or 

absent. Flowers yellowish; calyx-lobes subreniform; petals free, fleshy; stamen filaments 

united at the lower part; ovary conical, longitudinally striate, stigma not lobed. Fruits 

oblong-ellipsoid or oblong-obovoid, c. 1 x 0.5 cm, furfuraceous, crowned by persistent 

style. 

Distribution. Sri Lanka, Sumatra and Borneo. In Sabah, known from Mt. Kinabalu only; 

not recorded from Sarawak. 

Ecology. Submontane to montane forests at 1250-2500 m. 

The types of M. wallichiana and M suborbiculata have some differences which require to 

be substantiated by examining more material. There is no conclusive evidence for their 

distinction, although it is doubtful if they are the same species. 

149


8. PERROTTETIA Kunth 

(G.S. Perrotet, l793-1870, French botanist) 

Nov. Gen. Sp. 7 (1824) 73; Merrilll.e. (1921) 354; Ridley I.e. (1922) 454; Masamune I.e. 419; Ding 

Hou I.e. (1962) 288; Backer & BakhuizenJ I.e. 55; Kochummen & Whitmore I.e. 171; Anderson I.e. 

161; Ashton I.e. 112. Synonym: Caryospermum Blume I.e. (1850) 175. 

Shrubs or small trees. Leaves alternate. Inflorescences cymose, axillary. Flowers bisexual, 

rarely unisexual; calyx-lobes 5 or 4; petals similar to calyx-lobes in size and shape; 

stamens 5 or 4, inserted on the margin of the disc; ovary semi-inferior, mostly 2-celled, 

style 1; ovules two in each cell. Fruit a berry, globose, 2-4-seeded. Seeds covered with thin 

aril; endosperm thin; embryo small. 

Distribution. About 15 species; China, Formosa, Malesia and America. Only one species 


Ecology. Primary and secondary hill and montane forests to 2600 m. 

Perrottetia alpestris (Blume) Loes. Fig. 8. 

(Latin, alpestris = growing above the limit offorest growth; its habitat) 

in Engler & Prantll.e. 220; Merrilll.e. (1921) 354; Ridley I.e. (1922) 454; Masamune I.e. 419; Ding 

Hou I.e. (1962) 288; Kochummen & Whitmore I.e. 171; Ashton I.e. 161. Basionym: Celastrus 

alpestris Blume, Bijdr. (1826) 1145. Type: Blume, s.n., Java (BO, L). 

subsp. philippinensis (Vidal) Ding Hou 

I.e. (1962) 291. Basionym: Caryospermum philippinensis Vidal, Rev. PI. Vasc. Philip. (1886) 89. 

Type: Beeeari PB 7770, Sarawak (FI). 

Shrub or small tree to 12 m tall. Young twigs and petioles purplish when fresh, blackish on 

drying. Leaves thinly leathery, elliptic or oblong, 10-20 x 4-8 cm; base cuneate or rounded, 

sometimes subcuneate, margin prominently toothed, apex pointed; midrib flattened above; 

lateral veins 8-12 pairs, raised below, sunken or flattened above; intercostal veins 

reticulate, distinct below, faint above; petioles 7-15 mm long, not swollen. Flowers white 

or light greenish, 4-merous; calyx-lobes sparsely puberulous on both surfaces; petals 

puberulous on both surfaces; disc cup-shaped; stamen filaments longer than the disc; ovary 

conical, style short, stigma distinctly 2-lobed. Fruits globose or subglobose, 2.5-3.5 mm 

diameter, red. Seeds 3-4. 

Distribution. Borneo, Philippines and Celebes. In Sabah, collected from Kinabalu National 

Park, Kalabakan and Lamag. In Sarawak, reported from Dulit Range, Mulu National Park, 

Bt. Rawan, Kapit and Lubuk Antu. 

Ecology. Hill and montane forests to 2700 m, in open screes, usually on moist soils. 

This subspecies differs from the other two (alpestris and moluccana) by its 4-merous 

instead of 5-merous flowers. 

150


~~ 

~ 

16m~ 

.' .. ' ...-"')'i 

~ ~, 

~ 

'~-.~ 

,~ C 

A 

4cm 

] 'om 

}mm 

1 I 

2 mm 

4cm 

B 

Fig. 9. Sarawakodendron filamentosum. A, flowering leafy twig; B, inflorescence; C, flower with 

petal removed; D, longitudinal section through flower; E, cross section (left) and longitudinal section 

(right) of ovary; F, dehisced fruit; G, fruit valve with attached seeds. (All after FM I, 6 (1962) 931, 

fig. 4.) 

151


'om 1 

4 cm 

D 

A 

'------' 

1mm 

'-----------' 

1mm 

Fig. 10. Siphonodon celastrineus. A, flowering leafy twig; B, young flower with petals removed; C, 

longitudinal section through flower, with petals removed; D, fruiting twig. CA from SAN 55718, B-C 

after FM 1,6 (1962) 395, fig. 24, D from S. 39801.) 

152


9. SARAWAKODENDRONDingHou 

(Greek, dendron = a tree; Sarawak tree) 

Blumea 15 (1967) 139, I.e. (1969) 97, I.e. (1972) 930; Anderson I.e. 162. 

Small trees. Young twigs, petioles, midrib, leafblades andfloral parts with yellow pustules. 

Twigs without distinct stipular scars. Leaves alternate, entire; petioles not swollen. Stipules 

inconspicuous. Inflorescences axillary, with condensed decussate bracts. Flowers bisexual; 

calyx 5-lobed, lobes imbricate; petals 5, imbricate; disc slightly 5-angular; stamens 3, 

inserted at the base 9f ovary, anthers extrorse, transversely dehiscent; ovary semi-inferior, 

3-celled, 3-angled, stigma 3, ovules 8 in each cell, in 2 series, placentation axile. Fruit a 

capsule, dehiscing loculicidally into 3 valves. Seeds 6-8 in each cell, with endosperm; aril 

fleshy, cushion or caruncle-like at the chalazal end which bears 1.5-2-cm-Iong simple or 

dichotomously branched, thread-like appendages arising from the base offunicle. 

Distribution. A monotypic genus, endemic to Sarawak. 


Sarawakodendron filamentosum Ding HOll 

(Latin,filamentosus = thread-like; the appendages of the seed aril) 

Fig. 9. 

I.e. (1967) 141, I.e. (1969) 103, I.e. (1972) 932. Type: Ding Hou 333, Sarawak (holotype L; isotypes 

BO, SAR). 

Small tree to 10 m tall, 15 cm diameter. Leaves drying to greenish yellow, elliptic to 

oblong or oblanceolate, 10.5-29 x 4-10.5 cm; base cuneate, apex pointed; midrib flattened 

above; lateral veins 4-8 pairs, faintly raised on both surfaces; intercostal veins reticulate, 

faintly visible below; petioles 0.5-1.5 cm long. Inflorescences 1-2.5 cm long, unbranched, 

few-flowered, peduncles to '1.3 cm long, pedicels 1.5-2 cm long, articulated near base. 

Flowers pale orange, floral parts fleshy; calyx-lobes semi-orbicular, slightly toothed along 

the margin; petals suborbicular with distinct reticulate venation and light brownish stripes 

or dots; disc flat, slightly 5-angular; stamens reflexed at anthesis. Fruits narrow ellipsoid, 

3-angled, 6-8.5 x 2-3.5 cm, gradually narrowed towards both ends. Seeds 6-8 in each cell, 

2-2.5 x 0.5 cm; embryo narrow-lanceolate, c. 18 x 4 mm; cotyledons foliaceous, free. 

Distribution. Endemic to Sarawak. Uncommon, known by the following few collections: 

Ding Hou 133, S. 24506, S. 24897 and S. 24898 from the Nyabau FR, Bintulu; S. 18685 

from Bako National Park; S. 42715 from Sri Aman, and S. 51508 from Bayai, 2nd Div. 

Ecology. In mixed dipterocarp forest on yellow sandy humult ultisol soils and in kerangas 

forest. 

The genus has similar floral characters as that of Salacia and in fruit characters it has 

similarities with Kokoona and Lophopetalum. 

153


10. SIPHONODON Griff. 

(Greek, siphon os = tube, odontos = tooth; 

the hollow, columnar, toothed upper part of the ovary) 

Calc. J. Nat. Hist. 4 (1844) 246; Ding Hou, Blumea 12 (1963) 36, I.e. (1964) 394; Backer & 

BakhuizenJ I.e. 561; Kochummen & Whitmore I.e. 171; Cockburn I.e. 62; Anderson I.e. 162; Ashton 

I.e. 112; Whitmore, Tantra & Sutisna I.e. 45. Synonym: Capusia Lecompte, Bull. Mus. Hist. Nat. 

Paris 32 (1926) 95. 

Trees. Leaves spiral or alternate. Inflorescences axillary, cymose, sometimes only oneflowered. 

Flowers 5-merous; calyx-lobes imbricate; petals imbricate, larger than calyxlobes; 

stamens 5, sometimes alternating with 5 staminodes, united at lower part, anthers 

latrorse, connective distinct and broad; ovary semi-inferior, 3-celled, upper half hollow and 

with a style-like column arising from the bottom, ovules 1 in each cell, oblique or 

pendulous. Fruits drupaceous, with numerous bony I-seeded pyrenes. Seeds flat, with 

endosperm; cotyledons flat, free. 

Distribution. 7 species; SE Asia through Malesia to Australia; one species in Sabah and 


Siphon od on celastrineus Griff. 

(resembling Celastrus) 

Fig. 10. 

I.e. 247; Ding Hou I.e. (1964) 394; Backer & BakhuizenJ I.e. 561; Kochummen & Whitmore I.e. 

171; Cockburn I.e. 62; Anderson I.e. 162; Ashton I.e. 112; Whitmore, Tantra & Sutisna I.e. 45. Type: 

Griffith 9019, Penang (BO, K). Synonyms: S. pyriformis Merr. I.e. (1908) 240; Xanthophyllum 

subglobosum Elmer I.e. (1913) 1676. 

Tree to 35 m tall, 70 cm diameter; buttresses to 2 m high. Bark usually grey-brown, 

smooth to scaly; inner bark yellowish. Sapwood pale. Twigs brownish, zig-zag. Leaves 

drying greenish yellow, lanceolate to elliptic or oblong, 8-15 x 3.5-7 cm; base cuneate, 

margin toothed, apex pointed; midrib flattened above, yellOWish below on drying; lateral 

veins yellowish on drying; intercostal veins reticulate, distinct below, faint above; petioles 

5-8 mm long, drying yellOWish. Flowers cream-white; calyx-lobes almost rounded, 1-2 

mm long; petals ovate with obtuse apex; stamens c. 1 mm long, filaments flat, united near 

base; ovary subglobose to conical, occasionally with 5 ridges towards the apex. Fruits 

subglobose to obovoid, 3-6.5 x 2-6 cm. 

Distribution. India, Burma, Thailand, Indo-China, and Malesia. In Sabah, recorded from 

Lahad Datu, Ranau, Sandakan and Tawau. In Sarawak, collected from Belaga, Bau, Bt. 

Mersing, and Ulu Melinau. 

Ecology. Mixed dipterocarp forests on fertile clay soils, especially near limestone and on 

basic volcanic rock. Uncommon. Fruiting in March-November. 

This species can be confused with those of Xanthophyllum (Polygalaceae) but the toothed 

leaves at once distinguish this. 

154


CHRYSOBALANACEAE 

Ghillean T. Prance 

Royal Botanic Gardens, Kew, England 

R. Brown in Tuckey, Narr. Exp. Congo (1818) 433; King, J. As. Soc. Beng. 66,2 (1897) 275 (under 

Rosaceae); Ridley, F.MP 1 (1922) 665 (under Rosaceae); Merrill, EB (1921) 287, PEB (1929) 92 

(under Rosaceae); Masamune, EPB (1942) 324 (under Rosaceae); Backer & BackhuizenJ, FJ 1 

(1964) 521 (under Rosaceae); Prance & Whitmore, TFM 2 (1973) 321 (under Rosaceae); Anderson, 

CLTS (1980) 292 (under Rosaceae); Cockburn, TS 2 (1980) 79 (under Rosaceae); Corner, WSTM 2 

(1988) 615 (under Rosaceae); Prance & White, Phil. Trans. Roy. Soc. B. 320 (1988) 1; Prance, FM 

10,4 (1989) 635; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 298. 

Trees or shrubs. Leaves simple, entire, alternate, often coriaceous. Stipules small and 

caducous to large and persistent. Inflorescences racemose, paniculate or cymose; flowers 

bracteate and usually 2-bracteolate. Flowers actinomorphic (with a regular symmetry) to 

zygomorphic (with an irregular symmetry), bisexual or unisexual (in Parastemon, plant 

sometimes polygamo-dioecious), markedly perigynous; receptacle short to elongate; disc 

always present, forming a lining to the receptacle; calyx-lobes 5, imbricate, often unequal, 

erect or reflexed; petals 5, imbricate, usually caducous; stamens variable, 2-60, inserted on 

the margin of the surface of the disc, or basally adnate to it, forming a complete circle or, in 

zygomorphic flowers, unilateral, all fertile or some without anthers and then often reduced 

to small staminodes; filaments filiform, free, connate at the base or ligulately connate, 

included to far exserted; anthers small, dorsifixed, longitudinally dehiscent; gynoecium 

(ovary) basically ofthree carpels and gynobasic (inferior), but usually with only one carpel 

fully developed, attached to the base, middle or mouth of the receptacle-tube, sessile or with 

a short gynophore (stalk), pubescent or villous; carpel unilocular with two ovules or 

bilocular with one ovule in each locule, style filiform, arising from the receptacle._at the 

base o/the carpel, stigma distinctly or indistinctly 3-lobed, ovules erect, epitropous (with 

micropyle directed towards the base), tenuinucellate (with thin nucellus). Fruit a dry or 

fleshy drupe; endocarp various, thick or thin, fibrous or bony, often with a special 

mechanism for seedling escape, often densely hairy inside. Seeds erect, almost without 

endosperm; cotyledons pIano-convex, fleshy, sometimes ruminate. 

Distribution. A pantropical family of 17 genera and 520 species, of which over 400 in the 

Neotropics, 60 in Africa and 39 in Asia and the Pacific. In Sabah and Sarawak, the family 

is represented by 6 genera and 15 species all of which are trees. 

Ecology. Found mainly in lowlands both in swamp and well-drained forests, to 1500 m. 

The flowers are pollinated mainly by insects and in a few cases by bats, and the fruits are 

dispersed by birds, mammals and water. 

Uses. The wood is extremely hard and difficult to cut due to the presence of silica; therefore 

it is resistant to decay and to marine borers and is used for posts and marine piles. The 

155


wood of some species is used for general construction and firewood. The fruit of some 

species is edible. The fruit of Atuna racemosa subsp. racemosa is used as a putty for 

caulking boats and the oil from this species is used in the Pacific in hair dressings. 

Taxonomy. Although the family Chrysobalanaceae was described by Robert Brown in 1818 

(In: J.K. Tuckey, Narrative of an expedition to explore the River Zaire, Appendix 5: 433- 

434) it has been placed as a tribe or subfamily of Rosaceae in most of the older and widely 

used systems of classification such as those of Bentham and Hooker, Engler and Prantl and 

Hutchinson. This is in marked contrast to the opinions of nearly all workers with a detailed 

knowledge of the group, especially of its anatomy. The Chrysobalanaceae differs from the 

Rosaceae in the gynobasic style, the basal, erect ovules which are tenuinucellate, in the 

presence of abundant deposits of silica in the wood, stem and leaf and in many wood 

anatomical characters such as the large oblique vessels, the banded oblique vessels, the 

banded parenchyma which is composed of long strands. The authors of modern evolutionary 

systems such as Cronquist, Thorne and Takhtajan consider the Chrysobalanaceae to be a 

separate family but still placed in the Rosales. Dahlgren and Thorne (Ann. Missouri Bot. 

Gard. 71 (1984) 633-699) suggested a relationship between Chrysobalanaceae and Myrtales. 

Although the resemblances between Rosaceae and Chrysobalanaceae are superficial, the 

family is best left in the Rosales until further evidence such as molecular studies show that 

it should be placed elsewhere. Further details about the taxonomy of the group can be found 

in Prance and White's paper (l.c.). 

Key to genera 

(based on flowers) 

1. Flowers actinomorphic with ovary inserted in centre of receptacle; stamens included ... 2 

Flowers zygomorphic with ovary inserted laterally on receptacle; stamens exserted 

(except in Parinari) ..................................................................................................... 3 

2. Stamens 7-10, all fertile .............................................................................. 3. Licania 

Stamens 5, all fertile or 2 fertile and 3 sterile .......................................... 5. Parastemon 

3. Stamens 6-10, included. Leaves usually with stomatal crypts ....................... 6. Parinari 

Stamens 10-40, exserted. Leaves never with stomatal crypts ....................................... .4 

4. Stamens united into a ligule. Ovary unilocular.. ................................ 2. Kostermanthus 

Stamens free to base. Ovary bilocular.. ................................................... . 

5. Stamens 10-20. Inflorescence a raceme or sparsely branched contracted panicle ..... 1. Atuna 

Stamens 25-40. Inflorescence a many-flowered corymbose panicle ......... .4. Maranthes 

Key to genera 

(based on fruits) 

1. Fruits dehiscing by two lateral plates .......... . 

Fruits indehiscent or dehiscing by two basal stoppers ... 

156

CHRYSOBALANACEAE (PRANCE) 

2. Fruits bilocular (sometimes only 1 locule developing), 3-4 cm 10ng ........ .4. Maranthes 

Fruits unilocular, usually 1-1.5 cm long (2.5-3 cm inP. grandifructus) ........ 5. Parastemon 

3. Fruits dehiscing by two basal stoppers; exocarp lenticellate. Leaves usually with 

stomatal crypts beneath ................................................................................ 6. Parinari 

Fruits indehiscent or opening by longitudinal lines. Leaves without stomatal crypts ..... 4 

4. Fruits 1-1.3 cm long, unilocular, breaking open by longitudinal lines of weakness; 

exocarp smooth ....................................................................................................... 3. Licania 

Fruits 3-6 cm long, uni or bilocular, indehiscent; exocarp verrucose-crustaceous ......... 5 

5. Fruits unilocular. Stamens united into a ligule .................................. 2. Kostermanthus 

Fruits bilocular. Stamens free ............................................................................... 1. Atuna 

1. ATUNA Rafin. 

(an Amboinese vernacular name) 

merbatu (Malay) 

Sy1va Tellur. (1838) 153; Kosterrnans, Reinwardtia 7,5 (1969) 421; Prance & Whitmore I.c. 323; 

Anderson I.c. 292; Cockburn I.c. 81; Prance I.c. (1989) 665; Whitmore, Tantra & Sutisna I.c. 298. 

Synonyms: Cyclandrophora Hassk., Flora Beibl. 1 (1842) 47; Parinari subg. Cyclandrophora 

(Hassk.) Blume, Me1ang. Bot. 2 (1855) 10; Parinari auct. non AubI., Rid1ey I.c. (1922) 666. 

Small to large trees, ultimate shoots with complicated system of divaricate branching. 

Stipules large, prominently keeled, lateral, persistent or subpersistent. 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. Inflorescence a raceme, or sparsely branched, contracted panicle; bracts 

and bracteoles persistent, eglandular, not enclosing groups of flower-buds. Flowers bisexual; 

receptacle obconical to cylindrical, as long as or exceeding calyx-lobes, hollow, hairy 

inside throughout, throad 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 disc, filaments free, exserted; staminodes 

forming a barely visible denticular margin to throat; ovary inserted at mouth of receptacle 

tube, pilose on exterior, carpel bilocular with 1 ovule in each loculus. Fruits large; epicarp 

glabrous, densely verrucose-crustaceous; mesocarp transversely fibrous; endocarp hard, 

thick, shortly and sparsely hairy inside, breaking up irregularly at germination; cotyledons 

large and strongly ruminate; germination cryptocotylar (with hidden cotyfedon), eophylls 

alternate. 

Distribution. 8 species from Southern India throughout Malesia and to Fiji and Samoa in 

the Pacific. 3 species occur in Sabah and Sarawak. 

Ecology. Usually found on well-drained mixed dipterocarp forest to 600 m. 

Uses. Little used as a timber. 

157


Key to Atuna species 

1. Leaves broadly ovate, thickly coriaceous, cordate at base. Stamens c. 10 ... 1. A. cordata 

Leaves elliptic or oblong-lanceolate, chartaceous, rounded at base. Stamens 13-20 ...... 2 

2. Receptacle tube narrowly cylindrical. Leaves oblong-lanceolate. Stamens purple ........... 

............ 2. A. nannodes 

Receptacle turbinate-campanulate. Leaves elliptic to oblong-ovate. Stamens white ....... . 

......... 3. A. racemosa 

1. Atuna cordata Cockburn ex Prance 

(Latin, cordatus = heart-shaped; the leaf base) 

Fig. 1. 

Brittonia 39 (1987) 364; Cockburn I.e. 82; Prance I.e. (1989) 667. Type: Ahmad Talip SAN 47687, 

Sabah, Lahad Datu (holotype K; isotypes L, SAN). 

Tree to 40 m tall, the trunk often with thick buttresses. Bark smooth, grey-green, mottled 

white; inner bark hard, reddish brown. Sapwood ochre to red-brown, hard. Young branches 

glabrescent, inconspicuously lenticellate. Stipules to 1.7 cm long, very early caducous. 

Leaves coriaceous, broadly ovate, 4.5-12 x 3-9.5 cm; cordate at base, abruptly acuminate 

at apex, the acumen 1-3 mm long; glabrous and shiny above, glabrous beneath; midrib 

prominulous above, prominent beneath; lateral veins 9-12 pairs, lightly prominulous 

above, prominulous and glabrous beneath; petioles 1-3 mm long, thick, glabrous. 

Inflorescences of terminal and subterminal racemes, 4-8 cm long, borne in single or more 

often in paired branches, densely tomentellous on exterior, puberulous within; bracts and 

bracteoles ovate, tomentellous, early caducous. Flowers with receptacles 5-7 mm long, 

conical to campanulate, tomentellous on exterior, sessile; calyx-lobes slightly unequal, 

tomentellous on both surfaces; petals c. 7 mm long, obovate, glabrous; stamens 10, inserted 

on one side of ring, the filaments 10-12 mm long; ovary densely pilose, style slender, 

hirsutulous on lower portion. Fruits c. 6 x 5 cm, ovoid; epicarp verrucose-crustaceous; 

mesocarp c. 5 mm thick, fibrous, hard; endocarp thin. 

Distribution. Endemic to Sabah; recorded from Lahad Datu, Karamuak, Mt. Tawai and 

Mt. Silam. 

Ecology. Confined to hills and locally common on ultramafic rock to 1200 m. 

2. Atuna nannodes (Kosterm.) Kosterm. 

(Greek, nanno = dwarf; the comparatively small size of the tree) 

Reinwardtia 7, 5 (1969) 422; Prance & Whitmore I.e. 325; Prance I.e. (1989) 667. Basionym: 

Parinari nannodes Kosterm., Reinwardtia, 7, 1 (1965) 50. Type: Beeeari PB 2955, Sarawak, Mt. 

Matang (BO, K). Synonym: Cyclandrophora nannodes (Kosterm.) Kosterm. & Prance, Candollea 20 

(1965) 122. 

Tree to 20 m, usually smaller, unbutttessed. Bark smooth, dark grey. Young branches 

sparsely appressed hirsutulous-strigose, soon glabrous, obscurely lenticellate. Stipules narrowly 

lanceolate, acute, 6-12 mm long, strigose to glabrous, subpersistent. Leaves thinly coriaceous, 

oblong-lanceolate, 6.7-19 x 2.5-6 cm, glabrous on both surfaces, sometimes slightly 

158

CHR YSOBALAN ACEAE (PRANCE) 

lmm 

Fig. 1. Atuna cordata. A, flowering leafy twig; E, leaf lower surface; C, petal; D, flower m 

longitudinal section; E, ovary in cross section. (From SAN 47687.) 

159


bullate above; rounded at base, long slender acuminate at apex, the acumen 7-22 mm long; 

midrib prominulous above, prominent beneath; lateral 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. Flowers with cylindrical receptacle, 8-13 mm long, 

densely sericeous on exterior, sessile; calyx-lobes to 6 mm long, unequal, acute, sericeous 

on exterior; petals white, spathulate to ovate, 8-12 mm long, narrowed to the base; stamens 

18-20, black to purple, the filaments 10-15 mm long, slightly unilateral with tooth-like 

staminodes opposite; ovary pilose, style to 15 mm long, glabrous, stigma capitate. Fruits 

ellipsoid, 3-4 x 1.5 cm, slightly tapered to the base, crustaceous-verrucose on exterior; 

mesocarp 2-2.5 mm, fibrous, hard; endocarp thin. 

Distribution. Peninsular Malaysia, Borneo (Sabah, Sarawak, KalimantarI). In Sabah known by 

collections from localities in the Sandakan, Semporna, Tawau and Tongod districts. In 

Sarawak, recorded from Bau and Kuching areas, 1st Div. 

Ecology. Forest on well-drained soils to 500 ill. 

3. Atuna racemosa Rafin. 

(Latin, racemosus = with a raceme-like inflorescence) 

I.e. 153; Kostermans I.e. (1969) 422; Anderson I.e. 293; Cockbum I.e. 84; Prance I.e. (1989) 669. 

Type: Rumphius Herb. Amb. 1 (1741) pI. 66. 

Tree to 35 m tall often with short buttresses. Bark smooth, grey to black, mottled; inner 

bark orange-brown, red-brown to brown, hard. Sapwood white, ochre to red-brown; heartwood 

red-brown. Young branches glabrous, dark red-brown, smooth or obscurely lenticellate 

when dry. Stipules lanceolate, stiff, to 8-20 mm long, acute, glabrous to strigose, subpersistent. 

Leaves usually chartaceous, more rarely stiffly coriaceous, broadly ovate, elliptic, 

ovate-oblong, oblong or even lanceolate, 4.5-25(-35) x 2-11 cm; rounded, subcordate or 

sub cuneate at base, acuminate at apex, the acumen 3-25 mm long; glabrous on both 

surfaces when mature, sometimes sparsely strigose beneath on lower portion when young; 

midrib prominent on both surfaces; lateral veins 9-13 pairs, prominulous above, prominent 

beneath, straight or arcuate; the venation conspicuously papillose and often giving leaf a 

scabrous or beaded appearance; petioles slender or thick, 3-7 mm long, pilose or puberulous, 

glabrescent or glabrous. Inflorescences of axillary racemes or little-branched with 2-3 

racemose branches 011 short main peduncle, 5-15 cm long, the rachis and branches 

tomentellous or densely short sericeous; bracts and bracteoles ovate to oblong, acute, 3-8 

mm long, persistent or caducous. Flowers with receptacle turbinate-campanulate, 4-10 

mm long, tomentose to sericeous on exterior; pedicels 0.5-1 mm long; calyx-lobes ovate to 

ovate-oblong, 4-7 mm long, densely tomentellous on both surfaces or sericeous on exterior, 

tomentellous within; petals equal, ovate-oblong, c. la mm long, blue or white, caducous; 

stamens 13-20, pale blue, 8-15 mm long with tooth-like staminodes opposite; ovary pilose 

to densely villous, style equalling filaments, glabrous above, stigma smalL Fruits ellipsoid, 

suhglobase to slightly pyriform, 5-7.5 x 3.5-4.5 cm; epicarp crustaceous-verrucose; 

mesocarp fibrous, 5-11 mm thick; endocarp thin, 1-3 mm, densely pilose within. 

160



Leaves 10-25 (-35) cm long, usually elliptic, oblong or lanceolate but sometimes ovate, 

chartaceous or thickly coriaceous, the apex long finely acuminate, acumen 6-25 mm long; 

petioles thick. Flowers 10-17 mm long. Medium-sized to large tree often with fluted bole .... 

subsp. racemosa 

Synonyms: Cyclandrophora glaberrima Hassk. I.e. 47; Parinarium gIaberrimum (Hassk.) 

Hassk., Tijd. Nat. Ges. Phys. 10 (1843) 147, Merrilll.e. (1921) 290, I.e. (1929) 92, Masamune 

I.e. 321; P. seabrum Hassk. I.e. (1843) 147; P. elatum King I.e. 280; c. elata (King) Prance in 

Kostenn., Candollea 20 (1965) 122; Atuna elata (King) Kostenn. I.e. (1969) 42l. 

Thailand to Sumatra, Peninsular Malaysia, Singapore, Borneo (Sabah, Brunei, Sarawak), 

Philippines, Celebes, Moluccas, New Guinea, New Britain, and the Pacific (Admiralty, 

Caroline, Solomon Islands, Fiji, Samoa). In Sabah common and has been recorded in 

most districts. In Sarawak uncommon and collected only from Baram district, and 

known from ecological plots at Bt. Raya, Rejang, Bt. Iju, Balingian, and Bintulu. 

Found mainly in well-drained mixed dipterocarp forests, to 600 m, but also occurs on 

riverbanks and freshwater and brackish water swamps, can be very common. 

Leaves 4.5-12 cm long, usually ovate or oblong-ovate, subcoriaceous or coriaceous, the 

apex bluntly acuminate, acumen 3-10 mm long; petioles thin. Flowers 8-11 mm long. 

Large trees with cylindrical bole .............. . 

subsp. excelsa (Jack) Prance 

I.e. (1989) 670. Basionym: Petroearya exeeIsa Jack, Mal. Misc. 2 (7) (1822) 66. Type: 

Kostelmans & Anta 1136, Bangka, Lobok Besar (neotype K; isoneotypes A, BO, L, SING). 

Synonyms: Parinarium asperulum Miq., Fl. Ind. Bat., Suppl. (1861) 307; Cyclandrophora 

asperula (Miq.) Prance ex Kostenn., Candollea 20 (1965) 130; Cyelandrophora exeeIsa (Jack) 

Kostenn., Candollea 20 (1965) 128; Atuna exeeIsa (Jack) Kostenn. I. e. (1969) 422; P. villamilii 

Merr., Philip. J. Sc. 10 (1915) Bot. 308, Anderson I.e. 294; c. villamilii (Merr.) Prance ex 

Kostenn., Candollea 20 (1965) 126; A. villamilii (Merr.) Kostenn. I.e. (1969) 422. 

Sumatra, Peninsular Malaysia, Java, Borneo (Sabah, Sarawak, Kalimantan), N Celebes. In 

Sabah known from Beaufort, Lahad Datu, Lamag, Sandakan and Tawau districts. In 

Sarawak has been recorded in Lundu, Kuching, Serian, Miri and Kapit districts. 

Usually occurring in well-drained mixed dipterocarp forest, to 750 m, on ridges and 

hilhides, uncommon. 

Uses. The fruit (cotyledon) of subsp. racemosa is grated and made into a putty for caulking 

canoes, widely used in the Pacific islands. An oil extracted from the seeds is 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. 

2. KOSTERMANTHUS Prance 

(AJ.G.H. Kostermans, 1907-1994, botanist of the 

Forest Research Institute and Herbarium Bogoriense, Bogor, Indonesia) 

Brittonia 31 (1979) 91; Prance & Whitmore I.e. 327; Anderson I.e. 293; Cockburn I.e. 85; Prance I.e. 

(1989) 675; Whitmore, Tantra & Sutisna l. e. 299. Synonyms: Aeioa auet. non AubI., Kostennans, 

Reinwardtia 7, 1 (1965) 9; Parinari auet. non AubI.: quoad P. heteropetala et P. myriandra. tantum. 

161


mm 10 I 

-' 

:"mmj f F 

,,1\ , 

"1.\ 

'I 

o 

H 

@ 

::S"-F' 

Fig. 2. Kosterrnanthus heteropetalus. A, flowering leafy twig; B, fruit; C, leaf lower surface; D, 

flower; E, large petal; F, small petal; G, ovary and style; H, ovary in cross section; I, flower in 

longitudinal section. CA & C from Kosterrnans 13630, B from Elmer 21848, D-I from KEP 105046.) 

162

CHRYSOBALAl\IACEAE (PRANCE) 

Large trees, ultimate shoots riot 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 cymulcs proximally and singly 

inserted flowers distally; bracts and bracteoles small, suborbicular, persistent, eglandular, 

not enclosing groups of flower buds. Flowers bisexual, strongly zygomorphic; receptacle 

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 Iigulate, 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 disc; filaments united for half to three quarters of 

length into a strap; staminodes 5-8, inserted opposite stamens; ovary inserted laterally at 

mouth of receptacle, unilocular with 2 ovules. Fruits large, hard; epicarp glabrous, crustaceousverrucose; 

endocarp hard, thick, glabrous within, breaking irregularly on germination. 

Cotyledons slightly ruminate. 

Distribution. 2 species, one (K. malayanus) is confined to Peninsular Malaysia, the other 

(K. heteropetalus) is known from Sumatra, Borneo, the Philippines, and Celebes. 

Kostermanthus heteropetalus (Scort. ex King) Prance 

(Greek, hetero = uneven, petalon = petal; the unequal petals) 

Fig. 2. 

I.e. (1979) 91, I.e. (1989) 677; Anderson I.e. 293; Cockbum I.e. 85; Prance & White I.e. (198g) 152; 

Whitmore, Tantra & Sutisna I.e. 299. Basionym: Parinari heteropetalum Scort. ex King I.e. 283 .. 

Type: Seorteehini 2040, Perak (BO, SWG). Synonyms: Parinarium myriandrum Merr. I.e. (1929) 

93; Aeioa heteropetala (Scort. ex King) Kosterm., Reinwardtia 7, 1 (1965) 11. 

Tree to 35 m tall, 2 m diameter, often with low small buttresses. Bark with small fissures 

and brittle flakes, grey-brown; inner bark pale red-brown to ochre. Sapwood pale yellow; 

heartwood red-brown with a wavy junction between it and sapwood. Young branches 

glabrous, lenticellate. Stipules 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 x 2.5-6 cm; cuneate to rounded at base, bluntly acuminate at 

apex; glabrous on both surfaces, minutely papillose on venation of both surfaces giving a 

bead-like appearance; midrib prominulous above, prominent beneath; lateral veins 6-10 

pairs, arcuate, slender, prominent beneath; petioles 6-12 mm long, sometimes lightly alate 

from decurrent leaf margins, slightly flattened above, eglandular. Inflorescences littlebranched, 

to 10 cm long, the rachis and branches lightly tomentellous; bracts and bracteoles 

ovate, acute, to 3 mm long, caducous. Flowers with a receptacle broadly campanulate, 2-3 

mm long, tomentose on both surfaces; calyx-lobes fleshy, unequal, acute, to 7 mm long, 

pilose on both surfaces, reflexed in open flowers; petals white-tinged pink, fleshy, elliptic, 

concave, largest up to 15 mm long, tomentellous on exterior, enveloping staminal ligule, 

the others much smaller to 6 mm long; stamens 25-30 united into a unilateral ligule for 

two thirds of their length, to 12 mm long, glabrous; anthers pubescent; ovary densely 

pilose, style densely appressed pilose, stigma truncate. Fruits ovoid, unilocular, c. 4 x 3 

cm; epicarp glabrous, crustaceous; endocarp hard, thick. Cotyledons slightly ruminate, c. 

1.5 x 3 cm. 

163


0·········',;):· 

:";... ~.,":,.' ;" 

", Xi;:: 

":'.1 ..:: 

F ..,"/,.. 

/5m m 

10 [ 

mm 

'\\. 

~ 

G 

Fig. 3. Licania splendens. A, flowering leafy twig; 13, flower: C, flower in longitudinal section; D, 

ovary and style; E, petal; F, fruit in longitudinal section; G, fruit. (A-E froni S. 14958, F-G from 

Kostermans 6353.) 

164


Vernacular names. Sabah and Sarawak-merbatu, rasak batu (Malay). 

Distribution. Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei and 

Kalimantan), the Philippines (Mindanao), and Celebes. In Sabah found in Beaufort, 

Keningau, Kota Marudu, Mostyn, and Tongod districts in the west, and in Lahad Datu, 

Sandakan, and Tawau districts in the east. In Sarawak widespread. 

Ecology. Mixed dipterocarp forest on sandy clay soils, to 500 m. 

Uses. The fruit is eaten in Celebes and Sumatra. The timber is easy to cut and is red when 

freshly cut turning brown with age. It is little used because of its tendency to rot. 

3. LICANIA AubI. 

(intended to be an anagram of the local name in French Guiana-caligni) 

Hist. PI. Guiane Fr. 1 (1775) 119; Prance & Whitmore l.c. 328; Anderson I.c. 293; Cockbum l.c. 86; 

Prance I.e. (1989) 645. Synonym: Angelesia Korth., Ned. Kmidk. Arch. 3 (1854) 384. 

Small to large trees. Stipules small, free, caducous. Leaves glabrous on both surfaces, 

without stomatal crypts; petioles eglandular. Inflorescence a panicle of cymules; bracts and 

bracteoles to 1.5 mm long, membranous, eglandular, not enclosing groups of flower buds. 

Flowers bisexual; receptacle campanulate, slightly asymmetric, tomentose on exterior and 

interior; calyx-lobes 5, acute, unequal; petals 5, small, not exceeding the calyx-lobes, not 

clawed; stamens 7-] 0, all fertile, inserted on margin of disc; filaments glabrous, included, 

slightly united at base; ovary inserted at or near base of receptacle, pilose on exterior; 

carpel unilocular, with 2 ovules, style pubescent at base, the stigma capitate. Fruit a small, 

fleshy drupe, narrowed to a shortly stipitate base; epicarp smooth, not ridged, glabrous, 

not lenticellate; mesocarp thin, fleshy; endocarp thin, hard, bony, breaking up in longitudinal 

lines during germination, tomentose within. 

Distribution. 200 species in Neotropics, 1 species in West Africa and 3 species in Thailand 

and Malesia (Sumatra, Java, Borneo, the Philippines, and Papua New Guinea). In Sabah 

and Sarawak only 1 species recorded. 

Licania splendens (Korth.) Prance Fig. 3. 

(Latin, splendens = shining; the shiny surface of the dried leaves) 

Ft. Neotropica 9 (1972) 172, I.c. (1989) 646; Prance & Whitmore l.c. 328; Anderson l.c. 293; 

Cockbum I.c. 86. Basionym: Angelesia splendens Korth. I.c. 384, Merrilll.c. (1921) 290, I.c. C 1929) 

92, Masamune I.c. 324. Type: Korthals, S.n., Sumatra CL). 

Tree to 25 m tall. Bark smooth to scaly, flaking by small scales when old; inner bark redbrown. 

Sapwood pink, hard. Young branches sparsely lanate, soon glabrous. Stipules 

linear-lanceolate, to 3 mm long, caducous. Leaves 4-11 x 1.8;-4.2 cm, oblong; cuneate at 

165


10 

1 

mm 

J 

Fig. 4. Maranthes corymbosa. A, flowering leafy twig; B, leaf-base with glands; C, fruit; D, fruit in 

longitudinal section; E, fruit dehiscence; F, flower; G, flower in longitudinal section; H, petal; I, 

cross-section of ovary; J, ovary and style. (A-B and F-J from SAN A 1888, C-E from Podzorski 

SMHJ 816.) 

166

CHRYSOBALANACEAE(PRANCE) 

base, usually acuminate at apex; glabrous beneath; petioles 2-5 mm, canaliculate, glabrous 

when mature. Inflorescences 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, tomentose within; pedicels c. I 

mm long; calyx-lobes acute, tomentellous on both surfaces; petals pubescent on exterior; 

stamens 7-10 , slightly unilateral, the filaments glabrous; ovary at or near base of 

receptacle, unilocular, pilose on exterior. Fruits 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. 

Vernacular names. Sabah-sampaluan, tampaluan (Dusun). 

Distribution. Thailand, Peninsular Malaysia, Java, Borneo (Sabah, Sarawak, Brunei, 

Kalimantan), the Philippines. In Sabah common, recorded in most districts. In Sarawak, 

known from Bintulu, Kuching, Lundu and Serian districts. 

Ecology. Common tree of primary and secondary mixed dipterocarp forests in well-drained 

usually sandy soils on slopes and also in peat swamp and on seashores and in rocky places 

below 400 m. 

Uses. The timber is strong, durable and resistant to marine borers and is used for saltwater 

piles, railroad 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. 

4. MARANTHES Blurne 

(Greek, maraino = to wither; the long-persistent 

withered calyx and stamens below the fruit) 

Bijdr. (1825) 89; Kosterrnans, Candollea 20 (1965) 196; Prance, BoI. Soc. Brat. Ser. 2,40 (1966) 

183, I.e. (1989) 671; Prance & Whitmore I.e. 329; Anderson I.e. 293; Cockburn I.e. 88; Whitmore, 

Tantra & Sutisna I.e. 299. Synonyms: Parinari auet. non AubI.; Parinari sect. Sareostegia Benth. in 

Hooker, Niger Fl. (1849) 335,pro parte; Parinari subg. Sareostegia (Benth.) Miq., FI. Ind. Bat. 1, 1 

(1855) 355, pro parte; Parinari subg. Exitelia Blume I. c. (1855) 10. 

Medium-sized to large trees. Stipules deltate, intrapetiolar, stiff, caducous. Leaves glabrous 

on both surfaces when mature, 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 

eglandular, caducous, not enclosing flower buds in small group. Flowers bisexual; 

receptacle obconical, narrowed into pedicel, solid, almost completely filled with nectariferous 

tissue, short-tomentose to glabrous on exterior, glabrous within; calyx-lobes suborbicular, 

deeply concave, unequal; petals 5, not clawed; stamens 25-40, inserted on margin of disc, 

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 locule, 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 

167


within, with 2 lateral plates which break away on germination; germination phanerocotylar 

(with exposed cotyledons); cotyledons fleshy, pale green; cataphylls absent; first 2 

eophylls opposite, the others alternate or opposite. 

Distribution. 10 species in tropical Africa, 1 in Central America, and 1 (Maranthes corymbosa) 

widespread in Malesia, Australia and the western Pacific. 


Uses. Wood hard and durable, used for posts and house building. Fruits of most species 

edible. 

Maranthes corymhosa Blume Fig. 4. 

(Greek, corymbos = a cluster; the clustered iirllorescences) 

I.e. (1825) 89; Kostermans, Candollea 20 (1965) 107; Prance & Whitmore I.e. 330; Anderson I.e. 

295; Cockburn I.e. 88; Prance & White I.e. 127; Prance I.e. (1989) 673; Whitmore, Tantra & Sutisna 

I.e. 299. Type: Blume, s.n., Java, Prov. Krawang, near Tjiradja (L). Synonyms: Parinarium 

eorymbosum (Blume) Miq. I.e. (1855) 356, Merrill I.e. (1921) 290, Masamune I.e. 325; Exitelia 

eorymbosa (Blume) Blume, Fl. Jav. 1, Praef. (1828) 7; Parinarium griJjithianum Benth. in Hooker 

I.e. (1849) 334. 

Tree to 30 m tall, 1.5 m diameter, unbuttressed. Bark smooth, grey-brown, lenticellate, 

often mottled with patches of lichen; inner bark red, thin. Sapwood pale white to pink; 

heartwood red-brown. Young branches red-brown, minutely white-lenticellate, glabrous. 

Stipules 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 x 2.5-8 

cm; cuneate at base, acuminate at apex, the acumen 8-20 (-30) mm long; glabrous when 

mature but often sparsely caducous arachnoid-lanate when young, usually with 2 

conspicuous prominent glands at junction of petiole and decurrent lower surface; lateral 

veins 7-10 pairs, arcuate, prominulous on both surfaces; midrib plane above, prominulous 

beneath; petioles 4-9 mm long, glabrous when mature, flattened above. Inflorescences of 

flattened many-jlowered corymbose panicles; rachis and branches sparsely pilose, glabrescent; 

bracts and bracteoles ovate to lanceolate, sparsely pubescent, caducous. Flowers with 

receptacle turbinate, 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, glabrous, 3-6 mm long, caducous; stamens 25-35, 

inserted in several rows on one side of throat, with tooth-like staminodes opposite; ovary 

densely lanate and villous; stigma truncate. Fruits ellipsoid, 3-4 x 1.5-2 cm, tapered 

towards base; epicarp thin, glabrous on exterior when mature, sometimes lanate when 

young; endocarp c. 5 mm thick, densely lanate within, bilocular usually with seed in one 

locule only. Cotyledons pIano-convex. 

Vernacular names. Sabah-bangkawang (Dusun, Malay). Sarawak-merbatu (Malay). 

Distribution. S Thailand extending east to Solomon and Caroline Islands and Australia. In 

Malesia: Sumatra, Peninsular Malaysia, Java, Borneo (Sabah, Sarawak, Brunei, Kalimantan), 

168


the Philippines, Celebes, Moluccas, New Guinea, New Britain and Admiralty Islands. In 

Sabah, widespread but in Sarawak, uncommon in mixed dipterocarp forest in the northern 

parts. 

Ecology. In primary and secondary forests from the sea-coast to 1500 m on Mt. Kinabalu, 

Sabah. The fruit is eaten by many birds which probably disperse the seed and account for 

the wide distribution of this species. 

Uses. The wood is used for house building and for posts and the fruit is edible. 

5. PARASTEMON A. DC. 

(Greek, para = near, stemon = stamen; both fertile and 

sterile stamens are located near one another at one side of the flower) 

Ann. Sci. Nat. Bot. Ser. 2,18 (1842) 28; Miquell.e (1855) 359; Merrilll.e. (1921) 290; Ridley I.e. 

(1922) 672; Corner I.e. 617; Masamune I.e. 324; Anderson I.e. 293; Cockburn I.e. 90; Prance I.e. 

(1989) 648; Whitmore, Tantra & Sutisna I.e. 299. Synonyms: Diemenia Korth. I.e. (1854) 388; 

Triehoearya Miq. I.e. (1855) 537, pro parte. 

Trees or shrubs. Stipules small and triangular, caducous. Leaves glabrous on both surfaces, 

without stomatal crypts, with 2 small discoid glands at base of lamina; petioles eglandular. 

Inflorescences axillary or rarely terminal simple or sparsely branched racemes; bracts 

and bracteoles small, eglandular, not enclosing groups of flower buds. Flowers bisexual or 

unisexual; receptacle patelliform or shallowly cupuliform, shortly hairy within; calyx-lobes 

5, acute, subequal; petals 5, not exceeding calyx-lobes, not clawed; stamens 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 receptacle, 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 drupe, 1.5-3 cm long, with 2 large lateral plates which break away on germination 

to allow seedling escape; epicarp smooth, not lenticellate; endocarp thin, hard, bony, 

smooth on exterior, glabrous within. 

Distribution. 3 species in the Nicobar Islands, Sumatra, Peninsular Malaysia, Borneo, 

Moluccas, New Guinea, and Admiralty Islands. 2 species in Sabah and Sarawak. 

Ecology. Lowland and swamp forests. 

Key to Parastemon species 

Leaves with 5-6 pairs oflateral veins. Fruit 2.3-3.5 cm long ................. 1. P. grandifructus 

Leaves with 8-11 pairs oflateral veins. Fruit 1-1.5 cm long ....................... 2. P. urophyllus 

169

TREE FLORA OF SABAH AND SARAWAK VOL. \ (\995) 

10 I mm 

A 

§is 

§f> 

I'~ 

J 

I··· 

(' 

~ 

" 7 

o.'mm'H \~~: 

Fig. 5. Parastemon urophyllus. A, flowering leafy twig; B, leaf lower surface; C, male flower; D, 

male flower in section; E, female flower; F, female flower in section; G, ovary and style; H, petal; I 

fruit showing plate of dehiscence; J, interior of fruit; K, cotyledons. (A-B and E-H from Anderson S. 

9803, C-D from Carrick et al. JC/42, I-K from SAN 67203.) 

170


1. Parastemon grandifructus Prance 

(Latin, grandis = large, fructus = fruit) 

Brittonia 39 (1987) 366, I.e. (1989) 650. Type: Wright & Ismawi S. 32320, Sarawak (L). 

Tree to 30 m tall, 30 cm diameter; trunk lightly buttressed to 1 m high. Young branches 

glabrous. Stipules caducous. Leaves coriaceous, glabrous on both surfaces, elliptic to 

narrowly elliptic-oblong, 5-8.5 x 1.8-3.2 cm; cuneate at base, with a long-cuspidate 

acumen at apex, the tip 10-16 mm long; midrib prominent above, prominulous or plane 

beneath; lateral veins 5-6 pairs, prominulous above, plane beneath; petioles 5-8 mm long, 

glabrous, slightly canaliculate, slightly swollen at base. Inflorescences of axillary and 

terminal racemes, the rachis glabrous. Flowers (seen in fruiting specimens): calyx-lobes 5, 

acute, glabrous on exterior, glabrous within except for a few hairs around base; receptacle 

glabrous on exterior in fruiting condition; style persistent below fruits, the stigma bifid or 

trifid. Fruit ellipsoid, 2.3-3.5 x 1.3-1.5 cm; epicarp smooth, glabrous; mesocarp thin, c. 

0.25 mm; endocarp thin, hard, bony, c. 0.25 mm thick, glabrous within, opening by 2 

lateral plates 1.9-2 cm long. 

Vernacular names. Sabah--kayu ajung, mandailas (Dusun). Sarawak--ngilas (lban, in 

common with Xanthophyllum). 

Distribution. Endemic to Borneo (Sabah and Sarawak). In Sabah known from Beaufort 

Hill and Lumat Estate in the Beaufort district. In Sarawak found in Bau, Kapit, Kuching 

and Limbang districts. 

Ecology. Lowland forests, including swamp and heath forests, to 150 m. 

2. Parastemon urophyllus (Wall. ex A. DC.) A. DC. 

(Greek, aura = tail, phyllon = leaf; the cuspidate apex of the leaf) 

Fig. 5. 

I.e. 208; Miquell.e. (1855) 359; Merrilll.e. (1921) 290; Ridley I.e. (1922) 672; Corner I.e. 617; 

Masamune I.e. 324; Browne, FTSB (1955) 308; Anderson I.e. 293; Cockburn I.e. 90; Prance & 

Whitmore I.e. 331; Prance I.e. (1989) 649; Whitmore, Tantra & Sutisna I.e. 300. Basionym: Embelia 

urophylla Wall. ex A. DC., Trans. Linn. Soc. Lond. 17 (1837) 131. Type: Griffith, s.n., Malacca (K). 

Synonym: Parastemon spieatum Ridl., J. Str. Br. R. As. Soc. 75 (1917) 29, Merrilll.e. (1921) 290, 


Tree to 35 m tall often with small buttresses. Bark brownish, smooth, becoming slightly 

cracked and fissured with age. Young branches glabrous, waxy resinous when young. 

Stipules triangular, c. 1 mm long, caducous. Leaves thinly coriaceous, narrowly oblong, 

2.5-8 x l.4-2.5 cm; cuneate at base, cuspidate-acuminate at apex, the lip 5-15 mm; midrib 

plane above, prominulous beneath; lateral veins 8-11 pairs; petioles 4-5 mm long, 

canaliculate, glabrous. Inflorescences of axillary and rarely terminal racemes or occasionally 

slightly branched, 4-14 cm long, the rachis glabrous. Flowers unisexual (plant polygamodioecious), 

c. 1.5 mm long; receptacle broadly cupuliform to flattened saucer-shaped, 

glabrous on exterior, tomentose within; pedicels to 2 mm long; calyx-lobes acute, glabrous 

on exterior; petals 5; stamens 2fertile and 3 sterile staminodes opposite; ovary inserted at 

171


base of receptacle, pilose on exterior, unilocular, style pilose at base, glabrous above, the 

stigma trifid. Fruits e /lipsoi d, 1-1.5 cm long; epicarp smooth, glabrous; mesocarp thin, 

hard; endocarp thin, hard, bony, glabrous within, opening by 2 lateral plates. 

Veruacular names. Sabah-mendailas (Dusun). Sarawak-mengilas, gilas (Bidayuh). 

Distribution. Nicobar Islands, Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, 

Brunei, Kalimantan). In Sabah found in Beaufort, Keningau, Papar and Sipitang districts 

in the west, and in Lahad Datu and Sandakan districts in the east. In Sarawak widespread 

and recorded in most districts. 

Ecology. Characteristic component of peat swamp forest where ·it is a common large tree, 

but also wide-ranging into kerangas, more open scrub forest, and secondary forest on poor 

soils. 

Uses. The wood is hard and very heavy; grain straight or interlocked. It is used for general 

construction, posts and as a firewood. 

6. PARINARI AubI. 

(a vernacular name in French Guiana) 

merbatu (Malay) 

I.e. 204; MerrillI.e. (1921) 290, I.e. (1929) 92; Masamune I.e. 325; Backer & BakhuizenJ I.e. 521; 

Kosterrnans, Reinwardtia 7, 1 (1965) 147; Prance & Whitrnore I.e. 332; Anderson I.e. 294; Cockbum 

I.e. 90; Prance I.e. (1989) 654; Whitrnore, Tantra & Sutisna I.e. 300. Synonym: Parinarium Juss., 

Gen. PI. (1789) 342; Petroearya Schreb. in Linnaeus, Gen. PI. ed. 8, 1 (1789) 245; Parinarium subg. 

Petroearya (Schreb.) Miq. I.e. (1855) 352. 

Small or large trees or rarely shrubs. Stipules small to large, persistent or caducous. Leaves 

usually with stomatal crypts, filled with pubescence on lower surface or rarely glabrous, or 

lanate pubescent without crypts; petioles usually with 2 circular glands above. Inflorescence a 

many-flowered complex cyme or cymose panicle; bracts and bracteoles eglandular, usually 

concealing flower buds indiVidually and in small groups. Flowers bisexual; receptacle 

subcampulate 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 upper half of receptacle-tube below mouth, 

pilose on exterior, carpel bilocular with 1 ovule in each locule, style arcuate, included. 

Fruit a fleshy drupe; epicarp verrucose; endocarp thick, with a rough fibrous surface, with 

2 basal obturators for seedling escape. 

Distribution. Pantropical with 18 species in the Neotropics, 6 in Africa and 15 in tropical 

Asia, Malesia and the Pacific. 7 species in Sabah and Sarawak. 

Ecology. A wide range of lowland forest and one species in submontane forest. 

Uses. The wood is hard and heavy and is little used, the fruits of most species are edible. 

172

CHRYSOBALANACEAE(PRANCE) 

Key to Parinari species 

1. Stomatal crypts absent from leaf underside; leaf underside glabrous or with a persistent 

lanate pubescence and then with large persistent stipules 7-20 mm ............................. 2 

Stomatal crypts present on leaf underside; leaf underside lanate or at least pubescent in 

crypts; stipules usually small or iflarger then early caducous ...................................... .4 

2. Leafunderside glabrous; stipules small and caducous .................................................. 3 

Leaf underside densely lanate pubescent, but when removed no stomatal crypts present; 

stipules large and persistent, 7-18 mm 10ng ............................................... .4. P. elmeri 

3. Leaves elliptic to oblong or obovate-elliptic, 9.5-20.5 x 4.5-8.5 cm; lateral veins 11- 

16 pairs. Panicles large and silvery pubescent... .......................... 1. P. argenteo-sericea 

Leaves ovate, 5-9 x 2-45 cm; lateral veins 7-11 pairs. Panicles small and subsericeous 

brown pubescent.. ............................................................................... 2. P. canarioides 

4. Leaves with 20-33 pairs of lateral veins ............................................. 6. P. oblongifolia 

Leaves with 10-19 pairs oflateral veins ....................... . 

5. Petioles 14-20 mm long. Leaves with metallic sheen above .................... 5. P. metallica 

Petioles 3-10 mm long. Leaves without a metallic sheen ............................................. 6 

6. Leaves rigidly coriaceous, often broadest well below middle point; midrib and often 

lateral veins lightly impressed on upper surface. Receptacle c. 5 mm. Fruits ovoid, 7-8 

cm long ....................................................................................................... 7. P. rigida 

Leaves chartaceous, broadest at or above middle; midrib and lateral veins prominulous 

above. Receptacle 3-3.5 mm. Fruits ellipsoid, 3.5-4.5 cm long .................. 3. P. costata 

1. Parinari argenteo-sericea Kosterm. 

(Latin, argenteus = silvery, sericeus = silky; the pubescence of the flowers and inflorescence) 

Reinwardtia 7, 1 (1965) 47, 158; Cockburn I.e. 91; Prance I.e. (1989) 656; Whitmore, Tantra & 

Sutisna I.e. 300. Type: Wood SAN 16175, British North Borneo, Lahad Datu (holotype BO; isotypes 

A, BR!, K, KEP, L, SING). 

Tree to 35 m tall. Bark brown, lenticellate, hard; inner bark red, hard, c. 1.2 mm thick. 

Sapwood yellow. Young branches glabrous, dark purplish brown, with numerous lenticels. 

Stipules lanceolate, to 8 mm long, tomentose on exterior, early caducous. Leaves chartaceous, 

oblong, elliptic to subovate-elliptic, 9.5-10.5 x 4.5-8.5 cm, glabrous on both surfaces, 

without stomatal crypts beneath, usually 2 glands beneath at base near junction with 

midrib; rounded at base, acute to shortly acuminate at apex, the tip 7-10 mm long; midrib 

lightly impressed above except near base, prominent beneath; lateral veins 11-16 pairs, 

plane above, prominent beneath, erect-patent; petioles 5-9 mm long, eglandular, glabrous, 

rugulose. Inflorescence a lax, much-branched, terminal panicle, 9-15 cm long, the rachis 

and branches densely grey sericeous-tomentose; bracts and bracteoles ovate, acute, densely 

tomentellous on exterior, glabrous within except near apex, caducous. Flowers with 

173

TREE FLORA OF SABAI-! AND SARAWAK VOL. 1 (1995) 

receptacle campanulate, markedly gibbous, densely grey-tomentellous on exterior, 2-3 mm 

long, narrowly ovate, densely grey-tomentose on exterior, tomentellous within; petals 

spathulate, c. 2 mm long, caducous; fertile stamens 7-8, base forming a conspicuous fused 

ring with opposite tooth-like staminodes; ovary densely pilose, style pilose, stigma truncate. 

Fruits ovoid, 7-8 x 4.5-5.5 cm; epicarp densely lenticellate; mesocarp thin, fleshy; 

endocarp extremely hard and thick (1-8 cm thick), woody, granular, and very irregularly 

ridged, with 2 smalllocules in centre, densely lanate within. 

Distribution. Endemic to Sabah, known from a few collections from the Lahad Datu, 

Sandakan, and Tawau districts. 

Ecology. Lowland forest to 100 m and forest along rivers. 

2. Parinari canarioides Kosterm. 

(resembling the genus Canarium) 

New & Crit. Mal. PI. (For. Dept. Bur. of Planning Bogor, Indonesia) 3 (1955) 25; Reinwardtia 7,2 

(1965) 159; Anderson I.e. 294; Cockburn I.e. 93; Prance I.e. (1989) 656; Whitmore, Tantra & Sutisna 

I.e. 300. Type: Kostermans 7152, Kalimantan, Tanjong Bangko (holotype BO; isotypes A, BISH, 

BR!, CAL, CANB, K, L, LAE, IvIEL, NY, P, PNH, SAN, SING). 

Tree to 60 m tall, trunk often buttressed to 1.5-2.5 m. Bark smooth, irregularly fissured 

and flaking, dark red-brown; inner bark pale red-brown. Sapwood dull yellow to pink; 

heartwood brownish red. Young branches sparsely puberulous soon becoming glabrous, 

grey-brown. Stipules linear, acute to 5 mm, hirsute, early caducous, present on very young 

leaves only. Leaves chartaceous, ovate, 5-9 x 2-4.3 cm, glabrous on both surfaces when 

mature, without stomatal crypts beneath; rounded to subcordate at base, acuminate at apex, 

the tip 5-12 mm long; midrib lightly impressed above, prominent beneath, sparsely 

pubescent when young; lateral veins 7-11 pairs, plane to prominulous above, prominent 

beneath, arcuate; petioles 3-7 mm, glabrous when mature, eglandular or with small rather 

inconspicuous central glands. Inflorescences dense-flowered axillary panicles to 4.5 cm 

long, the rachis and branches tomentose; bracts and bracteoles persistent, ovate, puberulous 

on exterior, caducous. Flowers with receptacle campanulate, c. 3 mm long, tomentose on 

exterior; pedicels 1-2 mm long; calyx-lobes elliptic, concave, c. 2 mm, acute, sparsely 

puberulous on exterior, densely tomentellous on interior; petals elliptic, obtuse, c. 2 mm, 

tapered to the base; fertile stamens 7-8. Fruits ellipsoid, 3.5-5 x 1.5-2.5 cm; epicarp 

densely to sparsely lenticellate; mesocarp fleshy, c. 1 mm thick; endocarp c. 5 mm thick, 

hard, marbled, densely lanate within. 

Distribution. Sumatra, Borneo (Sabah, Sarawak, Brunei, Kalimantan), the Philippines 

(Palawan), and Celebes. In Sabah found in Kunak and Sandakan districts; widespread but 

scattered in Sarawak and Brunei. 

Ecology. Primary and secondary mixed dipterocarp forests on well-drained clay-rich soils 

to 800 m. 

Uses. Fruit edible. 

174


3. Parinari costata (Kunth) Blume 

(Latin; costatus = ribbed; the strongly veined leaf) 

I.e. (1855) 10; Miquell.e. (1855) 354; Merrilll.e. (1921) 290; Ridley I.e. (1922) 666; Masamune I.e. 

325; Kosterrnans, Reinwardtia 7, 2 (1965) 179; Prance & Whitmore I.e. 333; Anderson I.e. 294; 

Prance I.e. (1989) 663; Whitmore, Tantra & Sutisna I.e. 300. Basionym: Lepidoearpa eostata Korth. 

I.e. 387. Type: Korthals, s.n., Sumatra (L). 

Tree to 60 m tall, 90 cm diameter; trunk buttressed up to 2 m. Bark smooth to roughish, 

cracked, grey or brown, c. 0.5 mm; inner bark pale reddish to reddish brown with pale 

spots, 6-10 mm thick. Wood pale brown, darker towards centre. Young branches densely 

yellow-brown, appressed tomentellous becoming glabrous, with numerous small conspicuous 

lenticels. Stipules lanceolate, membranous, 3-7 mm long, pilose on exterior, early 

caducous. Leaves rigidly chartaceous, elliptic, sub ovate-elliptic, oblong-elliptic to oblonglanceolate, 

4-11.5 x 1.6-4.3 cm, glabrous above when mature but with sparse lanate 

covering when very young, with stomatal crypts filled with grey lanate pubescence 

beneath; rounded to subcuneate at base, acuminate at apex, the tip 3-5 mm long; midrib 

prominulous above, tomentellous towards base, prominent beneath; lateral veins 10-19 

pairs, arcuate, prominulous above, prominent beneath; intercostal veins rounded or only 

slightly flattened; petioles 4-9 mm long, slender, thickly tomentose or tomentellous when 

young, soon glabrous, usually eglandular or with 2 inconspicuous median glands. Inflorescences 

of predominantly axillary or terminal few-flowered lax or dense panicles to 8 cm 

long, the rachis and branches grey to brown appressed tomentellous or ferrugineousvillous 

pubescent; bracts and bracteoles lanceolate, c. 2 mm long, caducous. Flowers with 

receptacle campanulate, slightly gibbous, grey-brown pubescent on exterior, 3-3.5 mm 

long; pedicels 0.5-1 mm long; calyx-lobes ovate, acute, l.5-2 mm long, grey tomentellous 

on exterior; petals white, spathulate, 1.5-2 mm long, caducous, glabrous; stamens 7-8, 

with small tooth-like staminodes opposite, slightly unequal; style glabrous, stigma capitate. 

Fruits ellipsoid, to 3.5 x 4.5 cm; epicarp usually sparsely verrucose or lenticellate; 

mesocarp c. 2 mm, fleshy; endocarp hard, marbled, 3-5 mm thick, fibrous, densely lanate 

within. 


Inflorescences and flowers sparsely to densely grey or brown appressed pubescent. Lowland 

plants ............................................................. . 

subsp. costata 

Surnatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan). Mixed 

dipterocarp forests on well-drained soils, to 300 m. In Sabah and Sarawak uncommon. 

Inflorescences and flowers densely ferrugineous-villous pubescent. Usually at high altitudes 

subsp. rtIbiginosa (Ridl.) Prance 

Brittonia 39 (1987) 368, I.e. (1989) 663. Basionym: Parinarium eostatum var. rubiginosum 

Rid!. I.e. (1915) 143; Parinarium rubiginosum (Rid!.) Rid!. I.e. (1917) 29, I.e. (1922) 668. 

Type: Ridley 16016, Malaya, Padang (holotype K; isotype SING). Synonym: Parinari bieolor 

Merr.,Philip. J. Sc. 10 (1915) Bot. 309. 

Burma, Peninsular Malaysia, Borneo (Sabah, Sarawak, Kalimantan). Submontane 

forests at 750-1000 m. Uncommon; in Sabah known from Mt. Kinabalu (Clemens 

175


G 

E 

8 

Fig. 6. Parinari oblongifolia. A, flowering leafy twig; B, leaf lower surface; C, f1ower; D, 

longitudinal section of f1ower; E, cross-section of ovary; F, petal; G, ovary and style; H, fruit. (A-G 

from Enggoh 7249, H from SAN 69261.) 

176

50081), Sandakan district (Puasa 669), and in Sarawak from Similajau and Segan FR, 

Bintulu, 4th Div. 

Distribution. Burma, Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak and 

Kalimantan), and the Philippines. 

Ecology. Mixed dipterocarp to submontane forests on well-drained soils at 300-1000 m. 

Taxonomy. Three subspecies are recognised, two of which occur in Sabah and Sarawak (as 

above), and a third in the Philippines. 

4. Parinari elmeri Merr. . 

(A.D.E. Elmer, plant collector with the Bureau of Science, Manila, the Philippines) 

177 

Ecology. Mixed dipterocarp forests on well-drained soils to 900 m, including areas on 

uItramafic rock. 

Uses. The wood is used for supports of Than long houses. 

Tree to 30 m usually much smaller; trunk unbuttressed. Bark pale cream or grey and white 

mottled, roughened by green excrescences, soft, thin; inner bark orange, hard, c. 2.5 mm 

thick. Sapwood white, thin; heartwood straw coloured, hard. Young branches densely 

tomentellous soon becoming glabrous, obscurely lenticellate. Stipules lanceolate, acute, to 

18 mm long and 3 mm broad at base, lateral, tomentellous, perSistent. Leaves oblong to 

oblong-lanceolate, 5-18 x 1.5-7 cm, chartaceous to thinly coriaceous, glabrous above, 

densely pubescent beneath, without stomatal crypts; subcuneate at base, acuminate at apex, 

the tip 5-13 mm long; midrib plane or slightly impressed and pubescent above when 

young, prominent beneath; lateral veins 14-21 pairs, prominent beneath, curved at margin; 

intercostal veins more or less parallel forming ladder-like reticulation; petioles 1.5-6 mm 

long, tomentellous, 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, c. 2 mm long, ovate, persistent. 

Flowers with receptacle conical, gibbous, to 3 mm long, brown-Ianate on exterior, 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 the base; fertile stamens 7-9, with tooth-like 

staminodes opposite. Fruits oblong-ellipsoid, c. 6.7 x 3.7 cm; epicarp sparingly lenticellate. 

Distribution. Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan), and the 

Philippines. Uncommon; in Sabah known from the type collection and from Kelumpang 

Hill, Lahad Datu district (SAN 29329), and Tawau district. In Sarawak only two collections 

from G. Silantek, 85th mile, Sirnanggang Rd., 2nd Div. (S. 42636) and Mt. Matang 

(Clemens s.n.) are known, but has been recorded from ecological plots at Bt. Mersing, 

Anap, and Nyabau FR. 

I.e. (1929) 92; Masamune I.e. 325; Kostermans, Reinwardtia 7, 2 (1965) 161; Prance & Whitmore 

I.e. 335; Anderson I.e. 294; Cockburn I.e. 91; Prance I.e. (1989) 657; Whitmore, Tantra & Sutisna 

I.e. 300. Type: Elmer 20806, British North Borneo, Tawao, Elphinstone Province (holotype VC; 

isotypes BISH, BO, BR, C, DC, DS, F, GH, L, M, MO, P, S). 

CHRYSOBALANACEAE (PRANCE)

l.c. 309; Ridley l.c. (1922) 335; Anderson l.c. 294; Cockburn I.c. 93; Prance I.c. (1989) 659; 

Whitmore, Tantra & Sutisna I.c. 301. Type: Griffith, s.n., Malacca (K). Synonyms: Parinarium 

borneense Merr. l.c. (1929) 93, Masamune I.c. 325; Parinari gigantea Kosterrn., Reinwardtia 7, 2 

(1965) 182; Prance l.c. (1989) 660; sy1/. novo 

Tree to 40 m tall, trunk with low thick buttresses to 2 m. Bark smooth, grey to pale brown, 

very thin; inner bark reddish brown to brown, 6-12 mm thick. Sapwood white to pale 

yellow; heartwood reddish, hard. Young branches minutely tomentellous, with numerous 

pale, small, roundish lenticels. Stipules ovate to lanceolate, acute, 3-5 mm, pilose on 

exterior, early caducous. Leaves coriaceous, elliptic to oblong, 14-23 x 4-9 cm, glabrous 

above, with stomatal crypts, filled with grey lanate pubescence beneath; rounded to 

subcordate at base, shortly acuminate at apex, the tip 3-13 mm long; midrib plane above, 

glabrous when mature except at base, prominent, glabrescent beneath; lateral veins 23-35 

pairs, erect, plane above, flattened and prominent beneath; intercostal veins prominulous, 

parallel and more or less ladder-like beneath; petioles 9-17 mm long, thick-tomentellous 

when young, glabrescent, eglandular or glandular. Inflorescences of large, spreading 

178 

Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei). In Sabah found in Beaufort 

Hill FR, and in Sarawak in Semengoh Arboretum, Kuching, 1st Div., and Lambir Hills, 

Miri district, 4th Div. 

6. Parinari oblongifolia Hook.! Fig. 6. 

20 mm long, glabrescent, with inconspicuous glands near to lamina base, puberulous, 

glabrescent. Inflorescences of axillary little-branched panicles, 4-10 cm long, the rachis 

and branches densely brown tomentellous; bracts and bracteoles ovate, early caducous. 

Flowers with receptacle campanulate, slightly gibbous, 2-3 mm long, ferrugineous, 

pubescent on exterior; pedicels c. 0.5 mm long; calyx-lobes lanceolate, acute, c. 1 mm long, 

tomentellous; petals lanceolate, glabrous; stamens c. 8 with short tooth-like staminodes 

opposite; ovary densely pilose, style glabrous, equalling stamens, stigma truncate. Fruits 

not seen. 

4-9 cm, glabrous and shiny with metallic sheen above when dry, with dense stomatal crypts 

Reinwardtia 7, 1 (1965) 49, 160; Cockburn I.c. 91; Prance l.c. (1989) 660. Type: Ashton BRUN 

3267, Brunei, Andulau FR (holotype SAR; isotypes BO; K, L). 

Tree to 20 m tall, unbuttressed. Young branches appressed strigose soon becoming 

glabrous, conspicuously lenticellate. Stipules ovate-lanceolate, acute, 8-15 mm long, densely 

brown-tomentose, membranous, early caducous. Leaves thickly coriaceous, elliptic, 8-17 x 

filled with hairs; rounded or sub cuneate at base, apex rounded to shortly blunt-acuminate, 

the tip to 3 mm long; midrib plane above, prominent beneath; lateral veins 10-15 pairs, 

prominulous to plane above, prominent beneath, erect, curved only at margin; petioles 14- 

(Latin, metallicus = metal-like; the metallic sheen on the upper surface of dried leaves) 

Ecology. Mixed dipterocarp forests on deep sandy humult ultisol soils, to 300 m. 

(Latin, oblongus = rather long,folius = leaves; the leaf-shape) 

TREE FLORA OF SABAl-I AND SARAW AK VOL. 1'(1995) 

5, Parinari metallica Kosterm.

terminal panicles, 10-21 cm long, 7-12 cm broad, the rachis and branches yellow-grey 

tomentellous; bracts and bracteoles ovate, c. 3 mm long, early caducous. Flowers with 

receptacle campanulate, slightly gibbous, c. 3 mm long, densely grey-tomentose on 

exterior; pedicels 1-3 mm long; calyx-lobes ovate, acute, l.5-2 mm long, unequal, greytomentose; 

petals white to bluish, lanceolate to spathulate; narrowed towards the base, c. 2 

mm long, glabrous; stamens 7-10, with tooth-like staminodes opposite; ovary pilose, style 

glabrous, stigma truncate. Fruits ellipsoid, 5-9 x 3-4 cm; epicarp densely lenticellate; 

mesocarp 1.5-2 mm thick; endocarp hard, marbled, 7-13 mm thick, fibrous, densely lanate 

within. 

Distribution. Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan). 

In Sabah found in Lahad Datu, Sandakan, Semporna, and Tawau districts; in Sarawak only 

recorded from sterile ecological collections from Santubong to Miri. 

Ecology. Lowland rainforest, on river banks or in valleys, to 450 m. 

179 

Ecology. Heath and swamp forests, lovvlands to 1400 m. 

Distribution. Sumatra, Peninsular Malaysia, Borneo (known only from Lundu, Kuching 

and Miri districts in Sarawak). 

Reinwardtia 7, I (1965) 53, 163; Prance I.c. 660. Type: Sin clair & Kiah SFN 40773, Malay 

Peninsula, Trengganu (SING). Synonym: Parinari ashtonii Kostenn., Reinwardtia 7, I (1965) 53, 

Anderson I.c. 294. 

Tree to 30 m tall. Bark smooth. Young branches tomentellous soon becoming glabrous, 

obscurely lenticellate. Stipules caducous (not seen). Leaves rigidly coriaceOl.is, elliptic to 

oblong-ovate, 7.5-23 x 3-8 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; rounded or subcordate at base, shortly and broadly acuminate at 

apex, the tip 3-I 7 mm long; midrib plane or impressed for upper portion above, prominent 

and appressed pilose beneath when young; lateral veins 13-20 pairs, slightly impressed 

above, prominent beneath, slightly curved at margins only; intercostal venation flattened or 

rounded, parallel; petioles thick, 3-10 mm long, grey-pi lose 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 lanceolate. to 2 mm 

long, early caducous. Flowers with recf'ptacIe campanulate, slightly gibbous, c. 5 mm long, 

densely viIlous-tomentose on exterior; pedicels c. 1 mm long; calyx-lobes elongate 

triangular, 2-2.5 mm long; petals spathulate; stamens 6-8; ovary densely villous, style 

equalling stamens, stigma capitate. Fruits irregularly ellipsoid, c. 5 x 4 cm, tapered towards 

the base almost into a stipe; epicarp densely lenticellate; mesocarp thin, fleshy; endocarp 

thick, woody, marbled, lanate within. 

7. Parinari rigida Kosterm. 

(Latin, rigidus = stiff; the thick coriaceous leaves) 

CHRYSOBALANACEAE (PRANCE)

CLETHRACEAE 

A. Berhaman 




181 

Key to species 

1. Leaf lower surface with stellate hairs only on the midrib and veins, glabrous on the 

intervenium. Calyx-lobes 1.5-2 mm long .............................................. 1. C. canescens 

CLETHRA Gronov. ex L. 

(Latinised old Greek word - klethra = alder tree) 

kolintuhan (Dusun Ranau, Sabah) 

Gen. PI. ed. 5 (1753) 188, Sp. PI. (1753) 396; Merrilll.e. (1921) 460; Masamune I.e. 567; Sleumer, 

Bot. Jahrb. 87 (1967) 85, I.e. (1971) 139; Whitmore I.e. 27; Anderson I.e. 162; Whitmore, Tantra & 


Trees or shrubs. Leaves simple, spiral, crowded towards the end of the shoots; margin 

serrate with glandular teeth, more rarely entire; lateral veins and midrib sunken on upper 

side, prominent on the lower side; stipules none. Inflorescence a solitary terminal raceme, 

or (usually) a terminal raceme and several lower approximate racemes, from the axils of 

reduced or caducous leaves; bracts mostly caducous during anthesis, rarely subpersistent. 

Flowers bisexual; calyx-lobes 5(-6), persistent, alternate with the petals; petals 5(-6), 

generally free; stamens 10(-12) in 2 whorls of 5(-6), filaments adnate to the corolla at 

base, anthers dorsifixed, opening by apical valves; ovary superior, 3-celled, with axile 

placentation; ovules many, small, anatropous; style simple, projecting out of the flower, 

sometimes divided into three apical lobes, each lobe stigmatic at the top. Fruit a capsule, 3- 

loculed, enclosed by the persistent calyx. Seeds many, small; endosperm fleshy; embryo 

cylindrical. 

Taxonomy. A monograph of the genus has been published by Sleumer I.c. (1967). There 

are 10 species recorded for South East Asia. Only 3 species occur in Sabah and Sarawak; 

they are closely related species differing mainly in the details of their hair and flower size. 

Merrill, EB (1921) 460; Masamune, EPB (1942) 567; Sleumer, FM 1,7 (1971) 139; Whitmore, 

TFM 2 (1973) 27; Anderson, CLTS (1980) 162; Whitmore, Tantra & Sutisna, CLK 1 (1989) 45. 

A small mono generic family in the Ericales of about 65 species distributed in the 

temperate, tropical American and Asiatic-Malesian regions. 

TREE FLORA OF SABAH AND SARA W AK

1mm 

Nooteboom & Chai 2267.) 

4cm 

8 

1cm 

182 

TREE FLORA OF SABAH AND SARAW AK VOL. I (1995) 

Fig. 1. Clethra pachyphylla. A, fruiting leafy twig; B, detail of lower leaf surface; C, flower; D 

longitudinal section through flower; E, fruit; F, seeds. (All from SAN 65004 except C & D, fror

Leaf lower surface with stellate hairs on the midrib and veins, and a fine pale tomentum 

covering the entire surface of the intervenium. Calyx-lobes 2 mm or longer ................. 2 

2. Raceme slender (1-1.5 mm thick), axis covered by a mixture of large and tiny stellate 

hairs (the hairs shorter than 0.5 mm). Calyx-lobes 2-3 mm long ....... 2. C. longispicata 

Raceme more robust (1.5-2 mm thick), axis covered by a mixture of tufts of long hairs 

(typically 0.5-1 mm long) and tiny stellate hairs. Calyx-lobes 3-4 mm long ................. . 

..... 3. C. pachyphyIla 

1. Clethra canescens Reinw. ex Blume 

(Latin, canescens = turning grey; the pubescent leaves) 

Bijdr. (1826) 863; Sleumer I.e. (1967) 85, I.e. (1971) 145; Anderson I.e. 162; Whitmore, Tantra & 

Sutisna I.e. 46. Type: Reinwardt, s.n., 1821, Celebes, Minahasa, Mt. Klabat (L). 

183 

Taxanomy. Sleumer (1967 & 1971) recognises 5 varieties, viz. var. canescens, clementis, 

ledermannii, luzonica, and novoguinensis, of which only var. clementis occurs in Sabah 


Ecology. In montane forest, rarely in lowland forest. 

Distribution. Endemic to Borneo (Sabah, Sarawak and Kalimantan). In Sabah, var. clementis 

is locally common on Mt. Kinabalu; in Sarawak it has been recorded from Mt. Dulit 

(Richards 1772 & 1804) and the Baram River (Haviland 1828). 

Small tree to 10 m. Bark smooth, pale brown; inner bark pale yellow. Sapwood yellowish. 

Leaves chartaceous to thinly coriaceous, upper surface smooth and shiny, lower surface 

with stellate hairs only on the midrib and veins, glabrous on the intervenium, oblong to 

elliptic-obovate, 3-13(-15) x 1-3(-5.5) cm; base cuneate, apex acute to acuminate pointed; 

lateral veins 10-14 pairs, looping towards the margin; midrib sunken on upper side, 

glabrous to sparsely covered with stellate hairs and rusty tomentum on lower side; stalk 1- 

l.5(-2.2) cm long, glabrous to sparsely stellate-hairy. Inflorescence axis slender, 1-2 mm 

thick, covered with a fine rusty tomentum of both smaller and larger stellate or fascicled 

hairs, flowers densely arranged along the rachis; bracts subulate, hairy, c. 5 mm long. 

Flowers white, scented; calyx-lobes 1.5-2 mm, covered with stellate and fascicled hairs; 

petals white, spathulate, glabrous, fimbriate, 4-5 mm long; filaments glabrous, 3-3.5 mm 

long; anthers ob ovate, 0.5-0.8 mm long; ovary densely covered with straight hairs; style 

glabrous, 2.5-3.7 mm long. Fruit subglobose, 2-3 mm diameter. Seed irregularly ovoidangular, 

0.6-1 mm across. 

I.e. (1967) 86, I.e. (1971) 145. Basionym: Clethra clementis Merr., Philip. 1. Sc. Bot. 13 (1918) 104, 

I.e. (1921) 460; Masamune I.e. 567; Anderson I.e. 162. Type: Clemens 11148, British North Borneo, 

Mt. Kinabalu (PNH, destroyed). Synonym: Clethra eaneseens (non Reinw. ex Blume) Stapf, Trans. 

Linn. Soc. Bot. 42 (1914) 105. 

var. c1ementis (Merr.) Sleumer 

CLETHRACEAE (BERHAMAN)


CONNARACEAE 

Lesmy Tipot 



Hookerf, Fl. Brit. Ind. 2 (1879) 46; King, J. As. Soc. Beng. 66,2 (1897) 1; Ridley, FMP 1 (1922) 

544; Merrill, EB (1921) 291, PEB (1929) 94; Masamune, EPB (1942) 328; Leenhouts, FM 1, 5 

(1958) 495; Kochummen, TFM 3 (1978) 47; Keng, OFMSP (1978) 211; Anderson, CLST (1980) 

163; Ashton, MNDTS 2 (1988) 223; Whitmore, Tantra & Sutisna, CLK 1 (1989) 49. 

Mostly woody climbers or shrubs, rarely small trees. Leaves alternate or spirally arranged, 

without stipules, pinnate with terminal leaflet (imparipinnate), rarely unifoliolate; base of 

petioles and petiolules swollen; leaflets not always opposite, pinnately veined or 3-veined, 

base often slightly peltate. Inflorescences panicuiate, axillary or terminal, rarely on branches. 

Flowers 4-5-merous, bisexual or unisexual; sepals free or joined at base; petals free; 

stamens free or joined at base, in two whorls, inner (epipetalous) rudimentary; ovary 

superior, usually 5-carpellate (in Connarus and Ellipanthus only l-carpellate), ovules 2 in 

each carpel, orthotropous or anatropous, placentation basal or axillar. Fruits follicular, 

smooth, finely striate, or warty, reddish, stalked, with persistent calyx, splitting ventrally or 

dorsally, rarely circumscise at the base or indehiscent. Seeds one per carpel, with an 

arillode or sarcotesta, with or without endosperm; cotyledons thick, flat. 

Distribution. A pantropical family with 16 genera and c. 300-350 species, predominantly 

African. In Sabah and Sarawak, represented by 6 genera with at least 27 species. 

Ecology. Frequent in open areas, forest fringes and river banks in lowland rain forests. 

Taxonomy. The Connaraceae was placed under the Dilleniales by Hutchinson (Fam. F1. PI. 

1, ed. 2 (1959) 329). Keng (l.c.), however, included the Connaraceae in the Rosales, 

considered a more primitive order than the Dilleniales. The Connaraceae was once thought 

to be the link between the Rosales and Leguminosae but many recent workers believed an 

affinity to the Sapindaceae is more natural. Some species of the Connaraceae are often 

confused with those of Leguminosae, especially Derris. However, the absence of stipules in 

the former distinguishes them from the latter. 

Uses. Generally of no commercial value. Only the wood of Ellipanthus species is sometimes 

used locally for house posts and bridges. Some species are said to have medicinal properties. 

For instance, the roots of Agelaea macrophylla or akar malam, when boiled together with 

roots of tapang (Koompasia malaccensis), are used traditionally to treat weakness in 

infants, while young leaves of Rourea mimosoides arc used for treating wounds and boils. 

187


Key to genera 

1. Trees or treelets .......................................................................................................... 2 

Woody climbers or shrubs ............................................................................................ 3 

2. Leaves trifoliolate or pinnate .......... ........ 1. Connarus (in part) 

Leaves unifoliolate ............................. 

............ 2. Ellipanthus 

3. Leaves trifoliolate, upper surface ofleaflets with minute pits. Fruits usually warty ......... 

Agelaea Sol. ex Planch. 

Linnaea 23 (1850) 437; Ridley I.e. 552; Merrill I.e. (1929) 95; Leenhouts I.e. 500; 

Kochummen I.e. 48. Synonym: Cantanola Llanos, Mem. R. Ac. Cienc. Madr. 3, 2 (1859) 

505, Masamune I.e. 328. 

50 species, Mrica, Madagascar and Malesia; 4 species in Sabah and Sarawak. 

Woody climbers, rarely scrambling shrubs. Flowers bisexual, 5-merous; petals 

much longer than sepals; stamens usually 10. Fruit papillose or rugulose to warty. 

Seeds shiny black, partly covered with orange or yellow arillode. 

Mainly in lowland primary and secondary forests. A. borneensis is widely distributed 

in Sabah and Sarawak; A. insignis is very uncommon and in Sarawak is represented 

by a few collections from Baram and Kuching areas but in Sabah is recorded only 

from Tawau; A. macrophylla is endemic to Sarawak, whereas A. trinervis is endemic 

to Sabah. 

Leaves imparipinnate, upper surface ofleaflets not pitted. Fruits smooth .................... .4 

4. 

Leaflets with emarginate apex ................... . 

Roureopsis Planch. 

I.e. 423; Masamune I.e. 331; Leenhouts I.e. 505; Kochummen I.e. 52. Synonym: Taenioehlaena 

Hook.! in Bentham & Hooker!, Gen. PI. 1 (1862) 433, Masamune I.e. 332. 

10 species, West Mrica, Northern Burma, Southern China, Sumatra, Peninsular 

Malaysia, Java, and Borneo; 2 species in Sabah and Sarawak, viz. R. acutipetala 

and R. emarginata. 

Woody climbers, sometimes scandent shrubs. Inflorescences axillary, racemose or 

paniculate; bracts lanceolate, densely appressed hairy. Flowers bisexual, longstalked, 

usually 5-merous; petals linear; stamens usually 10. Fruits obliqueellipsoid, 

opening by a ventral slit, red, with persistent calyx. Seeds ellipsoid, 

hilum partly or entirely covered by a yellow fleshy arilloid. 

Apex ofleaflets not emarginate .. 

5. Fruits velvety hairy, pear-shaped. Leaflets usually more than 10 pairs ........... . 

Cnestis Juss. 

Gen. (1789) 374; Ridley I.e. 553; Merrilll.e. (1929) 96; Masamune I.e. 329; Leenhouts I.e. 

497; Kochummen I.e. 48. 

About 40 species, predominantly tropical Mrica and Madagascar. Two species in 


Woody climbers or scandent shrubs. Inflorescenses axillary or in fascicles from 

knobs on branches. Flowers 5-merous; stamens 10; carpels 5. Fruits velvety brown, 

188

Fruits glabrous, finely striate, pod-like or ellipsoid to ovoid. Leaflets less than 10 pairs 

6. Carpel only one per flower; calyx in fruit accrescent and hard. Fruit ellipsoid, base not 

stipitate .......................... . 

Rourea AubI. 

Hist. PI. Guiane 1 (1775) 467, t. 187; Ridley I.e. 549; Merrilll.e. (1929) 95; Leenhouts I.e. 

510; Kochununen I.e. 51. Synonym: Santaloides 1. ex Kuntze, Rev. Gen. PI. 1 (1891) 155 

(as Santalodes), Merrilll.e. (1921) 291, Masamune I.e. 331. 

.6 

1. CONNARUS L. 

(Greek, konaros, a plant name) 

Sp. PI. 2 (1753) 675, Gen. PI. ed. 5 (1754) 305; Bentham & Hooker f I.e. (1862) 432; King I.e. 

(1897) 2; Ridley I.e. 544; MerrillI.e. (1921) 291, I.e. (1929) 94; Masamune I.e. 329; Leenhouts I.e. 

525; Kochununen I.e. 49; Anderson I.e. 163; Ashton I.e. 223. 

Usually woody climbers, rarely shrubs or small trees. Leaves imparipinnate, same times 

trifa lia late, rarely unifoliolate; leaflets almost always pellucid-glandular punctate. Inflorescences 

terminal or axillary rarely on the branches or stems. Flowers bisexual, fragrant, 5- 

meraus; sepals, petals, stamens punctate, seen as dark gland-dots in herbarium materials; 

sepals thick and fleshy, not accrescent in fruit; petals nearly always hairy; stamens 10, 

joined at base; ovary l-carpellate, stigma capitate. Fruits pod-like, opening lengthwise, 

base narrowed into a stipe, with small persistent calyx, apex short-beaked or curved. Seed 

1, shiny black, basal part partially enveloped by a fleshy, yellow arillode. 

Distribution. About 100 species, pantropical. 13 species in Sabah and Sarawak, including 

two unnamed taxa (indicated as Connarus sp. A and Connarus sp. B in the key given 

below). Of these only one species, C. agamae, is a small tree. 

189 

About 100 species; Central and South America, Malesia, NE Australia and 

Melanesia; 4 species in Sabah and Sarawak. 

Woody climbers, shrubs, very rarely small trees. Leaves very variable in size and 

shape. Inflorescences axillary. Flowers 5-merous; sepals ovate, imbricate, margin 

ciliate; petals twice or more longer than sepals; stamens 10, joined at base. Fruits 

usually 1 per flower, ellipsoid to ovoid, splitting ventrally. Seeds with the testa 

partly or almost entirely fleshy or seeds enveloped by fleshy arillode. 

Mainly found in low altitude, in primary and secondary mixed dipterocarp 

rainforests, along forest edges, river banks, and roadsides. 

Carpels more than one per flower. Calyx in fruit not accrescent. Fruit pod-like, base 

narrowly stipitate ......................................................................... 1. Connarus (in part) 

CONNARACEAE (LESMY) 

pear-shaped, StIpltate. Seeds flattened bean-shaped, partially covered by a 

sarcotesta at the base. Found mainly in a primary and secondary mixed dipterocarp 

forests to 500 m.

Connarus sp. B. 

A climber. Leaflets 1-2 pairs, broadly ovate. Inflorescences terminal. 

One collection (s. 44912) from kerangas forest in Bukit Sadok, Ulu Sekrang, Sri 

Aman, Sarawak. 

Leaflet attachment not peltate; lateral veins less than 10 pairs, usually curved; petiolules 

much shorter ................................................................................................................ 6 

6. Leaflets drying reddish brown. Fruits obovoid; pericarp coarse, thick and woody, c. ~ 

mm thick. .................................................................................................................... . 

C. grandis Jack 

Mal. Mise. 2, 7 (1822) 40; Masamune I.e. 329; Leenhouts I.e. 529. Synonym: Connaru 

e/liptieus King I.e. 7, Merrilll.e. (1929) 95. 

Woody climber, rarely shrub. Branches glabrous. Leaflets 1-2 pairs, obovate t 

lanceolate. Inflorescences usually densely fulvous-tomentose. 

190 

. .. 3 

Leaflets hairy at least on the midrib below or at young stage ...... . . ............ 9 

4. Leaflets glabrous ...... , ..... . . ................... 5 

A climber. Leaves-trifoliolate, blade distinctly swollen at base, leatherly. Fruits 

borne on branches, smooth, drying chestnut brown. Seed I, arillode covering 

mostly the lower part. 

Mixed dipterocarp forest at 740 m. Collected once (s. 36403) from Bt. Goram, 

Song, Kapit District, Sarawak. 

Leaflets narrowly lanceolate, apex distinctly emarginate; veins not raised below, intercostal 

veins not scalariform .... 

Usually shrub, rarely small tree reaching 5 m tall and 5 cm diameter. Inflorescences 

on short condensed panicles, peduncles stout. Fruits obliquely obovoid, glabrous, 

stipitate, sparsely tomentose, slightly compressed, bright orange-red, drying brown, 

apex rounded. 

Primary mixed dipterocarp forest; collected several times from Bukit Raya, Kapit 

District, Sarawak. 

. ................................. 2 

. .... 4 

. ............... Connarus agamae 

Key to Connarus species 

Inflorescences terminal or almost so on leafY shoots ... 

Climbers, scandent or erect shrubs ...................... . 

3. Leaflets oblong-elliptic, apex shortly acuminate; all veins distinctly raised below, intercostal 

veins scalariform ..................................... . 

I. Small trees ........................................................ . 

2. Inflorescences on stems or on leafless twigs ..... . 


C. impressinervis B. C. Stone 

Malays. For. 43,2 (1980) 255. 

Connarus sp. A. 

5. Leaflet attachment distinctly peltate; lateral veins 16-18 pairs, straight and close together; 

petiolule 2.5-4 cm long .................................................................................... .

Primary and secondary mixed dipterocarp forests, also in open areas and along 

river banks. Distributed widely in Sabah and Sarawak, fairly common. 

Leaflets not drying as above. Fruits ellipsoid or spindle-shaped; pericarp thin, not 

coarse .......................................................................................................................... 7 

7. Leaflets drying pale green and purplish brown on midrib and lateral veins. Fruits not 

stipitate ......................................................................................................................... . 

C. lucens Schellenb. 

Bot. Jahrb. 59 (1924) 36; Leenhouts I.e. 539. 

Glabrous climber. Leaflets 3-4 pairs, obovate to oblong-lanceolate; lateral veins 

drying purplish brown. Inflorescences broadly branched. Fruits trapezoid to 

oblique-ellipsoid, slightly flattened, not stipitate, drying purplish brown. 

Recorded from Saribas and Kapit, Sarawak. 

Leaflets not drying as above. Fruit stipitate ......................................... . . ......... 8 

191 

9. Leaflet attachment usually subpeltate, lateral veins 10-18 pairs, straight to slightly 

curved at margin ........................................................................................................... . 

C. euphlebius Merr. 

J. Str. Br. R. As. Soc. 85 (1922) 200, I.e. (1929) 94; Masamune I.e. 329; Leenhouts I.e. 

527. 

Woody climber to 25 m high, sometimes a scandent shrub. Branchlets, petioles, 

rachises and petiolules densely ferruginous hairy. Leaflets 1-4 pairs, petiolules 2- 

5 mm long. Inflorescences densely fcrruginous-tomentose. Fruits obovoid and 

rather flattened, shortly stipitate; pericarp densely minutely ferruginous-tomentose. 

Generally lowland forests. Uncommon in Sabah and Sarawak, represented by only 

three collections. 

Fruits ellipsoid, more or less bulging, stipe 0.5-1 cm long. Reticulation of intercostal 

veins not conspicuous ................................................................................................... . 

C. winkleri Schellenb. 

I.e. (1924) 38; Masamune I.e. 330; Leenhouts I.e. 540. Synonym: C. paehyphyllus Merr. 

I.e. (1918) 71, I.e. (1921) 292, Masamune I.e. 330. 

Woody climber, to 20 m high. Leaflets 2-3 pairs. Inflorescences to 5 cm long, 

laxly branched. Sepals ovate; petals ovate-lanceolate, thinly tomentose on both 

surfaces. 

Generally lowland forests. Few collections from Sabah, and only one from Sarawak. 

8. Fruits obliquely spindle-shaped, shortly stipitate. Reticulation of intercostal veins 

conspicuous ................................................ . 

C. monocarpus L. 

Sp. PI. (1753) 675; Leenhouts I.e. 538 (ssp. malayensis). Synonyms: C. Jaleatus Blume, 

Mus. Bot. Lugd. Bat. 1 (1850) 266, Masamune I.e. 329; C. densiflorus Merr., Philip. J. Sc. 

13 (1918) Bot. 70, Merrilll.e. (1921) 292, I.e. (1929) 95. 

Woody climber. Leaflets 2-4 pairs. Inflorescences many-flowered. Sepals usually 

distinctly keeled, pubescent outside, glabrous inside. 

Uncommon; in Sabah once found along the seashore in Lahad Datu; in Sarawak, 

collected from forest edges and river banks, and once on limestone. 

CONNARACEAE (LESMY)

I.e. 39; Leenhouts I.e. 534. Synonyms: C. mutabilis B1ume I.e. 269; C. borneensis Merr. I.e. 

I.e. 329. 

192 

- - -- - -- -- ------ 

Philip. J. Sc. 4 (1909) Bot. 120; Leenhouts I.e. 53l. Synonym: C. stellatus Merr. I.e. (1909) 119, 


Woody climber or scandent shrub. Branchlets densely ferruginous-tomentose, later 

glabrous. Leaflets 2-5 pairs, tomentose. 

Usually occurs in open areas in the lowland mixed dipterocarp forests. 

12. Fruits oblique-ellipsoid, shortly stipitate; hairs branched, usually stellate ...................... . 

Woody climber or scandent shrub. Branches, leaves and inflorescences densely 

pubescent. Leaflets 1-4 pairs. Inflorescences 10-30 cm long, few-flowered. Fruits 

ellipsoid, not stipitate; pericarp rather thin, outside densely orange-brown 

pubescent, inside glabrous. 

Primary mixed dipterocarp and kerangas forests at low altitudes. Found mainly in 

the Kuching and Samarahan Divisions in Sarawak but only one record from 

Sabah. 

Leaflets not so. Fruit compressed. . .... 12 

11. Leaflets densely ferruginous-pubescent beneath. Fruits not compressed ......................... . 

Climber or scandent shrub. Branches, leaves, petioles, rachis and petiolules 

densely tomentose when young, later glabrescent. Inflorescences to 20 cm long, 

widely branched and densely tomentose. Fruits oblique-ellipsoid, glabrous; pericarp 

thin, minutely wrinkled outside, densely pubescent inside. 

Lowland forests and thickets. Widespread in Sabah and Sarawak. 

Leaflets equal in size. Fruits thinly or densely ferruginous-tomentose ........................ 11 

I.e. 38; Masamune I.e. -230; Leenhouts l.e 531. Synonym: C. plumoso-stellatus Merr. I.e. 

Leaflet attachment not subpeltate, lateral veins less than 10 pairs, curved and looped at 

margin ....................................................................................................................... 10 

Trans. Linn. Soc. 23 (1860) 72; Merrilll.e. (1921) 292, I.e. (1929) 95; Leenhouts I.e. 533. 

Synonym: C. hebephyllus King I.e. (1897) 5, Merrilll.e. (1921) 292. 

10. Leaflets progressively increasing in size from basal pair upwards. Fruits glabrous ........ . 


(1918) 72, I.e. (1921) 292. 

C. cullionensis Merr. 

C. odoratus Hook. f 

C. villosus Jack 

Fruits oblique-pyriform, stipitate, stipe to 1.5 cm long; hairs simple, not stellate ........... . 

C. semidecandrus Jack 

(1918) 69, I.e. (1921) 292, Masamune I.e. 329; C. Jaekiana Schellenb. I.e. 40, Masamune 

Large woody climber or scandent shrub. Leaflets 1-3 pairs, sometimes 5 pairs. 

Inflorescences terminal, to 35 cm long, broadly branched, many-flowered, minutely 

tomentose. Fruits compressed; pericarp thin, minutely ferruginous-pubescent, 

glabrescent outside, inside densely pubescent. 

Primary and secondary mixed dipterocarp forests especially in open areas along 

forest edges and river banks, also in clearings along beaches and sometimes in 

swampy habitats, on granite and limestone. Widely distributed in Sabah and 

Sarawak.


'om [ 

8 

Fig. 1. Connarus agamae. A, flowering leafy twig; B, flower, C, flower with one sepal and one petal 

removed; D, fruit. (A-C from SAN 16882, D from SAN 57255.) 

193


B 

[,mm 

A 

2cm 

Fig. 2. Ellipanthus tomentosus. A, fruiting leafy twig; B, flower. CA from S. 42098, B from SAN 

50543.) 

194


Connarus agamae Merr. Fig. 1. 

(Jose Agama, Deputy Conservator of Forests, British North Borneo, 1889-1982) 

I.c. (1918) 68, I.c. (1921) 291, I.c. (1929) 94; Masamune I.c. 329; Leenhouts I.c. 527. Type: Agama 

422, British North Borneo, Tawau (holotype K). 

Small tree, rarely reaching 20 m high and 20 cm diameter; sometimes shrub, rarely climber. 

Twigs thinly pubescent, glabrescent. Leaves with 1-2 pairs of leaflets, sometimes un i 

Joliolate especially below the inflorescence; leaflets oblong, slightly oblique, 11-20 x 4-10 

cm, terminal ones ovate, thinly papery, glabrous above, drying reddish brown, densely 

Jerruginous-pilose beneath especially on midrib; base rounded to acute, apex blunt; lateral 

veins 8-15 pairs, straight, strongly looped and closed toward the margin but not joining; 

intercostal veins reticulate, inconspicuous above. Inflorescences densely ferruginouspubescent. 

Flowers: sepals ovate, brown-pilose on both sides; petals linear, punctate on 

both sides; stamens all fertile, connate at base. Fruits obovate, 5 x 2-3.5 cm, beak acute at 

4/5 of the height; pericarp glabrous, shiny and slightly wrinkled, woody; densely fulvoustomentose 

inside; stipe 1-1.5 cm long. 

Distribution. Endemic to Borneo. In Sabah, recorded from Sandakan and Mostyn, but not 

yet recorded from Sarawak. 

Ecology. In primary mixed dipterocarp forest to 300 m. 

2. ELLIPANTHUS Hook.! 

(Greek, ellipes = defective; anthos = flower; 

referring to the defective or imperfect development of 5 of the 10 stamens) 

I.c. (1862) 434, I.c. (1879) 55; King I.c. 8; Ridley I.c. 548; MerrillI.c. (1921) 291, I.c. (1929) 96; 

Masamune I.c. 330; Leenhouts I.c. 520; Kochummen l.c. 50; Anderson I.c. 163; Ashton I.c. 224; 

Whitmore, Tantra & Sutisna I.c. 49. Synonym: Pseudellipanthus SchelIenb. in Mez, Bot. Arch. 1 

(1922) 314, Masamune I.c. 331. 

Shrubs or small trees reaching 25 m tall and 25 cm diameter. Twigs tomentose, at least 

when young. Leaves alternate, unifoliolate, with a prominent joint at the base. Inflorescences 

axillary, paniculate or clustered; bracts caducous, lanceolate, small. Flowers 4-5-merous, 

protandrous, unisexual (and then plants dioecious) or bisexual; sepals densely hairy outside, 

valvate in bud; petals free, imbricate in bud; stamens twice as many as petals, connate 

at base, episepalous ones well-developed, epipetalous ones rudimentary; ovary I-carpellate, 

flattened ovoid, style slender, stigmas disk-shaped to bilobed. Fruits densely tomentose, 

apex shortly pointed, base stipitate; pericarp woody; persistent calyx not accrescent. Seed 

one, ellipsoid, shiny black, covered by a yellowish orange arillode at base. 

Distribution. About 10 species; Africa, Madagascar, Sri Lanka, continental SE Asia and 

Malesia. 2 species in Sabah and Sarawak. 

Ecology. Lowland mixed dipterocarp forest and mixed swamp forest. 

195


Key to Ellipanthus species 

Flowers predominantly 4-merous, unisexual. Leaf-blade usually glabrous above, densely 

fermginous-tomentose beneath, base rounded ................................................. 1. E. beccarii 

Flowers predominantly 5-merous, usually bisexual. Leaf-blade usually tomentose above (at 

least on the midrib), glabrous or thinly fulvous-tomentose beneath, base cuneate or subcordate 

....................................................................................................... 2. E. tomentosus 

1. Ellipanthus beccarii Pierre 


Fl. eoch. 5 (1898) t. 378; Merrilll.e. (1921) 291; Leenhouts I.e. 524; Anderson I.e. 164; Ashton I.e. 

225; Whitrnore, Tantra & Sutisna I.e. 49. Type: Beeeari PB 296, Sarawak, Matang (K). 

Small dioecious tree or shmb, rarely reaching 10 m tall, 15 cm diameter. Leaves elliptic to 

ovate, sometimes lanceolate, glabrous above, densely ferruginous-tomentose beneath, 7-15 

x 3-5 cm; base rounded, peitate or not, apex acuminate; lateral veins 8-10 pairs, curved, 

looped and joined; petiole jointed at leaf-base, c. 1 cm long. Flowers unisexual, 4-merous. 

Fruits shortly stipitate, stipe to 1 cm long. Seed with a small cupular arilloid. 


Leaf-base not peltate ...................................... . 

var. beccarii Leenh. 

I.e. 524; Anderson I.e. 164; Whitrnore, Tantra & Sutisna I.e. 49. Synonyms: E. mindanaensis (non 

Merr.) Merr., 1. Str. Br. R. As. Soc. 76 (1917) 84, I.e. (1921) 291, I.e. (1929) 96; Pseudellipanthus 

beeearii (Pierre) Schellenb. I.e. (1922) 314; Diehapetalum tetramerum Ridl., Kew Bull. (1938) 

234. 

Known only in the Kuching area, Sarawak on leached yellow soils in mixed dipterocarp 

forest, and Kalimantan. 

Leaf-base peltate ..... . 

var. peitatus (Schellenb.) Leenh. 

I.e. 524; Anderson I.e. 164; Whitmore, Tantra & Sutisna I.e. 49. Basionym: Pseudellipanthus 

peltatus Schellenb. I.e. (1922) 314. 

Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan). In Sabah uncommon; in 

Sarawak frequent in mixed dipterocarp forest on fertile clay soils, particularly on basic 

volcanic rocks. 

Vernacular names. Sarawak--kelana (Kenyah), merinang (lban). 

2. Ellipanthus tomentosus Kurz Fig. 2. 

(Latin, tomentosus = thickly and evenly covered with short matted hairs; the leaves) 

J. As. Soc. Beng. 41, 2 (1872) 305; Hooker f I.e. 56; Leenhouts I.e. 521; Kochummen I.e. 50; 

Anderson I.e. 164; Ashton I.e. 225; Whitmore, Tantra & Sutisna I.c. 49. Type: Wallich Cat. 8551, 

Lower Burma, Moulmain (holotype K). 

196


subsp. tomentosus var. luzoniensis (Vidal) Leenh. 

I.e. 524; Whitmore, Tantra & Sutisna I.e. 49. Basionym: E. luzoniensis Vidal, Rev. PI. Vasc. Filip. 

(1886) 104. Synonyms: Connarus urdanetensis Elmer, Leafl. Philip. Bot. 7 (1915) 2594; E. 

burebidensis Elmer I.e. 2596; E. vidalii Elmer I.e. 2596; E. longifolius Merr., Philip. 1. Sc. 17 (1921) 

Bot. 262; E. urdanetensis (Elmer) Merr., En. Philip. 2 (1923) 241; E. sarawakensis Schellenb., Pfl. 

R. Heft 103 (1938) 185. 

Shrub or tree to 20 m tall, 25 cm diameter. Bark smooth, greyish brown; inner bark 

reddish brown. Sapwood pale yellow. Leaflets elliptic to lanceolate, 7-20 x 4-9 cm, 

chartaceous, glabrous or thinly fulvous-tomentose below; base cuneate, sometimes rounded 

or subcordate, apex acuminate; midrib tomentose above; lateral veins 5-7 pairs, looping 

near margin; intercostal veins conspicuous below, very faint above; petiole jointed at base, 

0.5-3 cm long. Flowers white, bisexual, 5-merous. Fruits stipitate, stipe c. 3.5 cm long, 

apex pointed. Seeds c. 5 mm long, covered with faintly lobed arilloid. 

Vernacular name. Sarawak--kelin (Melanau). 

Distribution. Borneo, the Philippines and Central Celebes. In Sarawak uncommon, distributed 

around Miri, Bintulu, Sibu, Sarikei and Kuching. In Sabah, recorded from Lahad Datu and 

Tawau. 

Ecology. Locally frequent in primary mixed dipterocarp forest and peat swamp forest. 

Uses. The wood is considerably hard and durable, used for local construction works such as 

bridges and house posts. 

197


CORNACEAE 

RC.K Chung 



Rid1ey, Fl\.1P 1 (1922) 889 (excluding Alangium, Aralidium and Nyssa); Merrill, EB (1921) 459, 

PEB (1929) 233; Danser, Blumea 1 (1934) 46; Backer & BakhuizenJ, FJ 2 (1965) 158; Matthew, 

Blumea 23 (1976) 51, FM 1, 8 (1977) 85; Kochummen, TFM 3 (1978) 53; Anderson, CLTS (1980) 

164; Whitmore, Tantra & Sutisna, CLK 1 (1989) 49. 

Trees, shrubs, or rarely herbs. Leaves opposite, subopposite or alternate, simple, entire or 

rarely serrate, pinnately veined, exstipulate, usually petiolate. Flowers regular (actinomorphic), 

4-5-merous, bisexual or sometimes unisexual, arranged in axillary or terminal 

cymes, panicles or heads; calyx-tube adnate to the ovary, 4-5-lobed or subtruncate, persistent in 

fruit; petals 4-5, free, valvate or imbricate in buds; stamens short and equal in number with 

the petals, 2-locular, dehiscing lengthwise; disc large, cushion-shaped at the top of the 

ovary, generally persistent in fruit; ovary inferior, 1-4-locular, ovule 1 in each locule, 

pendulous, with dorsal or ventral (in Curtisia and Mastixia) raphe, style simple or lobed. 

Fruit usually a drupe, rarely a berry (in Aucuba and Griselinia). Seeds usually 1-2, rarely 

4, with small elongate embryo embedded in copious oily endosperm. 

\ 

Distribution. About 8-15 genera and 100 species; mainly in the temperate zones and in the 

tropics (rare in Mrica and South America). In Sabah and Sarawak, represented by the 

genus Mastixia with 7 species. 

" 

Ecology. Insect-pollination is probably a general phenomenon in the family, except in 

Griselinia litoralis which has been reported to be wind-pollinated. 

Uses. In Sabah and Sarawak, very little is known about the utilization of members of the 

family. However, the fruit of Chamaepericlymenum suecicum is eaten by the Eskimos. In 

France, Corn us mas (Cornelian cherry) is grown as ornamentals, and the fruit is edible and 

used as a source of raw material for making "Vin de Cornoulle". The Assegay tree or Cape 

Lance wood (Curtisiafaginea) produces durable, red to brown timber and used for making 

furnitures in South Mrica. Aucuba japonica is a popular evergreen garden shrub widely 

cultivated in temperate and subtropical countries. 

Taxonomy. The taxonomic and systematic status of the Cornaceae remains controversial 

(Matthew I.c. 1976, l.c. 1977; Mabberley, The Plant Book (1987) 146; and Brummitt, 

Vascular Plant Families & Genera (1992) 543). In the Malesian region, Ridley (l.c.) and 

Keng (OFMSP, 1969) included Alangium, Aralidium, Mastixia and Nyssa in the 

Cornaceae, whereas van Steenis (Checklist Gen. Names Mal. Bot. (1987) 50) placed 

Alangium in its own family (Alangiaceae), Aralidium in the Araliaceae, and Nyssa in the 

Nyssaceae, leaving lviastixia as the sole representative of the family in the Malesian region. 

199

TREE FLORA OF SABAII AND SARAWAK VOL. 1 (1995) 

MASTIXIA Blume 

(Greek, mastix = whip; the whiplike apex ofthe petals) 

Bijdr. (1825) 654, Mus. Bot. Lugd. Bat. I (1850) 256; Bentham & Hooker J, Gen. PI. (1887) 950; 

Harms in EngI. & Prantl, Pfl. Fam. 3 (1898) 262; Wangerin in EngI., Pfl. Reich. 4, 229, 41 (1910) 

19; Ridley I.c. 889; Merrilll.c. (1921) 459, I.c. (1929) 233; Danser I.c. 47; Backer & BakhuizenJ 

I.c. 159; Matthew l.c. (1976) 51, l.c. (1977) 85; Kochummen l.c. 53; Whitmore, Tantra & Sutisna 

I.c. 49; Ng, MFFSS I (1991) 47. 

Trees, usually without buttresses. Bark grey to grey-brown, smooth with horizontal rings, 

rarely cracked to shallowly fissured, often exuding white resin when bruished; inner bark 

orange-yellow, gritty, granular, with strong sme!t of sugarcane water. Sapwood soft, 

yellowish white or white. Leaves alternate, sub opposite or opposite, margin entire; midrib 

sunken on adaxial surface, prominent below; lateral veins usually distinct on the abaxial 

surface. Inflorescences terminal or sometimes axillary cymose-panicles. Flowers bisexual, 

in triads, sessile, subtended by tiny, persistent bracts; calyx-tube obconical or barrel-shaped 

or cup-shaped, lobes spreading or not, broader than long or sometimes appear as minute 

sharp tips, persistent in fruit; petals thick, concave, valvate in bud, inflexed and strongly 

connate at the upper parts; stamens attached below the disc, abutting on and alternating 

with disc-lobes, opposite the calyx-lobes, filaments subulate, flattened and tapered toward 

its upper part, anthers cordate, dorsifixed, introrse-Iatrorse; ovary turbinate, one-locular, 

surmounted by a fleshy, lobed and grooved disc which is sometimes persistent in fruit; style 

very short, stout, ribbed; stigma punctiform, sometimes bifid or 4-5-lobed, reflexed, some 

persistent in fruit. Fruit a drupe, ovoid, ellipsoid or oblong, surmounted by calyx-tube and 

crowned by the persistent disc (the exposed part of the fruit); pericarp (exocarp and 

mesocarp) thin or thick, turning to dark purple or blue when ripe; fruit-wall formed by 

calyx-tube and pericarp; endocarp stony. Seed 1, ovoid or ellipsoid, with membranous 

testa; endosperm large and U-shaped in transverse section; embryo small and straight, 

cotyledons thin and foliaceous; germination epigeal. 

Distribution. About 13 species; from the western Ghats and Sri Lanka, NE India, Bhutan, 

Burma, Thailand, Indo-China, S Yunnan, Hainan, through Malesia to New Britain and the 

Solomon Islands. 7 species (including 2 endemic) are found in Sabah and Sarawak. 

Ecology. Found mainly in valleys, slopes or ridges of primary and secondary mixed dipterocarp 

forests, often in moist habitats, from sea-level to 2200 m. 

Uses. Although the trees may reach a considerable size, the scattered occurrence does not 

contribute to their general use as timber. In addition the timber has little commercial value 

and therefore only used for packing cases and temporary constructions. 

Taxonomy. Mastixia is strictly a SE Asian genus. It was included in the Cornaceae by 

Bentham & Hooker.f (I.c.) and Hutchinson (Genera of Flowering Plants 2 (1967) 42), 

whereas Harms (l.c.) and Wangerin (I.c.) included it in a distinct subfamily. Wangerin 

distinguished two subgenera, viz. Tetramastixia and Pentamastixia. Matthew (1976 & 

1977), on the other hand, established two subgenera, Manglesia (2 species) and Mastixia 

(11 species). He recognised two series, the Oppositae and Alternae within the sub genus 

Mastixia based on a single character, viz. whether the first branches of the inflorescence are 

200

CORNACEAE (CHUNG) 

opposite (or subopposite) or alternate. In the absence of flower and fruit and due to the 

presence of resin in the bole and on the cut ends of logs, the genus can be easily confused 

with some species of the Dipterocarpaceae that have smooth or cracked bark, such as 

Vatica species. However, in Mastixia the inner bark is thick and gritty with a strong 

sugarcane smell and the wood is soft. Foresters sometimes confuse this genus with those of 

the Lauraceae (medang) because of the gritty inner bark and strong aromatic smell; 

however medang has no resin. 

Key to Mastixia species 

1. Leaf surface velvety hairy below, if glabrescent then midrib hairy ...... . ..2 

Leaf surface glabrous ............................... 

... 3 

2. Leaves alternate. Twigs and stalks distinctly grooved ...................... .4. M. macrocarpa 

Leaves opposite. Twigs and stalks not so ............................................ 7. M. trichotoma 

3. Leaves glaucous below ........ M~~a 

Leaves not glaucous below ...... . ..................................... 4 

4. Leaf-margin wavy when dry; stalks 2-3 cm long ............................... 2. M. eugenioides 

Leaf-margin not wavy when dry; stalks shorter than 2 cm ............................................ 5 

5. Leaves thickly leathery; margin curled inwards. Twigs dark brown to black when dry 

....... 5. M. pentandra 

Leaves thinly leathery; margin not curled inwards. Twigs grey or grey-brown when dry 

.................................................................................................................................... 6 

6. Leaf lateral veins looping near margin. Flower buds up to 3 mm diameter; petals 

densely hairy on the abaxial surface ...................................................... 1. M. cuspidata 

Leaf lateral veins not looping near margin. Flower buds up to 2 mm diameter; petals 

glabrous on the abaxial surface ............................................................... 6. M. rostdta 

1. Mastixia cuspidata Blume 

(Latin, cuspidatus = sharp-pointed; the leaf apex) 

I.e. (1850) 256; Miquel, Fl. Ind. Bat. I, 1 (1855) 772; Ridley I.e. 891; Merrilll.e. (1921) 459; Danser 

I.e. 55 (excl. var. margarethae); Matthew I.e. (1976) 79, I.e. (1977) 95; Kochummen I.e. 54; 

Whitmore, Tantra & Sutisna I.e 49. Type: Korthals. s.n., Sumatra, West Coast (lectotype L; 

iso1ectotype U). Synonyms: Mastixia pentandra Blume var. euspidata (Blume) Miq. I.e. (1858) 

1095; Mastixia braeteata Clarke in Hooker f. Fl. Brit. Ind. 2 (1879) 746. 

Tree to 40 m tall, and 40 cm diameter. Bark greyish to chocolate-brown, smooth to 

shallowly fissured; inner bark yellowish to brownish, mottled. Sapwood yellowish to 

brownish. Twigs subglabrous, grey-brown. Leaves alternate, sub opposite or opposite, stalk 

slender. 0.5-1.5 cm long; blades narrowly obovate, elliptic or oblong, 3.5-9 x 1.5-3.5 cm, 

thinly leathery, glabrous; base "Cuneate, apex cuspidate with a tip 0.5-1(-1.5) cm long, 

201

TREE FLORA OF SABAH AND SARAW AI< VOL. I (1995) 

oblique; lateral veins (4-)5 pairs, curving near the margin and joining with next one to 

form looped intramarginal veins, sunken above; intercostal veins faint to inconspicuous 

below, inconspicuous above. Inflorescences to 4 cm long, subglabrous to puberulous. 

Flowers 5-merous, green to yellow, buds to 3 mm diameter; calyx-tube densely silky-hairy, 

with 5 lobes, broader than long, subglabrous; petals 1.5-2 x 0.5-1 mm, densely silky-hairy 

on abaxial surface; stamens 5, filament 1-3.2 mm long; disc yellowish. Fruits oblong, 1.5- 

3 x 0.5-1.3 cm; fruit-wall thin; persistent disc well-exposed,; persistent calyx-lobes 

inconspicuous. Seeds ellipsoid, 1.5-2.5 x 0.4-1 cm. 

Vernacular name. Sarawak-biansu gunong (lban). 

Distribution. Sumatra, Peninsular Malaysia and throughout Borneo. Uncommon in Sabah 

and Sarawak. In Sabah, recorded from Sipitang, Sandakan, Lahad Datu, and Tawau. In 

Sarawak, known from Mt. Matang (lst Div.), Sri Aman (2nd Div.), Mt. Dulit near Long 

Kapa (4th Div.), and Belaga (7th Div.). 

Ecology. In primary and secondary mixed dipterocarp forests, to 900 m. Flowering in 

February-March and July-October; and fruiting in January, March, July-August and 

October. 

Sterile specimens of small-leaved M cuspidata are difficult to distinguish from M rostrata 

ssp. caudatifolia. 

2. Mastixia eugenioides Matthew 

(leaves resembling that of Eugenia) 

I.c. (1976) 73, l.c. (1977) 93; Whitmore, Tantra & Sutisna I.c. 50. Type: Beccari PB 2033, Sarawak, 

Mt. Matang (holotype FI; isotype L). 

Tree to 30 m tall and 30 cm diameter. Bark greyish to yellowish brown, smooth; inner 

bark yellowish brown. Twigs glabrous. Leaves opposite, stalk green to greenish yellow, 

stout, 1.5-3 cm long; blades elliptic to oblong-elliptic, 4-15 x 2-5.5 cm, thickly leathery, 

glabrous; base cuneate, margin wavy after drying, apex acuminate to caudate; lateral veins 

5-7 pairs, flattened above; intermediate lateral veins and intercostal veins prominent 

below. Inflorescences to 8 cm long, glabrous. Flowers 4-merous, buds about 1.5-2 mm 

diameter; calyx-tube glabrous, with 4 lobes, broader than long, glabrous; petals 4, 1-1.5 x 

0.5-0.8 mm, glabrous on abaxial surface; stamens 4, filament 0.6-1 mm. Fruits green 

turning to purple when ripe, ovoid (young stage) to oblong, 2-2.5 x 1-1.5 cm; fruit-wall 

thin; persistent disc not well-exposed; persistent calyx-lobes inconspicuous. Seed ellipsoid, 

1.9-2.4 x 0.8-1.1 cm. 

Vernacular name. Sarawak-sempetan (Kenyah). 

Distribution. Endemic to Borneo (Sarawak, Sabah, Brunei, and Kalimantan). In Sarawak, 

recorded from Bukit Senyandang Lingga (2nd Div.), Lambir Hills National Park; Similajau 

FR (4th Div.), Lawas (5th Div.), and Kapit (7th Div.). In Sabah, known from Tenom, 

Sandakan and Tawau. 

202


Ecology. In primary mixed dipterocarp forests, from lowlands up to 1200 m. Flowering in 

July-August, fruiting in September-December. 

3. Mastixia gIauca Matthew 

(Greek, glaukos = bluish-grey; the lower surface of the leaves) 

I.e. (1976) 76, I.e. (1977) 95; Whitmore, Tantra & Sutisna I.e. 50. Type: Hj. Bujang S. 13481, 

Sa~awak, 1st Div., Kuching, G. Santubong East (ho1otype L; isotypes K, SAR). 

Tree to 15 ill tall, 25 cm diameter. Twigs yellowish, glabrous. Leaves alternate, stalk stout, 

2-3.5 cm long; blades obovate, 7-16 x 4-8.5 cm, thickly leathery, glaucous and waxy 

below, glabrous; base obtuse, margin wavy when dry, apex apiculate; lateral veins 4-5 

pairs with intermediate lateral veins faint to inconspicuous; intercostal veins inconspicuous. 

Inflorescences subglabrous to puberulous. Flowers 4(-5)-merous, greenish yellow; 

buds 2.8-3.2 mm diameter; calyx-tube puberulous, with 4(-5) lobes, as long as wide, 

puberulous; petals 4(-5), 1.6-2.1 x 0.6-1.2 mm, hairy on abaxial surface; stamens 4-5, 

filament 0.9-1.2 mm long. Fruits & seeds unknown. 

Distribution. Endemic to Sarawak, and so far only known from Mt. Santubong. 

Ecology. In mixed dipterocarp forest at about 90 m. Flowering in April-May. 

4. Mastixia macrocarpa Matthew 

(Greek, macros = large, carpos = fruit) 

I.e. (1976) 75, I.e. (1977) 94; Whitmore, Tantra & Sutisna I.e. 50. Type: Benang S. 24745, Sarawak, 

Miri, Bakam Road (ho1otype A; isotypes BO, K, KEP, L, MEL, NY, SAN, SAR, SING). 

Tree to 20 m tall, 20 cm diameter. Twigs covered with sticky resin when fresh, rusty brown, 

rough-hairy and distinctly grooved. Leaves alternate, stalk stout, 4-6 cm long, grooved, 

hairy like the twigs; blades elliptic-oblong to oblong, 13-26 x 5.5-13 cm, thinly leathery, 

villous below; base cuneate to asymmetric, apex acute to acuminate; lateral veins 7-10 

pairs, flatten above, prominent below; intercostal veins prominent, scalariform or 

reticulate. Inflorescences to 9 cm long, densely hairy. Flowers 5-merous, densely hairy, 

buds c. 4 mm diameter; calyx-tube densely hairy, with 5 lobes, broader than the length, 

villous; petals 5, 1.8-2.3 x 0.8-1.2 mm, velvety on abaxial surface; stamen 5, filament 1- 

1.5 mm long. Fruits pale green, oblong-ovoid, c. 4 x 2 cm; fruit-wall thin; persistent disc 

not well-exposed; persistent calyx-lobes prominent, 4-5 mm long. Seeds ellipsoid, c. 3.8 x 

1.8 cm. 

Distribution. Borneo, Philippines (Luzon). In Sarawak known from Miri, Bakam Road; 

not yet recorded from Sabah, Brunei, and Kalimantan. 

Ecology. In mixed dipterocarp forest, at about 75 m. Flowering in October, fruiting in 

June. 

203


5. Mastixia pentandra Blume 

(Greek, penta = five, -andrus = male: with 5 stamens) 

I.e. (1825) 65.4; Danser I.e. 49; Backer & Bakhuizenj I.e. 159; Matthew I.e. (1976) 80, I.e. (1977) 

95; Whitrnore, Tantra & Sutisna I.e. 50. Type: Blume, s.n. (= Leiden no. 901, 169-375), Java 

(lectotype L; isolectotypes NY, W). 

sllbsp. scortechinii (King) Matthew 

I.e. (1976) 80, I.e. (1977) 54. Basionym: Mastixia seorteehinii King, J. As. Soc. Beng. 71, 1 (1902) 

72. Type: Seorteehini 1971, Perak (lectotype K; isolectotypes BM, CAL, G, L, P). Synonyms: 

Mastixia megaearpa Rid!. I.e. 891; Mastixia parvifolia Hallier j, Beih. Bot. Centralb!. 34,2 (1916) 

41, Merrilll.e. (1921)459. 

Tree to 40 m tall, 80 cm diameter; buttresses short. Bark grey-brown, smooth to shallowly 

fissured with horizontal rings and lenticels in rows; inner bark yellowish brown to dark 

yellow, with strong smell of sugarcane. Sapwood pale yellow to brownish. Twigs glabrous, 

dark brown to black. Leaves alternate, spiral or sub opposite, stalk stout, 1-2 cm long; 

blades obovate to oblong or elliptic, 5-12 x 2.5-5 cm, thickly leathery, glabrous; base 

cuneate, margin curled inwards, apex acute or acuminate but sometimes caudate; lateral 

veins 4-6 pairs, flattened above; intercostal veins and reticulations faint to inconspicuous 

on both surfaces. Inflorescences to 8 cm long, puberulous to villous. Flowers (4-)5- 

merous, greenish or yellowish white, buds c. 2 mm diameter; calyx-tube puberulous, with 

(4-)5 lobes, as long as wide; petals (4-)5, 1.2-1.5 x 0.8-1 mm, hairy on the abaxial 

surface; stamens (4-)5, filament 0.5-0.7 mm long. Fruits green, ripening purple to bluish 

black, ovoid to oblong, 1.6-3.5 x 0.8-1.2 cm; fruit-wall thick; persistent disc well-exposed 

and sometimes bulging; persistent calyx-lobes inconspicuous. Seeds ovoid, 0.9-1.2 x 0.5- 

0.8cm. 

Vernacular names. Sabah and Sarawak-kaju wulu, medang surungan (Malay). 

Distribution. Thailand, Sumatra, Peninsular Malaysia, Borneo (Sabah and Kalimantan), 

and Celebes. In Sabah recorded only from Mt. Kinabalu and Mesilau Hill. 

Ecology. Uncommon, distributed from lowland to submontane forests to 1500 m. Flowering 

and fruiting in January-December 

Taxonomy. In Matthew's treatment, Mastixia pentandra was segregated into six 

subspecies, viz. pentandra, moluccana, chinensis, cambodiana, philippinensis and 

scortechinii. Ofthese, only M. pentandra subsp. scortechinii is known from Sabah. 

6. Mastixia rostrata Blume Fig. 1. 

(Latin, rostratus = with a beak, narrowed into a slender tip or point; the leaf apex) 

I.e. (1850) 258; Danser I.e. 52; Backer & Bakhuizenj I.e. 159; Matthew I.e. (1976) 73, I.e. (1977) 

94; Anderson I.e. 164; Whitmore, Tantra & Sutisna I.e. 50. Type: Blume, s.n. (= Leiden no. 901, 

169-384), Java (lectotype L). 

204


subsp. caudatifolia (Merr.) Matthew 

I.e. (1976) 74, I.e. (1977) 94. Basionym: Mastixia eaudatifolia Merr.l.e. (1929) 233. Type: Elmer 

21584, British North Borneo, Tawau, Elphinstone Pro~. (holotype VC; isotypes A, BM, BO, GH, 

HBG, K, L, NY, P, SING, V, VC, VS). Synonyms: M margarethae Wangerin, Fedde Rep. 4 (1907) 

335; M. cuspidata Blume var. margarathae (Wangerin) Hallier f l.c. 41. 

Tree to 30 m tall and 50 cm in diameter. Bark greyish to chocolate-brown, smooth to 

occasionally shallowly fissured; inner bark yellowish to pale orange-yellow, fibrous, soft. 

Sapwood yellowish. Twigs glabrous, grey or grey-brown. Leaves alternate, sub opposite or 

opposite, stalk slender, to 1(-1.5) cm long; blades elliptic-oblong to elliptic, 4-8(-10) x 2- 

5 cm, papery to thinly leathery, glabrous; base cuneate, margin not curled inward, apex 

caudate with the tip to 1.5 cm long; lateral veins 4-6 pairs, prominent below, sunken 

above, not looping toward leaf margin; intercostal veins faint to inconspicuous below. 

Inflorescences to 6 cm long, subglabrous. Flowers 4-merous, green-yellow, buds to 1 mm 

diameter; calyx-tube glabrous, with 4 lobes, broader than long, glabrous; petals 4, 1.1-1.3 

x 0.7-0.9 mm, glabrous on the abaxial surface; stamens 4, filament 1-1.3 mm long; disc 

yellowish. Fruits ovoid to oblong, 1.5-2.2 x 0.5-1 cm; fruit-wall thick; persistent disc not 

well-exposed; persistent calyx-lobes inconspicuous. Seeds ellipsoid, 1.3-2 x 0.6-0.8 cm. 

Vernacular name. Sarawak-patoli entelit (!ban). 

Distribution. Sumatra, Peninsular Malaysia (new record), and Borneo. Widely distributed 

in Sabah and the Wand C parts of Sarawak. 

Ecology. In primary mixed dipterocarp forest on clay-rich soils, from lowland to 1600 m. 

Flowering in June to October, fruiting in August to March. 

Taxonomy. In Matthew's treatment, Mastixia rostrata was segregated into two subspecies, 

viz. rostrata and caudatifolia. Of these, only M. rostrata subsp. caudatifolia is known from 


7. Mastixia trichotoma Blume 

(Greek, tri == three, tomos == part; the 3-branched inflorescence) 

I.e. (1825) 655, King I.e. 72; Wangerin I.e. (1910) 24; Merrill I.e. (1921) 459; Danser I.e. 57; 

Matthew I.e. (1976) 68, I.e. (1977) 92; Kochummen I.e. 54; Anderson I.e. 164; Whitmore, Tantra & 

Sutisna I.e. 50. Type: Blume, S.n., West Java, Mt. Salak (lectotype L; isolectotypes BM, W). 

Synonyms: Mastixia laxa Blume I.e. (1850) 257 (including var. angustifolia); Mastixia 

aeuminatissima Blume I.e. (1850) 258; Mastixia eaesia Blume I.e. (1850) 258; Mastixia kimanilla 

Blume I. e. (1850) 258. 

Tree to 40 m tall and 50 cm diameter. Bark yellowish grey to grey-brown, smooth to 

shallowly fissured; inner bark yellowish brown to pale brown. Sapwood pale white. Twigs 

not distinctly grooved, yellowish brown to pale brown, puberulous to woolly. Leaves 

opposite, stalk stout or slender, 1-3.5 cm; blades ovate, elliptic, lanceolate to oblong, 5-24 

x 2.5-12 cm, thinly to thickly leathery, subglabrous to velvety hairy below; base acute, 

205


cuneate, obtuse or attenuate, apex acute to acuminate; lateral veins 5-15 pairs, sunken 

above, prominent below; intercostal veins reticulate, faintly visible to prominent below. 

Inflorescences to 15 cm long, puberulous to woolly. Flowers 4-5-merous, green to 

yellowish green; buds 1-2.5 mm diameter; calyx-tube puberulous to villous, with 4 or 5 

lobes, as long as wide, puberulous to villous; petals 4 or 5, puberulous to villous on the 

abaxial surface; stamens 4 or 5. Fruits ovoid to ellipsoid, 1.5-3 x 0.6-1.5 cm, fruit-wall 

thin; persistent disc well-exposed and bulging or not; persistent calyx-lobes inconspicuous 

to slightly prominent. Seeds ovoid to ellipsoid, 1.3-2.9 x 0.4-1.4 cm. 


1. Inflorescence villous to woolly. Lateral veins looping to form marginal veins; 

intercostal veins distinct and prominent below ............................................................. 2 

Inflorescence sub glabrous to puberulous. Lateral veins not looping; intercostal veins 

not prominent below .................................................................................................... 3 

2. Twigs woolly. Leaves villous to woolly, thickly leathery. Flowers 4-merous .................. . 

var. maingayi (Clarke) Danser 

l.c. 63; Matthew l.c. (1976) 70, I.c. (1977) 93; Kochummen I.c. 54. Basionym: M. maingayi 

Clarke I.c. 746. Type: Maingay 2680 = Kew Distr. 711, Singapore (holotype K; isotypes 

BM, GH, L). Synonyms: M. maingayi Clarke var. subtomentosa King I.c. 75; M. propinqua 

Ridl., J. Fed. Mal. St. Mus. 4,1 (1909) 25. 

Twigs woolly by short yellowish brown hairs. Leaf-stalk stout, 2-3.5 cm, woolly 

brown-hairy. Leaves 8.5-22 x 3.5-10.5 cm, thickly leathery, abaxial surface 

woolly brown-hairy, adaxial surface glabrous except the midrib and veins; lateral 

veins 5-6 pairs, prominently sunken above and raised below, forming marginal 

loops below; intercostal veins prominently sunken above and raised below. 

Inflorescences villous to woolly. Flowers 4-merous. Fruits with well-exposed 

persistent disc; persistent calyx-lobes 1.5-2 mm long. 

Sumatra, Peninsular Malaysia, Borneo. In Sabah recorded from Lahad Datu, 

Ranau, Sipitang, and Tawau districts. In Sarawak, recorded from only Kapit (3rd 

Div.) and Ulu Lawas (5th Div.). In primary and secondary mixed dipterocarp 

forests, to 1500 m. 

Twigs subglabrous to velutinous. Leaves glabrous to subglabrous. Flowers 5-merous ..... 

var. korthalsiana (Wangerin) Danser 

I.c. 63; Matthew l.c. (1976) 70, I.c. (1977) 93. Basionym: M. korthalsiana Wangerin l.c. 

(1907) 335. Type: Korthals, s.n., E Borneo, Bandjermasin, G. Sakoembang (L). 

Twigs subglabrous to velutinous. Leaf-stalks slender, 1.5-2 cm. Leaves 7-13 x 3- 

5 cm, glabrous to sub glabrous; lateral veins 5-6 pairs, forming looped marginal 

veins below; intercostal veins prominent below. Inflorescences velutinous to woolly. 

Flowers 5-merous. Fruits with well-exposed persistent disc; persistent calyx-lobes 

to 1 mm long. 

Sumatra and Borneo (Sabah and Kalimantan). In Sabah, recorded from Kota 

Belud, Lahad Datu, Sandakan, Sipitang, and Tawau districts. No record from 

Sarawak. In primary mixed dipterocarp forest, from low altitude to 1100 m. 

206


l,m" 

B 

"" [ , E 

c 

p 

[ 'cr'~~ 

..(', " 

Cl:,' :~i' 

lmm "',~ D 

," 

'~~~?.!:~" 

Fig, L lv£astixia rostrata subsp, caudatifolia, A, flowering leafy twig; B, flower bud; C, open flower; 

D, open t10wer with petals and stamens removed; E, fruit; F, fruit in longitudinal section; G, fruit in 

cross section, (A-D from SAN 30572, E-G from S, 36376,) 

207


3. Leaves.9-::-24 x 4-12 cm, margin not wavy after drying, under surface not pale yellow 

when dry.' Twigs and leaves (sub)glabrous. Fruits with persistent disc well-exposed and 

bulgin,g .......................................................................................................................... 

~i:. rhynchocarpa Danser. 

i.'9. 64; Matthew I.e. (1976) 71, I.e. (1977) 93. Type: Endert 4769, E Borneo, East Coast, 

near Long Hoet, 150 m (lectotype L; isolectotypes A, BO, K). Synonyms: Mastixia 

triehotoma Blume var. beneuluana Danser I.e. 64; Mastixia triehotoma Blume var. 

simalurana Danser I.e. 65. 

Twigs subglabrous. Leaf-stalks stout, 1.5-2 cm long. Leaves 9-24 x 4-12 cm, 

thickly leathery, subglabrous, under surface not pale yellow when dry; lateral 

veins 5-15 pairs, not looping near the margin; intercostal veins faintly visible 

below. Inflorescences subglabrous to puberulous. Flowers 4-merous. Fruits 

crowned by bulging, well-exposed persistent disc; persistent calyx-lobes to 0.5 mm 

long. 

Sumatra, Java, Borneo, Celebes, and the Moluccas. In Sabah widely distributed. In 

Sarawak in Ulu Mujong (3rd Div.), Lawas (5th Div.), and Kapit (7th Div.). In 

primary mixed dipterocarp forests, to 1800 m. 

Leaves 7.5-15 x 2.5-6 cm, margin wavy after drying, under surface pale yellow when 

dry. Twigs and leaves covered with powdery tomentum. Fruits not so ............... 

var. c1arkeana (King) Danser 

I.e. 62; Matthew I.e. (1976) 72, I.e. (1977) 93; Kochummen I.e. 54. Basionym: Mastixia 

clarkeana King I.e. 72. Type: Seorteehini 869, Perak (lectotype K; isolectotypes CAL, G, L, 

P). Synonyms: Mastixia clarkeana King var. maerophylla King I.e. 72; Mastixia 

korthalsiana Wangerin var. maerophylla Wangerin I.e. (1907) 336; Vitex premnoides 

Elmer I.e. 2874; Mastixia premnoides (Elmer) Hallier f I.e. 42; Mastixia triehotoma 

Blume var. tenuis Danser I.e. 6l. 

Twigs covered with pale brown powdery tomentum. Leaf-stalks slender, 1-1.5 cm 

long. Leaves elliptic, 7.5-15 x 2.5-6 cm, under surface pale yellow on drying; 

lateral veins 5-7 pairs, not looping near the margin; intercostal veins faintly 

visible below. Inflorescences subglabrous to puberulous. Flowers 4-merous. Fruits 

without well-exposed persistent disc; persistent calyx-lobes prominent, 0.5-1 mm 

long. 

Pattani (peninsular Thailand), Surnatra, Peninsular Malaysia, Borneo, and Mindanao 

(Philippines). Uncommon in Borneo, recorded only from Kuching and Mt. 

Penrissen (lst. Div.) in Sarawak. No record from Sabah. In primary mixed 

dipterocarp forest, from low altitude to 1100 m. 

Vernacular names. Sabah-bantis (Dusun Kinabatangan), medang kanigara (Ma1ay). 

Sarawak-itan beruang (Kelabit), medang kanigara (Malay), priabu (Murut). 

Taxonomy. Of the five varieties recognised by Matthew I.e., four occur in Sabah and 


208


DATISCACEAE 

E.J.F. Campbell-Gasis 

clo Forest Research Centre, 



Merrill, EB (1921) 414; Masamune, EPB (1942) 507; van Steenis, FM I, 4 (1953) 382; Browne, 

FTSB (1955) 82; Backer & BakhuizenJ, FI 1 (1963) 313; Smythies, CST (1965) 43; Burgess, TS 

(1966) 86; Whitmore, TFM 2 (1973) 29; Cockburn, TS 1 (1976) 73; Anderson, CLTS (1980) 165; 

Ashton, MNDTS 2 (1988) 234; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 52. 

Trees, shrubs or herbs. Leaves spiral, simple, without stipules; blade palmately veined, 

domatia/glands present. Flowers in axillary and/or terminal elongated spikes or panicles, 

unisexual, regular, parts not overlapping; male: sepals 4-9, free and unequal or connate in 

a lobed tube; petals 4-9 or 0, free; stamens 4-9 on the sepals, filaments often long, anthers 

basifixed, 2-celled, longitudinally dehiscent; female: sepals 3-8, often joined above the 

ovary or free; petals 0; staminodes 0; ovary inferior, i-celled with 3-8 parietal placentas 

and numerous ovules; styles 3-8 opposite calyx-lobes, thick, free, mostly adnate to the 

margin of the calyx, simple or bifid, stigma club-like or capitate. Fruits capsular, wall 

papery, opening at apex with slits and/or splitting laterally. Seeds very numerous, minute; 

endoperm none; embryo straight, cylindric. 

Distribution. 3 genera, with 4 species: Datisca (2 species, extra-Malesian), Octomeles (1 

species, Indomalesian) and Tetrameles (1 species, Indomalesian). Only Octomeles is found 


Taxonomy. The systematic position of the Datiscaceae remains controversial. Several 

authors (e.g., Whitmore I.c.; Mabberley, PB (1987) 172) maintain the Datiscaceae as most 

closely related to the Begoniaceae and Cucurbitaceae, in the Cucurbitales. Mabberley I. c 

and others (e.g., Willis, DFPF (1973) 1140) suggest that both Octomeles and Tetrameles 

should be taken out of the Datiscaceae and put in the Tetramelaceae (Warb.) Airy-Shaw. 

OCTOMELES Miq. 

(Greek, octo = eight, melos = parts, limbs; the 8-merous flowers) 

Fl. Ind. Bat., Suppl. (1861) 133, 336; van Steenis I.e. 382; Burgess I.e. 86; Cockbum I.e. 73; 

Anderson I.e. 165; Wong, DMT (1984) 25; Ashton I.e. 234; Whitmore, Tantra & Sutisna I.e. 53. 

Evergreen trees with thick twigs sharply 3-angular at apex. Leaves roundish cordate, 5-7(- 

9)-veined; stalk long, 5-angled; lower surface with domatial glands. Flowers sessile, green, 

209


5-8-merous, dioecious, solitary; male: calyx-tube short with free triangular lobes; petals 

triangular; stamens strongly incurved in bud, filaments thick, anthers large, kidney-shaped, 

curved; female: sepals joined to form a tube; ovary apex cup-shaped by the thick calyx-tube; 

styles fleshy, inserted in the calyx-rim opposite the free triangular calyx-lobes, stigma 

capitate. Fruit a barrel-shaped capsule crowned with persistent styles, the styles and 

exocarp drop away irregularly leaving a pale membranous endocarp behind; endocarp 

splitting from the top downwards, the segments spreading stellately, persistent. Seeds 

spindle-shaped, with narrower tail and thickened head. 

Octomeles sumatrana Miq. 


Fig. 1. 

I.c. 133, 336; van Steenis I.c. 382; Browne I.c. 82; Smythies I.c. 43; Cockbum I.c. 73; Anderson l.c. 

165; Ashton I.c. 234; Whitmore, Tantra & Sutisna l.c. 53. Type: Teijsmann, s.n., Sumatra, Prov. 

Palembang (holotype L; isotype K). 

Tree to 70 m tall and 3 m diameter; bole columnar; buttresses steep, to 10 m high, spreading 

to 5 m away from the tree. Branches radial, ascending, pagoda-like; crown monopodia!. 

Bark pale brownish cream or grey, thin, hooped (branch-scars), dot-like lenticels visible; 

inner bark pinkish cream, soft, fibrous, thick. Sapwood cream, exudate slight, clear. Twigs 

stout, glabrous, with prominent large round leaf-scars subtended by 5 almost papery sharp 

ridges. Leaves broadly ovate to round, 12-33 x 16-29 cm, drying rather papery, with 

scattered white hairs below; base heart-shaped, often deeply to 16 cm, margin toothed or 

wavy, apex acute to acuminate; midrib raised above and below; lateral veins 7-10 pairs, the 

basal 1-2 pairs originating from the stalk insertion, giving a somewhat 3-veined or 

palmately veined appearance, mostly parallel and looping near margin, raised above and 

below; intercostal veins net-like and flat; midrib and lateral veins dark red on both 

surfaces; obscure dark domatia present at apex, at the junctions of the midrib and lateral 

veins and alongside smaller veins (obvious in dry leaves); stalk glabrous, 3-5 mm thick and 

6-32 cm long, smooth on upper side, 5-ridged on lower side, grooved near base for 1-2 cm. 

Male flowers in spikes 20-60 cm long, c. (5-)8 mm across and 7 mm long; sepals connate 

for most of the length, lepidote, lobes free, triangular, 1.5-2 mm; petals triangular, 

alternate with calyx-lobes, c. 3 mm long; stamens 4-10 mm long, anthers 2-3 mm. Female 

flowers in shorter spikes 8-35 cm long, c. 5 mm across and 5-10 mm long; calyx-tube 2-4 

mm high; ovary 1-2 mm long, styles fleshy, triangular, 2-3 mm long, stigma thick, 1-2 

mm. Fruits 8-12 mm long, with pale brown endocarp. Seeds thicker in the middle, c. 0.8 

mm long, numerous. 

Vernacular names. Sabah-binuang (Malay). Sarawak-benuang (lban), binong (Bidayuh), 

lemeng (Berawan, Punan, Tutoh). 

Distribution. Sumatra, Borneo, Philippines, Celebes and New Guinea. In Sabah, common 

on the east coast and interior. In Sarawak, in lowlands and hills in all districts. 

Ecology. Often associated with riverine and alluvial deposits but also found in primary and 

secondary mixed dipterocarp forests on well-drained fertile friable soils below 600 m on the 

east coast of Sabah and to 800 m on the west coast and in Sarawak. It is more typically 

210

DA TISCACEAE (CAMPBELL-GASIS) 

------' 

A 

H_ r 

Fig. 1. Octomeles sumatrana. A, fruiting leafy twig; B, female flower; C, male flower. CA from SAN 

67376, B from Kokawa & Hotta 1426, C from SAN 67406.) 

211


found growing in disturbed habitats such as logged-over forest and after cultivation where 

it can grow extremely fast, often in gregarious even-aged stands. Flowering and fruiting from 

November to April in Sabah and Sarawak. Flowers probably wind-pollinated and the seeds 

probably wind-dispersed. 

Silviculture. Binuang is one of the fastest growing trees and a strong light-demander. It 

takes only about 4 months"from sowing to planting out in the field. Seed viability, however, 

soon decreases after collection. Its ability to grow on low hilly sites and in temporarily 

flooded areas, quick crown development, and self-pruning abilities, make it a suitable 

plantation tree. For optimal growth, however, the seedlings/saplings would have to be 

planted at sufficiently wide spacings to accommodate the huge crowns of the mature trees. 

Timber strength & wood anatomy. Binuang timber is a soft, light hardwood of the obeche 

(Triploehiton scleroxylon - Sterculiaceae) type with long fibres and a low density. The 

sapwood is almost white but of drab appearance and the heartwood is pale brown or pinkish 

brown. The wood has an interlocked grain and the texture is coarse (Burgess I.e., Wong 

I.e.). The pores are rather large, oval, mostly open, evenly distributed, with broader rays 

distinct, almost white and the cut end surfaces are dull (Browne I.e.). The dry weight is 

270-465 kg/m 3 air-dried. The wood seasons slowly and is not durable. There is a tendency 

for both brittle-heart and fungal stain to develop, and damage by pinhole-borer beetles and 

powder-post beetles is sometimes present but treatment with preservatives gives moderate 

resistance (Burgess I.e., Wong I.e.). 

Uses. The timber works easily with machine and hand tools but the timber can crumble or 

give a woolly finish if the tool cutters are not kept sharp. It also stains and polishes 

satisfactorily. Burgess I.e. and Wong I.e. suggest that the wood can be used for cores, backs 

to plywood, and is a useful utility timber for packing cases, concrete shuttering, matchboxes 

and other temporary purposes. It is a potentially important tree for wood chip and pulping 

and as a shade tree or for stabilising river banks but its potential is not yet exploited in 


Notes. The seedlings and saplings have usually larger leaves with a cordate base and a few 

coarse teeth, a distinctly opposite and decussate branching, and characteristically angled 

twigs which make them easy to identify. Some young Maearanga species (Euphorbiaceae) 

have similar leaves but these can be distinguished easily as the twigs of this genus do not 

have papery ridges. Older trees can sometimes be confused with Neolamarekia eadamba 

(Roxb.) Bosser (Rubiaceae), but Oetomeles sumatrana has radial branching in distinct tiers 

due to its episodic growth pattern whereas N eadamba produces radial branches continuously 

up the stem. The leaves of 0. sumatrana are always dotted with holes due to insect damage 

unlike the leaves of N eadamba. It has been noted by van Steenis I. e. as being a "preferred" 

tree for nesting bees like the other lofty tree species, Koompassia exeelsa (Becc.) Taub. 

(Leguminosae). 

212


GOODENIACEAE 

K.M. Wong 


Forestry Department, Sabah 

Merrill, EB (1921) 586; Masamune, EPB (1942) 724; Leenhouts, FM 1, 5 (1957) 335, FM 1, 5 

(1958) 567, FM 1, 6 (1972) 949, FM 1, 7 (1976) 827, FM 1, 9 (1982) 566. 

Small trees, shrubs or herbs, or scrambling-climbing plants. Leaves spirally arranged or 

opposite, or whorled, simple, pinnately veined; leafaxils often with hair tufts; stipules 

none. Inflorescences cymose, bracteate. Flowers 5-merous, bisexual, protandrous (male 

parts maturing first); calyx tubular with distinct lobes; corolla members fused into one or 

two broad lip-like structures in the open flower, the lobes with thin membranous margins; 

stamens 5, free, borne on the calyx, filaments linear, anthers basifixed and introrse, 2- 

celled; disc none; ovary 2-celled or imperfectly I-celled, style cylindrical, stigma surrounded 

by a cup-shaped structure (indusium) with ciliate margin; ovules I-many, placentation 

axile or basal. Fruits capsular or drupaceous. Seeds I-many, with endosperm. 

Distribution. 16 genera, about 430 species, mostly Australian. In Malesia, 5 genera with 

11 species. In Sabah and Sarawak, 5 species of Scaevola are documented. 

Ecology. The family seems confined to or adapted to poor soils. In Malesia its species occur 

in rather open sites in beach or coastal areas, or on ultramafic soils and in low-stature 

montane forests. 

Taxonomy. The Goodeniaceae are closely related to Lobelia and related genera in the 

Campanulaceae. 

SCAEVOLAL. 

(the name given to Gajus Mucius around 507 B.C., who burnt his own hand in 

fearlessness, to which is likened the white, 5-lobed corolla which turns brown) 

Mant. (1771) 145; Merrilll.e. 586; Masamune I.e. 724; Leenhouts I.e. (1957) 339, I.e. (1958) 567, 

I.e. (1972) 951. Synonym: Temminekia de Vriese, Ned. Kruidk. Arch. 1,2 (1851) 141. 

Shrubs or small trees, or scrambling-climbing plants. Leaves spirally arranged or opposite, 

or whorled, entire or crenate to dentate, the margins and tip provided with glands; leafstalks 

with broad insertion on the shoots. Flowers in axillary, bracteate cymes; calyx-tube 

fused to the ovary; corolla members all fused to form a broad lip-like structure in the open 

flower, the free lobes with membranous margins and fimbriate at their base; ovary inferior 

213


or semi-inferior, 1-2-celled; ovules 1 per ovary cell. Fruits drupaceous, with a hard stone. 

Seeds 1 or 2. 

Distribution. About 130 species, predominantly Australian, one species reaching to Taiwan 

and Hainan in south China, a few in east Malesia (New Guinea), and two widely distributed, 

chiefly littoral species (s. sericea in the lndo-Pacific region, S. plumieri in the lndo 

Atlantic region). In Sabah and Sarawak, 5 species. 

Key to Scaevola species 

l. Leaves in whorls offouf.. ..................................................................... 5. S. verticillata 

Leaves spirally arranged .............................................................................................. 2 

2. Leaves sessile to subsessile, the blades decurrent more or less all the way to the nodes 

or the stalks exceedingly short and inconspicuous ........................................................ 3 

Leaves distinctly stalked, decurrent basal part of the blades very narrow and inconspicuous 

........................................................................................................................... 4 

3. Plants of sandy, rocky or cliff sites beside the sea or the coast. Leaves exceeding 10 cm 

long, with long-decurrent bases. Cymes (2-)3-5(-6)-times branched, often 6-10 cm 

long. Calyx-lobes linear to lanceolate, 2-3 mm long (sometimes elongating to 5-7 mm 

long); corolla with a fused part 15-20 mm long and free lobes 5-10 mm long. Fruits 

broad-obovoid to globose, 8-12 mm long, strongly ridged .......................... 4. S. sericea 

Plants of high elevation on mountains (known only on Mt Kinabalu, Sabah). Leaves 1- 

7 cm long, more or less abruptly sessile. Cymes 1-2-times branched, not exceeding 3 

cm long. Calyx-lobes triangular, less than 1 mm long; corolla with a fused part 4-6 

mm long and free lobes 4-6 mm long. Fruits narrow-obovoid, 3.5-5 mm long, slightly 

ridged ......................................................................................................... 1. S. chanii 

4. Plants ofultramafic substrates (in Borneo known only in Sabah). Leaves 8-15 cm long, 

distinctly crenate-dentate on the margins. Cymes 3-6-times branched, 5-9 cm long, 

the first pair of bracts conspicuously larger and leaf-like, peduncle 2-3.5 cm long. 

Flowers in the forks of cyme branches, stalked ...................................... 2. S. micrantha 

Plants so far known only on limestone (in Sarawak). Leaves not exceeding 8 cm long, 

only minutely and inconspicuously crenate on the margins. Cymes 1-2-times branched, 

not exceeding 2.5 cm long, the first pair of bracts tiny triangular structures, 

peduncle to 1 cm long only. Flowers in the forks of cyme branches, sessile ............... . 

.............................................................................................................. 3. S. muluensis 

1. Scaevola chanii W ong Fig. 1. 

(C.L. Chan, orchidologist and botanical illustrator) 

Sandakania 3 (1993) 12. Type: Chan, lamal & Wong WKM 2360, Sabah, Mt. Kinabalu, sununit 

trail, 2900 ill alt. (holotype SAN; isotypes K, KEP, L, SAR, SNP). 

Shrub or treelet, to 2-3 m high. Leaves spirally arranged, ob ovate, 1-7 x 0.6-3 cm, margin 

minutely crenate-dentate, coriaceous; glabrous except for small tufts of silvery white hairs 

214

GOODENIACEAE (WON G) 

5 

cm 

o 

d~~ 

. flldlH8stiJindlP 

-i,L.ieJy 1. ce; ~ --.. 

'9d~ /('( 

c j ,~ 

Al~ 

B 

Fig. 1. Scaevola chanii. A, flowering leafy twig; B, detail ofleaftip and margin showing glands; C, 

flower; D, indusiate stigma. (All from Chan, lamal & Wong WKM 2360.) 

215


in the leafaxils, and tiny silvery round scales all over the young leaves; lateral veins 4-8 

pairs, inconspicuous; stalks very short, only 1-2 mm long at most, inconspicuous. Inflorescences 

1.5-2.5 cm long, branched only 1-2-times, peduncle 0.7-2 cm long; first pair of 

bracts triangular-linear, small; glabrous except for tufts of pale hairs in the bract axils. 

Flowers sessile; calyx cup obovoid, glabrous, the lobes triangular, less than 1 mm long, 

pale pilose on the margins; corolla sparsely pale hairy on the outside, the fused part 4-6 

mm long, the free lobes 4-6 mm long, pale pilose inside; stamens with filaments c. 3 mm 

long, anthers 0.7-1 mm long; style 6-8 mm long; indusium around the stigma glabrous on 

the surface, densely pale hairy on the margin. Fruits ob ovoid, 3.5-5 mm long, slightly 

ridged. 

Distribution. Recorded only from Mt. Kinabalu, Sabah. So far endemic. 

Ecology. Upper montane forest on ultramafic soil, at 2500-3000 m. 

2. Scaevola micrantha C. Presl 

(Greek, micros = small, anthos = flower) 

ReI. Haenk. 12 (1835) 58; Leenhouts I.c. (1957) 342. Type: Haenke 124, Philippines (W). 

Synonyms: Temminckia micrantha (C. Presl) de Vriese I.c. (1851) 145; S. pedunculata MeIT., 

Philip. J. Se., Bot. 5 (1910) 251, ineI. var. mollis, I.c. (1921) 586, Masamune I c. 724; S. merrillii 

Elmer, Leafl. Philip. Bot. 4 (1912) 149l. 

Shrub or small tree to 10 m tall, 5 cm diameter. Bark dark grey to greenish grey, smooth to 

slightly flaky. Leaves spirally arranged, obovate to oblanceolate, 8-15 x 2-5.5 cm, margin 

crenate to distinctly dentate, chartaceous to thinly coriaceous; subglabrous (with scattered 

hairs on the stalk, leaf base and midrib on the lower side) to scantily short-hairy to velvety 

on the lower side, the leafaxils with tufts of silvery white hairs; lateral veins 6-11 pairs, 

inconspicuous; stalks 4-17 mm long, distinct. Inflorescences 5-9 cm long, branched 3-6- 

times, peduncle 2-3.5 cm long; first pair of bracts ovate and leaf-like, much larger than 

subsequent bracts; glabrous to pale velvety all over. Flowers on stalks 0.5-1.5 mm long; 

calyx-cup obovoid, glabrous to densely pale pilose, the lobes triangular, c. 0.5 mm long, 

short-hairy on the margins to pale pilose all over; corolla scantily to densely pale pilose all 

over the outside, the fused part 5-7 mm long, the free lobes 4-5 mm long, pale pilose 

inside; stamens with filaments c. 4 mm long, anthers c. l.5 mm long; style 6-7 mm long; 

indusium around the stigma scantily hairy to pale pilose on the surface, densely pale-hairy 

on the margin. Fruits obovoid, 5-6 mm long, slightly ridged. 

Distribution. Borneo and the Philippines, including Palawan Is. In Borneo, known only 

from Sabah (Kinabalu-Ranau area, Telupid, Beluran, Mt. Tawai and Mt. Danum). The 

Celebes collection (Lam 3266) from Talaud Is. attributed to this species by Leenhouts I.c. 

(1957) appears to be a different species; it has entirely glabrous leaves, I-mm-long calyx 

teeth, linear and longer (6-8 mm) inflorescence bracts, and only 2-3-times branched 

inflorescences. 

Ecology. Recorded only from ultramafic substrates, from the lowlands to submontane forest 

at c. 1500 m. 

216

GOODENIACEAE (WONG) 

3. Scaevola muluensis Wong 

(ofMt. Mulu, Sarawak) 

Sandakania 3 (1993) 15. Type: Argent & Jermy 1011, Sara:wak, 4th Division, Gunong Api (holotype 

SAN; isotypes E, L, SAR). 

Shrub or treelet to c. 1 m high. Leaves spirally arranged, obovate, 4-6.5 x 2-3 cm; margin 

only minutely and inconspicuously crenate; thinly coriaceous; glabrous except for scattered 

short pale hairs on the stalk and tufts of short pale hairs in the leafaxils; lateral veins 5-6 

pairs, slightly elevated on both surfaces when dry; stalks 5-13 mm long, distinct. Inflorescences 

1-2 cm long, branched only 1-2-times, peduncle to 1 cm long only; first pair ofbracts tiny 

triangular structures; glabrous except for scanty pale hairs in the bract axils. Flowers 

sessile; calyx-cup obovoid, glabrous, the lobes triangular, c. 0.5 mm long, glabrous to 

sparsely short-hairy on the margins; corolla sparsely pale-hairy on the outside, the fused 

part 6-6.5 mm long, the free lobes 4.5-5 mm long, pale pilose inside; stamens with 

filaments c. 4 mm long, anthers c. 1 mm long; style 6-6.5 mm long; indusium around the 

stigma with pale long hairs all over, densely pale hairy on the margin. Fruits obovoid, 4-5 

mm long, slightly ridged, ripening black. 

Distribution. Known from two collections (830907 and Argent & Jermy lOll) between 

1000 ill and 1600 m on the limestone of Gunong (Mt.) Api in the Gunong Mulu National 

Park in Sarawak's 4th Division. 

Ecology. Probably restricted to limestone, found on exposed ridges and cliff faces. 

4. Scaevola sericea Vahl 

(Latin, sericeus = silky with long hairs; the leaves) 

Symb. Bot. 2 (1791) 37; Leenhouts, I.e. (1957) 339. Type: G. Forster, s.n., Pacific, Niue Is. (C). 

Synonyms: Lobelia plumieri (non L.) Burm./, Fl. Ind. (1768) 189; S. koenigii Vahl, Symb. Bot. 3 

(1794) 36; S. taceada Roxb., Hort. Beng. (1814) 15, Leenhouts I.e. (1972) 951; S. plumieri (non 

Vahl) Blume, Bijdr. (1826) 730; S. lesehenaultii DC., Prod. 7 (1839) 506; S. maeroealyx de Vriese 

I.e. (1850) 138; S. pi/iplena Miq., Fl. Ind. Bat. 2 (1857) 581; S. fruteseens (non Lobelia fruteseens 

Mill.) Krause, Pfl. R. Heft 54 (1912) 125, Merrilll.e. (1921) 586, Masamune I.e. 724. 

Shrub or small tree to 10 m tall and 10 cm diameter. Bark yellowish green to grey, smooth. 

Leaves spirally arranged, obovate, 12-26 x 5-10 cm; base decurrent to the node, margin 

crenate to subentire, apex obtuse, rounded; thickly succulent to thinly coriaceous; glabrous 

to scantily hairy to tomentose on both sides, the leafaxils with dense tufts of pale longhairs; 

lateral veins 8-12 pairs, slightly elevated on upper side; stalks nil or very short, 

hardly 1-2 mm long. Inflorescences (4-)6-10 cm long, branched (2-)3-5(-6)-times, 

peduncle c. 1 cm long; first pair of bracts linear-triangular; glabrous to densely short-hairy 

all over. Flowers sessile on the slender cyme branches; calyx cup obovoid-ellipsoid, 

glabrous to scantily to densely pale hairy, the lobes linear to lanceolate, 2-3 mm, elongating to 

5-7 mm long, scantily to densely short-hairy all over; corolla scantily to densely pale-hairy 

all over the outside, the fused part 15-20 mm long, thefree lobes 5-8(-10) mm long, pale 

pilose inside; stamens with filaments 10-15 mm long, anthers 2-3 mm long; style 15-20 

217


mm long, indusium around the stigma scantily to densely hairy all over, densely pale-hairy 

on the margin. Fruits broadly obovoid to subglobose, 8-12 x 10-15 mm, conspicuously 

ribbed, fleshy, stone to 8 mm across. 

Vernacular name. Sarawak-butun laut (Lundu Malay). 

Distribution. Madagascar, SE Asia, throughout Malesia, tropical Australia, Micronesia, 

Melanesia, to Hawaii. In Borneo common along the sea and coast in Sabah, Sarawak, 


Ecology. Common on sea-shores and sandy sites behind the shore, on rocky cliffs and even 

some sandstone hills on the coast (e.g., Trig Hill in Sandakan). Flowers and fruits all year 

round. The stones, with a corky outer layer, are buoyant and adapted to dispersal by sea 

currents. The seeds remain viable after long periods in sea water, but will germinate only 

with fresh water as after being washed ashore on a rainy day (Lesko & Walker, Ecology 50 

(1969) 730). 

Uses. The pith ("taccada") is used for making Malayan rice-paper and crafting artificial 

objects of decoration, such as flowers (Burkill I.c.). Mamit (S. 35139, Sarawak, 1st 

Division, Lundu) records that the liquid extracted from the fruit can be applied directly to 

cure sore eyes. 

5. Scaevola verticillata Leenh. 

(Latin, verticillatus = whorled; the leaves) 

Blumea 12 (1964) 317, f. 1, I.e. (1972) 951. Type: Meijer SAN 22818, Sabah, Ranau, Mt. 

Tambuyokon (ho1otype L; isotypes K, SAN, SAR). 

Shrub or treelet. Leaves in whorls of four, obovate, 2-5.5 x 0.5-2 cm; margin minutely 

crenate and recurved strongly; thickly succulent to coriaceous; densely tomentose but later 

glabrescent on upper side, densely woolly tomentose on lower side and stalk, leafaxils with 

tufts of pale long-hair; lateral veins 8-12 pairs, inconspicuous to slightly elevated on upper 

side; stalks very short, 3-6 mm long. Inflorescences 1.5-2 cm long, branched only 2-4- 

times, peduncle to 0.7-1.2 cm long only, the branches very short; first pair of bracts 

slightly larger and broader than subsequent bracts; densely hairy all over. Flowers sessile; 

calyx-cup obovoid, scantily to densely short-hairy all over, the lobes of different sizes and 

shapes (one ovate and c. 2.5 mm long, one triangular and c. 1 mm long, three rounded and 

c. 0.5 mm long), scantily short-hairy on the surface, densely long-hairy on the margin; 

corolla densely woolly tomentose on the outside, the fused part 5-6 mm long, the free lobes 

5-6 mm long, pale pilose inside; stamens with filaments 4.5-5 mm long, anthers c. 1 mm 

long; style 6.5-7 mm long; indusium around the stigma with pale long hairs around the 

base, densely pale-hairy on the margin. Fruits obovoid-ellipsoid, 3-3.5 mm long, slightly 

ridged, glabrescent except for the hairy persistent calyx-lobes. 

Distribution. Recorded only from Mt. Tambuyukon near Mt. Kinabalu, Sabah. Probably 

endemic. 

Ecology. Found in stunted subalpine vegetation at around 2500 m, on ultramafic substrate. 

218

HYPERICACEAE 

K.M. Wong 




Ecology. In Sabah and Sarawak, species of Cratoxylum are mainly lowland plants; a few 

species occur also in lower montane areas. In general these are all fast-growing trees 

commonly found in forest fringes, gaps and disturbed habitats. The light seeds are winddispersed. 

Hypericum petiolulatum is a widespread species which in Borneo occurs only on 

Mount Kinabalu. H. japonicum occurs in lowland marshy places and rice fields. 

Uses. Cratoxylum timber is distinguished into two types, one heavier and classified as 

derum in the trade, the other lighter and known as geronggang. C. cochinchinense, C. 

formosurn and C. maingayi produce derum timber, which is moderately hard and 

moderate ly heavy to heavy; the timber is sometimes known as serungan batu (Sabah) or 

entemu (.Sarawak). C. arborescens and C. glaucum produce geronggang timber, a light 

hardwood sometimes called serungan in Sabah. Derum is suitable for heavy and medium 

construction under cover, for tool handles and for wooden pallets. Geronggang has been 

found sui table for veneers and plywood, flooring, interior works and light to medium 

219 

Trees, shrubs or herbs. Leaves simple, opposite, entire, without stipules, often with translucent 

or dark glandular dots. Inflorescences terminal and sometimes also axillary, rarely 

axillary only, cymose, thyrsoid or paniculate, I-many-flowered. Flowers bisexual, radially 

symmetrical, in some species with short-styled and long-styled forms; sepals 5, free or 

partially ·fused, imbricate, often with linear glands, persistent in the fruit; petals 5, free, 

imbricate, sometimes with scale-like nectar-bearing appendages at the base, glabrous; 

stamens 5-many, fused and grouped into 3-5 fascicles, anthers dorsifixed, staminodal 

fascicles sometimes present; ovary superior, 3-5-celled or just I-celled, styles (2-)3-5, free 

or partially fused; ovules 2-many; placentas 2-5, parietal. Fruits capsular, splitting septicidally 

or loculicidally. Seeds I-many, unwinged or winged; endosperm none. 

Distribution. Seven genera and about 550 species, cosmopolitan except for the Arctic 

regions_ In Malesia, including Borneo, bnly two genera (Cratoxylum and Hypericum) 

occur. In Sabah and Sarawak, 6 species of Cratoxylum (shrubs and trees) and 2 species of 

Hypericum (herbs) are found. 

Merrill, EB (1921) 392 (as part of Guttiferae); Masamune, EPB (1942) 477, 482 (as part of 

Guttiferae); Browne, FTSB (1955) 195; Smythies, CST (1965) 68; Burgess, TBS (1966) 311; 

Kochummen, TFM 2 (1973) 248; Robson, FM 1, 8 (1974) 1; Shea, TS 1 (1976) 142; Anderson, 

CLTS (1980) 217; Corner, WSTM 1 (1988) 364; Ashton, MNDTS 2 (1988) 295; Whitmore, Tantra 

& Sutisna, eLK 1 (1989) 181. 


eRA TOXYLUM Blume 

(Greek, kratos strong, xylon = wood; referring to the hard and durable timber) 

derum, geronggang (Malay) 

Verh. Bat. Gen. 9 (1823) 174; Merrilll.c. 392; Masamune l.c. 477; Browne I.c. 195; Smythies I.c. 

68; Burgess I.c. 311; Gogclein, Blumea 15 (1967) 453; Kochummen I.c. 248; Robson I.c. 4; Shea I.c. 

144; Anderson I.c. 217; Comcr I.c. 364; Ashton l.c. 295; Whitmore, Tantra & Sutisna I.c. 181. 

Shrubs or small to medium-sized trees. Bark exuding a yellow resinous sap which drie~ 

black. Leaves with translucent glandular dots. Flowers basically 5-merous; sepals 5 

coriaceous, with longitudinal glandular lines or dots, persistent and often elongating il 

fruit; petals 5, white or pink to crimson, with longitudinal glandular lines or dots, in som, 

species with an inner basalnectariferous scale-like appendage; stamen fascicles 3, the tw 

larger attached to sepals, the smaller third attached to a petal, each with many stamem 

filaments white to crimson, anthers white to crimson; (sterile) staminodal fascicles 3, ShOl 

Shrubs or trees. Bark exuding yellow resinous sap which dries black. Flowers white to 

reddish pink; petals sometimes with basal scale-like appendages. Fruits loculicidally 

dehiscent (splitting between the internal partitions). Seeds winged .................... Cratoxylum 

About 400 species. Most tropical and temperate regions, but notably absent from the 

Amazon basin and uncommon in Australasia. 

Small trees, shrubs or herbs; in Borneo the two species are herbs. If. petiolulatum 

Hook. J & Thorns. ex Dyer occurs from Nepal through Indo-China to Sumatra and 

Borneo; in Borneo it is known only from Mount Kinabalu. If. japonicum has 

naturalised in marshy places and rice fields in Borneo; it is distributed from Japan and 

Korea through China, Sri Lanka to Malesia, Australasia and Hawaii. 

Herbs. Bark without resinous sap. Flowers yellow to orange; petals without scale-like 

appendages. Fruits septicidally dehiscent (splitting along the internal partitions). Seeds 

without wings ...................................................................................................................... . 

Taxonom:y. Three tribes are recognised within the family: Vismieae (3 genera, Africa and 

America), Crato~'Yleae (3 genera, Madagascar, Indo-Malesia, E Asia, NE America), and 

Hypericeae (HypericulII, mostly montane tropical and temperate areas). Phytochemically, 

the Hypericaceae are closely related to the Guttiferae; there are also many basic morphological 

and anatomical similarities between the two families. 

construction (Wong (1982), DMT). Geronggang logs float in water. Browne i.c. and 

Burgess /. c. review the wood characteristics. 

Key to genera 

220 


Gen. PI. cd. 5 (1754) 341; Robson l.c. 14. 

Hypericum L.

HYPERICACEAE (WONG) 

and altlernating with the stamen fascicles; ovary incompletely 3-celled; styles 3, free, linear; 

stigmas small, knob-like; ovules 4-many, attached to the basal part of the placenta, 

ascending. Fruits ellipsoid, coriaceous to woody, 3-valved, splitting loculicidally. Seeds 4- 

many on each placenta, linear to ovoid, winged all round or only on one side. 

Distribution. 6 species, from India through S China to Malesia (absent in the Moluccas 

and Ne-w Guinea); all present in Borneo. 

Ecology. Rare in primary forest, and more common in gaps, forest fringes and disturbed 

habitats in the lowlands. C. arborescens and C. formosum can occur on well-drained as 

well as swampy substrates. C. glaucum is found mostly in peat swamps, freshwater 

swamps, and wet heath forests on white sands. C. maingayi is restricted to limestone. The 

species are deciduous or semi-deciduous except for the evergreen C. arborescens and C. 

glaucum. C. formosum and C. maingayi flower on the bare branches; the other species bear 

inflorescences with the expanded new foliage or on mature leafy branches. 

Key to Cratoxylum species 

1. Leaf lateral veins inconspicuous, rather straight from midrib to very near the margin 

where a smooth-running marginal vein occurs. Seeds with wing all round (section 

]sopterygium) .............................................................................................................. 2 

Leaf lateral veins distinct and prominent at least on lower leaf surface, arching at least 

2-3 mm away from the margin, not joining at all, or joining to form conspicuous veinloops. 

Seeds with wing on one side only ....................................................................... 3 

2. Leaves not glaucous beneath, apex with at least a short tip, acuminate or caudate; leafstalks 

at least 4 mm long. Seeds 10-18 in each fruit cell .................... l. C. arborescens 

Leaves glaucous beneath, apex rounded, blunt or notched; leaf-stalks 1-3 mm long. 

Seeds 4-8 per fruit celL ......................................................................... 4. C. glaucum 

3. Leaf lateral veins not or only weakly joining to form vein-loops. Inflorescences developing 

on leafy branches, either terminal panicles or short cymes of a few flowers each at the 

ends of shoots or in leafaxils. Petals crimson to dark red or orange, without nectary 

scale (section Cratoxylum) .......................................................................................... .4 

Leaf lateral veins joining strongly to form conspicuous vein-loops towards the margin. 

Inflorescences developing on leafless parts of branches, of single axillary flowers or on 

short axillary shoots and so appearing clustered. Petals white to pale lilac, with a basal 

nectary-bearing scale (section Tridesmos) .......................... .......................................... 5 

4. Lateral veins on lower leaf-surface distinctly raised and rather coarse. Inflorescence a 

terminal panicle of many flowers. Persistent sepals less than half the capsule length 

......................................................................................................... 6. C. sumatranum 

Lateral veins on lower leaf-surface slightly raised and rather fine. Inflorescence a short 

I-few-flowered cyme, terminal or axillary. Persistent sepals more than half the capsule 

length ........................................................................................... 2. C. cochinchinense 

221


A I",\ j 

>f-l 

Fig. 1. Cratoxylum cochinchinense. A, leafy twig with young inflorescence; B, fruting leafy twig. CA 

from SAN 89495, B from SAN 134599.) 

222


5. Mature leaves typically at least 6 cm long, lateral veins more than 7 pairs, stalks 6-10 

mm long. Persistent sepals to only a third the capsule length ................ 3. C. formosum 

Mature leaves typically not exceeding 4.5 cm long, lateral veins only 5-6 pairs, stalks 

to (Jnly 5 mm long. Persistent sepals nearly half the capsule length ........ 5. C. maingayi 

1. Cratoxylum arborescens (Vahl) Blume 

(Latin, arbor = tree; the habit) 

Mus. Bot. Lugd. Bat. 2 (1852) 17; Merrill/.e. 392; Masamune I.e. 477; Browne I.e. 195; Smythies 

I.e. 68; Kochummen I.e. 249; Shea I.e. 144; Anderson I.e. 217; Corner I.e. 365; Ashton I.e. 296; 

Whitmore, Tantra & Sutisna I.e. 181. Basionym: Hypericum arboreseens Vah1, Symb. Bot. 2 (1791) 

86, t. 43 . Type: Konig, s.n., 1778, Malacca (C). 

Tree, sometimes a shrub, to 60 m tall and 120 cm diameter; buttresses if present to 1 m 

high. Bark grey to dark brown or reddish brown, smooth to fissured-cracking and papery 

scaly; inner bark pink to pale brown, finely laminated. Sapwood pale yellow. Leaves 

elliptic to obovate, 5-14 x 2-6.5 cm, coriaceous; base cuneate, apex cuspidate to slightly 

caudate; midrib flat to sunken on upper side; lateral veins 30-50 pairs, fine, indistinct, 

running straight to margin where there is a clear marginal vein; stalk 6-10 mm. Flowers 

arranged in a panicle; petals pink to crimson, with punctate glands, basal scale small. 

Fruits 7-9 x 3-4 mm, persistent sepals to half or more than half the capsule length. Seeds 

10-18 per locule, each with a wing all round. 

Vernacular names. Sabah---geronggang, serungan (general). Sarawak--dat (Kayan, Punan 

Tutoh), di'it (Melanau Oya), geronggang (Bidayuh), idat (Melanau Oya), kata mudung 

(Kenyah), labakan (Kelabit), manat (Melanau Rejang), mertilan (lban), serungan labakan 

(Murut, Kenyah), tat (Kenyah). Brunei-geroking (Belait), geronggang (Belait, Dusun, 

Than), labakan (Murut), madak (Tutong). Browne I.e. coins the name geronggang gajah. 

Distribution. S Burma, Sumatra, Peninsular Malaysia, Borneo. In Sabah and Sarawak in 

all districts. Also in Brunei and Kalimantan. 

Ecology. Lowlands principally, also up to lower montane forest at 1400 m. Primary and 

secondary forests, including kerangas (heath) and peat swamp forest, sometimes gregarious, 

and may form thickets in exploited forest areas. 

Uses. The species provides much of the geronggang timber in Sabah and Sarawak. 

2. Cratoxylum cochinchinense (Lour.) Blume 

( ofIndo-China) 

Fig. 1. 

I.e. (1852) 17; Masamune I.e. 477; Shea I.e. 145; Kochummen I.e. 251; Anderson I.e. 217; Corner I.e. 

365; Ashton I.e. 299; Whitmore, Tantra & Sutisna I.e. 181. Basionym: Hypericum eoehinehinense 

Louf., Fl. Cochin. (1790) 472. Type: Loureiro, s.n., "in sy1vis Cochinchinae" (BM). Synonyms: C. 

polyanthum Korth., Verh. Nat. Gesch. Bot. (1842) 175, t. 36; C. myrtifolium Blume I.e. (1852) 17; 

C. ligustrinum (Spach) Blume I.e. (1852) 16. 

223

TREE FLORA OF SABAH AND SARAWAK VOL. 1(1995) 

Tree, sometimes a shrub, to 30 m tall and 65 cm diameter; trunk sometimes with small 

buttresses and sometimes spiny. Bark pale brown to reddish brown, smooth to flaky or 

papery; inner bark greenish yellow to pink, thin. Sapwood white to yellowish. Leaves 

elliptic, 3.5-11 x 1-4 cm, glaucous beneath, chartaceous to thinly coriaceous; base cuneate, 

apex acute; midrib sunken on upper side; lateral veins 9-12 pairs, fine, distinct, not joining 

to form a marginal vein; stalks 2-3 (-10) mm. Flowers I-several in short axillary cymes; 

petals crimson, with linear glands, without basal scale. Fruits 8-12 x 4-5 mm, persistent 

sepals more than half the capsule length. Seeds 5-8 per locule, each with a wing only on 

one side. 

Vernacular names. Sabah-selangan biabas (Suluk). Brunei-machit laling (Belait), 

merlilan (Than), serungan (Brunei Malay), taikakang (Kedayan). 

Distribution. Burma, Indo-China, South China, Sumatra, Peninsular Malaysia, Borneo, 

and the Philippines. In Sabah and Sarawak in all districts. Also in Brunei and Kalimantan. 

Ecology. Lowland and hill forest, in primary or secondary forest including kerangas 

(heath) forest. 

3. Cratoxylum formosum (Jack) Dyer 

(Latin,formosus = beautiful; the flowers) 

in HookerJ, Fl. Br. Ind. 1 (1874) 258; Merrilll.c. 392; Masamune l.c. 477; She a I.c. 146; Browne 

I.c. 197;Andersonl.c. 217; Cornerl.c. 365,Pl. 80; Kochummenl.c. 251; Ashtonl.c. 299; Whitmore, 

Tantra & Sutisna l.c. 181. Basionym: Elodea formosa Jack, Mal. Mise. 2 (1822) 24. Type: Jack, 

S.n., Sumatra (K). 

Tree, sometimes a shrub, to 20 m tall and 30 cm diameter, trunk sometimes spiny. Bark 

grey to red-brown or black, smooth to fissured or scaly; inner bark yellowish. Sapwood 

pale yellowish. Leaves broadly elliptic, 4-15 x 2.5-8 cm, glaucous beneath, chartaceous to 

coriaceous; base cuneate, apex rounded to acute or shortly tipped; midrib sunken on upper 

side; lateral veins 8-16 pairs, coarse, raised, distinct, arching and forming vein-loops 

towards the margin; stalks 5-10 mm long. Flowers single, axillary, opposite; petals white 

to pale lilac, with punctate glands, basal scale large. Fruits 10-16 x 4-6 mm, persistent 

sepals to only a third of the capsule length. Seeds 7-17 per locule, each with a wing on 

one side only. 

Vernacular names. Sabah---geronggang, serungan (generally). Sarawak--dat tetong (Punan 

Tutoh), entemu (Than), kajo jelan (Berawan), melan (Kenyah), mirinos (Bidayoh Sadok), 

nyalin bahe (Kayan), patok tilan (Than), raja tugag (Nidayoh Padawan), sidodot (Bidayoh 

Bau). 

Distribution. Indo-China, S Andarnan Is., Sumatra, Peninsular Malaysia, Java, Borneo, 

Philippines, Celebes. In Sabah and Sarawak in all districts. Also in Brunei and Kalimantan. 

Ecology. Principally lowlands, in primary or secondary forests, including peat swamps. 

Recorded on sea shores, kerangas (heath forest) on white sand, sandstone, alluvium. 

224


limesto ne and ultramafic soils. Trees of this species can be fully deciduous and the 

inflorescences develop on bare parts of branches before and as a new flush of leaves 

develops. Ashton (BRUN 5554) noted this species as flowering gregariously in riverbank 

forest in Brunei. 

4. Cratoxylum gIaucum Korth. 

(Latin, glaucus = pale blue-green; the lower leaf surface) 

I.c. 176; Merrilll.c. 392; Masamune l.c. 477; Anderson I.c. 217; Ashton I.c. 298; Whitmore, Tantra 

& Sutisna l.c. 181. Type: Muller L 904, Karrau, Doeson (lectotype BM; isolectotype W). Synonym: 

C. procerum Diels, Bot. Jahrb. (1926) 311. 

Small tree or shrub, to 10 m tall, rarely to 25 m tall and 45 cm diameter. Bark reddish 

brown, flaky. Leaves broadly elliptic, 2-5 x 1.5-3 cm, glaucous beneath, coriaceous; base 

cuneate, apex rounded, blunt or notched; midrib flat to sunken on upper side; lateral veins 

10-14 pairs, fine, indistinct, running straight toward the margin where there is a clear 

marginal vein; stalks 1-3 mm long. Flowers in a panicle or thyrse; petals crimson, with 

punctate glands, basal scale small. Fruits 7-10 x 3-4 mm, persistent sepals to half the 

capsule length. Seeds 4-8 per locule, each with a wing all round. 

Vernacular names. Sarawak--geronggang (Iban), geronggang lompong (Iban), kirap 

(Bidayoh), pidang (Kenyah), timau (Iban). Brunei--geronggang timau (Iban). Browne I.c. 

coins the name geronggang puteh. 

Distribution. Sumatra (Lingga, Bangka and Billiton), Peninsular Malaysia (Johore), 

Karimata and Natuna Islands, Borneo (Wand SE). In Sarawak in coastal areas throughout 

the state. Also in Brunei and Kalimantan. 

Ecology. Found in lowland primary and secondary kerangas (heath) forest on white sand, 

in peat swamps (especially those dominated by the dipterocarps Dryobalanops rappa and 

Shorea albida), and also freshwater swamps. Also documented at 1100 m in mossy 

kerangas forest on sandstone ridges. 

5. Cratoxylum maingayi Dyer 

(Ae. Maingay, 1836-1869, botanist of the East India Company) 

I.c. 258; Browne I.c. 197; Anderson l.c. 217; Corner I.c. 367; Kochummen I.c. 251; Ashton l.c. 300; 

Whitmore, Tantra & Sutisna I.c. 181. Type: Maingay, s.n., Penang (holotype BM; isotype L). 

Synonym: C. cochinchinense var. calcareum Ridl., Kew Bull. (1938) 115. 

Tree or shrub, to 10 m tall and 10 cm diameter, often crooked. Bark brown, smooth to 

fissured or flaky. Leaves broadly elliptic, 1.5-4.5(-5.5) x 0.8-3 cm, coriaceous; base 

cuneate, apex rounded, notched or obtuse with a short tip; midrib flat to sunken on upper 

side; lateral veins 5-6 pairs, raised, distinct, arching and forming vein-loops towards the 

margin; stalks to 5 mm long. Flowers single, axillary, opposite; petals pale pink, with 

225


punctate glands, basal scale large. Fruits 6-15 x 3-5.5 mm, persistent sepals nearly half 

the capsule length. Seeds 5-6 per locule, each with a wing on one side only. 

Vernacular names. Sarawak---gerunggang (Iban), patok ti/an (Iban). 

Distribution. Indo-China, Sumatra, Peninsular Malaysia, Borneo. In Sarawak, documented 

from the 1st Div. near Kuching only; also reported in Kalimantan. 

Ecology. Lowlands, known only from limestone substrates. Trees of this species are often 

bare of leaves for a short time, when the inflorescences develop on bare parts of branches. 

6. Cratoxylum sumatranum (Jack) Blume 


l.c. (1852) 16; Kochummen l.c. 251; Shea I.c. 146; Anderson I.c. 218; Ashton I.c. 301; Whitmore, 

Tantra & Sutisna I.c. 181. Basionym: Elodea sumatrana Jack I.c. 22. Type: Jack, s.n., Sumatra, 

Tello Dalam (K). Synonyms: C. hypericum (Blume) MeIT. I.c. 392; C. celebicum Blume I.c. (1852) 

16. 

Tree or shrub, to 35 m tall and 60 cm diameter, trunk sometimes with buttresses to 1 m 

high. Bark grey to pale or dark brown, cracking-fissured to scaly; inner bark pale yellow to 

reddish, soft. Sapwood pale yellow to white. Leaves ovate-elliptic, 4.5-17 x l.5-4.5 cm, 

chartaceous to thinly coriaceous; base cuneate, apex acute; midrib flat to sunken on upper 

side; lateral veins 9-22 pairs, coarse, raised, distinct, not joining to form marginal vein; 

stalks 2-3 mm long. Flowers in a panicle; petals crimson, with linear glands, basal scale 

absent. Fruits 7-10 x 3-5 mm, persistent sepals less than half the capsule length. Seed 3- 

10 per locule, each with a wing on one side only. 

Vernacular names. Sabah-serungan (Malay). Brunei-serungan mampat (Malay). 

Distribution. Southeast Asia, Sumatra, Java, Borneo, Philippines, Celebes, Lesser Sunda 

Is. In Sabah and Sarawak in all districts. Also in Brunei and Kalimantan. 

Ecology. Lowlands, in primary and secondary forest, documented on sandstone, white sand 

in kerangas (heath) forest, and also on ultramafic soils. Rarely found up to 1300 ID. 

Taxonomy. The Bornean taxon is subsp. sumatranum; subsp. blancoi (Blume) Gogelein is 

restricted to the Philippines. 

226


ILLICIACEAE 

Richard M.K.Saunders 

Department of Ecology & Biodiversity, 

University of Hong Kong 

AC. Smith, Sargentia 7 (1947) 8; Hutchinson, Fam. Fl. PI. 1, 2nd ed. (1959) 125; Gen. FI. PI. 1 

(1964) 57; Ng, TFM 2 (1973) 253. 

The family contains only one genus, Illicium, which has been the source of considerable 

discussion regarding its evolutionary relationhips. Its affinities with the Magnoliales have 

long been recognised, and this has been reflected historically by its classification in both the 

families Magnoliaceae (e.g., Bentham & Hooker, Gen. PI. 1 (1862) 16) and Winteraceae (e.g., 

Ridley, FMP 1 (1922) 18). The most recent comprehensive treatment of the genus is the 

monograph by A.C. Smith (Sargentia 7 (1947) 1), who proposed that it should be isolated 

as the family Illiciaceae on the basis of various morphological and anatomical criteria, 

discussed in detail by Bailey & Nast (J. Am. Arb. 26 (1945) 37; I.c. 29 (1948) 77), and Keng 

(Bot. Bull. Acad. Sin. 6 (1965) 61). The IIliciaceae bears the closest relationship to the 

Schisandraceae, a small family of scrambling and twining woody vines. The isolated 

evolutionary position of these two families has been recognised more recently by their 

classification as the sole members of the order Illiciales (e.g., Takhtajan, Bot. Rev. 46 (1980) 

225; Cronquist, Integr. Syst. FI. PI. 1981); it is generally agreed, however, that the Illiciales are 

derived from the Magnoliales, and are probably from the same stock as the Winteraceae. 

ILLICIUM L. 

(Latin, illicere = an attractant, probably in reference to the presence of aromatic oils) 

Syst. Nat. ed. 10 (1759) 1050; Stapf, Trans. Linn. Soc. London Ser. 2,4 (1894) 128; Ridley, FMP 1 

(1922) 18 (under Winteraceae); AC. Smith l.c. 10; Ng, I.c. 253. 

Shrubs or small to medium-sized trees; evergreen, glabrous, aromatic with scattered ethereal 

oil cells. Leaves simple, entire, alternate often clustered to give appearance of whorls of 

3-6 at distal nodes; exstipulate; petioles with groove on adaxial surface; lamina papery to 

leathery, venation pinnate; base generally attenuate, decurrent, apex generally acuminate. 

Flowers solitary or in clusters of 2-3, mostly axillary, sometimes borne on the stem; 

bisexual, regular; perianth not differentiated into sepals and petals, parts numerous (7- 

33), free, overlapping, spirally arranged, white, cream, pink, red or purplish; androecium of 

(4-) numerous (up to c. 50) stamens, spirally arranged in one to several series, filaments 

short, thick; anthers basifixed, introrse-lateral, dehiscing by longitudinal slits; gynoecium 

of (5-) 7-15(-21), free, superior carpels, arranged in a single whorl, obliquely attached to 

receptacle; carpel unilocular, with single, near-basal, anatropous, bitegmic (with 2 layers 

of integument), crassinucellar (with a thick nucellus) ovule, stigma dry, nonpapillate, decurrent. 

227


3cm 

2mm 

lcm 

Fig. 1. Illicium stapfii. A, leafy twig; B, flower with proximal perianth-parts and stamens removed; 

C, stamens; D, lateral view of carpel; E, fruit; F, seed. (A from Carr 26441, B from Beaman 9959, 

C-D from Vermeulen & Duitermaat 1002, E-F from Anderson 4554.) 

228

ILLICIACEAE (SAUNDERS) 

Fruits of single-seeded follicles, star-shaped, green (ripening red), splitting along ventral 

edge oI each segment when ripe. Seeds solitary in each segment, glossy, brown, with 

copious oily endosperm. 

Distribution. A medium-sized genus, with a disjunct distribution in Southeastern North 

America (5 species) and eastern Asia (c. 35 species). In Asia the distribution extends from 

southern Japan to northern Sumatra and Borneo, and from Assam to the Philippines, and 

the centre of diversity lies in northern Burma and southern China (Yunnan, Guangxi and 

Guangdong). In Borneo, it is restricted to the cooler montane regions, where two species, 1. 

kinabaluense and 1. stapfii, are known. 

Ecology. Vegetative growth is markedly discontinuous, with periods of dormancy of 

vegetative buds alternating with active growth (Ng I.e.). The resumption of growth involves 

the rapid elongation of a bud to produce a stem several centimetres long, which bears small 

caducous leaves; apical "pseudowhorls" of leaves are then produced, consisting of alternately 

arranged normal leaves that are tightly clustered together. Studies in the reproductive 

biology of North American species have shown that they are pollinated by a wide variety of 

small insects, but primarily Diptera, and that a gametophytic self-incompatibility system 

operates (Thien et al., Amer. 1. Bot. 70 (1983) 719; White & Thien, J. Elisha Mitchell Sci. 

Soc. 101 (1985) 15). A system of ballistic seed dispersal (autochory) is apparent in the 

genus (Roberts & Haynes, PI. Syst. EvoI. 143 (1983) 227), although its efficacy has not 

been demonstrated in the Bornean species. 

Uses. The fruit of 1. verum Hook. f from China and Indo-China is the source of the spice 

Chinese Star Anise, used for flavouring food and liqueurs. Although this species does not 

occur in Borneo, the spice has been imported extensively from China and is traded in 

Malaysia as bunga lawang or adas china (Burkill, EPMP 2 (1966) 1244). The fruits of the 

Japanese species 1. anisatum L. (syn. 1. religiosum Sieb. & Zucc.) are poisonous, although 

small quantities can be used for flavouring, and are sometimes retailed in SE Asia (Burkill 

I.e.). Other Illicium species have various reported medicinal properties, often as stomachics, 

carminatives, stimulants or vermifuges (Perry, MPESE (1980) 180). The timber is of very 

limited value due to the small size of the trees. 

Taxonomy. The last comprehensive revision of the genus was the monograph by A.C. 

Smith (I. c.), in which 42 species were recognised. These species were classified into two 

sections, viz. sect. Badiana Spach (which includes the type species and should therefore 

bear the autonym sect. Illicium), and sect. Cymbostemon (Spach) A.c. Srn.; the Bornean 

species belong to the latter section. Smith recognised three species in Borneo (1. kinabaluense, 

1. stapfii and 1. cauliflorum); the acceptance of these species was based on the examination 

of only 11 herbarium sheets, however, and evidently requires re-evaluation. Only two 

species are accepted in the present treatment. 

Key to Illicium species 

Leaves (5-)6-7.5(-11) x (1.5-)2-3(-4) cm. Stamens 7-8. Carpels 8 ......... 1. I. kinabaluense 

Leaves (6-)10-11.5(-16.5) x (2-)4-5(-8.5) cm. Stamens 9-14 (-20). Carpels 8-l3 ................ . 

.......................................................................................................................... . 2. I. stapfii 

229


1. Illicium kinabaluense A. C. Srn. 

(of Mt. Kinabalu, Sabah) 

I.e. 61. Type: 1. & M.S. Clemens 50154, British North Borneo, Mt. Kinabalu, Penibukan (holotype 

A; isotype L). 

Small tree, to 15 m tall, 45 cm diameter. Leaves clustered in ''pseudowhorls'' of up to 6 

leaves; lamina elliptic, c. (5-)6-7.5(-11) x (1.5-)2-3(-4) cm; base attenuate, margins slightly 

revolute, apex (short-)acuminate; midrib slightly impressed above and prominent below; 

lateral veins 5-8 per side; petioles 9-17 mm long, grooved on adaxial surface. Flowers 

axillary or subterminal, generally solitary; pedicels 10-20 mm long at an thesis; perianth 

parts 8-10, pink, red or purplish, outermost ones 4.5-6 x 3-3.5 mm, largest 4-7 x 3-5 

mm, innermost 3.5-5 x 2 mm; androecium of 7-8 stamens, uniseriate, stamens l.7-2.5 

mm long; gynoecium of8 carpels, carpels l.6-2 mm long. 

Vernacular name. Sabah-longugan (Dusun). 

Distribution. Restricted to Mt. Kinabalu, Sabah. 

Ecology. Primary forests at 1200-2000 m. 

2. Illicium stapfii Merr. Fig. 1. 

(Otto Stapf, 1857-1933, botanist at Kew in charge of "Indian" collections) 

Philip. J. Sci. Bot. 13 (1918) 67, EB (1921) 252; Masamune, EPB (1942) 278; A.C. Smith I.e. 65; 

Anderson, CLTS (1980) 346. Type: MS. Clemens 10995, British North Borneo, Mt. Kinabalu, 

Marai Parai Spur (PNH). Synonyms: lllieium sp. Stapf I.e. 128; 1. eauliflorum Merr., Sarawak Mus. 

J. 3 (1928) 522, Masamune I.e. 277. 

Medium-sized tree, to 25 m tall, 80 cm diameter. Leaves clustered in ''pseudowhorls'' of up 

to 6 leaves; lamina elliptic, (6-)10-11.5(-16.5) x (2-)4-5(-8.5) cm; base obtuse to attenuate, 

margins markedly revolute, apex (short-) acuminate, acute or obtuse; midrib markedly 

impressed above and prominent below; lateral veins 5-12 per side; petioles 10-35 mm 

long, grooved on adaxial surface. Flowers axillary or subterminal, occaSionally borne on 

the bare branches, solitary; pedicels up to 40 mm long at anthesis; perianth parts 9-15, 

pink, red or purplish, outermost ones 5-8.5 x 4-5 mm, largest 5-9.5 x 3.5-6 mm, innermost 

(3.5-)7-7.5 x 2.5-4.5 mm; androecium of 9-14(-20) stamens, uniseriate, stamens 2-3.5 

mm long; gynoecium of 8-13 carpels, carpels 2-5 mm long. 

Vernacular names. Sarawak--ala (Than), bowlong (Kenyah). 

Distribution. Endemic to Borneo. Montane regions, from Mt. Kinabalu (Sabah) in the 

north to the Linau-Balui Plateau (Sarawak) in the south; also reported from northern 

Kalimantan. 

Ecology. Primary forests at 800-2000 m. 

230

ILLICIACEAE (SAUNDERS) 

Taxonomy. Merrill (I.e. 1918) and AC. Smith (I.e.) recognised 1. eauliflorum as a distinct 

species largely on the basis of its "cauliflorous" habit and greater number of carpels and 

stamens. Only the holotype was available to these authors for examination (E. Mj6berg 

J J 4, Mt. Murud, Sarawak (UC)). This specimen is recorded to have flowered at a height of 

only 1 rn, and would more correctly be referred to as "ramiflorous". The study of numerous 

collections of 1. stapfii now available indicates that no clear distinction can be drawn 

between ramiflory and the formation of flowers on young growth, suggesting that it is a 

poor taxonomic character for the genus. The greater number of carpels and stamens in 1. 

eauliflorum is furthermore of little significance, since these characters are considerably 

more variable in 1. stapfii than observed by Smith (l.e.). There do not appear to be any 

convincing criteria for retaining 1. eauliflorum as a distinct species. 

231


JUGLANDACEAE 

E.J.F. Campbell-Gasis 

c/o Forest Research Centre, 



Lesch. ex Blume, Bijdr. 10 (1825) 528; Blume, Ft. Jav. Jugl. (1829) 5; Merrill, EB (1921) 210, 

Enum. Philip. PI. 2 (1923) 23; Ridley, FMP 3 (1924) 368; Masamune, EPB (1942) 231; Dilmy, 

Rimba Indonesia 4 (1955) 29; Jacobs, FM 1,6 (1960) 143; Backer & BakhuizenJ, FJ 2 (1965) 158; 

Burgess, TBS (1966) 321; Manning, Bull. Torrey Bot. Club 93 (1966) 34; Whitmore, TFM I (1972) 

233; Cockburn, TFS 1 (1976) 151; Anderson, CLTS (1980) 220; Ashton, MNDTS 2 (1988) 303; 

Corner, WSTM 1 (1988) 367; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 185. 

Trees with aromatic tissues; leaves, young twigs and reproductive parts mostly lepidote 

with golden yellow glandular scales. Stipules none. Leaves spirally arranged, paripinnate; 

leaflet blade often slightly asymmetric. Inflorescence a panicle of catkins, axillary or 

occasionally terminal, with flowers of one sex or both sexes in the same inflorescence; male 

catkins usually pendulous; female catkins erect at least initially. Flowers unisexual, in axils 

of 3-lobed bracts; perianth 4-lobed; male: perianth often reduced or irregular, stamens 4- 

13 (in Asiatic species) and sessile or on short filaments, anthers longitudinally dehiscing; 

female: perianth lobes in 2 whorls, partly connate with the 2-carpellate ovary; carpels 1- 

locular with an incomplete transverse septum, ovule 1; style 1, short or absent; stigmas 2(- 

4) in Engelhardia, persistent, papillose, or 4, sessile. Fruit a drupe or in Engelhardia a 

small nut, attached to an enlarged wing-like 3-4-Iobed bract. Seed 1, without endosperm; 

cotyledons often much contorted. 

Distribution. 7 genera and about 60 species, chiefly in north temperate regions in both the 

Old and New World, with a few species in the tropics. In Sabah and Sarawak, represented 

by one genus, Engelhardia, with 9 species. 

Uses. Walnut (Juglans) and hickory (Carya) are important north temperate timber genera. 

The family is also well known as the source of edible nuts, walnuts (Juglam;) and pecan 

(Cmya). In Sabah and Sarawak, the timber of Engelhardia is too rme to be of much 

commercial importance (see under genus). 

Taxonomy. The family seems most closely allied to the Myricaceae \\"ithin the Amentiferae. 

ENGELHARDIA Lesch. ex Blume 

(N. Engclhardt, 1761-1831. a Governor in Java) 

sal1sal1glal1g (Iban. Sarawak), lal1sal1g/al1g (Malay_ Sarcl\\"ak) 

I. e. (1825) 528. I. e. (1829) :5, "ElIgelhardlia"~ Mcrrill I. e. (1921) 210, I. c. (1 923) 23~ Ridky I. c. 3()8~ 

Masamunc I.e. 23 L Dihu\" I.e. 29~ Jacobs I.e. 143~ Backcr & Bakhuizcn/ I.c. 158~ Burgcss I.c. 32 L 

233


Manning l.c. 34; Whitmore I.c. 233; Coekbum l.c. 151; Anderson I.e. 220; Ashton I.e. 303; Corner 

I.e. 367; Whitmore, Tantra & Sutisna l.e.185; Campbell-Gasis, Sandakania 3 (1993) 1. Synonyms: 

"Engelhardtia" C. DC., Ann. Se. Nat. 4, 18 (1862) 35; Pterilema Reinw., Syll. 2 (1826) 13; 

Oreomunnea Oerst., Yid. Medd. Nat. For. Kjobenh. (1856) 52. 

Small to medium-sized evergreen trees to 35 m (exceptionally to 50 m) tall, to 150 cm 

diameter; bole columnar at first but later often of poor form with short flutes; buttresses 

thin, small, but sometimes large, steep, thick and branching; crown generally oblong to 

hemispherical, rather obtuse, with somewhat twisted branches. Bark from pale grey 

through brown-grey to red-brown, smooth to fissured, peeling off in small, rectangular, 

papery flakes, often purplish; inner bark from yellow to pinkish to brown, fibrous. Sapwood 

white to yellow, soft to hard. Young twigs, leaves and inflorescences covered in golden or 

orange scurfy scales and pale yellow and orange hairs (visible with a XI0 hand lens). Twigs 

often with conspicuous leaf-scars and cream to orange lenticels. Leaves often with a small 

protuberance at the apex of the rachis; leaflets 2-7 pairs, alternate to subopposite, sessile 

or stalked; base often unequal; veins often faintly looping near margin; above glabrous to 

scaly, below variously scaly and hairy, vein axils often with tufted domatia (visible with a 

XlO hand lens). Inflorescences so far known with flowers of only one sex. Flowers 4- 

merous, tiny, often yellow or green, perianth inconspicuous; male: stamens 4-13, attached 

to the fused perianth-bract structure; female: perianth partly connate with the ovary and 

fused to a small one-sided 3-lobed bract, ovule erect and conical with a broad base. Fruits 

attached to a 3-4-lobed bract; bract membranous, dry; the abaxial and lateral wings with 

midribs and reticulate venation, the abaxial wing the largest, a smaller adaxial bract often 

present; nut small, attached to base of bract; pericarp set with scales or hairs, or both. 

Distribution. 12 species, 3 of which are found in the Americas (Mexico, Costa Rica and 

Guatemala), the other 9 in the Old World from the western Himalayas to south China and 

extending southwards across Malesia to New Guinea. All 9 species are found in Sabah and 


Ecology. Occurring as scattered individuals in lowland mixed dipterocarp forest below 

2000 m, on leached acid yellow soils, but becoming common though never dominant at 

higher elevations (up to 2700 m) in oak-laurel forest on more fertile soils and on ultramafic 

and basaltic soils. Often shortly deciduous and then flowering. The flowers in catkins 

suggest wind pollination but the male catkins of E. roxburghiana have been noted to have a 

slight fragrance and may therefore be insect-pollinated also. The fruits are at least partially 

dispersed by wind. The seedlings often have a terminal leaflet which, as the trees become 

older, is reduced until all that is left in the adult is a small terminal protuberance. Saplings 

and pole-sized trees tend to have leaves with more pairs of leaflets, which are usually 

thinner, longer acuminate, sometimes serrate and often more densely hairy than those of 

adult trees. Sterile trees may be mistaken for sapindaceous species because of the similarity 

in their pinnate leaves. 

Uses. The wood is locally used for timber but is of inferior quality, being not durable and 

only of use under cover. The wood is rather soft to moderately hard, light to moderately 

heavy, with a straight grain, shallowly interlocked, or slightly wavy. The texture is 

moderately fine to slightly coarse and even. The timber is easy to work. The bark, containing 

tannin-like compounds, is occasionally used for fish-poison in Sumatra. 

234

JUGLANDACEAE (CAMPBELL-GASIS) 

Key to Engelhardia species 

(based on shoot and leaflet characters) 

1. Leaflet margins distinctly toothed in some part ............................................................ 2 

Leaflet margins entire or wavy, not toothed .............................................................. 3 

2. Distal leaflets often twice as large as the lower ones; stalk nil to 1 mm, rarely more. 

Leaflet base acute or wedge-shaped; intercostal veins and areoles mostly invisible on 

the upper surface in dried specimens ......................................................... 8. E. serrata 

Distal leaflets sometimes larger than lower ones but not up to twice as large; stalk 

longer, to 4 mm. Leaflet base rounded to subcordate; intercostal veins and areoles 

distinct on the upper surface in dried specimens ...................................... 1. E. apoensis 

3. Leaflets coriaceous, elliptic-ob ovate (or if lanceolate-falcate then with veins impressed 

on the upper side), and not hairy on the lower side ...................................................... .4 

Leaflets chartaceous, or lanceolate-falcate, or with hairy veins on the lower side (the 

veins not impressed on the upper side) .......................................... , .............................. 6 

4. Leaflet intercostal venation coarse, the areolation strongly prominent under XI0 

magnification .................................................................................. 3. E. kinabaluensis 

Leaflet intercostal venation finer, the areolation only faintly visible under XIO 

magnification .............................................................................................................. 5 

5. Shoot-apices and leaf rachises brown scurfY hairy ....................................... 6. E. rigida 

Shoot-apices and leaf rachises not scurfY hairy ................................. S. E. mersingensis 

6. Leaflets not decreasing markedly in size from apex to base of leaf rachis; veins on 

lower side hairy ......................................................................................... 9. E. spicata 

Leaflets decreasing in size by one third to half from apex of leaf rachis; veins on lower 

side minutely scaly to sub glabrous or apparently glabrous ............................................ 7 

7. Leaflets elliptic, papery, base only slightly unequal, margin not thickened; stalk almost 

nil or only to 1 mm long ..................................................................... 2. E. danumensis 

Leaflets ovate to lanceolate or falcate, leathery, base very unequal, margin thickened; 

stalk 1-10 mm long ..................................................................................................... 8 

8. Leaflets to 11 cm long, usually 4-5 pairs per leaf; stalk 1-4 mm long ........................... . 

................................................................................ 4. E. mendalomensis 

Leaflets usually larger, to 16 cm long, usually 2-3 pairs per leaf; stalk longer, to 10 

mm ................................................................................................. 7. E. roxburghiana 

Key to Engelhardia species 

(based on fruit characters) 

1. Nut 6-14 mm long with a conspicuously narrowed apex .......................... 1. E. apoensis 

Nut shorter, to 6 mm long, apex rounded ..................................................................... 2 

235


«- [ 

1: 

Fig. 1. Engelhardia danumensis. Fruiting leafy twig. (From SAN 85066.) 

236


2. Adaxial lobe ofbract not developed, the nut fully exposed ........................................... 3 

Adaxial lobe ofbract well-developed, hiding most of the nut... ................................... .4 

3. Nut-stalk 4-8 mm long. Nut covered with scales only. Perianth-lobes broadly ovate 

..... 7. E. roxburghiana 

Nut-stalk shorter, to 3 mm long. Nut hairy. Perianth-lobes linear ..... 5. E. mersingensis 

4. Perianth of inconspicuous tiny lobes adnate to the style. E. spicata 

Perianth of ovate to linear lobes at the apex of the .5 

5. Nut set with stiff, loose, irritant long hairs ................... : ............................. 8. E. serrata 

Nut indumentum not so ............................................................................................... 6 

6. Adaxial lobe ofbract cleft in the middle ....................................................................... 7 

Adaxial lobe ofbract not cleft in the middle ................................................................. 8 

7. Adaxial lobe ofbract only shallowly cleft. Perianth-lobes broadly ovate .... 2. E. danumensis 

Adaxial lobe ofbract deeply cleft to the base. Perianth-lobes linear ....... 3. E. kinabaluensis 

8. Adaxial lobe ofbract brbicular... ................................................................. 6. E. rigida 

Adaxial lobe ofbract subtruncate, the margin sometimes wavy ..... 4. E. mendalomensis 

1. Engelhardia apoensis Elmer ex Nagel 

(of Mount Apo in the Philippines) 

Bot Jahrb. 50 (1914) 477; E1mer, Leafl. Philip. Bot. 7 (191S) 2693; Merrilll.c. (1923) 23; Jacobs I.c. 

lSl; Burgess I.c. 321; Manning I.c. 48; Whitmore I.c. 234; Cockburn I.c. lS2; Ashton I.c. 304; 

Corner I.c. 368; Whitmore, Tantra & Sutisna I.c. 18S. Type: Elmer 11744, Philippines, Mindanao, 

Mount Apo (holotype NY; isotypes BM, BO, GB, L, P). 

Tree to 50 m tall and 150 cm diameter; bole often straight; buttresses to 4 m tall and 1.5 m 

out; crown spreading to dense. Bark grey-brown to reddish brown; inner bark pink with 

white wedges to purplish and reddish brown, soft. Sapwood pale yellow to white. Twigs, 

leaves and inflorescences set with golden to orange scales and pale yellow hairs. Leaf 

rachis (8-)13-23(-32) cm long, 3.5-11 cm to first leaflet, thickened at base, densely to 

occasionally covered in scales and occasional hairs; leaflets (3-)4-5( -7) pairs, alternate, 

ovate to lanceolate, (5-)9-13(-16) x (2.2-)3-4(-6) cm, gradually decreasing a little in size 

towards base, drying orange-brown to dark chocolate-brown below, thinly leathery, upper 

side with occasional scales and hairs, lower side with occasional to dense scales, only 

midrib and lamina hairy; base rounded to subcordate, unequal, margin toothed irregularly, 

at least in lower half, often inrolling, apex acute; tufted domatia occasional to frequent in 

vein axils with midrib; lateral veins 9-16(-18) pairs, above and below visible; intercostal 

veins net- to ladder-like, above visible, below sometimes invisible; petiolule (1-)2-3(-4) 

mm long, with dense to occasional scales and hairs. Inflorescences axillary, scaly and 

hairy; male catkins 9-20 cm long, peduncle densely scaly and hairy; stamens 9, anthers 

scaly and hairy; fruiting catkins (8-)12-19(-30) cm long, peduncle and rachis scaly and 

hairy. Fruits sessile to 3-mm-stalked; bracts with hairs and scales, abaxial wings 55-75 x 

12-15 mm, lateral wings 28-35 x 8 mm, adaxial lobes entire to shallowly 4-lobed, hiding 

top of nut; nuts ovoid, 6-14 x 5-6 mm, densely covered in long, soft hairs, without scales; 

perianth-lobes adnate to the style, tiny, inconspicuous, not hiding the stigmas. 

237


Distribution. Peninsular Malaysia, Borneo, and the Philippines. In Sabah, an uncommon 

tree known on Mount Kinabalu only (e.g., SAN 27565, SAN 22408); uncommon also in 

Sarawak, known only on Mount Dulit (Richards 1521). Also in E Kalimantan. 

Ecology. Primary mixed dipterocarp forest from 700 to 1300 m, on fertile and ultramafic 

soils. 

2. Engelhardia danumensis Carnpbell-Gasis 

(of the Danum Valley area in Sabah) 

Fig. 1. 

Sandakania 3 (1993) 6. Type: Cockbum SAN 85066, Sabah, Lahad Datu, Ulu Segama Forest Reserve, 

Ulu Sungai Segama (holotype SAN; isotypes A, Ba, FHO, K, KEP, L, SAR, SING). 

Tree to 35 m tall and 60 cm diameter; buttresses thin, plank-like, c. 1 m tall and 2 m out. 

Bark dark chocolate-brown; inner bark orange-brown, cut wood and bark smelling like 

coconut water. Sapwood white, hard. Twigs and leaves covered with golden scales but 

without hairs. Leaf rachis 4-10 cm long, 2.8-4.2 cm to first leaflet, blackish, not thickened 

at base, scales occasional; leaflets 2-3 pairs, alternate to subopposite, elliptic, 2.4-7 x 1. 5- 

1.5 cm, decreasing to about half the size towards base, drying dark brown to blackish 

below, thin, papery; upper side glabrous, lower side with occasional to scattered scales all 

over; base acute to wedge-shaped, subequal, margin entire, undulate, marginal vein not 

thickened, apex acute to shortly acuminate, acumen to 5 mm long; tufted domatia 

occasional in vein axils with the midrib; lateral veins 8-12 pairs, raised on both sides, 

conspicuous; intercostal veins net-like, obscure on both sides, areoles obscure; petiolule 

absent to 1 mm long, glabrous or with occasional scales. Inflorescences axillary, flowers 

not known; fruiting catkins 14-18 cm long, peduncle and rachis slender, angular, with 

golden scales and hairs. Fruits sessile to I-mm-stalked; bracts covered in scales and a few 

hairs, abaxial wings 19-30 x 6-9 mm, lateral wings 8-11 x 3-4mm, adaxial lobe shallowly 

cleft in the middle, often hiding the top of the nut; nuts globose c. 3 mm across, covered in 

golden scales and short golden hairs; perianth-lobes on the nut apex ovate, free from style; 

not hiding the stigmas. 

Distribution. Endemic to Sabah. Uncommon, and known only from the type collection. 


Vegetatively it approaches E. rigida but the leaflets almost blackish on drying, have a thin 

and papery texture, the lower leaflets decrease in size by half, and the lamina is set with 

scales but never hairs. Also, the nut in this species differs in being set with both scales and 

. short hairs. 

3. Engelhardia kinabaluensis Carnpbell-Gasis 

(of Mount Kinabalu in Sabah) 

l.c. 7. Type: Gibot SAN 66824, Sabah, Ranau, Mamui Copper Mine (holotype SAN). 

Tree to 24 m tall and 35 cm diameter. Bark grey to blackish; inner bark yellow to pale 

white, exudate yellowish. Sapwood cream. Twigs, leaves and inflorescences set with golden 

238


scales but without hairs. Leaf rachis 3.7-8 cm long, black, 1.5-3.5 cm to first leaflet, 

thickened at base, occasionally with scales; leaflets 2-3 pairs, subopposite, elliptic to 

obovate orjalcate, 1.5-6.2 x 0.7-2.8 cm, not decreasing in size towards base, drying greybrown 

to orange-brown below, leathery, upper ~ide glabrous, lower side glabrous to 

sparsely scaly all over; base acute to round, very unequal, margin entire and wavy with 

thickened marginal vein, apex acute to shortly acuminate, acumen to 2 mm long; tufted 

domati:Cl absent to rare in vein axils with midrib; lateral veins 5-7 pairs, flat to occasionally 

impressed above, often obscure; intercostal veins net-like, coarse, areoles visible and 

strongly prominent under XlO magnification; petiolule 2-3 mm, glabrous or occasionally 

with scales. Inflorescences axillary; flowers not known; fruiting catkins 7-10 cm long, the 

rachis 5caly and hairy. Fruit stalk 1-2 mm long; bract occasionally scaly, hairs absent, 

abaxial wings 20-24 x 5-6 mm, lateral wings 8-12 x 2-3 mm, adaxial lobes deeply cleft in 

the middle, not hiding the top of the nut; nuts transversely ellipsoid, 2-3 x 3-4 mm, with 

golden scales and short, golden hairs; perianth-lobes on the nut apex linear, free from 

style, not hiding the stigmas. 

Distribution. Uncommon. In Sabah only known on or near Mount Kinabalu (at the Mamut 

Copper Mine and in the Mesilau Area ofKundasang; e.g., SAN 74274, SAN 111603, besides the 

type). 

Ecology. Lowland mixed dipterocarp forest and submontane forest from 200 m to 1500 m. 

Vegetatively, it approaches E. rigida but the leaflets with their thickly, leathery texture, the 

cover of scales on the leaflets and nuts, and the lack of hairs on the leaflets distinguish it 

from that species. 

4. Engelhardia mendalomensis Campbell-Gasis 

(of Mendalom Forest Reserve in Sabah) 

I. c. 7. Type: Fidi/is SAN 116714, Sabah, Tenom District, Mendalom Forest Reserve (holotype SAN). 

Tree to 35 m tall and 80 cm diameter; buttresses to 3 m high and 4 m out. Bark pale 

brown-grey; inner bark pale yellow to orange-brown. Sapwood cream-white to yellowish. 

Twigs and leaves covered in golden scales, without hairs. Leaf rachis 6-14 cm long, 2.2-6 

cm to first leaflet, blackish, thickened at base, occasionally with scales and scant hairs; 

leaflets (3-)4-5 pairs, subopposite, ovate to lanceolate or falcate, 4-11 x 2-4 cm, often 

decreaSing to about two-thirds to half the size towards base, drying dark brown to blackish 

below, thinly leathery, occasionally with scales but no hairs all over on both sides; base 

acute or rarely round, very unequal, margin entire, slightly wavy, with thickened marginal 

vein, apex acute to long-acuminate, acumen to 10 mm long; tufted domatia occasional to 

absent in vein axils with midrib; lateral veins 7-11 pairs, above raised but rather obscure, 

below raised, conspicuous; intercostal veins net-like, areoles on both ~ides obscure; 

petiolules 1-4 mm long with scattered scales. Inflorescences axillary, flowers not known; 

fruiting catkins 13-30 cm, peduncle and rachis quite stout, angular, drying blackish, scaly 

and hairy. Fruits sessile to I-mm-stalked; bracts with scales and hairs, abaxial wing 21-50 

x 8-10 mm, lateral wings 11-18 x 4-6 mm, adaxial lobes sub truncate (the margin 

sometimes wavy), often hiding the top of the nut; nuts transversely ellipsoid, 3-4 x 4-5 

mm, set with golden scales and scattered, short golden hairs; perianth-lobes on the nut 

apex ovate, free from style, not hiding the stigmas. 

239


Distribution. Uncommon. In Sabah, only known from the type collection. 

Ecology. Primary mixed dipterocarp forest from sea-level to 550 m. 

Vegetatively it approaches E. roxburghiana but differs in having four to five pairs of 

leaflets drying dark brown to blackish, with fewer pairs of veins and being less densely set 

with scales. Also, the sessile to I-mm-pedicellate nuts set with both hairs and scales 

distinguish it from that species. 

5. Engelhardia mersingensis Campbell-Gasis 

(of Mount Mersing in Sarawak) 

l.c. 8. Type: Ashton S. 16724, Sarawak, Bukit Mersing, Ulu Anap (holotype SAR; isotypes A, BO, 

K, KEP, L, MEL, SAN, SING). 

Tree to 30 m tall and 70 cm diameter; buttresses tall, sinuate and branching. Bark pale 

orange-brown, thin, powdery. Twigs, leaves and inflorescences set with golden and orange 

scales, but without hairs. Leaf rachis 4.5-14 cm long, 2.5-4.5 cm to first leaflet, black, 

thickened at base, occasionally with scales, without hairs; leaflets 3-4 pairs, alternate to 

opposite, elliptic to obovate, 4.5-11 x 2.5-4.8 cm, drying orange-brown to blackish below, 

chartaceous, upper side glabrous, lower side occasionally scaly all over; base acute to 

round, subequal to very unequal, margin entire, slightly wavy, with thickened marginal 

vein, apex acute to acuminate, acumen to 9 mm long; tufted domatia occasionally in vein 

axils with midrib; lateral veins 7-12 pairs, impressed or keeled above, below raised; 

intercostal veins net-like, areoles only slightly visible under XIO magnification; petiolules 

1-2 mm long, with scattered scales. Inflorescences terminal or axillary, flowers not 

known; fruiting catkins 9.5-20 cm long, peduncle and rachis quite stout, angular, drying 

dark brown or blackish, scaly and hairy. Fruits sessile to 0.5-mm-stalked; bracts glabrous 

to scaly, without hairs, abaxial wings 20-31 x 7-10 mm, lateral wings 9-16 x 4-6 mm, 

adaxial lobes not developed and not hiding the nut; nuts transversely ellipsoid, 3 x 4-6 

mm, with scales and a few short hairs present; perianth-lobes on nut apex linear, free from 

style, not hiding the stigmas. 

Distribution. Uncommon. In Sarawak known only from the type collection. Also in East 

Kalimantan (on Gunung Beratus). 

Ecology. Primary mixed dipterocarp to submontane forest on basaltic soils, at 600-1000 m. 

Vegetatively, this species approaches E. rigida but the larger leaflets set with scales but not 

hairs, fewer pairs of lateral veins, and a sessile nut with scales and short hairs distinguish it 

from that species. 

6. Engelhardia rigid a Lesch. ex Blume 

(Latin, rigidus = stiff; the leaflets) 

I.c. (1825) 528, I.c. (1829) 13, t. 3; Jacobs l.c. 148; Backer & BakhuizenJ l.c. 158; Burgess I.c. 321; 

Manning l.c. 40, 45; Cockbum I.c. 152; Anderson I.c. 220; Ashton I.c. 304; Whitmore, Tantra & 

Sutisna I.c. 185. Type: Blume 1958, Java (ho1otype L; isotypes K, P). Synonyms: E. subsimplicifolia 

240


Merr., Govt. Lab. Publ. Philip. 34 (1906) 6, I.e. (1923) 24; E. rigida var. subsimplieifolia (Merr.) 

Manning I.e. 40; E. lepidota Schltr., BotJahrb. 50 (1913) 66; E. zambalensis Elmer I.e. 3195. 

Tree to 50 m tall, and 90 cm in diameter; buttresses to 3 m high and 2 m out. Bark pale 

brown-grey to reddish brown, soft; inner bark white, orange-yellow to brownish, soft. 

Sapwood white. Twigs, leaves and inflorescences set with golden to orange scales and pale 

yellow hairs. Leaf rachis (1-)2-6(-21) cm long, 1.5-2.5 cm to first leaflet, blackish, 

thickened at base, sometimes densely scaly and hairy; leaflets typically 2-3 pairs, opposite, 

ovate to elliptic, 1.5-7.1(-16.5) x 1.4-4(-7.5) cm, often drying orange-brown to brown 

below, thickly leathery, often somewhat bullate; base acute to wedge-shaped, subequal, 

margin entire, with thickened marginal vein, sometimes inrolled, apex shortly acuminate, 

acumen 1-2 mm long; tufted domatia occasional to frequent in vein axils with midrib; 

midrib on upper and lower sides glabrous to densely scaly and hairy; lateral veins 6-9 

pairs, distinct on both sides; intercostal veins ladder-like, indistinct on both sides; petiolules 

rarely absent to 3(-5) mm long, with scant to dense scales and sometimes hairs. 

Inflorescences axillary on leafy and older leafless twigs, scaly and hairy; male catkins 1.5- 

6 cm long, perianth-lobes narrowly triangular to completely reduced, stamens (3-)4-6( -7) 

with anthers equal or unequal, 0.5-1 mm long, more or less white hairy; fruiting catkins 

(6-)10-15(-22) cm long, peduncle and rachis slender, round, more or less densely scaly 

and hairy. Fruits 0.5-1-mm-stalked; bracts scaly and hairy, abaxial wings 20-60 x 3-8 

mm, lateral wings 12-14 x 3-4 mm, adaxial lobes orbicular or absent so the top of the nut 

is sometimes exposed; nuts globose, c. 3.5 x 4 mm, densely covered by long, soft hairs, but 

without scales; perianth-lobes on the nut apex ovate, free from style, not hiding the 

stigmas. 

Distribution. Sumatra, Java, Borneo, the Philippines, Celebes, Moluccas, New Guinea. In 

Sabah uncommon, known only on Mount Kinabalu; in Sarawak uncommon in lowlands 

and mountains. Also in Brunei and Kalimantan. 

Ecology. Primary mixed dipterocarp forest from sea-level to 2000(-2500)m on both 

leached yellow, sandy and ultramafic soils. 

Uses. The wood is used for canoes and general building work in Sumatra. 

Taxonomy. Manning (l.c.) distinguished two vanetres of this species in Borneo, var. 

subsimplicijolia and var. rigida. Var. subsimpliciJolia was said to differ from the other in 

having fewer pairs of leaflets and shorter fruiting catkins and fruits. The type specimen of 

E. rigida var. rigida unfortunately does not have any fruits but otherwise looks like var. 

subsimplicijolia with one extra pair of leaflets. The other specimens of E. rigida var. rigida 

named by Manning had all the characters which Manning ascribed to var. subsimpliciJolia, 

including fewer leaflets, shorter fruiting catkins and smaller fruits. It is felt that E. rigida, 

at least in Borneo, cannot be separated into these two varieties. 

7. Engelhardia roxburghiana Wall. 

(W. Roxburgh, 1751-1815, a Scottish botanist) 

PI. As. Rar. 2 (1831) 85, t. 199; Jacobs I.e. 154; Burgess I.e. 321; Manning I.e. 38,48; Whitmore I.e. 

235; Cockbum I.e. 154; Anderson I.e. 220; Ashton I.e. 305; Corner I.e. 369; Whitmore, Tantra & 

241


Sutisna l.c. 185. Type: Wallich Cat. 4942, Malaya (holotype CAL). Synonyms: luglans pterococea 

Roxb., Hort. Beng. (1814) 68; E. chrysolepis Hanee, Ann. Se. Nat. 4, 15 (1861) 227; E. wallichiana 

Lind1. ex Wal1., Cat. (1831/32) no. 4942; E. pterococca (Roxb.) Kuntze, Rev. Gen. PI. 2 (1891) 637; 

E. spicata var. formosana Hay, Fl. Mont. Form. 6 (1908) 199; E. formosana Hay, le. P1. Form. 6 

(1916) 61; E./enzelii Merr., Lingn. Se. J. 7 (1931) 300. 

Tree to 35 m tall and 70 cm diameter; buttresses to 2 m high, thin, branching; crown dense 

or spreading. Bark fawn to dark chocolate-brown or black; inner bark yellow to reddish 

brown, soft, moist; exudate clear, white. Sapwood white to pale yellow. Twigs often. with 

crusts of white eXlj,date, together with leaves and inflorescences set with golden to orange 

scales and pale yellow to orange hairs. Leaf rachis (5.5-)6-14(-l7) cm long, 3.4-9.5 cm 

to first leaflet, thickened at base, scantily to densely scaly and hairy; leaflets 2-3(-4) pairs, 

alternate to subopposite, ovate to lanceolate, (5-)10-16(-23) x (1.4-)3.5-5(-8) cm, gradually 

decreasing in size towards base, drying greenish to greyish brown or brown below, 

leathery; base acute to wedge-shaped, very unequal, often somewhat decurrent, margin 

entire, slightly wavy, with thickened marginal veins, sometimes inrolled, apex longacuminate, 

acumen 1O-l7 mm long; tufted domatia absent to rare in the vein axils with 

the midrib; midrib and veins above with occasional to scattered scales, below with 

occasional to dense scales but without hairs; lateral veins 8-12 pairs, above raised to flat, 

below raised; intercostal veins ladder-like, distinct above, indistinct below, areoles distinct 

above and below; petiolules (1-)6-1O( -12) mm long thickened at base, sparsely to densely 

scaly and hairy. Inflorescences terminal on lateral twigs, paniculate; male catkins 8-10 per 

inflorescence, 9-10 cm long, peduncle slender, scaly but without hairs; perianth-lobes 1- 

1. 5 mm across; stamens 8-12, inserted (2-)3 at the base of each perianth-lobe, anthers 

equal, c. 0.5 mm long, filaments equal to unequal, 0.3-0.5 mm long, glabrous, faintly 

sweet scented; fruiting catkins 10-17(-23) cm long, peduncle and rachis slender, scaly and 

densely hairy. Fruit stalk 4-5(-8) mm long; bracts scaly but without hairs, abaxial wings 

28-45(-55) x 5-8 mm, lateral wings 21 x 5-6 mm, adaxial lobes not developed so nut is 

visible; nuts globose, 4-5 x 4-5 mm, with scales but without hairs; perianth-lobes at the 

nut apex ovate, enclosing the 4 sessile stigmas. 

Distribution. India to Indo-China, Sumatra, Peninsular Malaysia, and Borneo. In Sabah 

uncommon, only known from the Crocker Range (SAN 44313); in Sarawak scattered 

throughout mixed dipterocarp and submontane forests (s. 28588, S. 32811, Nooteboom & 

Chai 2153), locally abundant as on the summit ridges of the basaltic Bt. Mersing and the 

Tau Range, where it grows together with E. mersingensis. Also in Brunei at Ulu Mendamit. 

Ecology. Primary mixed dipterocarp forest in hilly country, to 1750 m, on clay-rich, relatively 

fertile soils. 

8. Engelhardia serrata Blume 

(Latin, serratus = toothed like a saw; the leaflet margin) 

I.c. (1829) 14, t. 4; Jaeobs I.c. 150; Backer & BakhuizenJ I.c. 158; Burgess I.e. 321; Manning I.e. 

40, 45; Whitmore l.c. 236; Coekburn l.c. 152; Anderson l.c. 220; Ashton l.c. 305; Corner I.e. 370; 

Whitmore, Tantra & Sutisna I.c. 185. Type: Blume 2221, Java, Mt. Salak (holotype L). Synonyms: 

242

JUGLAl'lDACEAE (CAMPBELL-GASIS) 

E. palembanica Miq., Fl. Ned. Ind., Suppl. (1861) 346, 139; E. parvifolia C. DC. I.e. 34; E. 

nudiflora Hook. f, Fl. Br. Ind. 5 (1888) 596; E. serrata Blume var. nudiflora (Hook.f) Manning I.e. 

46; E. permierophylla Elmer, Leafl. Philip. Bot. 9 (1934) 3194. 

Tree to 30 (-45) m tall, 120 cm diameter; buttresses to 3-4 m high and 3 m out. Bark light 

to dark grey to reddish brown; inner bark from pinkish brown to orange- and reddish 

brown, soft. Sapwood yellowish to white. Twigs, leaves and inflorescences set with golden 

and orange scales and pale yellow hairs. Leaf rachis (2.6-)5-18(-24) cm long, 1.2-3.1 cm 

to first leaflet, slightly thickened at base, densely to scantily covered with scales and hairs; 

leaflets (2-)3-6( -9) pairs, alternate to subopposite, oblong to lanceolate in young trees to 

ob ovate in older trees, (1-)2-15(-20.5) x (0.5-)1-4(-8) cm, often decreasing to about half 

the size towards base, drying from ochre-brown to dark brown below, chartaceous in young 

trees to leathery in older trees, upper side with only the midrib and veins sparsely to 

densely scaly and hairy, lower side with scattered to dense scales and sparse hairs all over; 

base acute to wedge-shaped, rarely rounded, subequal, margin regularly to irregularly 

toothed at least in upper third, apex short- to long-acuminate, acumen 1-5 mm long; tufted 

domatia frequent in vein axils with midrib; lateral veins 4-15 pairs, fewer in older trees, 

above distinct or not, raised to flat; intercostal veins net-like, above and below distinct or 

not; petiolules absent to J (-3) mm long, occasionally with dense scales and hairs. Inflorescences 

axillary on leafy or older leafless twigs, scaly and hairy; male catkins 2-3 per 

inflorescence, 3-13 cm long, perianth-lobes more or less cCJmpletely reduced, stamens (3-) 

5-7, anthers sometimes unequal and 0.5-1 mm long, filaments sessile to 0.3 mm long and 

sparsely hirsute; fruiting catkins 9-14(-30) cm long, peduncle and rachis stout, angular, 

densely covered with scales and hairs. Fruits sessile to 1-mm-stalked; bracts glabrous or 

scaly and hairy; abaxial wings 16-40 x 6-11 mm, lateral wings 6-12 x 3-6 mm, adaxial 

lobe fritled to shallowly 2-3-lobed, often hiding the top of the nut; nuts globose, c. 3 mm 

across, sometimes with a few scales, densely covered in stiff, long hairs; perianth-lobes on 

nut apex ovate, free from style, not hiding the stigmas. 

Vernacular names. Sabah-pusing-pusing (Malay). Sarawak--entalun (Iban), momon 

(Malay), owl (Murut), tepanga (Kenyah). 

Distribution. Burma, Thailand, Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo, 

Philippines, and the Moluccas. In Sabah, not uncommon in the lowlands but also on the 

Crocker Range. In Sarawak, scattered in lowland and submontane forests. Also in Brunei 

and Kalimantan. 

Ecology. Primary mixed dipterocarp forest, on sandy or clayey soils from sea-level to 2200 

m; rarely in secondary forest. 

Taxonomy. Manning I.c. recognised four varieties, viz. var. serrata, parvifolia (C. DC.) 

Manning, nudiflora (Hook. f) Manning, and cambodica Manning, of which the first three 

occur in Sabah and Sarawak. These taxa seem to share the characters described above, and 

with limited material available for study, it is impossible to say whether the varietal status 

proposed by Manning is truly justified. 

243


9. Engelhardia spicata Lesch. ex Blume 

(Latin, spicatus = bearing a spike; the inflorescence) 

I.e. (1825) 528; Merrill/.e. (1921) 210, I.e. (1923) 24; Ridley I.e. 368; Masamune I.e. 231; Jacobs 

. I.e. 151; Backer & BakhuizenJ I.e. 158; Burgess I.e. nl; Manning I.e. 41; Whitmore I.e. 236; 

Cockburn I.e. 152; Corner I.e. 370; Whitmore, Tantra & Sutisna I.e. 186. Type: Blume, s.n., Java 

(holotype L; isotypes K, NY, P). Synonym: E. philippinensis C. DC. I.e. 34, t. 2,/ 15. 

var. aceriflora (Reinw.) Koords. & Valeton 

Bijdr. Booms. Java 5 (1900) 167. Basionym: Pterilema aeeriflorum Reinw., SylI. PI. Nov. Soc. 

Rolisb. 2 (1826) 13. Type: Reinwardt, s.n., Java (holotype L). Synonyms: E. aeeriflora (Reinw.) 

Blume I.e. (1829) 11, t. 2 and 5B; E. pteroeoeea Roxb. ex Kuntze var. aeeriflora (Reinw.) Kuntze 

I.e. 637. 

Tree to 40 m tall, 280 cm diameter; buttresses sometimes present, to 3 m tall; crown dense, 

rounded. Bark light brown-grey. Twigs with pale golden scales and hairs. Leaf rachis 

(5.5-)10-30 cm long, to 5.5 cm to first leaflet, slightly thickened at base, brown or 

blackish, glabrous to scaly and hairy; leaflets (2-)3-5( -7) pairs, subopposite to opposite, 

oblong to lanceolate, (6-)4-16(-30) x (3-)1.5-6(-8) cm, not decreasing in size towards 

base, leathery; upper and lower sides glabrous to sparsely scaly to very hirsute, or both 

scaly and hairy, especially on the midrib and veins on the lower side; base rounded to 

sub cordate, very unequal, margin entire, wavy, apex shortly acuminate, tip obtuse to acute; 

tufted domatia frequent in the vein axils with the midrib and often also in the axils of veins 

which join near the margin, visible to the unaided eye; lateral veins 11-18 pairs, above 

distinct, raised, below often indistinct; intercostal veins indistinct above and below; 

petiolules absent to 3 mm long. Inflorescences paniculate, axillary on leafy or older 

leafless twigs; male (4-)9-17 cm long; perianth-lobes present or reduced, very irregular, 

lobes to 2 mm long; stamens 6-14, anthers equal or unequal, (sub)sessile, hirsute; fruiting 

catkins (8-)19-30(-40) cm long, peduncle and rachis angular, glabrous or scaly. Fruits 

sessile to I-mm-stalked; abaxial wings 20-60 x 7-15 mm, lateral wings 20-25 x 5-8 mm, 

wings often very irregular, adaxial lobes frilled to shallowly 4-5-lobed, often hiding the 

top of the nut; nuts globose to transversely ellipsoid, 3-6 x 3-8 mm, with long, soft, dense 

hairs, without scales; perianth-lobes tiny, inconspicuous, adnate to style, not hiding the 

stigmas. 

Distribution. India to Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, 

and Lesser Sunda Islands. In Sabah, uncommon on Mount Kinabalu (e.g., SAN 42865, SAN 

6 J 785); not reported from Sarawak or Brunei. Also in East Kalimantan. 

Ecology. Primary forest, more common on mountains, from sea-level to 2500 m. 

Taxonomy. Koorders & Valeton I.c. and Manning I.c. recognised three varieties, namely 

var. acerifolia (Reinw.) Koord. & Valeton, colebrookeana (Lindl. ex Wall.) Kuntze, and 

spicata, of which only var. acerifolia is so far found in Sabah. 

244


MONIMIACEAE 

P.C.Yii 


Kuching, Malaysia 

and 

Lesmy Tipot 



HookerJ, Fl. Br. Ind. 5 (1890) 114; Merrill, EB (1921) 271, PEB (1929) 77; Ridley, FMP 3 (1924) 

73; Masamune, EPB (1942) 305; Backer & BakhuizenJ, FJ 1 (1963) 116; Anderson, CLTS (1980) 

253; Philipson, Blumea 28 (1982) 77, Blumea 30 (1985) 389, FM I, 10 (1986) 255; Ng, TFM 4 

(1989) 261; Whitmore, Tantra & Sutisna, CLK2, 1 (1990)235. 

Trees or shrubs, rarely woody climbers. Leaves simple, decussate or rarely alternate or in 

whorls of three, usually with round oil-cells in the lamina, bearing simple or stellate hairs 

or glabrous. Stipules absent. Inflorescences cymes, solitary or fascicled, terminal or axillary, 

rarely on the trunk. Flowers unisexual or bisexual, regular; receptacles usually welldeveloped 

or rarely reduced, round, urceolate or bell-shaped; tepals inconspicuous, rarely 

developing as distinct sepals and petals, decussate, in whorls or spirals. Male flowers with 

few to many stamens arranged in whorls, spirally or irregularly, filaments strap-shaped, 

anthers 2-4-loculed, opening through slits or valves. Female flowers with or without 

staminodes; carpels few to many, sessile or stalked (stipitate), free or immersed in the 

receptacles; ovule solitary, anatropous, erect or pendulous, with a thick nucellus (crassinucellar), 

bitegmic or unitegmic. Fruits achenes or drupes, rarely plumose, usually enclosed by the 

persistent receptacle, stalked or sessile, set free by splitting of the receptacles. Seed one; 

endosperm copious, oily; embryo straight; cotyledons appressed or divergent. 

Distribution. About 33 genera and 320 species, mostly in the warmer parts of the southern 

hemisphere (Malesia, Australia, SW Pacific, islands in the western Indian Ocean and S 

America). In Sabah and Sarawak, represented by 2 genera with 3 species. 

Taxonomy. The two genera (Kibara and Matthaea) occurring in Sabah and Sarawak are 

included by PhiIipson (I.e. (1986) 261) in the subfamily Mollinedieae and characterised by 

the male flowers having globose or flask-shaped receptacles and by drupaceous fruits borne 

in heads. For- a detailed account of the systematic position of the family see Philipson's 

account (I.e. 1986). 

Key to genera 

Leaves ovate to broadly elliptic; lateral veins diminished and not joined toward the margin. 

Stamens 6-9, arranged in 2 series, anthers opening by a single longitudinal slit. Fruit-stalk 

shorter than 8 mm ................................................................................................ 1. Kibara 

245

occasionally on limestone hills, sandy and coral beaches. 

Key to Kibara species 

Leaves broadly ovate to elliptic-oblong, base rounded or subcordate, margins entire or 

toothed, apex distinctly acuminate ................................................................ 1. K coriacea 

Leaves narrowly to broadly elliptic, base narrowly to broadly cuneate, margins entire, apex 

obtuse .................................. , ......................................................... , .................. 2. K. obtusa 

1. Kibara coriacea (Blume) Tulasne 

(Latin, coriaceus = leathery; the leaves) 

Arch. Mus. Hist. Nat. Paris 7 (1855) 404; Hooker J I.e. 114; Rid1ey I.e. 75; Merrilll.e. (1929) 77; 

Masamune I.e. 305; Backer & BakhuizenJ I.e. 117; Philipson I.e. (1985) 406, I.e. (1986) 298; Ng 

246 

Distribution. About 40 species, in Peninsular Thailand, Nicobar Islands, Malesia and 

Ecology. Mostly understorey shrubs and small trees in rain forest from sea-level to 2800 m; 

thickened and grandular; carpels many, free on the inside of the receptacles; style very 

short; ovule 1. Fruit a sessile or short-stalked drupe enclosed by enlarged receptacle. Seeds 

coniform; seed-coat membranous; embryo small. 

flowers with a minute opening surrounded by 5 decussate pairs of tepals, the inner pairs 

flat or cup-shaped receptacles. Male flowers usually smaller than the females, with a 

minute opening surrounded by 2-4 decussate pairs of tepals; stamens 6-9, arranged in 2 

series, with an outer series of 4(-5) larger stamens, and an inner series of 4 smaller, often 

infertile stamens, anthers opening by a single slit, with a filament or subsessile. Female 

surrounded by scale leaves. Leaves simple, decussate; blades entire or toothed, usually with 

short, tufted, soft, brown hairs, gradually becoming glabrous; lateral veins arched and 

diminishing toward the margins. Inflorescences terminal or axillary cymes (racemose in 

the non-Bornean K. streimannii), paniculate or fasciculate; pedicels thickening distally into 

Trees or shrubs with aromatic smell. Terminal vegetative buds conical or pyramidal, 

Gen. PI. (1837) 314; Hooker J I.e. 114; MerrillI.e. (1921) 271, I.e. (1929) 77; Rid1ey I.e. 73; 

Masamune I.e. 305; Backer & BakhuizenJ I.e. 117; Anderson I.e. 253; Phi1ipson I.e. (1985) 389, I.e. 

(1986) 287; Ng I.e. 261; Whitmore, Tantra & Sutisna I.e. 235. Synonyms: Brongniartia Blume, 

Bijdr. 9 (1825) 455 (non Knuth); Sciadiearpus Hassk., Flora 25, 3 (1842) 20; Sareodiseus Griff., 

Not. PI. As. 4 (1854) 380. 

Leaves narrowly lanceolate to lanceolate-oblong, lateral veins looped and joined at the 

margin. Stamens usually 4, free, anthers opening by two longitudinal slits. Fruit-stalk 

longer than 8 mm ............................................................................................. 2. Matthaea 

Queensland (Australia). Only 2 species occur in Sabah and Sarawak. 

( a Sundanese plant name). 

1. KIBARA Endl. 

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

MONIMIACEAE (YII & LESMY) 

I.e. 261; Whitmore, Tantra & Sutisna I.e. 235. Basionym: Brongniartia eoriaeea Blume I.e. (1825) 

436. Type: Blume, s.n., Java (L). Synonyms: K. blumei Steud., Nomencl. Bot. (1840) 846; 

Seiadiearpus brongniartii Hassk. I.e. 20; Sareodiseus ehloranthiformis Griff. I.e. 350; K. ehartaeea 

Blume, Mus. Bot. Lugd. Bat. 2 (1856) 89; K. euspidata Blume l.c. (1856) 89, Merrilll.e. (1921) 

271, Masamune I.e. 305; K. tomentosa and maerophylla lR. Perkins, Bot. Jahrb. 25 (1898) 571; K. 

triehantha JR. Perkins I.e. (1898) 572; K. serrulata lR. Perkins I.e. (1898) 575; K. angustifolia lR. 

Perkins I.e. (1898) 577; K. motleyi lR. Perkins, Bot. Jahrb. 45 (1911) 424, Merrilll.e. (1921) 272, 

Masamune l.c. 305; K. grandifolia Merr., Philip. Govt. Lab. Bur. Bull. 29 (1905) 15; K. ellipsoidea 

Merr., Philip. J. Sc. 1 (1906) Suppl. 56; K. mollis Merr., Phi1ip. 1 Sc. 3 (1908) Bot. 225. 

Shrub or tree, to 15 m tall, 15 cm diameter. Bark smooth, pale grey; inner bark pale 

yellow. Twigs slightly pubescent, gradually become glabrous. Leaves broadly ovate to 

elliptic-oblong, 6-26.6 x 4-18.5 cm, leathery or papery, glabrous or sparsely to rather 

densely hairy beneath; base cuneate, rounded or subcordate, margin entire or toothed 

toward the apex, apex distinctly acuminate; midrib and lateral veins prominent beneath; 

lateral veins arched, ascending andjoining near the margins; stalk 5-25 mm long, slightly 

channelled above, pubescent or glabrous. Inflorescence a terminal or axillary, simple 3- 

flowered or compound cyme, with the male flowers at the proximal and the female flowers 

at the distal parts. Male flowers rounded, 1.5-2 mm across, hairy; tepals 6-8 with rounded 

apex; stamens 4 in outer series and 4 smaller ones in the inner series, filaments strapshaped. 

Female flowers larger than male flowers, rounded, about 3-5 mm across; tepals 

about 6 with swollen pendulous glands within the minute opening. Fruit an ovoid drupe, c. 

2 x 1.5 cm, ripening deep blue, purple or black, on short swollen orange stalk; in clusters 

of 3-13. Seed coat membranous, orange when dried. 

Vernacular names. Sabah-ambibiliw, labak (Dusun). 

Distribution. Throughout Malesia; in Sabah and Sarawak widespread. 

Ecology. Lowland rain forests including swamp, coral beach, limestone hill, mixed dipterocarp 

forests, and lower montane forests from sea-level to 1600 m. 

Uses. The fruit is said to be edible and the leaves are used as flavouring in meat dishes. 

2. Kibara obtusa Blume Fig. 1. 

(Latin, obtusus = blunt or rounded; the leaf apex) 

I.e. (1856) 89; Philipson I.e. (1985) 409, I.e. (1986) 300. Type: Blume, s.n., Celebes (holotype L). 

Synonym: K. depauperata Merr., Philip. Govt. Lab. Bur. Bull. 35 (1906) 13. 

Tree to 20 m tall, 20 cm diameter. Bark scaly to shallowly fissured, pale yellow; inner bark 

dull orange. Young twigs with short stiff-hairs. Leaves narrowly to broadly elliptiC, 7-16.5 

x 3.3-10 cm; base cuneate, margin entire, apex obtuse or rounded; midrib and lateral veins 

prominent beneath, glabrous or with sparse stiff-hairs; stalk 10-18 mm long, pubescent or 

glabrous. Inflorescence a terminal or axillary, simple or compound cyme, to about 70 mm 

long, with a pair of small bracteoles. Male flowers obovoid, c. 2 mm across; tepals 4, 

minute; stamens usually 4 in the outer series and 2 smaller, infertile ones in the inner 

247


series. Female flowers larger than male flowers, globose, about 2.5-3 mm across; tepals 4, 

apex obtuse, with 4 swollen glands projecting among the carpels; carpels about 13, hairy, 

with blunt stigmas. Fruit an ovoid drupe, 17-24 x 10-12 mm, ripening black, seated on a 

short orange stalk. Seed coat membranous, orange ~n colour when dried. 

Distribution. Borneo, Philippines, Celebes, and W New Guinea. In Sabah, uncommon and 

only known from 4 collections, 3 from Lahad Datu (SAN 29844, SAN 31104, SAN 33382) 

and one from Semporna (SAN 46055). Not yet reported in Sarawak. 

Ecology. Primary rain forest from sea-level to 700 m. 

2. MATTHAEA Blume 

(Matteo de S. Guiseppe, 1617-1691, an Italian Missionary and Botanist in India) 

I.e. (.1856) 89; Hooker f I.e. 115; Merrill I.e. (1921) 272; Ridley I.e. 73; Masamune I.e. 305; 

Anderson I.e. 254; Philipson I.e. (1982) 77, I.e. (1986) 319; Ng I.e. 263. 

Trees or shrubs. Leaves simple, opposite, entire or sub-serrate, leathery. Inflorescences 

axillary, rarely terminal cymes, much shorter than leaves. Male receptacle subglobose; 

tepals 4; stamens 4, free, filaments short, anthers opening by 2 longitudinal slits. Female 

receptacle depressed globose; tepals 4, without apical pore or gap but upper half abscissing 

as a calyptra at anthesis to reveal numerous carpels. Fruits long-stalked drupes, fleshy; 

one or more on the enlarged receptacle. 

Distribution. 6 species, all Malesian: Sumatra, Peninsular Malaysia, Anambas Island, Borneo, 

Celebes, Philippines, and N Moluccas. Only 1 species has been recorded in Sarawak. 

Ecology. Understorey trees in lowland and submontane forests to about 1700 m. 

Matthaea sancta Blume Fig. 2. 

(Latin, sanctus = holy; alluding to the religious work of Matteo de S. Guiseppe) 

I.e. (1856) 90; Hooker f I.e, 115; Merrill I.e. (1921) 272; Ridley I.e. 73; Masamune I.e. 305; 

Anderson I.e. 264; Philipson I.e. (1982) 82, I.e. (1986) 323; Ng I.e. 263. Type: Blume, s.n., "Sumatra 

and Borneo" (holotype L). Synonyms: M. latifolia JR. Perkins I.e. (1898) 563; M. ealophylla J.R. 

Perkins I.e. (1898) 563, Merrill/.e. (1921) 272, Masamune I.e. 305; M. ellipsoidea Merr. ex JR. 

Perkins I.e. (1911) 423. 

Shrub or small tree, to 10 m tall, 10 cm diameter. Twigs green, glabrous. Leaves papery, 

glabrous, lanceolate-oblong to oblong, 15.5-29 x 5.5-9.5 cm; base broadly cuneate, truncate or 

rounded, margins entire or slightly toothed at the upper part, apex acuminate; lateral veins 

arched, ascending and looping far from the margins, impressed on the upper surface, 

prominently raised beneath; stalk 1.5-3 cm long, glabrous. Inflorescences axillary cymes. 

Male flowers depressed globose, 2-3 mm across, opening by a small gap between a pair of 

248

MONIMIACEAE (YII & LESMY) 

".[ 

H 

G 

2 cm 

40mm 

Fig. 1. Kibara obtusa. A, leafy twig with young axillary inflorescence; B, inflorescence; C, male 

flower; D, stamen; E, female flower; F, section of female flower; G, carpel; H, fruits; I, section of 

fruit. (A-G from SAN 29844, H-I from SAN 3Jl04.) 

249


] 2cm 

2cm [ 

F 

Fig. 2. Matthaea sancta. A, leafy twig with female inflorescence; B, male inflorescence; C, male 

flower in longitudinal section; D, female flower before anthesis; E, carpel; F, fruits. (From Steven 

261.) 

250

MONIMIACEAE (YIl & LESMY) 

lip-like perianth-lobes; stamens 4. Female flowers similar to males but much larger, c. 10 

mm across, on 1-4 pedicels arising in clusters of 3-5 from short peduncles; without any 

apical pore or gap but the upper half abscissing as a calyptra at anthesis to reveal numerous 

sessile carpels which are closely packed on a dish"like receptacle. Fruit a drupe, ellipsoid, 

c. 2.5 x 1.5 cm, with thin bony endocarp, ripening blue or purple, attached in clusters of up 

to 18 to an enlarged orange receptacle. Seed coat light brown when dried. 

Distribution. Sumatra, Peninsular Malaysia, Anambas Is., Borneo, Philippines, and Celebes. 

Uncommon in Sabah but widespread in Sarawak. 

Ecology. Lowland rain forests, including limestone hill and mixed dipterocarp forests on 

clay-rich soils, and lower montane forest, from sea-level to 1400 m. 

251


NYSSACEAE 

p. C. Yii 


Kuching, Sarawak 

Wangerin in Engler, Pfl. R. 41, 2 (1910) 1; Wasscher, Blumea 1 (1935) 343, FM 1,4 (1948) 29; 

Masamune, EPB (1942) 517; Hutchinson, Fam. Fl. PI. 1 (1959) 175, Gen. FI. PI. 2 (1967) 50; Eyde, 

J. Am. Arb. 44,1 (1963); Backer & BakhuizenJ, FJ 2 (1965) 161; Kochummen, TFM 1 (1972) 346; 

Cockbum, IS 2 (1980) 61; Mabberley, PB (1987) 404; Whitmore, Iantra & Sutisna, CLK 2, 1 

(1990) 277. 

Trees or shrubs. Leaves simple, spirally arranged, without stipules. Flowers more or less 

regular, male or hermaphrodite, solitary or more often in axillary heads or condensed racemes; 

calyx 5-toothed or absent; corolla composed of 5(-10) imbricate petals; nectary disc 

present; stamens (8-)10(-15), usually in 2 whorls, the outer whorl opposite the petals in 

male flowers or as many as and alternate with the petals in hermaphrodite flowers, anthers 

dorsifixed, dehiscing lengthwise; ovary injerior, 1-2-loculed, with 1-2 basally united styles, 

ovule 1, pendulous, anatropous. Fruit a drupe. Seeds with scanty, oilyendosperm; embryo 

straight. 

Distribution. 3 genera, with 8-10 species, distributed in N America, Mexico, E and SE 

Asia. In Sabah and Sarawak, represented by 1 genus (Nyssa) with 1 species (N. javanica). 

Taxonomy. The family is closely related to the Cornaceae with which it has been frequently 

included in previous publications (Ridley, FMP 1 (1922) 889; Keng, OFMSP (1978) 219; 

Brummitt, Vasc. PI. Fam. & Gen (1992) 543). It can be distinguished from the Cornaceae 

by the following characters: male flowers with tepaloid sepals and petals, stamens 8-10 

arranged in two whorls; hermaphrodite flowers solitary or in heads, sepals and petals well 

developed; style 1, rarely 2, each with 1 stigma; fruit a drupe. 

NYSSAL. 

(a legendary town in India where Bacchus, 

the Greek God of Wine, was brought up by nymphs) 

Sp. PI. (1753) 1058; Ridley I.e. 895 (under Comaceae); Wasscher I.e. (1935) 343, I.e. (1948) 29; 

Masamune I.e. 517; Backer & Bakhuizen J I.e. 161; Kochununen I.e. 346; Cockbum I.e. 61; 

Whitmore, Tantra & Sutisna I.e. 277. Synonyms: Agathisanthes and Ceratostaehys Blume, Bijdr. 

(1825) 644; Agathidanthes Hassk., Cat. Hort. Bog. (1844) 254; Daphniphyllopsis Kurz, J. As. Soc. 

Beng. 44, 2 (1875) 201. 

253


50mm[ 

8 

c 

2mm] 

2cm [ 

F 

~ 

~ 

D 

2mm 

'---------' 

E 

Fig. 1. Nyssa javanica. A, fruiting leafy twig; B, inflorescence; C, male flower; D-E, female flowers; 

F, fruits. CA and F from S. 35531. B-E from SAN 42783.) 

254

NYSSACEAE (YII) 

Dioecious trees or shrubs. Leaves simple, spirally arranged, without stipules. Flowers 

male or hermaphrodite, often in heads, in the axils of a bract and with 2 bracteoles. Male 

flowers in axillary heads or short racemes; calyx campanulate, rim smooth or 5-toothed; 

petals 5, imbricate in bud, alternate with the calyx-lobes; stamens 8-16, in 2 alternating 

whorls; anthers dorsifixed, opening lengthwise; disc pulvinate; ovary and style rudimentary. 

Hermaphrodite flowers in 2-1O-flowered, axillary, stalked heads; calyx entire or 5-toothed; 

petals 5-8; stamens of the inner whorl partly sterile; ovary I-locular, connate with the 

calyx, style with 2 appressed, curving, often torulose branches, stigmatose inside. Fruits 

drupaceous, ovoid to ellipsoid-oblong. 

Distribution. About 6 species, 4 in the Atlantic N America, 1 in China, and 1 (N javanica) 

widespread from India to W Malesia. 

Nyssa javanica (Blume) Wangerin 

(of Java) 

Fig. 1. 

in Engler I.c. 15; Wasscher l.c. (1935) 344, I.e. (1948) 29; Masamune I.c. 517; Kochummen I.c. 346; 

Cockburn I.c. 61; Whitmore, Tantra & Sutisna I.c. 277. Basionym: Agathisanthes javanica Blume 

I.c. 645. Type: Blume, s.n., Java (holotype L; isotype K). Synonyms: Agathidanthes javanica Hassk. 

I.c. 254; Nyssa sessiliflora Hook.! & Th., Gen. PI. 1 (1867) 952; !lex daphniphylloides Kurz, J. As. 

Soc. Beng. 39,2 (1870) 72; Daphniphyllopsis capitata Kurz I.e. (1875) 201; Nyssa arborea Koord., 

Exk. Fl. Jav. 2 (1912) 731; Nyssa bifida Craib, Kew Bull. (1913) 69. 

Tree to 50 m tall, 90 cm in diameter, sometimes with very low buttresses. Bark grey, 

smooth to slightly flaky, with prominent corky lenticels; inner bark dull yellow or pale 

brown, fibrous or laminated, staining dark blue after slash. Sapwood heavy, yellowish 

white. Twigs green, with large scattered lenticels and leaf-scars, often covered with brown 

tomentum when young, gradually turning glabrous. Leaves typically crowded toward the 

end of the twigs; stalk 1.5-4 cm long, hairy; blades slightly glaucous below, thinly leathery, 

oblong-lanceolate to obovate, 5-19 x 2-7 cm; base gradually narrowed toward stalk, 

margin entire to slightly wavy, apex abruptly pointed; midrib flushed reddish, slightly 

hairy; lateral veins 8-11 pairs, hairy. Flowers in stalked, roundish, axillary heads 12-18 

mm across, stalk slightly angUlar, 10-50 mm long, glabrous or slightly hairy, bracteate; 

bracts perSistent in hermaphrodite flowers. Male flowers in 20-40-flowered heads; calyx 

bell-shaped, 4-5-toothed; petals 4-5, free, overlapping, re curved, shortly hairy; stamens 

8-10 in 2 whorls. Hermaphrodite flowers in 3-9( -I8)-flowered heads; calyx campanulate, 

densely appressed-sericeous, lobes 4-5, irregular, rounded or almost absent; petals 4-5; 

stamens 8-10, inner whorl sterile; ovary I-chambered, style with 2 curving branches. 

Fruits ellipsoid drupes, slightly flattened, to 22 x 15 mm, crowned by small persistent 

calyx. Seed one, stone-like, grooved on one side and knobby on the other. 

Vernacular name. Sarawak-terang bulu (Kelabit). 

Distribution. Sumatra, Peninsular Malaysia, Java and Borneo. In Sabah, the species is 

known only from Mt. Kinabalu (SAN 22427, SAN 23507, SAN 42755, SAN 42785, SAN 

60599, SAN 62216 and SAN 62220); in Sarawak, one collection so far (s. 35531), from 

Kelabit Highlands, Bario. 

255


Key to genera 

1. Shrubs to medium-sized trees. Stamens 10; carpels 5-10, free. Fruit a I-seeded drupe ........ 2 

Shrubs. Stamens 5; carpels 2-5, united. Fruit a capsule or berry with more than one 

seed ............................................................................................................................. 3 

2. Leaves with di.stinct marginal veins, margin toothed; stipules united. Anthers opening 

by apical pores ............................................................................................ 3. Gomphia 

Leaves without marginal vein, margin entire; stipules free. Anthers opening by 

longitudinal slits ............................................................................... 1. Brackenridgea 

3. Leaf margin toothed. Ovary 5-celled. Anthers opening by apical pores. Fruit a berry 

.. 2. Euthemis 

Leaf margin entire or toothed. Ovary l-celled. Anthers opening by longitudinal slits. 

Fruit a capsule ............................................................................................................ .4 

4. Branches hollow. Leaves shiny, lateral veins invisible. Seeds winged ........ 4. Schuurmansia 

Not as above ................................................................................................................ 5 

5. Stem coarsely bristly with persistent stipules and bracts. Inflorescences axillary, oneflowered 

........ 

Neckia Korth. 

Ned. Kruidk. Arch. 1 (1848) 358; Merrill I.c. 388; Ridley I.c. 134; Masamune I.c. 469; 

Kanis I.c. (1968) 69, I.c. (1971) 110. 

Monotypic genus (N. serrata Korth.) confined to Sumatra, Peninsular Malaysia, 

Borneo (Sabah, Brunei, Sarawak, Kalimantan) and the Philippines. Lowland to 

submontane forests to 1200 m. 

Understorey shrubs. Stipules comb-shaped. Leaves spiral, blade with toothed 

margin. Inflorescences axillary; rachis thread-like, 1-5 cm long; bracts many; 

only single flower developing; pedicels jointed. Flowers 5-merous; stamens 5, 

anthers opening by longitudinal slits, staminodes many; carpels 3, united, ovary 1- 

celled, with distinct style and stigma. Fruit a capsule, splitting into 3. Seed not 

winged, pitted. 

Stem not so. Inflorescences terminal, with many flowers .............................................. 6 

6. 

Leaves oblanceolate, subsessile. Staminodes 

Indovethia Boerl. 

Feestbllndel P.J. Veth (1894) 89; Merrilll.c. 388; Masamune l.c. 469; Kanis I.c. (1968) 72, 

I.c. (1971) 112; Anderson i.c. 283. 

Monotypic genus (1. calophylla Boerl.). Central Sumatra and NW Borneo. Uncommon, 

in moist shady places in lowland forest. 

Shrublets. Stipules comb-shaped. Leaves spiral, margin entire or toothed. Flowers: 

sepals 5, persistent in fruit; petals 5, free; stamens 5, anthers opening by longitudinal 

slits, staminodes 10; carpels 3, united, ovary l-celled, with distinct style and 

stigma. Fruit a capsule, splitting into 3 including style. Seed not winged, surface 

pitted. 

Leaves linear with distinct petiole. Staminodes many ...................... 5. Schuurmansiella 

258

1. BRACKENRIDGEA A. Gray 

(W.D. Brackenridge, 1810-1893, the Scots-American botanist and horticulturist) 

New Gen. PI. (1853) 5; Merrilll.e. 387; Masmnune I.e. 468; Furtado, Gard. Bull. Sing. 19 (1962) 

181; Kanis I.e. (1968) 41, I.e. (1971) 101; Ng I.e. 256; Coekburn I.e. 64; Anderson I.e. 282; 

Whitmore, Tantra & Sutisna I.e. 278. 

Small to medium-sized trees. Stipules small, often divided, falling off early. Leaves 

alternate-distichous, glossy above; lateral veins strongly curved to the apex, often some of 

the lower ones parallel to the margin. Inflorescences thyrsoid of condensed cymes; rachis 

often growing vegetativeIy after flowering; bracts many, at base of inflorescences, falling 

259 

1. Brackenridgea hookeri (Planch.) A. Gray Fig. 1. 

(W.J. Hooker, 1785-1865, sometime Director of the Kew Botanic Gardens) 

I.e. 6; Furtado I.e. 182; Kanis I.e. (1968) 45, I.e. (1971) 102; Ng I.e. 256; Anderson I.e. 283; 

Whitmore, Tantra & Sutisna I.e. 278. Basionym: Gomphia hookeri Planeh. in Hooker, Land. 1. Bot. 

6 (1847) 3. Type: Phi/ipso S.n.. Penang (K). Synonyms: Gomphia corymbosa (King) Ridl., J. Str. Br. 

R. As. Soc. 54 (1910) 36 p.p. excl. typus; Braekenridgea dentieulata Furtado I.e. 183. 

Key to Brackenridgea species 

Inflorescence a many-flowered cyme; pedicels in two or more tiers .. .1. B. hookeri 

Inflorescence a 3(-5)-flowered cyme; pedicels in one tier only ....................... 2. B. palustris 

off early leaving distinct ring-like scars; pedicels jointed at base, filiform. Flowers 

arranged in 1-4 tiers; sepals 5, accrescent, fleshy and red in fruit; petals 5(-lO), white or 

yellow; stamens lO (or many), anthers dehiscing by longitudinal slits; carpels 5(-lO),free, 

but sharing a common style; ovules camptotropous or epitropous; stigma small. Fruit a 

drupe, 1-2(-5), greenish becoming almost black on ripening, surrounded by persistent 

sepals. 

Distribution. About 8 species, tropical E Africa, Madagascar, Malesia; 2 species in Sabah 


Ecology. Everwet tropical areas to lOOO m. Dispersal is mainly by birds because of the 

conspicuous black fruits on the red torus and calyx. Flowering and vegetative growth tend 

to occur at alternative intervals during the growth of the stem. After a period of leaf 

production, the terminal bud is covered with closely spaced scale-leaves, some of which 

bear inflorescences in their axils. When flowering and fruiting is completed the terminal 

bud resumes vegetative activity to bear foliage leaves again. This pattern of growth can be 

traced through the series of alternating leaf-scars and peduncle-bases along the twig. 

Taxonomy. Two sections are recognised in the genus. In section Brackenridgea, all flowers 

in a cyme open simultaneously; the corolla is 5-merous, white; there are lO stamens in one 

whorl; and 5 carpels. In section Notochnella, the flowers of a cyme open successively; the 

corolla is yellow and irregular; there are many stamens in more than 1 whorl; and 5-10 

carpels. Section Notochnella is confined to the Philippines. 

OCHNACEAE (KOCHUMMEN)

Ecology. Usually on lowlands, especially in peat swamp and kerangas forests, rarely in 

submontane forests to 1000 m. 

2. EUTHEMIS Jack 

(Greek, eu = good, themis = law; the even thickness and symmetry of the leaves) 

Mal. Misc. 1, 5 (1821) 5; Merrill/.e. 388; Ridley I.e. (1922) 367; Masamune I.e. 469; Kanis I.e. 

(1968) 62, I.e. (1971) 108; Ng I.e. 257; Anderson I.e. 283; Ashton I.e. 62. 

Shrubs. Stipules falling off early. Leaves spiral, with numerous close, almost parallel 

lateral veins; margin thick-rimmed, distinctly or faintly toothed. Inflorescences terminal 

and axillary racemes. Flowers: sepals persistent in fruit; petals 5, white or pink; 

staminodes 0-5; stamens 5, anthers opening by apical pores; carpels 5, united, ovary 5- 

260 

Distribution. Sumatra, Peninsular Malaysia, Borneo, Philippines, Celebes. Uncommon in 


faint; petiole 0.5-1 cm long. Flowersfascicled in one tier only, pedicel jointed. Fruits to 8 

Tree to 30 m tall, 40 cm diameter. Bark brown to reddish brown, smooth to scaly, inner 

bark reddish brown. Sapwood white to pale yellow. Leaves elliptic to oblong, 5.5-10 x 2-4 

cm; base cuneate, apex pointed; midrib raised above; lateral veins of lower half curving 

upwards, running parallel to the margin towards apex; intercostal veins invisible to very 

278. Type: Beeeari PB 3472, Borneo (holotype FI; isotypes A, K, P). Synonyms: Gomphia hookeri 

(Planch.) A. Gray var. eorymbosa King, J. As. Soc. Beng. 62 (1893) 233; B. serrulata Bartelli I.e. 

163, Merrilll.e. 387, Masamune I.e. 469; Gomphia eorymbosa (King) Ridl. I.e. (1910) 33,p.p., FMP 

1 (1922) 367. 

Malpigia 15 (1901) 165; Merrilll.e. 387; Masamune I.e. 468; Furtado I.e. 182; Kanis, I.e. (1968) 46, 

I.e. (1971) 102; Ng I.e. 256; Cockburn I.e. 64; Anderson I.e. 283; Whitmore, Tantra & Sutisna I.e. 

Distribution. India, Andamans, Thailand, Peninsular Malaysia and Borneo. Uncommon 

and of scattered occurrence in Sabah and Sarawak. 

Ecology. Lowland mixed dipterocarp forest on sandy humult ultisols to submontane forests 

curving upwards to almost to leaf apex, running parallel to margins; petiole 0.6-1 cm 

Tree to 30 m tall, 60 cm diameter. Bark reddish brown, smooth to slightly fissured and 

scaly; inner bark reddish brown. Leaves elliptic to oblong, rarely obovate, 6-17 x 2.5-7.5 

cm; base cuneate, apex pointed; midrib raised above; lateral veins of lower half of leaf 

long. Flowers clustered in two or more tiers; pedicel 1-2 cm long. Fruits 6-9 x 5.5-6 mm. 

to 1000 m, including kerangas and dry hillocks in swampy forests. 

(Latin, palustris = inhabiting boggy or marshy ground) 


2. Brackenridgea palustris Bartelli 

x6mm.

261 

Fig. 1. Brackenridgea hookeri. A, fruiting leafy twig; B, fruit; C, fruit in longitudinal section. (All 

from S. 27231.) 

'------- - 

10mm 

A 

[ "m 

OCHNACEAE (KOCHUMMEN)

262 

1mm l 

] 2cm 

1./ 

Fig. 2. Euthemis /eucocarpa (A-B) and E. minor (C-D). A, fruiting leafy twig; B, dehisced fruit; C, 

flower bud; D, open male flower. (A-B from SAN 84753, C-D after FM l, 7 (1972) 109, fig. 4.) 

TREE FLORA OF SABAH AND SARAW AK VOL. 1 (1995)

OCHNACEAE (KOCHUMMEN) 

celled, ovules 2 per cell, pendulous, axile; style distinct, stigma minute. Fruit a berry with 

5 pyrenes. Seeds 1-2 per cell. 

Distribution. 2 species; SW Cambodia, Sumatra, Peninsular Malaysia, Borneo. 

Ecology _ Kerangas forests, on low ridges, in peat swamp forests and also on ridges with 

poor sandy soils, below 1250 m. 

Uses. E. Ieueocarpa roots are used in traditional medicine in Peninsular Malaysia and in 

Brunei; the fruits are used to treat eye diseases. 

Key to Euthemis species 

Leaves 8-40 cm long; apex pointed; margin distinctly toothed; petiole 2-5 cm long. 

Mature fruit white ...................................................................................... 1. E. leucocarpa 

Leaves 4-15 cm long; apex rounded to slightly notched, with short mucro; margin faintly 

toothed; petiole to 1.5 cm. Mature fruit red ......................................................... 2. E. minor 

1. Euthemis leucocarpa Jack 

(Greek, leueo == white, karpos== fruit) 

Fig.2A-B. 

I.e. 16; Merrilll.e. (1921) 388; Rid1ey, FMP 1 (1922) 368; Masamune I.e. 469; Kanis I.e. (1968) 62, 

I.e. (1971) 108, Ng I.c. 257; Anderson fc. 283. Type: Wallich 2516, Singapore (K, neotype). 

Synonym: E. robusta Hook.! in Rid1ey I.c. 368. 

Shrub to 2 m tall. Stipules ovate. Leaves oblong to linear-oblong, 8-40 x 2-10 cm; base 

tapered, margin distinctly toothed, apex acute; midrib raised above; lateral veins numerous, 

1-2 mm apart; petiole winged, 2-5 cm long. Inflorescences in branched racemes. Fruits 

globose, 6-10 mm in diameter, white when mature. 

Distribution. Cambodia, Sumatra, Peninsular Malaysia, and Borneo. Widely distributed in 


Ecology. Lowland to submontane forests to 1000 m, on poor soils. In Sarawak frequent in 

mixed swamp and kerangas forests. 

2. Euthemis minor Jack Fig.2C-D 

(Latin, minor == small; the smaller size compared to E. Ieueoearpa) 

I.e. 18; Merrilll.e. (1921) 388; Rid1ey l.c. (1922) 368; Masamune I.e. 469; Kanis I.e. (1968) 65, I.e. 

(1971) 108; Ng I.c. 257; Cockburn I.e. 62; Anderson I.e. 283. Type: Wallieh 2517, Singapore (K, 

neotype). Synonyms: E. obtusifolia Hook.!, Trans. Linn. Soc. 23 (1862) 163, Merrilll.e. (1921) 

389, Masamune l.c. 469; E. ciliata Pearson, Kew Bull. (1906) 3; E. haekenbergii Die1s, Bot. Jahrb. 

60 (1926) 310. 

263

D 

O 

'1'~~~" 

8 mm 

.. ~-~. 

264 

E 

Fig. 3. Gomphia serrata. A, flowering leafy twig; B, open flower; C, infructescence; D, fruit; E, fruit 

in longitudinal section. (A-B from S. 38696, C-E from S. 32942.) 

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)


Shrub or treelet to 5 m tall. Twigs black. Leaves oblong to oblonceolate, 4-15 x l.5-4 cm; 

base tapered, margin faintly toothed, apex rounded with a short point; midrib raised above; 

lateral veins c. 1 mm apart, very faint to invisible; petiole 0.5-l.5 cm long. Inflorescence 

usually an unbranched raceme. Flowers sepals and petals pinkish when fresh. Fruits 

globose, 4-6 mm in diameter, with 5 prominent ridges. 

Distribution. Sumatra, Peninsular Malaysia, Borneo. Common in Sabah and Sarawak. 

Ecology. Lowland to submontane forests to 1250 m; locally abundant in Sarawak in kerangas 

and padang alan forest. 

3. GOMPHIA Schreb. 

(Greek, gomphos = a thorn or spike; 

the form of the flower base during fruit development) 

Gen. PI. ed. 8 (1784) 291; Ridley I.e. (1922) 365; Kanis, Taxon 16 (1967) 420, I.e. (1968) 51, i.e. 

(1971) 105; Ng I.e. 258; Cockburn I.e. 65; Anderson I.e. 283; Whitmore, Tantra & Sutisna I.e. 278. 

Synonyms: Campyloeerum Tiegh., Bull. Mus. Hist. Nat. Paris (1902) 546; Meesia Gaertn., Fruct. & 

Sem. PI. 1 (1788) 344. 

Shrubs or small to medium-sized trees. Stipules united, falling off early. Leaves spiral, 

margin firmly toothed; lateral veins numerous, close and almost parallel, with 2-3 distinct 

inframarginal veins; petiole to 3 mm long. Inflorescences in terminal and axillary panicles, 

peduncle perSistent. Flowers: sepals 5, enlarged and persistent in fruit; petals 5; stamens 

10, anthers opening by apical pores; gynophore columnar, 5-ribbed; carpels 5, free. Fruit 

a drupe. Seed one. 

Distribution. Mainly African; 1 species in SW India, Sri Lanka, E Thailand, Indo-China, 

Hainan and W Malesia. 

Ecology. Confined to everwet tropical and moderately dry monsoon areas from lowlands to 

1500 m. Dispersal is presumed to be by birds. 

Gomphia serrata (Gaertn.) Kanis 

(Latin, serratus = with teeth; the leaf margin) 

Fig. 3. 

I.e. (1967) 418, I.e. (1968) 53, I.e. (1971) 105; Ng I.e. 258; Cockburn I.e. 65; Anderson I.e. 283; 

Whitmore, Tantra & Sutisna I.e. 278. Basionym: Meesia serrata Gaertn. I.e. (1788) 344. Type: 

Koenig 25, Ceylon (L). Synonyms: Gomphia sumatrana Jack I.e. 29; G. mierophylla Ridl. I.e. 

(1922) 369; G. oblongifolia Ridl., Kew Bull. (1925) 281; Ouratia angustifolia (Vahl) Baill. ex 

Laness, Rev. Gen. PI. (1891) 106, Merrilll.e. 387, Masamune I.e. 470; 0. bomeensis Bartelli I.e. 

158, Merrilll.e. 387, Masamune I.e. 470; O. neriifolia Bartelli I.e. 158, Merrilll.e. 387, Masamune 

I.e. 470; O. beeeariana Bartelli I.e. 159, Merrilll.e. 387, Masamune I.e. 470; O. megaearpa Ridl., 

Kew Bull. (1930) 76, Masamune I.e. 470. 

Shrub or medium-sized tree, to 20 m tall, 40 cm diameter; bole slightly fluted. Bark dark 

grey-brown, scaly; inner bark pink, fibrous. Sapwood pale. Leaves shiny above, very 

265


Fig. 4. Schuurmansia elegans. Fruiting leafy twig. (From SAN 132717.) 

266


variable in shape and size, elliptic, oblong, or obovate, 5.5-33 x 2-7.5 cm; base cuneate, 

margin faint to distinctly toothed, apex pointed; midrib raised above; lateral veins 

numerous, close, almost parallel, very faint to almost invisible on both surfaces, forming 1- 

3 distinct wavy intramarginal veins; intercostal veins finely reticulate, very faint; petiole to 

3 mm long. Inflorescences axillary and terminal panicles. Flowers: pedicel c. 1 cm long; 

sepals 5, tinged pink, enlarged in fruit; petals 5, yellow or cream, obliquely obovate to 

broadly spathulate; stamens 10, subsessile or with very short filaments, anthers opening 

with 2 apical pores, obovoid; carpels 5, free, with a single style. Fruits kidney-shaped, 1- 

2(-5), yellowish green, ripening purplish, 0.5-1 x 0.4-0.6 cm, with persistent sepals. Seed 

one. 

Vernacular names. Sabah--antimagas gimbaan (Murut), kolambang (Dusun), majangmajang 

(Kadazan), quintalai (Kadazan). Sarawak--aam (Kenyah), chinaga-Iampong, 

keladang, kelutak (!ban), ladin (Malay). 

Distribution. SW Peninsular India, Sri Lanka, E Thailand, Indo-China, Hainan, Malesia. 

Common and widely distributed in Sabah and Sarawak. 

Ecology. Lowland forests on well-drained infertile organic soils, including mixed dipterocarp 

and kerangas forests, to submontane forest to 1500 m, sometimes occurring by stream 

banks, rarely on limestone forests. Flowering and fruiting samples have been collected 

throughout the year. As this species is found in a number of ecological habitats there is 

considerable variation in its morphological characters. This variation is more pronounced 

in Sarawak. In the more exposed habitats, on cliffs and on poor kerangas soils, this species 

has very small leaves. 

Uses. Roots and leaves are bitter and are decocted in S India for a stomachic and antiemetic 

tonic. Young branch tissue is used to treat toothache in Cambodia. 

4. SCHUURMANSIA Blume 

(l Schuurmans Stekhoven, 1972-1855, 

horticulturist at the Leiden University Botanical Garden) 

Mus. Bot. Lugd. Bat. 1 (1850) 177; Masamune I.e. 470; Kanis I.e. (1968) 74, I.e. (1971) 115; 

Anderson l.c. 283; Whitmore, Tantra & Sutisna I.e. 278. 

Shrubs or small trees. Twigs hollow. Stipules entire, narrow, lanceolate. Leaves spiral, 

shiny, margin entire or toothed, glandular; lateral veins invisible. Inflorescences in terminal 

panicles. Flowers: sepals 5; petals 5; stamens 5, anthers opening by longitudinal slits, 

staminodes many in one or two whorls; carpels 3, united, ovary 1-celled. Fruit a capsule, 

splitting into 3. Seeds winged. 

Distribution. 3 species; Borneo, Philippines, Celebes, Moluccas, New Guinea, Solomon 

Islands, Bismarks; 1 species in Borneo. 

Ecology. Pioneer plants from sea-level to 3000 m. 

267

Distribution. Monotypic, endemic to Borneo (Sarawak). 

Ecology. Mainly lowlands, especially in kerangas forests on poor soils and on sandstone 

cliffs near sea, rarely to 600 m. 

Schuurmansiella angustifolia (Hook. f) Hall. f 

(Latin, angustus = narrow,folium = leaf) 

l.c. 345; Merrilli.c. (1921) 387; Masamune i.c. 470; Kanis l.c. (1968) 73, I.c. (1971) 114; Anderson 

i.c. 283. Basionym: Schuurmansia angustifolia Hook.! i.c. 157. Type: Lobb, s.n., Sarawak (K). 

Shrub to 10 m tall. Stipules to 12 x 0.5 mm. Leaves linear-oblong, 8-17 x 0.7-1.45 cm; 

base tapered, apex pointed; midrib raised above; lateral veins invisible to very faint, numerous, 

close; petiole 0.5-1 cm long. Flowers: petals white with pink base; stamens purplish; ovary 

ovoid, style short, purplish, stigma capitate. Fruits ellipsoid, c. 8.5 x 3 mm. 

268 

'- ---.----- 

Fig. 5. 

Shrubs. Stipules needle-like. Leaves spiral, margin finely toothed. Inflorescences terminal 

racemes. Flowers: sepals 5, petals 5, staminodes many, stamens 5, anthers opening by 

longitudinal slits; carpels 3, united, ovary l-celled, style short, stigma capitate. Fruit a capsule, 

Small tree to 5 m tall. Leaves obovate to oblong, 10-30 x 2.5-10 cm, shiny, surface of 

Distribution. Borneo, Philippines, Celebes, Moluccas, New Guinea. In Sabah and Sarawak 

uncommon, known by only 2 collections (Sabah, SAN A 3636; Sarawak, Nooteboom 1517). 

Rec. Trav. Bot. Neerl. 10 (1913) 344; Merrilli.c. 387; Masamune i.c. 470; Kanis I.c. (1968) 73, I.c. 

(1971) 113; Anderson I. c. 283. 

Fruits fusiform, c. 2.5 x 0.5 cm, with pointed tip and persistent sepals. Seeds c. 3 mm long 

278. Type: Zippelius, s.n., Amboina (L). Synonyms: S. parviflora Ridl., Trans. Linn. Soc. 2, Bot. 9 

(1916) 18; S. borneensis Ridl., Kew Bull. (1930) 77, Masamune i.c. 470. 

dried leaves finely reticulate; base tapered, apex rounded to blunt; lateral veins many, 

close, almost parallel, very faint on both surfaces; petiole 5-6 cm long. Flowers yellow. 

l.c. 178; Kanis i.c. (1968) 75, i.c. (1971) 115; Anderson i.c. 238; Whitmore, Tantra & Sutisna i.c. 

Fig. 4. 

5. SCHUURMANSIELLA Hall.! 

Ecology. Lowlands and hills including limestones, to 900 m. 


(Latin, elegans = elegant; the growth habit) 

Schuurmansia elegans Blume 

splitting into 3. Seeds hairy. 

with slender wings.


5mm 

c 

Fig. 5. Schuurmansiella angustifolia. A, flowering leafy twig; B, dehisced fruit; C, open flower. CA 

& C from S. 21431, B from Anderson 8368.) 

269


Distribution. Endemic to western Sarawak. 

Ecology. Lowlands from sea-level to 600 m, mainly in kerangas forest. 

270


OLACACEAE 

Lesmy Tipot 



Hooker f, Fl. Brit. Ind. 1 (1875) 572 (under 0Iacineae); King, J. As. Soc. Beng. 64, 2 (1895) 108; 

Merrill, EB (1921) 242; Ridley, FMP 1 (1922) 419; Sleumer in Engler & Prantl, Pfl. Fam. 2, 16b 

(1935) 5, Blumea 26 (1980) 145, FM 1, 10 (1984) 1; Masamune, EPB (1942) 258; Backer & 

BakhuizenJ, FJ 2 (1965) 63; Smythies, CST (1965) 113; Whitmore, TFM 2 (1973) 299; Keng, 

OFMSP (1978) 212; Anderson, CLTS (1980) 284; Cockburn, TS 2 (1980) 65; Corner, WSTM 2 

(1988) 600; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 279. 

Trees, shrubs or climbers. Leaves spirally arranged, rarely distichous, simple, entire, 

stalked, without stipules, when dry often with parchment-like and/or finely tuberculate 

surface, pinnately veined with the lateral veins rather distant. Inflorescences axillary 

cymes, racemes or branched spikes. Flowers bisexual, rarely unisexual, mostly radially 

symmetrical, rarely heterostylus; calyx often cup-like, shortly 3-7-1obed or dentate, sometimes 

accrescent; petals 3-7, free or joined below; stamens as many or twice the number of 

petals; disc present; ovary mostly superior, rarely inferior, 3-5-loculed, ovule solitary in 

each locule; style usually 1, stigma 3-5-lobed. Fruit a drupe; exocarp thin or fleshy; endocarp 

crustaceous to woody. Seeds one per fruit, with thin testa and abundant endosperm. 

Distribution. Pantropical, 27 genera with c. 170 species. In Sabah and Sarawak, represented 

by 8 genera with 9 species of trees, shrubs and climbers. 

Ecology. Found in a wide range of habitats, from open and sandy seashores to secondary 

and primary mixed dipterocarp forests. Occasionally in submontane forests. 

Uses. Most species are small to medium-sized trees and are, therefore, commercially not 

important, except for Scorodocarpus borneensis which produces a dark red timber. Species 

of Anacolosa, Strombosia and Ochanostachys are occasionally used in local construction 

works. The wood of Ximenia americana can be used as a substitute for sandal wood, 

whereas the fruit kernel is said to have medicinal properties. Fruits of Scorodocarpus 

borneensis are edible and the young leaves are taken as vegetable. 

Taxonomy. Engler (Syllabus, 1924) recognised Olacaceae as the most primitive family in 

the order Santalales. Ridley I. c., following Hooker f I. c., on the other hand, treated 

Olacaceae together with Icacinaceae under the order Olacineae. Hutchinson (Fam. FI. PI. 1 

(1964) 329) placed Olacaceae together with Opiliaceae and a few other families in the 

Olacales, an order comparatively more primitive than the Santales. Keng I.c., Sleumer I.c. 

and Whitmore I. c. followed Engler's classification, a stand accepted in the present 

account. The Santalales consists of a small group of families showing a tendeI).cy toward 

271


2cm 

,mmI 

B 

c 

2 cm 

Fig. 1. Anacolosa frutescens. A, flowering leafy twig; B, longitudinal section of flower bud; C, fruit. 

(FromS. 29191.) 

272

OLACACEAE (LESMY) 

hemiparasitism, reduction of the flowers and adaptation to the climbing and epiphytic 

habit. In this account, the nomenclature follows that of Sleumer's revision (1984), in which 

he has considerably reduced the number of species of Strombosia and Olax recognised by 

earlier workers. 

Key to genera 

l. 

Slender woody climbers, often with axillary tendrils ........................ . 

Erythropalum Blume 

Bijdr. (1826) 921; Merrill, PEB (1929) 58; Masamune I.e. 258; Sleumer I.e. (1980) 151, 

I.e. (1984) 17. 

One species, E. scan dens Blume, distributed from the Himalayas to Assam, 

Bengal, Burma, Indo-China, Thailand and Malesia. In Sabah, recorded from 

Kinabatangan, Lahad Datu, Ranau, Tambunan, Tawau, Tenom and Mt. Kinabalu 

at c. 1200 m. In Sarawak, in Melinau and Bukit Jebong in Bau district. 

Fruit drupaceous, ellipsoid, ripening reddish, entirely enclosed by the persistent 

calyx which finally splits from top downwards into 3-6 re flexed segments. In open 

areas and hillsides. 

Trees, shrubs or scandent shrubs, without axillary tendrils ........................................... 2 

2. Leaves usually mucronate at apex. Twigs armed with thorns .................. . 

XimeniaL. 

Sp. Pl. (1753) 1193; Masamune I.e. 259; Sleumer l.c. (1980) 166, I.c. (1984) 10. 

Eight species in the tropics and subtropics. I species, X americana L., in Borneo 

and has been recorded from Teluk Tambak, Santubong near Kuching, Lubok 

Jayau and Tatau near Bintulu in Sarawak; and from Lahad Datu, Sandakan and 

Semporna in Sabah. 

Low-branching shrub, deciduous in dry season. Usually on sandy seashores and in 

dry forest fringes, sometimes on stony ground. 

Leaves not mucronate. Twigs without thorns ............................................................ 3 

3. Leaves distichous or subdistichous. Calyx well-developed, enclosing fruit or flat and 

fleshy at maturity ....................................................................................................... .4 

Leaves not distichous. Calyx not developed as above .................................................... 5 

4. Leaves elliptic, both surfaces parchment-like. Fruit seated on an enlarged flattish 

calyx ....................................................................................................... 2. Harmandia 

Leaves ovate, elliptic to oblong, surface not parchment-like. Fruit enclosed in the 

enlarged calyx .............................................................................................................. . 

Olax L. 

Sp. Pl. (1753) 34; Merrilll.e. (1921) 242; Masamune l.c. 259; Sleumer I.c. (1984) 8. 

About 40 species, Old World Tropics. I species, 0. imbricata Roxb., distributed in 

India, Sumatra, Peninsular Malaysia, Borneo, Java, New Guinea and Solomon 

Islands. In Sabah, recorded from Lamag, Mt. Lotung, Bongawan, Papar, Sipitang, 

and Sandakan. In Sarawak, in Lawas, Limbang, Sibu, Kapit and Kuching. 

273


Scandent shrub or climber. On open sandy ground including coasted areas, and 

occasionally on rocky ground along streamsides. 

5. Leaves usually with fine pellucid-dots visible against strong light.. ............................. 6 

Leaves not so ............................................................................................................. 7 

6. Young twigs zig-zag. Petals entirely free ............................................. 5. Strombosia 

Young twigs not zig-zag. Petals fused in the lower haIL ......................... 1. Anacolosa 

7. All parts reeking of garlic smell when fresh. Leaves broadly elliptic, c. 20 cm long. 

Bark fissured or thinly flaky, inner bark without any sap .................. 4. Scorodocarpus 

Garlic smell absent. Leaves elliptic, rather small. Bark dippled, distantly scaly, inner 

bark with a slight white sap ............................................................... 3. Ochanostachys 

1. ANACOLOSA Blurne 

(Greek, anakolos = knotted; the calyx-cup rim) 

Mus. Bot. Lugd. Bat. 1 (1850) 250, t. 46; Merrilll.e. (1921) 242; Rid1ey I.e. (1922) 424; Sleumer in 

Eng1er & Prant1l.e. (1935) 20, I.e. (1980) 146, I.e. (1984) 23; Masamune I.e. 258; Whitmore I.e. 

300; Anderson I.e. 283; Whitmore, Tantra & Sutisna I.e. 279. 

Trees or shrubs. Leaves spiral, pinnately veined. Flowers bisexual, in axillary cymes or 

fascicles; calyx cup-shaped, (5-)6(-7)-dentate or minutely toothed; petals (5-)6(-7), inserted 

on the margin of the cupular disc, fused in the lower part, fleshy and concave below; 

stamens (5-)6(-7), filaments short, anthers ovoid; disc hypogenous, fused with the ovarywall, 

6-denticulate or furrowed; ovary superior, incompletely 2(-3)-celled below, l-celled 

above, style short and thickened at base; ovules 2(-3), unitegmic, pendant, placentation 

central. Fruit a drupe, surrounded by the enlarged disc, tipped by the remains of the style, 

seated on the small persistent calyx; pericarp thin, fleshy; endocarp thin, crustaceous. 

Seeds: embryo minute; endosperm starchy and oily. 

Distribution. 15 species, in the Old World tropics, of which 3 species occur in Malesia. In 

Sabah and Sarawak, one species. 

Anacolosa frutescens (Blume) Blume 

(Latin,frutex = a shrub; the habit) 

Fig. 1. 

I.e. (1850) 251, t. 46; Masamune I.e. 258; Sleumer I.e. (1980) 146, I.e. (1984) 25; Anderson I.e. 283; 

Whitmore, Tantra & Sutisna I.e. 279. Basionym: Stemonurus fruteseens Blume I.e. (1826) 649. 

Type: Elume "2168", Java (ho1otype L; isotypes P, U). Synonyms: Anaeolosa heptandra Maingay ex 

Mast. in Hooker f I.e. 581; A. arborea Koord. & Va1eton, Bull. Inst. Bot. Buitz. 2 (1899) 9; A. 

luzoniensis Merr., Philip. J. Se. 4, Bot. (1909) 253; Anaeolosa sp. Merr. I.e. (1921) 242; Salaeia 

bartletti Ridl., Kew Bull. (1938) 239. 

Erect shrub or small slender tree, to 30 m tall and 30 cm diameter. Bark smooth to papery 

or mottled, greenish grey; inner bark reddish. Sapwood yellow to pale red. Twigs greyish 

274


brown or whitish. Leaves very variable in shape, from elliptic to elliptic-oblong or 

lanceolate, 9-18 x 6-9.5 cm, papery to leathery, glossy above when fresh, brownish and 

dull in dry state, usually with tiny warts or tubercles on both surfaces, or with fine 

pellucid-dots visible against strong light; base cuneate, slightly asymmetric, margin entire, 

apex acuminate, blunt or rounded; lateral veins 5-7 pairs, ascending, rather distant, lowest 

pair close to the base, faint on upper surface, conspicuous and slightly raised below; petiole 

stout, 5-12 mm long. Flowers in axillary clusters of 5-15, pedicels 2-5 mm; calyx shortly 

5-7-lobed, glabrous or pale rusty-puberulous, c. 3 mm in diameter; petals 6, ovatelanceolate, 

glabrous or rarely puberulous outside, 2-3 x 1-1.5 mm; stamens 6, anthers 

hairy; ovary with 2-3 pendulous ovules. Fruits obovoid to oblong, with persistent calyx, 

green, ripening yellow to orange, c. l.2 x 0.8 cm. Seeds: endosperm copious. 

Vernacular names. Sabah-salangugapit (Dusun, Tawau). Sarawak-belian landak (Malay, 

Iban),jerit (Iban), ladit (Melanau). 

Distribution. Burma, Andaman and Nicobar Is., Thailand, Sumatra, Peninsular Malaysia, 

Java, Borneo, Philippines, Celebes and the Moluccas. In Sabah, recorded from Kalabakan, 

Keningau, Kota BeIud, Lahad Datu, Ranau, Sandakan, Sepulut and Tawau. In Sarawak, 

known from Kuching, Sibu, Sarikei, Kapit, Bintulu, Miri to Baram. 

Ecology. Common on hillslopes of primary mixed dipterocarp forests on sandy humult 

uItisols, sometimes in heath and peat swamp forests. Once recorded from limestone hill, 

and once from submontane forest at about 1100 m. 

Uses. The wood is pale reddish brown, considerably hard and heavy, and sometimes used 

locally for house-posts though not very durable. 

2. HARMANDIA Pierre ex Baill. 

(Jules Harmans, 1845-1921, French colonial officer) 

Bull. Linn. Soc. Paris 2 (1889) 770; Sleumer I.e. (1935) 30, I.e. (1980), I.e. (1984) 9; Whitmore I.e. 

301; Whitmore, Tantra & Sutisna I.e. 279. 

Trees monoecious. Leaves distichous, pinnately veined. Inflorescences short, axillary 

racemes. Flowers unisexual; calyx at anthesis disc-form, shortly 4-dented, much accrescent 

at maturity forming a frill below the fruit; petals 4 in male, 6-8 in female flowers, connate 

to an urceolate tube at base, free at the upper part; disc extra-staminal, annular. Male 

flowers: stamens 4, epipetalous, filaments fused into a tube with the free anthers on top; 

ovary rudimentary. Female flowers: staminodal tube without anthers; ovary pyramidal, 1- 

celled, with 2 pendant ovules, placentation basal; style short-conical, stigma 3, sessile. 

Fruit a drupe, concrescent with the much enlarged calyx below; pericarp fleshy; endocarp 

thin and woody. Seeds: endosperm oily; embryo excentric. 

Distribution. Monotypic; Indo-China and W Malesia (Sumatra, Peninsular Malaysia and 

Borneo). 

275


[2 cm 

B 

A 

Fig. 2. Harmandia mekongensis. A, leafy twig; B, fruit. (From SAN 107911.) 

276


Ecology. Usually found in lowland forests. 

Harmandia mekongensis Pierre ex Baill. 

(of Mekong River in Laos) 

Fig. 2. 

I.c. 770; Sleumer I.c. (1980) 153, I.c. (1984) 9. Type: Harmand 1322, SE Laos (holotype P; isotypes 

BM, K, L, SING). Synonym: Harmandia kunstleri King I.c. 100. 

Medium-sized tree to 30 m tall, 30 cm diameter. Bark thin, pale brown to whitish, flaky; 

inner bark pale yellow, granular. Sapwood pale yellow. Twigs slender, dark brown to 

black, striate and slightly zig-zag. Leaves elliptic, 7-8.5 x 3-4 cm, subcoriaceous, 

parchment-like especially on lower surface, deep green when fresh; base cuneate to slightly 

rounded, margin recurved, apex pointed; lateral veins 4-6 pairs, rather inconspicuous on 

both surfaces; reticulations very faint; petiole 8-10 mm long. Flowers pale green to 

whitish; calyx cupular at bud stage, accrescent in frUit; petals c. 2 mm, connate below, 

forming an urceolate corolla. Male flowers: stamens 4, filaments connate to a fleshy tube, 

c. l.5 mm; anthers cordate, c. 0.5 mm; pistillodes present. Female flowers: staminodal 

tube without anthers; ovary conical, tapering to a short style; stigmas 3, sessile. Fruits 

ovoid-ellipsoid, green, maturing black, c. 3 x l.5 cm, with an enlarged pale red and fleshy 

calyx below. Seeds: pericarp fleshy, c. 0.5 mm thick; endocarp woody, c. 0.5 mm thick. 

Distribution. Indo-China (Laos, Annam) and W Malesia (Sumatra, Peninsular Malaysia 

and Borneo). In Sabah very uncommon, represented by only two collections from Keningau 

(SAN 107911) and Nabawan (SAN 118776). Not yet recorded from Sarawak. 

Ecology. In lowland and hill primary and disturbed forests to about 300 m. 

3. OCHANOSTACHYS Mast. 

(Greek, okanon = shield strap, stachus = spike; the inflorescence) 

in Hooker f I.c. 576; Merrilll.c. (1921) 242, I.c. (1929) 58; Ridley I.c. (1922) 422; Masamune I.c. 

259; Whitmore l.c. 302; Anderson l.c. 284; Sleumer l.c. (1980) 153, I.c. (1984) 12; Whitmore, Tantra 

& Sutisna I.c. 279. Synonym: Petalinia Becc., Malesia 1 (1883) 257. 

Trees. Leaves spiral. Flowers bisexual, on simple or branched axillary spikes; calyx small, 

cup-shaped, 4-5-lobed, not accrescent; petals 4-5, almost free; stamens 8-15, epipetalous, 

(1-)2(-3) on each petal, filaments joined at base, anthers globular to subglobular; disc 

hypogynous, fleshy, rather inconspicuous; ovary superior, incompletely 3-4-chambered, 

style short, stigma 3-lobed; ovules bitegmic, pendulous, placentation free basal. Fruit a 

drupe; pericarp thin; endocarp woody. Seeds: endosperm starchy; embryo small, apical. 

Distribution. Monotypic; confined to W Malesian region (Sumatra, Bangka, Lingga Island, 

Peninsular Malaysia, and Borneo). 

Ecology. Lowland rain forest. 

277


Ochanostachys amentacea Mast. 

(Latin, amentum = a catkin; the inflorescence) 

Fig. 3. 

in Hooker f I.e. 577; Merrill, I.e. (1921) 242, I.e. (1929) 58; Ridley, Kew Bull. (1931) 35, incl. var. 

ru/a Stapf ex Rid!.; Masamune I.e. 259; Browne, FTSB (1955) 251; Smythies I.e. 113; Burgess, TBS 

(1966) 420; Whitmore I.e. 302; Anderson I.e. 284; Cockbum I.e. 67; SIeumer I.e. (1980) 153, I.e. 

(1984) 14; Whitmore, Tantra & Sutisna I.e. 278. Type: Maingay 384, Malacca (lectotype K; 

isolectotype P). Synonyms: Petalinia baneana Becc. I.e. 258; Oehanostaehys baneana (Becc.) 

Valeton, Crit. Overz. OIacin. (1886) 104. 

Small to medium-sized tree, to 30 m tall, 70 cm diameter; bole straight to (often) rather 

poorly formed, at base shortly fluted or buttressed. Bark grey-brown to reddish brown, 

shedding off into thin, irregular scales, exposing lighter coloured patches, giving a dippled 

appearance; inner bark finely fibrous, yellowish brown, with discrete droplets of white 

latex. Sapwood hard, brownish yellow. Leaves usually glabrous, thinly leathery, ovateelliptic 

to elliptic-oblong, 6-13 x 3-7 cm, green and shiny above, yellowish green beneath 

when fresh, rather dull and brownish when dry, usually with blackish dots on both 

surfaces; base cuneate or rounded, margin entire, apex acute to acuminate; lateral veins 4- 

6 pairs, rather distant, sunken above prominent below; intercostal veins scalariform,very 

fine and rather faint; stalk slender, l.5-2.5 cm, with a faint, darker, 5 mm, apical knee. 

Inflorescences simple or branched spikes to 12 cm long. Flowers green, in opposite, wellspaced 

clusters; calyx 4-5-toothed, c. 1 mm; petals (3-)4(-5), ovate to ovate-oblong, c. 2.5 

x l.5 mm; ovary depressed ovoid, striate, glabrous, style short, cylindrical. Fruits 

sub globose, to 2 cm across, on a slender peduncle of c. 2 mm; pericarp thin, exuding a 

milky gum; endocarp woody. Seeds subglobose. 

Vernacular names. Sabah-tanggal (Dusun). Sarawak-imah (Bidayuh Bau), petaling 

(Malay), sagad berauh (Murud), santikal (lban). 

Distribution. Sumatra, Bangka, Peninsular Malaysia and Borneo. Common and widespread 

throughout Sabah and Sarawak. 

Ecology. Main canopy tree in primary and secondary mixed dipterocarp forest, on hillsides 

and ridges on loamy usually clay-rich soils, to 800 m, occasionally in heath forest. 

Uses. Produces a yellowish brown, hard, heavy, and reasonably durable wood, useful for 

house construction and logging railways. 

4. SCORODOCARPUS Becc. 

(Greek, skorodon = garlic, karpos = fruit; the strong garlic smell in the fruit) 

Nuov. Giom. Bot. Ita!. 9 (1877) 274, t.II, fI2-17; Merrilll.c. (1921) 242; Ridley I.c. (1922) 424, 

Masamune l.c. 259; Smythies l.c. 113; Whitmore I.c. 303; Anderson I.e. 284; Cockbum l.c. 67; 

SIeumer I.e. (1980) 160, I.e. (1984) 15; Whitmore, Tantra & Sutisna I.e. 279. 

Garlic-smelling tree, with straight bole, without buttresses. Leaves spiral. Flowers bisexual, in 

short racemes; calyx small, cup-shaped, narrow~y 4-5-toothed, not enlarged in fruit; petals 

278


'mm I 

c 

[ 'mm 

B 

Fig. 3. Ochanostachys amentacea. A, flowering leafy twig; B, flower bud; C, flower bud in section; 

D, fruit. (From SAN 61173.) 

279


D 

~ 

2cm 

'--------' 

2cm 

[ I mm 

B 

[ Imm 

c 

Fig. 4. Scorodocarpus bomeensis. A, flowering leafy twig; B, flower; C, longitudinal section of 

flower; D, fruit. (From S. 36079.) 

280


4-5, hairy within; stamens 8-10, in pairs, joined halfway down, and attached to the base of 

petal; ovary superior, imperfectly 3-4-celled, style elongate conical, stigma 3-4-lobed, 

ovule 1 in each cell, pendant, placentation free. Fruit a drupe; pericarp thin, fleshy; 

endocarp woody. Seeds: endosperm fleshy, containing starch and tannin. 

Distribution. Monotypic; S Thailand, Sumatra, Lingga Island, Peninsular Malaysia and 

Borneo. 

Ecology. Common in lowland forest. 

Scorodocarpus borneensis (Baill.) Becc. 


Fig. 4. 

I.e. 274, t. 11, f. 12-17; Merrilll.e. (1921) 242; Ridley I.e. (1922) 424; Masamune I.e. 259; Brown 

I.e. 280; Smythies I.e. 422; Whitmore I.e. 303; Anderson I.e. 284; Cockburn I.e. 67; Sleumer I.e. 

(1980) 160, I.e. (1984) 15; Whitmore, Tantra & Sutisna I.e. 279. Basionym: Ximenia borneensis 

Baill., Andansonia 11 (1874) 27l. Type: Eeeeari PE 1581, Sarawak, Mt. Matang (holotype FI; 

isbtypes P, W). 

Medium-sized to large tree, to 40 m tall, 80 cm diameter; all parts with garlic smell 

especially when crushed or cut; crown small, dense; bole straight, to 25 m, occasionally 

with small, low buttresses. Bark grey to dark red or brown, fissured and thinly 

rectangularly flaky; inner bark purplish red, inwards with coarse orange flecks. Sapwood 

hard, yellow to reddish brown. Leaves coriaceous, shiny green above when fresh, dull 

olive-green when dry, glabrous, elliptic, 10-22 x 4-9 cm; base cuneate to rounded, margin 

entire, apex acuminate; lateral veins 5-6 pairs, distant, curving upwards towards margin, 

flat above, very prominent beneath; reticulation rather faint; petiole 1.5-2 cm, thickened 

distally or almost at the leaf-base. Inflorescences racemose, to 4 cm, rusty to greyish 

puberulous. Flowers white, c. 1.5 cm long, solitary or grouped in clusters of 2-3 along the 

rachis; pedicels c. 1.5-2 mm; calyx small with wavy edges; petals white or creamy white, 

8-10 x 2 mm, woolly inside, reflexed; anthers 3-4 mm; ovary yellowish green, tapering 

toward the thickish white style. Fruits globose, green, c. 5 cm across; peduncle 1 cm; 

pericarp thin and fleshy; endocarp woody with numerous vertical fibre-like hard strands. 

Seeds subglobose. 

Vernacular names. Sabah-bawang hutan (Malay). Sarawak-sagan-berauh (Murut) , 

sindok, sindu (Iban), troduh (Bidayuh Bau). 

Distribution. Peninsular Thailand, Sumatra, Lingga Is., Peninsular Malaysia, Borneo. In 

Sabah and Sarawak widespread. Also occurs in Brunei and Kalimantan. 

Ecology. Commonly found on slopes in primary and secondary mixed dipterocarp forest on 

clay loam soils, and occasionally in seasonally flooded alluvial forest. 

Silviculture. Bawang hutan requires well-drained soils for good growth under normal 

conditions. It is evergreen but slow-growing. Fruiting usually occurs during June 

September. 

281

TREE FLORA OF SABAH A,1\ID SARAWAK VOL. 1 (1995) 

2cm 

"m[ 

[lmm 

c 

B 

Fig. 5. Strombosia javanica. A, flowering leafy twig; B, flowers; C, longitudinal section of flower. 

(FromS.34250.) 

282


Uses. Produces a dark brown, moderately durable, medium hardwood timber useful for 

piling, making bridges and house-posts. The fruits are boiled and eaten by many 

communities in Borneo (s. 36079 and SAN 10463). The young leaves are also cooked as a 

vegetable in Sarawak. 

5. STROMBOSIA Blume 

(Greek, strombos = conical or pear-shaped; the fruit) 

I.c. (1826) 1154; Merrill/.c. (1921) 242; Rid1ey I.c. 425; Sleumer in Eng1er & Prantll.c. (1935) 21, 

I.c. (1980) 163, I.c. (1984) 19; Masamune I.c. 259; Whitmore I.c. 305; Anderson I.c. 284; Whitmore, 

Tantra & Sutisna I.c. 280. Synonym: Lavallea Bai1l., Adansonia 2 (1862) 361. 

Shrubs or trees. Young twigs distinctly zig-zag, smooth, light coloured, old ones often black. 

Leaves spirally arranged, pinnately veined, glabrous; lateral veins rather distant, ascending. 

Flowers bisexual, in sessile or shortly stalked cymes or clusters; calyx cup-shaped, 5- 

lobed, accrescent and adnate to the pericarp almost to the top of the mature frUit; petals 5, 

free, inner parts hairy; stamens 5, adnate to the petals, filaments flat, with simple hairs, 

anthers dorsifixed; disc prominent, 5-lobed; ovary partly sunken into the receptacle, almost 

entirely covered by the 5-lobed fleshy disc, 3-5-chambered; ovules 3-5 per locule, 

anatropous, placentation free central; style short, stigmas subglobular, obscurely 3-5- 

lobed. Fruits drupaceous, crowned by the persistent calyx and style base; pericarp thinfleshy; 

mesocarp crustaceous or woody. Seeds: embryo small, apical; endosperm fleshy and 

oily. 

Distribution. About 12 species, c. 9 of which are confined to tropical Africa; the rest occur 

in India, Sri Lanka, Burma, Thailand, and Malesia (Sumatra, Peninsular Malaysia, Java, 

Borneo, the Philippines and N Moluccas). In Sabah and Sarawak, represented by 2 species. 

Ecology. Generally lowland forest. 

Key to Strombosia species 

Leaves smooth on upper surface, not pellucid-punctulate; intercostal veins usually visible. 

Flowers in cymose inflorescences. Fruits with truncate apex, apiculate by the persistent, 

hard style-base ................................................................................................ 1. S. javanica 

Leaves parchment-like on both surfaces, distinctly pellucid-punctulate; intercostal veins 

seldom visible Flowers in clusters. Fruits with obtuse-rounded apex, remains of style-base 

inconspicuous ................................................................................................ 2. S. ceylanica 

1. Strombosia javanica Blume 


Fig. 5. 

I.c. (1826) 1155; Merrill I.c. (1921) 242; Rid1ey I.e. 425; Masamune I.e. 259; Browne I.e. 282; 

Whitmore I.c. 306; Sleumer I.e. (1980) 164, I.e. (1984) 21; Whitmore, Tantra & Sutisna I.e. 280. 

Type: Blume, s.n., West Java, Mt. Sa1ak (ho1otype L; isotypes K, P). 

283


Tree to 40 m tall, 100 cm diameter; bole straight, often with knobs; crown dense and 

narrow. Bark yellowish grey, shallowly irregularly fissured; inner bark pinkish, fibrous. 

Sapwood pale yellow. Leaves elliptic-oblong, (10-)12-18(-24) x 4-8 cm, thick 

membranous to subcoriaceous, drying yellowish olivaceous or brownish, smooth and shiny 

above, without pellucid-dots; base obtuse to rounded, margin entire, apex subacuminate; 

lateral veins 5-7 pairs, conspicuous and raised below, flat above; petiole slightly swollen 

distally, 1.5-2 cm. Flowers solitary or in 3-7-flowered fascicles; calyx disc-shaped, 4-5 

angular, teeth obscure, c. 3 mm; petals greenish white, reflexed at apex, densely hairy 

inside, glabrous outside, ovate-lanceolate, 8-10 x 2-3 mm; filaments ciliate at the apex, 

anthers ovate-oblong, c. 0.5 mm; ovary de.eply 5-furrowed lengthwise; style columnar; 

stigma obscurely 5-lobed. Fruits ovoid, 2-3.5 x 1.5-2 cm, green, with thin and fleshy 

pericarp and woody endocarp, apex truncate, crowned by the remains of the calyx, disc and 

style. 

Vernacular name. Sarawak-belian landak (!ban). 

Distribution. Burma, S Thailand, Sumatra, Peninsular Malaysia, Java and Borneo 

(Sarawak, Brunei and Kalimantan). In Sarawak uncommon (e.g., S. 34250, S. 35255 from 

Mt. Gading, Lundu, and S. 1476 from Marudi). Not yet recorded in Sabah. 

Ecology. Scattered but locally common in mixed dipterocarp forest and secondary forest to 

600 m, on clay-rich fertile soils. 

Uses. Produces a moderately hard, durable, light yellowish timber, used locally for house 

construction. 

2. Strombosia ceylanica Gardner 

(ofCeylon) 

Calc. 1. Nat. Hist. 6 (1845) 350; Sleumer I.e. (1980) 165, I.e. (1984) 22; Whitmore, Tantra & Sutisna 

I.e. 280. Type: Gardner, s.n., Ceylon, Hantane (holotype K; isotype BM). Synonyms: S. lucida 

Teijsm. & Binn. ex Valeton I.e. 93; Anaeolosa maingayi Mast. in Hooker f I.e. 580; S. maingayi 

(Mast.) Whitmore, Gard. Bull. Sing. 26 (1973) 285; S. rotundifolia King I.e. lO2; S. multiflora King 

I.e. lO2; S. latifolia Stapf, KewBull. (1906) 71, Merrilll.e. (1921) 242, Masamune I.e. 259. 

Small to medium-sized tree, 40 m tall, 60 cm diameter; crown dense; bole straight, closely 

branched, sometimes buttressed. Bark grey to brown, peeling off in scroll-shaped patches 

or scales; inner bark reddish brown. Sapwood hard, orange-brown. Leaves subcoriaceous, 

elliptic to ovate-oblong, 8-15(-25) x 3-7(-11) cm, shiny above when fresh, drying dull or 

greenish brown or glaucous green, conspicuously parchment-like, distinctly pellucidpunctulate; 

base cuneate to rounded, margin entire, apex shortly acuminate; lateral veins 

5-8 pairs, ascending, curved, faint above, slightly raised beneath; intercostal veins 

invisible. Flowers in clusters or fascicles of 3-6, almost sessile or with 1-2 mm pedicels, 

arising from small woody-warts; calyx 5-10bed, lobes ovate, obtuse, ciliate; petals 5, 

greenish white, oblong, hairy inside, glabrous outside; anthers ovate; ovary semi-inferior, 

disc conical, faintly 5-lobed; style filiform, 2-4 mm. Fruits roundish to subglobose, c. 2- 

2.5 cm across, attenuate at base at maturity, apex with the persistent tiny style; pericarp 

284


thinly fleshy, pink to purple, rugose or tuberculate; endocarp thinly woody. Seeds c. 1.2 cm 

across. 

Vernacular name. Sarawak-belian landak (Iban,.Malay, Melanau). 

Distribution. SW India, Sri Lanka, Sumatra, Peninsular Malaysia, Java, and Borneo. 

Widespread in Sarawak, from Limbang, Baram, Miri, Marudi, Bintulu, Kapit, Sri Aman to 

Kuching. In Sabah, recorded from Beaufort, Beluran, Lahad Datu, Sandakan and Tenom. 

Ecology. In lowland mixed dipterocarp and secondary forest on leached sandy humult 

soils; also collected in hill forest up to 1000 m. 

Uses. The timber is hard, heavy but only moderately durable. The wood is yellowish brown, 

having rather fine and even texture, useful for house construction. 

285


OXALIDACEAE 

R.C.K. Chung 



Merrill, EB (1921) 311, PEB (1929) 111; Ridley, FMP 1 (1922) 329; Masamune, EPB (1942) 355; 

Backer & BakhuizenJ, FJ 1 (1963) 244; Veldkamp, FM 1,7 (1971) 151; Cockburn, TFM 1 (1972) 

347, TS 2 (1980) 69; Anderson, CLTS (1980) 286; Corner, WSTM 2 (1988) 605; Whitmore, Tantra 

& Sutisna, CLK 2, 1 (1990) 282. 

Trees, shrubs, climbers or herbs. Leaves alternate or spiral, compound with a terminal 

leajlet or simple, pinnately veined, stipules sometimes present, stalk usually with a basal 

joint. Inflorescences axillary, pseudoterminal or on the branches, racemes, panicles or 

cymes, 1- to many~flowered. Flowers bisexual, regular (actinomorphic), 5-merous, heterotristylous 

(with short, medium and long-styled forms), heterodistylous (with short and longstyled 

forms) or homostylous (with all styles of similar length), stalk jointed; sepals 5, free 

or fused at the base, persistent; petals 5, clawed, often faIling off early; stamens 10, 

arranged in two whorls, 5 opposite and 5 alternate with petals, connate at the base into an 

annulus, persistent, of which 5 are sometimes sterile (staminodes); anther dorsifixed, 

versatile, dehiscing extrorsely by longitudinal slits; disc absent; ovary superior, 5-locular; 

ovules anatropous, bitegmic, tenuinucelIate (with a thin nucelIus, Averrhoa) or crassinucelIate 

(with a thick nucelIus, .oxalis and Biophytum), 1 or more per locule; placentation 

axile; styles 5, of the same or different length, terminal, free, erect or recurved. Fruit a dry 

5-valved capsule or a berry. Seeds 1 to many per fruit, sometimes arillate (except for 

species of Sarcotheca and Averrhoa bilimbi); endospermjleshy; embryo straight. 

Distribution. 6 genera with about 890 species, mostly in tropical or subtropical areas. In 

Malesia, 5 genera with 29 species, of which 14 are endemic. In Sabah and Sarawak, 

represented by 5 genera with 12 species, of which only 5 are wild and 2 are cultivated trees. 

Ecology. Found in a wide variety of habitats, from lowland primary and secondary forests 

to montane forests, to about 2200 m. 

Uses. Species of Averrhoa are cultivated for its edible fruit, and that of Oxalis are 

cultivated as ornamentals. The wood of the ligneous Oxalidaceae is not utilised as timber. 

In Peninsular Malaysia, the fruits of Sarcotheca are edible. 

Taxonomy. Hutchinson (Fam. F1. PI. 1 (1959) 356, 497; Gen. F1. PI. 2 (1967) 610) 

considered the herbaceous taxa as the Oxalidaceae sensu stricto (in the Geraniales), and 

accommodated the arboreous taxa in the Lepidobotryaceae (Malpighiales) and Averrhoaceae 

(Rutales) respectively. Recent work indicates that Sarcotheca and Dapania (in the 

Lepidobotryaceae) should be associated with Averrhoa rather than with Lepidobotrys. 

287


Therefore, the Averrhoaceae would be better retained as a group in the Oxalidaceae. 

Species of Sarcotheca have sometimes been confused with that of Rourea (Connaraceae). 

However, Rourea differs from Sarcotheca in having free carpels, 2-collateral ovules, seeds 

with an aril, and a dry, indehiscent, l-celled and I-seeded fruit. 

Key to genera 

1. Herbs. ............. 2 

Climbers or trees ........ ................. \" ................. 3 

2. Leaves trifoliolate ........... 

Oxalis L. 

Sp. PI. (1753) 433, Gen. PI. ed. 5 (1754) 198; Ridley I.e. 330; Backer & Bakhuizenf I.e. 

245; Veldkamp I.e. 153. 

7 species in Malesia; 2 species (0. corniculata and 0. corymbosa) in Sabah and 


Herbs. Leaves trifoliolate; leaflets usually obcordate. Inflorescences cymose to 

pseudoumbellate, one- to many-flowered. Sepals shortly connate at base; petals 

connivent above the claw. Capsules dehiscing loculicidally, the valves remaining 

attached to the central axis. Seeds usually few, ejaculatory apparatus and testa 

smooth.· 

Leaves pinnate, with many leaflets ................................................................ . 

Biophytum DC. 

Prod. 1 (1824) 689; Edgewick & Hookerf, FI. Br.lnd. 1 (1874) 436; Ridley I.e. 330; van 

Steenis, Bull. Jard. Bot. Btzg 3,18 (1950) 449; Backer & Bakhuizenf I.e. 246; Veldkamp 

I.e. 159. 

7 species in Malesia; 1 species (B. sensitivum) in Sabah & Sarawak. 

Herbs or usually sympodially branched dwarf shrubs. Stipules bristle-like 

(setaceous). Leaves paripinnate, in tufts at the end of the stems or branches; 

leaflets opposite. Flowers terminal, usually in pedunculate, bracteate pseudoumbels, 

heterodistylous, heterotristylous or homostylous; sepals free; petals twisted, 

connivent above the claw. Capsules split loculicidally to the base, forming a 5- 

rayed star-shaped fruit without leaving a columella. Seeds 1-6 per cell, ejaculatory 

apparatus and aril white, thin. 

3. 

Climbers. Leaves unifoliolate ....................................................... . 

Dapania Korth. 

Ned. Kruidk. Arch. 3 (1854) 381; 2; Ridley I.e. 334; Veldkamp, Blumea 15 (1967) 523, I.e. 

(1971) 166. 

2 species in Malesia (Sumatra, Peninsular Malaysia, Borneo) with one species, D. 

grandifolia endemic to NE Sabah and SE Kalimantan. 

Common woody climbers in primary lowland forests. Leaves alternate, estipulate; 

stalks jointed in the middle. Inflorescences racemose, on the branches or axillary, 

solitary to fascicled. Flowers androdioecious; sepals 5, connate at the lower half, 

margins ciliate; petals 5, free, imbricate. Capsules fleshy, loculicidally dehiscent. 

Seeds hard; aril enveloping the seed. 

Trees. Leaves pinnate, trifoliolate, or unifoliolate ...................... . . ................. .4 

288

OXALIDACEAE (CHUNG) 

4. Leaves imparipinnate with 3-19 pairs of leaflets. Stalks of leaf and leaflet not jointed ... 

Averrhoa L. 

I.c. (1753) 428, I.c (1754) 196; Ridley I.c. 331; Backer & Bakhuizen/ I.c. 247; Veldkamp 

I.c. (1971) 174; Corner I.c. 605; Whitmore, Tantra & Sutisna I.c. 282. 

Two species (A. bilimbi andA. carambola) of unknown origin, currently cultivated 

throughout the humid tropics and subtropics. 

Shrubs or small trees, cultivated as fruit trees. Stipules absent. Leaves spirally 

arranged or clustered at the end of branches, imparipinnate; leaflets entire, subopposite, 

subsessile, 3 or more pairs. inflorescences panicles, axillary or borne on 

the trunk. Flowers heterodistylous or heterotristylous; sepals shortly connate at 

base; petals twisted, free or connivent above the claw, pink to dark red; stamens 

all fertile or 5 fertile and 5 sterile; ovary syncarpous with 3-7 superimposed 

pendulous ovules in each of the 5 locules. Fruit a large juicy berry. Seeds 

numerous, elliptic, flat; aril, if present, fleshy, attached to the entire adaxial placenta, 

2-lobed, and enveloping the seed; cotyledons thin, flat; germination epigeal. 

Both species prefer a climate with a dry season, but also do well in wetter climates. 

A. carambola has been cultivated successfully in frost-free subtropics up to 30 0 S in 

Australia and 32°N in Israel. For healthy growth, both species require adequate 

moisture and well-drained soils. In Malaysia, A. carambola has been planted and 

thrives well in well-drained peat soils. Vernacular name: belimbing (Malay). 

Leaves unifoliolate or trifoliolate. Stalks of leaf and leaflet jointed ............... Sarcotheca 

SARCOTHECA Blume 

(Greek, sarcos = fleshy; theca = container; the fleshy fruit) 

tabarus (Dusun, Sabah), piang (Iban, Sarawak) 

Mus. Bot. Lugd. Bat. 1 (1850) 241; Ridley I.c. 332; Merrilll.c. (1929) Ill; Veldkamp I.c. (1967) 

527, I.c. (1971) 168; Cockburn I.c. (1972) 348, I.c. (1980) 70; Anderson I.c. 286; Whitmore, Tantra 

& Sutisna I.c. 282; Ng, Mal. For. Rec. 34 (1992) 470. Synonyms: Roucheria Miq. I.c. (1859) 136; 

Connaropsis Planch. ex Hook./, Trans. Linn. Soc. 23 (1860) 166. 

Trees or shrubs; bole deeply fluted. Bark reddish or reddish brown, scaly or dippled; inner 

bark reddish. Sapwood pale. Stipules absent. Leaves alternate, unifoliolate or trifoliolate; 

margin entire; leaf-stalks jointed, proximal parts and leaflet stalks swollen and wrinkled. 

Inflorescences axillary or terminal panicles, one to several together. Flowers heterodistylous, 

arranged in more or less stalked cymes, scattered along a simple or sparsely 

branched rachis; cymes subtended by small caducous bracts; sepals unequal, persistent 

(except in S. diversifolia), imbricate, shortly connate at base, abaxial surface strigose with 

appressed hairs; petals twisted, imbricate, free at base, usually fused above and falling off 

as a single unit, adaxial surface with minute papillae in the upper half; stamens all fertile; 

ovary glabrous to hairy, ovules 2 in each locule. Fruits fleshy, red at least when dry, 

usually 5-ridged. Seeds flat, without aril, testa reddish, smooth to wrinkled, hard; cotyledons 

thin, flat; germination epigeal. 

289


Distribution. 11 species in W Malesia (Sumatra, Peninsular Malaysia, Borneo, Celebes). In 

Sabah and Sarawak, 5 species are recorded. 

Ecology. Primary and secondary forest on poor soils at low altitudes. 

Uses. Species of Sarcotheca produce a light, easily worked timber, which is sometimes 

used for roofing and interior work, but it lacks both strength and durability. The fruit, 

although sour, is eaten in curry dishes, and cooked with vegetables and sweets. It is said to 

be a remedy for cough. 

Key to Sarcotheca species 

1. Leaves trifoliolate; terminal leaflet larger than lateral leaflets. Mature fruits greenish 

yellow when fresh, red when dry ......................................................... 1. S. diversifolia 

Leaves unifoliolate. Mature fruits red when fresh ......................................................... 2 

2. Young twigs and abaxial surface of leaf-blades densely brownish velvety hairy; leaflet 

margin curled inwards ............................................................................ 4. S. ochracea 

Young twigs and abaxial surface of leaf-blades glabrous to sparsely hairy; leaflet 

margin not curled inwards ........................................................................................... 3 

3. Young twigs 4-angled. Leaves oblong to oblanceolate, 14-27 x 4-8 cm; lateral and 

intercostal veins distinctly prominent on the lower surface ................ 3. S. macrophylla 

Young twigs terete (rounded). Leaves elliptic, oblong or lanceolate, 4.5-15 x 2-6 cm; 

lateral and intercostal veins prominent to inconspicuous on the lower surface .............. 4 

4. Leaf blades glaucous below, elliptic or oblong; veins not drying reddish; stalks long 

(stalkblade ratio 1 :4.5). Panicles short and compact. Sepals dark red to crimson .......... . 

. . . . . 2. S. glauca 

Leafblades not glaucous below, oblong to lanceolate; veins drying reddish; stalks short 

(stalkblade ratio 1:10). Panicles long and lax. Sepals rusty ................. 5. S. rubrinervis 

1. Sarcotheca diversifolia (Miq.) Hallier f 

(Latin, diversifolius = with leaves of different shapes) 

Med. Rijksherb. Leiden 1 (1911) 2; Ve1dkamp I.c. (1967) 529, I.c. (1971) 170; Cockburn l.c. (1980) 

71; Whitmore, Tantra & Sutisna I.c. 282. Basionym: Rourea diversifolia Miq., Fl. Ind. Bat. 1 Suppl. 

(1860) 528. Type: Teijsmann HE 707 (= ? 706), West Sumatra, Sibolga, Morsala Island (holotype L; 

isotypes BO, CAL, K). Synonyms: Connaropsis diversifolia (Miq.) Kurz, J. As. Soc. Beng. 39, 2 

(1870) 69, excl. syn. C. griffithii; Santa lodes diversifolium (Miq.) o. Kuntze, Rev. Gen. PI. 1(1891) 

155; Connaropsis acuminata Pears., Kew Bull. (1906) 2; S. acuminata (Pears.) Hallier f, Beih. Bot. 

Centralbl. 34,2 (1917) 27; S. subtriplinervis Hallierf I.c. (1917) 27; Connaropsis grandiflora Ridl., 

KewBull. (1930) 75. 

Small to medium-sized tree to 30 m tall, 50 cm diameter, sometimes with equal buttresses 

to 2 m tall. Bark smooth, pock-marked or occasionally scaly, reddish brown; inner bark 

290

generally light red or pink. Sapwood white, pale yellow or sometimes pinkish. Twigs 

rounded. Leaves tri/oliolate, stalks 5-17 mm long, 1-2.5 mm thick, rachis 8-31 mm long, 

0.75-2 mm thick; leaflets ovate to elliptic or lanceolate, lateral leaflets 2.5-10.5 x 1-4 

cm, sometime falling off early but leaving a scdr, terminal leaflet often twice the size of the 

other leaflets, papery to thinly leathery, glabrous; base acute to broadly rounded, apex acute 

to caudate; lateral veins 2-5 pairs, prominent on both surfaces; intercostal veins net-like; 

leaflet stalk 3.5-8 mm long, 0.75-2 mm thick. Inflorescences 1-4 panicles together, 

shorter than the subtending leaf, loosely branched; 1-9 cm long, rusty hairy, becoming 

glabrous at mature state. Flower stalk with proximal part 2.5-5 mm long, distal part 1-3 

mm long; sepals purplish, ovate to oblong, 2.5-5 x 1.5-3 mm, apex acute to emarginate, 

291 

Small to medium-sized tree to 21 m tall, 30 cm diameter; no buttresses. Bark smooth to 

slightly fissured, occasionally scaly, often warty; inner bark yellow-brown, sometimes with 

a pink tinge. Sapwood usually white or pale yellow, rarely pink. Twigs rounded. Leaves 

unifoliolate, stalk 7-30 mm long, 0.5-1 mm thick, leaflet stalk 3-6 mm long, 1-1.5 mm 

thick; blades elliptic to oblong, 4.5-11 x 1.5-5 cm, slightly larger on sterile twigs, papery 

to thinly leathery, glabrous, glaucous, dull, grey-green on the abaxial surface, sometimes 

shiny on the adaxial surface; base broadly cuneate to rounded or subcordate, apex acute, 

acuminate or caudate; lateral veins 5-7 pairs, more or less looping and joining near the 

margin; intercostal veins net-like. Inflorescences 1-2 panicles together, erect, compact, 

usually many-flowered, 2-13 cm long, covered with minute rusty scale-like indumentum. 

I.e. (1911) 2; Merrilll.e. (1921) 312; Masamune I.e. 355; Veldkamp I.e. (1967) 535, I.e. (1971) 172; 

Cockbum I.e. (1980) 71; Anderson I.e. 286; Whitmore, Tantra & Sutisna I.e. 282. Basionym: 

Connaropsis glauea Hook.f I.e. (1860) 166. Type: Lobb 1857, "Borneo" (holotype K). 

2. Sarcotheca glauca (Hook.f) HaIIier f 

(Greek, glaukos = bluish grey; the lower leaflet surface) 

Fig. 1. 

Vernacular names. Sabah--ibajantan (Suluk), tabarus (general). Sarawak-buah piang (lban 

and Malay), jiwang (Punan). Brunei:-kerapa-kerapa, perapan macas, tabaus, tebarus (Tutong 

Dusun). General-belimbing bulat (Malay). 

Distribution. Sumatra and Borneo. Common in the coastal area of Sabah and throughout 


Ecology. Primary mixed dipterocarp forest and secondary forest from lowlands to 900 m, 

on leached yellow, especially humult sandy ultisols, occasionally in mixed swamp forest but 

very rare in the Rejang delta. Flowering in February-November, fruiting in March-June 

and September-December. 

ab axially puberulous to glabrous, mostly falling off, seldom persistent in fruit; petals pink 

to red, obovate-oblong to obovate-lanceolate, 8-10 x 2-3.5 mm, apex rounded to 

emarginate, 1-2 mm long clawed; filaments in short-styled form 2.5-3.5 mm and 3.5-4.5 

mm, in long-styled form 1.5-2.5 mm and 2.5-3 mm; ovary ellipsoid, puberulous to 

glabrous, styles in short-styled form 0.5-1 mm long, in long-styled form 2.5-4 mm long. 

Fruits green when young, greenish yellow when mature, ellipsoid, glabrous, 12-20 x 9-15 

mm. Seeds none to 7 per fruit, 6-7.5 x 4.5-5 mm, testa smooth. 

OXALIDACEAE (CHUNG)

unifoliolate, stalk 5-12 mm long, 1.5-3 mm thick, leaflet stalk 3-9 mm long, 1-3 mm 

thick; blades oblong to oblanceolate, 15-26 x 3-9 cm, thinly leathery, adaxial surface 

glabrous, abaxial surface puberulous on the veins; base rounded to truncate or subcordate, 

apex acuminate to cuspidate; midrib puberulous, sunken above; lateral veins 6-l3 pairs, 

distinctly prominent on the abaxial surface; intercostal veins net-like, distinctly prominent. 

Inflorescences 1-4 panicles together, slender, lax, pendulous; 12-85 cm long, brownpuberulous. 

Flower stalk with proximal part to 2 mm long, distal part to 1 mm long; sepals 

obovate to ovate, 1.5-2.5 x 1-1.5 mm, apex rounded, abaxially rusty hairy, puberulous in 

fruit; petals dark red, obovate-oblong to oblanceolate, 3-5 x 1-2 mm, apex obtuse to 

emarginate, to 0.5-mm-long-clawed; filaments in short-styled form 1-2 mm and 2-3 mm 

long, in long-styled form 1-1.5 mm and 1.5-2.0 mm long; ovary subglobose to ellipsoid, 

densely hairy, styles in short-styled form 0.25-0.75 mm long, in long-styled form 1-2 mm 

long. Fruits dark red, subglobose, 6-11 x 5-8 mm, glabrescent, glossy. Seeds to 9 x 4 mm, 

testa wrinkled. 

Vernacular names. Sarawak-belimbing manik (Bakumpai.-Dayak),piang (!ban). 

292 

174; Whitmore, Tantra & Sutisna I.e. 282. Type: Maller, s.n., "Borneo" (lectotype L; isolectotype 

BO). Synonym: Roueheria maerophylla Miq. I.e. (1859) l36. 

Small tree to 15 m tall, 10 cm diameter; no buttresses. Bark smooth, pale brown; inner 

bark reddish. Sapwood white. Twigs 4-angled when young and rounded when old . .Leaves 

I.e. (1850) 242; MerrillI.e. (1921) 312; Masamune I.e. 355; Veldkamp I.e. (1967) 541, I.e. (1971) 

Ecology. Primary and secondary lowland forests, often on hill slopes. In Sarawak, 

uncommon in mixed swamp forest, occasional or widespread in kerangas forest and mixed 

dipterocarp forest on sandy humult ultisols. Flowering in February-April and September 

December, fruiting in October-May. 

(around Papar). Sarawak--aremajuh (Dayak), medang, piang (!ban), tulang payung piang 

(Malay). Brunei--asam piai (Dusun), kandis daham (Kedayan). 

Flower stalk with proximal part 2-5.5 mm long, distal part 1.5-2 mm long; sepals dark 

red to crimson, ovate to oblong, 1. 5-3 x 0.5-1. 5 mm, apex rounded, minutely hairy on the 

abaxial surface, glabrescent in fruit; petals red, darker.at apex, oblong to lanceolate, 3.5-6 

x 1-2 mm, apex rounded to slightly emarginate, 0.5-1.5-mm-Iong-clawed; filaments in 

short-styled form 2-2.5 mm and 2.5-3.5 mm, in. long-styled form 1-1.5 mm and 1.5-2 

mm long; ovary ellipsoid, rusty hairy, styles in short-styled form 0.5-1 mm long, in longstyled 

form 2.5-3 mm long. Fruits bright pink to dark red, turning black when old, 

ellipsoid to nearly round, 7.5-12.5 x 5-11 mm. Seeds 1-9 per fruit, 6-7 x 3.5-4 mm, testa 

smooth. 

kandis daham (around Mempakul), rangkas-rangkas (Dusun Kinabatangan), t-izos-izos 

Distribution. NW Borneo (Sabah, Sarawak and Brunei). Common in Sabah and Sarawak. 

No record from Kalimantan. 

Vernacular names. Sabah--asam daham (around Sandakan), gitan gizu (Dusun Rungus), 


3. Sarcotheca macrophylla Blume 

(Greek, makros = large, phullon = leaf)


F 

5mm 

3cm 

, 

4cm 

!cm 

H 

G 

r 

~ 

K~ 

J 

L 

2mm 

!cm 

Fig. 1. Sarcotheca glauca. A, flowering leafy twig; B, upper portion of twig; C, flower; D, ovary of 

short-styled flower; E, short-styled flower with sepals and petals removed; F, longitudinal-section of 

short-styled flower, G, long-styled flower with sepals and petals removed; H, ovary of long-styled 

flower; I, infructescence; J, dehiscing fruit; K, basal view of mature fruit; L, cross-section of fruit. CA 

& B from SAN A 1195, C from SAN 72350, D-F from SAN 33562, G-H from SAN 33612, I-L from 

ChewCWL 1423.) 

293


Distribution. Borneo (Sarawak and Kalimantan). Only one collection, Beccari PB 3166 

reported from Marop, Sarawak. 

Ecology. Primary and secondary forest on sandy soils at low altitude. Flowering and 

fruiting in January-May. 

4. Sarcotheca ochracea Hallier j 

(Greek, ochre = yellow to yellowish brown; the dry leaves) 

l.c. (1917) 28; Merrilll.c. (1921) 312; Masamune l.c. 356; Veldkamp I.c. (1967) 541, I.c. (1971) 

173; Anderson l.c. 286; Whitmore, Tantra & Sutisna I.c. 282. Type: Haviland 2343, "Borneo" 

(holotype L; isotypes K, SING). 

Small tree to 10 m tall, 10 cm diameter; no buttresses. Bark brownish; inner bark pale 

brown. Twigs rounded. Leaves unifoliolate, stalk 8-24 mm long, 1.5-2 mm thick, leaflet 

stalk 3-6 mm long, 1.5-3 mm thick; blades elliptic to oblong, 7.5-19.5 x 4-10 cm, 

adaxial surface glabrous, abaxial surface rusty-pubescent to velvety; base obtuse to rounded, 

margin slightly wavy and curved towards the abaxial surface, apex acuminate; lateral veins 

6-9 pairs, abaxial surface prominent; intercostal veins net-like, prominent. Inflorescences 

1-2 panicles together, erect to pendulous, compact to elongate, branched i.n comb-like way, 

many-flowered, 7-64 cm long, rusty-velvety. Flower stalk with proximal part 4-8 mm 

long, distal part 0.5-1 mm long; sepals reddish brown, obovate, 2.5-3 x 2-3 mm, apex 

rounded to retuse, rusty-puberulous on the abaxial surface, puberulous in fruit; petals 

scarlet, darker at apex, ob ovate to lanceolate, 6-8 x l.5-2 mm, apex obtuse to rounded, 

1.5-2.0-mm-long-clawed; filaments in short-styled form c. 2.5 mm and 3.5 mm long, in 

long-styled form 1.5-2 mm and 2-2.5 mm long; ovary subglobose, rusty hairy, styles in 

short-styled form 0.5-1 mm long, in long-styled form 2-2.5 mm long. Fruits bright red, 

subglobose to oblong with rounded apex, 8-15 x 6-15 mm. Seeds 1-2 per fruit, 5-7 x 3- 

4.5 mm, testa wrinkled. 

Vernacular names. Sarawak-ikor mata (lban), pechi mata (general). 

Distribution. Borneo. In Sarawak uncommon (Miri, Bintulu and Tatau). 

Ecology. In mixed dipterocarp forest on pale yellow infertile clay soils. Flowering and 

fruiting in April-October. 

5. Sarcotheca rubrinervis Hallierj 

(Latin, ruber = red, nervus = nerve) 

I.c. (1917) 29; Merrilll.c. (1921) 312; Masamune I.c. 356; Veldkamp I.c. (1967) 539, I.e. (1971) 

173; Cockburn l.c. (1980) 73; Whitmore, Tantra & Sutisna I.c. 283. Type: Amdjah 1082, E Borneo 

(holotype L; isotype BO). Synonym: S. oblongifolia Merr. I.c. (1929) Ill. 

Small to medium-sized tree to 20 m tall, 30 cm diameter; no buttresses. Bark smooth, thin, 

reddish to reddish brown; inner bark yellow or yellowish brown, sometimes with a pink 

294


tinge, fibrous. Sapwood white or pale yellow. Twigs rounded. Leaves unifoliolate, stalk 6- 

20 mm long, 1.5-2 mm thick, leaflet stalk 2-4 mm long, 1.5-2 mm thick; blades oblong to 

lanceolate, 6.5-15 x 2-6 cm, glabrous, abaxial surface not or sometimes slightly glaucous; 

base obtuse to rounded, apex acute to acuminate; lateral veins 4-11 pairs, often with 

reddish tinge, more or less looping and joining near the margin; intercostal veins irregular, 

the more prominent ones running from vein to vein or from vein to midrib, fainter ones 

net-like. Inflorescences 1-2 panicles together, elongate, lax, pendulous, 6-36 cm long, 

rusty puberulous. Flower stalk with proximal part 3-5 mm long, distal part up to 1 mm 

long; sepals ovate to lanceolate, 1.5-2.5 x 1-1.5 mm, apex acute to emarginate, minutely 

rusty hairy at base, puberulous in fruit; petals red or rarely white, lanceolate, twice as long 

as sepals, apex rounded to emarginate, 0.5-1-mm-Iong-clawed; filaments in short-styled 

form 1.5-2 mm and 2.5-3 mm long, in long-styled form 0.5-1 mm and 1-1.5 mm long; 

ovary subglobose, densely hairy, styles in short-styled form c. 0.5 mm long, in long-styled 

form 1.5-2 mm long. Fruits pink to bright red, subglobose, 6-10 x 5-10 mm, glaucous to 

glossy. Seeds to 6.5 x 5 mm, testa smooth to more or less wrinkled. 

Vernacular names. Sabah--asem-asem (Dusun Kinabatangan), iba talon (Bajau), ira 

prumpuan (Suluk), lampyos (Dusun Penampang), pinggoh, pinguh (Kalabakan), tabarus 

(general). 

Distribution. Borneo (Sabah and Kalimantan). In Sabah only known from Tawau. 

Ecology. Primary and secondary forests near rivers on loamy soils of flat to undulating 

lands. Flowering and fruiting in March-December. 

295


PITTOSPORACEAE 

John B. Sugau 




Merrill, EB (1921) 287; Masamune, EPB (1942) 324; Bakker & van Steenis, FM 1, 5 (1957) 345; 

Whitmore, TFM 2 (1973) 309; Anderson, CLTS (1980) 286; Ashton, :MNDTS 2 (1988) 325; Corner, 

WSTM 2 (1988) 607; Sugau, Sandakania 4 (1994) 41. 

Small to medium-sized trees, shrubs, or climbers; indumentum if present consisting of 

simple hairs; bark resinous. Leaves spirally arranged, often crowded towards the end of 

the twigs, simple, mostly entire, pinnately veined; stipules absent. Inflorescence a few- to 

many-flowered, axillary or terminal, bracteate corymbose panicle, raceme or thyrse, sometimes 

on bare branches, often also as fascicles of flowers. Flowers regular, mostly 5-merous, 

bisexual or functionally unisexual; bracteoles often 2; sepals 5, free or connate at the base, 

overlapping; petals 5, often narrowed at the base and loosely joined as a tube, overlapping, 

caducous; stamens 5, inserted on the sepals, erect, free, or slightly connivent at the base, 

anthers 2-celled, introrse, basifixed, dehiscing lengthwise (or by pores); ovary superior, 

sometimes shortly stalked, mostly I-chambered, sometimes completely or incompletely 2- 

chambered, placentas. 2-6, basal or on the wall, style simple, stigma simple and thickened, 

or lobed, ovules numerous, anatropous with one integument. Fruit a berry or a loculicidal, 

2- or 6-valved capsule. Seedsfew to many per fruit, embedded in a resinous or oily pulp or 

flUid, seldom winged, testa thin, smooth; endosperm copious, hard; embryo small, close to 

the hilum; cotyledons small, 2-5. 

Distribution. 9 genera restricted to the Old World, chiefly Australian; 6 genera are completely 

restricted to Australia; and 2 in E Malesia and Australia; 1 (Pittosporum) is widely 

distributed and the sole representative of the family in Sabah and Sarawak. 

Ecology. Members of the family can be found in everwet or dry regions, from tropical to 

warm-temperate areas. 

Uses. Several genera have species that are grown as ornamentals; it has been suggested 

(Bakker & van Steenis I.c.) that a few Pittosporum species might be used for re-afforestation of 

bare lands, on account of their adaptability to such sites. 

PITTOSPORUM Banks ex Sol. 

(Greek, pitta = resin, spora = seed; the resinous seed) 

in Gaertner, Fruct. 1 (1788) 286; Merrilll.e. 287; Masamune I.e. 324; Bakker & van Steenis I.e. 345; 

Whitmore I.e. 309; Anderson I.e. 286; Ashton I.e. 325; Corner I.e. 607; Sugau I.e. 41. 

297

TREE FLORA OF SABAI-I AND SARAW AK VOL. 1 (1995) 

Trees or shrubs, with rhythmic growth; terminal bud protected by scale leaves. Leaves 

entire, often hairy when young. Flowers few to many, in fascicles, thyrses or cO/ymbs, 

seldom solitary, white or pale yellow, fragrant; sepals free or more or less connate; petals 

generally ligulate, free or united in the lower part, the free segments spreading or recurved; 

stamen filaments slender in male flowers, somewhat shorter and slightly broadened at the 

base in female flowers, anthers generally oblong in male flowers, sagittate and smaller in 

female flowers; ovary sessile or stalked, glabrous or hairy, I-chambered, slender in male 

flowers but stout in female flowers; style in female flowers short and with a 2-5-lobed 

capitate stigma, in male flowers longer and with obscure stigmas; placentas 2-5, basal or 

on the ovary wall, ovules few to many. Fruit a loculicidally dehiscent, 2(-6)-valved capsule, 

usually ripening orange. Seeds I to many per fruit, covered by a resinous viscous fluid; 

germination epigeal. 

Distribution. About 100 species, from warm-temperate to tropical areas, from Africa, 

South and East Asia and Australasia, to the Pacific Islands. 6 species are documented for 

Borneo, all found in Sabah, while only 2 occur in Sarawak. 

Ecology. Species of Pittosporum are generally understorey shrubs or trees of the rain forest, 

occurring from near sea-level to the subalpine zone on Mt. Kinabalu. 

Key to Pittosporum species 

(based mainly on leaf characters) 

1. Leaves very narrowly elliptic to linear, to 1.1 cm wide ..................... 2. P. Iinearifolium 

Leaves elliptic, oblanceolate or obovate, typically more than 1.5 cm wide .................... 2 

2. Leaves persistently thickly rusty hairy on the lower side, and conspicuously bullate on 

the upper side; margins markedly recurved when dry ............................. 6. P. silamense 

Leaves glabrous or, if hairy, only when very young, and not bullate or only finely so on 

the upper side; margins not or only weakly recurved when dry ..................................... 3 

3. Lateral veins finely impressed above. Sepals 6-7 mm 10ng ............... 3. P. longisepalum 

Lateral veins flat above. Sepals less than 5 mm 10ng .................................................... 4 

4. Leaves markedly obovate, the apex abruptly acuminate to cuspidate. Inflorescence 

borne on bare branches ................................................................................................ 5 

Leaves elliptic, or if oblanceolate or obovate, then the apex gradually acuminate. 

Inflorescence borne in the leafy portions ofbranches ......................... l. P. ferrugineum 

5. Leaf-stalks 8-18 mm long. Petals 9-12 mm long. Infructescence-stalks not conpicuous 

or to 0.5 cm long. Mature fruits to 1.7-2 cm long ............................... 5. P. resiniferum 

Leaf-stalks, 15-25 mm long. Petals only 4-7 mm long. Infructescence-stalks 1-3 cm 

long. Mature fruits 1-1.5 cm long ....................................................... 4. P. ramiflorum 

298

PITTOSPORACEAE (SUGAU) 

Key to Pittosporum species 

(based on flower and fruit characters) 

l. Inflorescences mainly on bare branches .................................................................... 2 

Inflorescences among leaves on the branches ............................................................... 3 

2. Petals 4-7 mm long. Infructescence-stalks 1-3 cm long. Fruit 1-l.5 cm long ............... . 

........................................................................................................... 4. P. ramiflorum 

Petals 9-12 mm long. Infructescence-stalks inconspicuous or to 0.5 cm long. Fruits 

1.7-2 cm long ................................................................................... 5. P. resiniferum 

3. Seeds attached to the fruit-wall to halfway to the fruit-apex ............... 1. P. ferrugineum 

Seeds attached to the basal portion of the fruit-wall ..................................................... .4 

4. Fruits, at least when young (and ovary), glabrous, or at most slightly hairy at the base 

........................................................................................................ 2. P. Iinearifolium 

Fruits, at least when young (and ovary), hairy all over.. ............................................... 5 

5. Infructescence-stalks 0.8-l.2 cm long. Seeds to 11 per fruiL ................ 6. P. silamense 

Infructescence-stalks 2-3 cm long. Seeds 1-4 per fruiL .................. 3. P. longisepalum 

1. Pittosporum ferrugineum Aiton 

(Latin,ferrugineus = rusty; the brown leaf tomentum) 

Hort. Kew ed. L., 2 (1811) 27; MerrillI.c. 287; Masamune I.c. 324; Bakker & van Steenis I.c. 345; 

Whitmore I.c. 309; Anderson l.c. 286; Ashton I.c. 322; Corner I.c. 607. Type: Hort: Kew (from a 

cultivated plant in the Royal Botanic Gardens Kew; native of New Guinea and introduced before 

1787 by the Right Hon. the Earl of Tankerville), fide Aiton (BM). Synonyms: Itea javanica Blume, 

Bijdr. (1926) 863; Pseuditea javanica (Blume) Hassk., Flora 25, 2 (1842) Beibl. 30; Pittosporum 

javanicum (Blume) Blume, Mus. Bot. Lugd. Bat. 1 (1850) 159; Pittosporum rufescens Turcz., Bull. 

Soc. (Imp.) Nat. Mosc. 27, 2 (1854) 367; Pittosporum nativitatis Bakker in Andrews, Monogr. 

Christm. Isl. (1960) 171; Pittosporum versteeghii Merr. & Perry,J. Am. Arb. 21 (1940) 177. 

Small to medium-sized tree, to 15 m tall, occasionally reaching 20 m tall, 25 cm diameter. 

Bark brownish; inner bark yellowish. Sapwood pale yellow, soft. Young shoot more or less 

densely covered with caducous rusty brown hairs. Leaves typically elliptic, rarely 

oblanceolate or obovate, 6-12 x 2-4 cm, papery, pale brown on both surfaces; base cuneate, 

margin plane and slightly undulate, apex narrowly acute to gradually acuminate; midrib 

sharp on the lower side but depressed on the upper side; lateral veins 5-9 pairs, flat above; 

intercostal veins reticulate, fine; leaf-stalks 8~ 15 mm long, slender and grooved on the 

upper side; Inflorescence a many-flowered thyrse, l.5-5 cm long, borne among leaves on 

the branches, or terminal; peduncle 1-5 cm long, rusty-hairy. Flowers pale yellow, sepals 

lanceolate to linear, 2-4 mm long, generally rusty hairy; petals ligulate, 6-8 x 1 mm; 

stamens 3-4 mm long, filaments 2.5-3 mm long, anthers 0.5-1 mm long; ovary ellipsoid 

or cylindric, sessile, densely rusty hairy, style 1-l.5 mm long, glabrous, stigma often 

scarcely capitate in male flowers, 2-10bed in female flowers. Fruits capsular, 1-16 in an 

infructescence, often broader than long or subglobose, 0.5-1 cm long, 2-valved, generally 

notched and mucronate apically, narrowed towards the often hairy base, rugose, ripening 

299


on the lower side, not bullate on the upper side; midrib impressed on the upper side, raised 

on the lower side; lateral veins 5-7 pairs, finely impressed on the upper side, raised on the 

lower side; intercostal veins reticulate, very fine; stalk c. 5 mm long, slender. Inflorescences 3- 

flowered cymes, terminal on leafy shoots; peduncle 2-3 cm long, hairy. Flowers with 

sepals free, linear-lanceolate, 6-7 x 1-1.5 mm, hairy, acute, longer than the petals in bud; 

petals narrowly oblanceolate, c. 8 x 1-2 mm, acute; stamens 4-5 cm long, filaments c. 3 

mm long, anthers c. 2 mm long, apiculate; ovary ellipsoid, subsessile, densely hairy to 1 

mm long from the base, stipe with 5 furrows, c. 2 x 1-1.5 mm. Fruits subglobose, c. 0.7 

cm across, 2-valved, mucronate, rugose; infructescence-stalks 2-3 cm long, hairy. Seeds 1- 

4 per fruit, c. 4 x 3 mm, attached on a placenta at the base of the fruit. 

Distribution. Sabah (Mt. Kinabalu) and Central Celebes. 

Ecology. Mainly in submontane rain forest, at 1200-1500 m. 

4. Pittosporum ramiflorum (Zoll. & Moritzi)Zoll. ex Miq. 

(Latin, rami = branches,florum = flowers; the inflorescences on the bare branches) 

Fl. Ind. Bat. 1,2 (1858) 122; Bakker & van Steenis I.e. 345. Basionym: Glyaspermum ramiflorum 

Zoll. & Moritzi, Nat. Geneesk. Areh. Ned. Ind. 2 (1845) 11. Type: Zollinger 2139, Java, Gede (P). 

Synonyms: Pitttosporum ramiflorum var. parviflorum K. Sehum. in K. Sehumann & Hollrung, Fl. 

Kais. Wilh. Land (1889) 70; P. comptum K. Sehum. & Lauterb., Fl. Sehutzgeb. (1900) 338; P. 

clementis MeIT., Philip. J. Se. 3 (1908) Bot. 137. 

Shrub or tree, (3-)5-20(-30)m tall. Leaves spirally arranged or in distinct clusters along 

the twigs, obovate, 5-18 x 2.5-6 cm, plane, subcoriaceous, glabrous when mature; base 

broadly cuneate, margin entire, apex abruptly acuminate, acumen for 0.5-1. 5 cm long; 

midrib flattened on the upper side but raised on the lower side; lateral veins 5-8 pairs, 

flattened on the upper side but somewhat raised on the lower side; intercostal veins 

reticulate, fine; stalks 15-25 mm long, slender and grooved on the upper side. Inflorescences 

densely branched and many-flowered, mainly on the bare parts of branches; peduncles 1-3 

cm long, hairy; bracts ovate, c. 1-l.5 mm, caducous; pedicels to 6 mm. Flowers white, 

sepals united at their base in a shallow cup, 1.5-2 mm high, otherwise free, with rounded 

apices; petals narrowly elliptic, (4-)5-6(-7) x (1-)2(-3) mm, rounded; stamens 4-5 mm 

long, filaments 3-5 mm long, anthers oblong, 0.7-2 mm long; ovary ellipsoid or obovoid, 

2-3 x 1-1.5 mm, subsessile, brownish hairy, style 1-3 mm long, stigma 2-lobed. Fruits 

varying in number, on woody stalks, ellipsoid to obovoid, compressed, 1-1.5 cm long, 2- 

valved, with an abrupt tip, valves hard, 1-2 mm thick, wrinkled; infructescence-stalks 1-3 

cm long, hairy. Seeds many per fruit, attached to the fruit-wall from the base up to apex of 

the fruit, very irregular in shape to roundish, flat, 2-3 mm diameter. 

Distribution. Java, Borneo, Southern Philippines (Negros, Cebu, Mindano and Bohol), 

Celebes, the Moluccas (Ambon, Buru, Ceram), New Guinea, Solomon Islands. In Sabah 

only found on Mt. Kinabalu; not yet recorded from Sarawak. 

Ecology. An understorey tree in rain forest, sometimes also riverine, occasionally to 3200 

m in montane forest. 

302

PITTOSPORACEAE (SUGAU) 

Kew Bull. (1894) 344; Bakker & van Steenis I. c. 345. Type: Holmes, s.n., Philippines (leetotype K, 

here designated). Synonyms: P. epiphyticum Merr. l.c. (1908) 138; P. accuminatissimum Merr., 

Philip. J. Se. 14 (1919) 402. 

Small to medium-sized tree, 2-10 m tall, 10-20 cm diameter. Bark thin, whitish to dark 

brown; inner bark creamy pink, exudate sticky. Sapwood pale white. Leaves spirally 

arranged, markedly obovate, 9-18 x 2.5-5 cm, glabrous; base narrowly cuneate, margin 

sligtly re curved when dry, rather thick, apex abruptly cuspidate; midrib slender, prominent 

on the lower side, flattened on the upper side, pale brown; lateral veins 7-10 pairs, 

flattened on upper side, raised on the lower side; intercostal veins reticulate, fine; stalks 8- 

303 

Small tree, bole c. 5 cm diameter. Bark smooth, brownish; inner bark yellowish. Sapwood 

yellowish. Leaves generally elliptic, 5.5-10 x 2.5-4 cm, conspicuously bullate on the 

upper side, persistently thickly rusty-hairy on the lower Side; base cuneate, margins 

markedly recurved when dry, rather thick, apex narrowly cuneate to acuminate; midrib 

sunken on the upper side, raised on the lower side; lateral veins 6-9 pairs, deeply 

impresiSed on the upper side, raised on the lower side; stalks 8-18 mm long, slender, 

grooved on the upper side. Inflorescence a many-flowered thyrse. Flowers unknown. 

Sandak:ania 4 (1994) 41. Type: Joseph et al. SAN 120894, Sabah, Lahad Datu, Mt. Silam (holotype 

SAN). 

6. Pittosporum silamense 1. B. Sugau 

(ofML Silam, Sabah) 

Ecology. In montane forest, sometimes also at lower localities. It seems to be a lightdemanding 

species. 

Distri bution. Sabah (Mt. Kinabalu) and the Philippines (Luzon, Mindoro, Panay, Catanduanes, 

Leyte, and Mindanao). 

Vernacular name. Sabah-bedung (Dusun Ranau). 

18 mm long, slender, grooved on the upper side. Inflorescence a false umbel, 1-1.5 cm 

long, borne on bare branches below the leaves; peduncles not conspicuous or only to about 

0.5 cm long, slightly hairy. Flowers whitish; sepals united at their lower half into a shallow 

cup, 1-2 mm high, apices rounded, 1-2 mm long; petals narrowly oblong, 9-12 x 1.5-2 

mm; stamens 7-9 mm long, filaments slender, 6-7 mm long, anthers l.5-2 mm long; 

ovary ellipsoid, 4-5 x 1.5-2 mm, sessile, densely hairy, style 2-2.5 mm long, stigma 

thickened. Fruits 1-6 in an infructescence, ripening orange, capsule globose to ellipsoid, 

1.7-2 cm long, very hard when dry, 2-valved, notched and with an abrupt tip, rounded to 

cordate at the base, glabrous, rugose; infructescence-stalks inconspicuous or to about 0.5 

cm long, hairy to glabrous. Seeds many per fruit, attached to the fruit-wall from the base to 

the apex of the fruit. 

5. Pittosporum resiniferum Hems!. 

(Latin, resinifer'= producing resin; the capsule)

304 

sub globose, 0.8-1.1 cm long, 2-valved, notched and with an abrupt tip, base narrowed to 

about 2 mm long, covered with rusty hair; infructescence-stalk 0.8-1.2 cm long, hairy. 

Seeds to 11 per fruit, attached to a placenta at the base offruit, black. 

Fruits 10-18 in an infructescence, greenish to blue when unripe, black when ripe, capsule 

Distribution. Endemic to Sabah. Apparently restricted to Mt. Silam in Lahad Datu district. 

Ecology. Hill forest, about 800 m, on ultramafic soil. 

TREE FLORA OF SABAI-I AND SARAWAK VOL. 1 (1995)


RHAMNACEAE 

Carsten Schirarend 

Botanischer Garten und Botanisches Museum Berlin-Dahlem, 

Berlin, Germany 

Ridley, FMP 1 (1922) 461; Masamune, EPB (1942) 434; Suessenguth in Engler & Prantl, Pfl. Fam. 

20 d (1953) 7; Backer & BakhuizenJ, FJ 2 (1965) 80; Anderson, CLTS (1980) 289; Keng, OFMSP 

(1983) 203; Corner, WSTM 2 (1988) 609; Latiff, TFM 4 (1989) 297; Whitmore, Tantra & Sutisna, 

CLK 2, 1 (1990) 293. 

Deciduous or evergreen, often thorny trees, shrubs, woody climbers or very rarely herbs. 

Leaves simple, alternate or rarely opposite, pinnately veined or 3-veined. Stipules generally 

present, small, deciduous or often transformed into straight to recurved spines. Inflorescences 

basically cymose, cymes mostly axillary, sessile or peduncled, or reduced to manyto 

few-flowered fascicles. Flowers radially symmetrical, 4- or 5-merous, rarely 3- or 6- or 

7-merous, bisexual or unisexual (plants monoecious and/or dioecious); hypanthium 

resembling a calyx-tube, patelliform or hemispherical to tubular; sepal 4 or 5, valvate in 

bud, triangular, erect to more or less recurved during anthesis, often medially keeled within; 

petals 4 or 5, rarely absent, usually smaller than the alternate sepals, concave or hooded 

(cucullate), rarely almost flat, often shortly clawed, enfolding the epipetalous stamens; 

stamens 4 or 5, in a single series opposite the petals, filaments thin, adnate to the base of 

the petals, anthers minute, versatile, generally 2-thecate, dehiscing by longitudinal slits; 

disc intrastaminal, nectariferous, thin to more or less fleshy, entire or lobed, glabrous or 

rarely pubescent, free from the ovary or adnate around the base; ovary superior or halfinferior, 

2-3(-5)-loculate, with 1 ovule in each locule; ovules anatropous, basal and erect. 

Fruit a capsule or a dry or fleshy drupe with 1-3 dehiscent or indehiscent stones (endocarpids), 

often winged. Seeds with scanty fleshy endosperm or rarely without endosperm; 

embryo large, oily, straight or rarely bent. 

Distribution. A cosmopolitan family of about 45 genera and 800 to 1000 species. In Sabah 

and Sarawak, represented by 8 native genera with 21 species of which only 4 species in 4 

genera are trees, the others are either erect or straggling shrubs or woody climbers. 

Ecology. Tropical rain forest to moderately arid areas and from near sea-level to elevation 

and latitude near the tree limit. 

Uses. A rather limited number of Rhamnaceae species furnish useful and valuable timber of 

local importance. Some West Indian species of Colubrina, Krugiodendron and Reynosia 

yield a hard and heavy wood sold as "Ironwood" Ornamental plants are restricted to the 

North American genus Ceanothus, of which several species are planted for their showy 

inflorescences, and the European Paliurus spina-christi, which is widely used as protective 

and ornamental hedge-plants. Edible fruits. are furnished by the Mediterranean jujubes 

305


(Ziziphus), which are widely cultivated in Africa and Asia. Further economic products of 

rhamnaceous plants are dyes (Rhamnus), laxatives and diuretics (Frangula, Paliurus, 

Rhamnus), soap (Colubrina, Ziziphus) and substitute for tea (Sageretia). None of these uses 

is, however, reported for taxa occurring in Sabah and Sarawak. 

Taxonomy. Suessenguth (l.c.) recognised 58 genera and grouped them into five tribes, 

mainly distinguished by ovary and fruit characters. The 21 genera of the largest tribe 

Rhamneae extend over nearly the whole area of the family, except extratropical South 

America. In Sabah and Sarawak, they are represented by Alphitonia, Colubrina and 

Rhamnus. The tribe Zizipheae comprises 20 genera, and is mainly centered in subtropical 

and tropical regions of the world, namely in the West Indies and Central America, East 

Asia and Indo-Malesia. Berchemia and Ziziphus are the two genera native to Borneo. The 

tribe Colletieae are mostly restricted to the extratropical South America, with one genus 

(Adolphia) in Mexico and California and another genus (Discaria) extending to New 

Zealand and Australia. The predominantly climbing or even herbaceous tribe Gouanieae 

are confined to a narrow pantropical belt. Out of the five genera recognised, only Gouania 

is widely distributed throughout Malesia, including Sabah and Sarawak. Smythea and 

Ventilago, the only two genera of the tribe Ventilagineae, are restricted to the Paleotropics, 

and both occur in Sabah and Sarawak. 

Key to genera 

1. Small to medium-sized trees or shrubs; fruits never winged ................ : ........................ 2 

Woody climbers with winged fruits .............................................................................. 6 

2. Leaves 3-veined from the base; stipules often transformed into straight or recurved 

spines .......................................................................................................... 4. Ziziphus 

Leaves pinnately veined; stipules never transformed into spines .............. . 

3. Fruit a single-stoned fleshy drupe .............. . 

Berchemia Necker 

Elem. bot. (1790) 122; Backer & BakhuizenJ I.e. 83. 

22 species; North America (1 species), Minor Asia, Indo-China, New Caledonia, 

Malesia; 1 species in Sabah and Sarawak. 

Erect or straggling shrubs. Leaves membranous, with numerous pairs of slightly 

curved, parallel lateral veins. Inflorescences raceme~like thyrses. Flowers bisexual 

or unisexual by abortion. Fruit an ellipsoid to obovoid, single-stoned fleshy drupe. 

Fruit a 2-4-stoned capsule or drupe; epicarp dry or more or less fleshy .... . .. .4 

4. Fruit a 3-locular, capsule ................................ ..2. Colubrina 

Fruit indehiscent... ...... . ......... 5 

5. Epicarp mealy or corky; endocarpids splitting ventrally. Seeds distinctly arillate .......... . 

...................................... 1. Alphitonia 

Epicarp fleshy; endocarpids indehiscent. Seeds never arillate ..................... 3. Rhamnus 

306

RHAMNACEAE (SCHIRAREND) 

6. Branches often with tendrils. Fruit with three rounded commissural wings .. 

Gouania Jacq. 

Select. Stirp. Amer. Hist. (1763) 263; Ridley l.c. 468; Masamune I.c. 434; Backer & 

BakhuizenJ I. c. 85. 

About 50 species; tropical America and Africa, Madagascar, India, Indo-China, 

Malesia, Australia, Polynesia; 3 species in Sabah and Sarawak. 

Woody climbers with solitary tendrils. Leaves membranous to coriaceous, ovate or 

cordate, pinnately veined, rarely 3-veined, margin entire to serrate. Inflorescence a 

multiflowered, axillary or terminal, spike- or raceme-like thyrse. Flowers bisexual 

or polygamous. Fruit a 3-locular schizocarp, splitting septicidally .into three 2- 

winged indehiscent endocarpids. 

Branches never with tendrils. Fruit with one apical wing ... . .. 7 

7. Fruit a globose indehiscent structure, bearing a long (at least twice as long as the fruit) 

apical wing. 

VentiIago Gaertn. 

Fruct. Sem. PI. 1,1 (1788) 233; Ridley I.c. 465; Masamune I.c. 434; Backer & BakhuizenJ 

I.c. 8I. 

About 40 species; Africa, Madagascar, India, Indo-China, Malesia, Australia; 4 


Leaves chartaceous to coriaceous, elliptic to oblong-ovate, pinnately veined, 

margin entire to serrate. Inflorescences axillary clusters or shortly stalked cymes 

which are often combined into racemes or panicles. Flowers bisexual, rarely 

apetalous. 

Fruit a compressed, septicidally dehiscing capsule bearing a small apical wing ............. . 

Smythea Seem. 

Bonplandia 9 (1861) 255; Ridley I.c. 468; Masamune I.c. 434; Backer & BakhuizenJ I.c. 

81. 

7 species; Indo-China, Malesia, Polynesia; 1 species in Sabah and Sarawak. 

Leaves membranous to chartaceous, elliptic to ovate, pinnately veined, often somewhat 

asymmetric. Flowers in axillary fascicles, bisexual. 

1. ALPHITONIA Reiss. ex End!. 

(Greek, alphiton = baked barley meal; the mealy nature of the ripe fruit exocarp) 

Gen PI. (1840) 1098; Braid, Kew Bull. (1925) 168; Masamune I.c. 434; Anderson I.c. 289; Whitmore 

Tantra & Sutisua I.c. 293. 

Evergreen trees or shrubs; buds and young stems often more or less densely brownish 

pubescent. Bark and wood of several species with rather strong smell of sarsaparilla. Leaves 

alternate, ovate to obovate or elliptic to lanceolate, rarely almost cordate, chartaceous to 

coriaceous, venation pinnate; blade usually glabrous above, whitish or rusty pubescent 

underneath; stipules small, deciduous. Inflorescences many-flowered, di- or trichotomous, 

axillary or rarely terminal cymes. Flowers 5-7-merous; hypanthium patelliform to hemi- 

307


sperical; sepals 5, triangular, keeled within; petals 5, clawed, cucullate; stamens 5; disc 

mainly thick, nectariferous; ovary inferior, immersed in the stout disc, 2-3-locular, style 

short, 2-3-lobed, basally often densely whitish to yellowish pubescent or glabrous. Fruits 

drupaceous, globose or broadly ovoid; mesocarp thick, mealy; endocarp splitting longitudinally 

into 2 to 3, hard and coriaceous endocarpids, at the base surrounded by the persistent 

hypanthium. Seeds more or less completely enclosed by a loose, membranous, reddish 

brown arit; testa smooth, crustaceous; embryo straight; endosperm cartilaginous. 

Distribution. About 10 species; Malesia, Polynesia and Australia. One species in Sabah 


Uses. The wood of the tree species in Australia, Malesia and most of the Polynesian Island 

is said to be of considerable value (Braid I. c. 170), and used for axe-handles, piling, and 

cabinet-making. 

Alphitonia excelsa (Fenzl) Reiss. ex Endl. 

(Latin, excelsus = lofty, high; the tree) 

Fig. 1. 

I.c. 1098; Maiden, For. Fl. N.S.w. (1903) 38; Braid I.c. 177; Masamune l.c. 434; Anderson I.c. 289; 

Whitmore, Tantra & Sutisna l.c. 293. Basionym: Colubrina excelsa Fenzl in Endl., Enum. PI. (1837) 

20. Type: Cunningham, s.n., Australia, Queensland, Brisbane, Moreton Bay (K). Synonyms: 

Rhamnus incanus Roxb., Fl. Ind. (1832) 603; Alphitonia incana (Roxb.) Kurz ex Hoogland, Kew 

Bull. 15 (1961) 33; A. philippinensis Braid I.c. 183. 

Small to medium-sized tree rarely to 30 m high; bole straight to 20 m high. Bark and wood 

with a rather strong smell of wintergreen. Bark smooth, grey to brownish; inner bark thin, 

greenish, more or less straw, cambium yellow. Sapwood whitish; heartwood reddish 

brown. Young twigs, axillary buds, stipules, petioles and lower surface of the leaves more 

or less densely yellowish to whitish pubescent. Stipules minute, acute triangular, early 

deciduous. Leaves alternate, chartaceous to subcoriaceous, ovate-lanceolate or elliptic, 6- 

20(-30) x 3-6(-10) cm; base acute or obtuse to rounded, rarely cordate, margin entire, 

apex acute to shortly acuminate; midrib prominent below, impressed above; lateral veins 

10-14 pairs; petioles 5-10 mm long. Inflorescences terminal or axillary cymes. Flowers 

about 4-5 mm in diameter, light green to yellowish, 5-merous, rarely 6- or 7-merous; 

hypanthium shallow hemispherical; sepals about 1.5 mm long; petals erect to slightly 

recurved; stamens about 1. 5 mm long; disc shallow, lining the hypanthium, ring-shaped, 

glandulous; ovary obconical, whitish tomentose; style conical. Fruits globose to broadly 

ovoid, to 15 mm in diameter, greenish when young, black when ripe. Seeds often permanently 

attached to the receptacle after dehiscence of the fruit. 

Vernacular names. Sabah--balik angin (Dusun, Malay), pati yata (Dusun), pokudata 

(Dusun). 

Distribution. A widely distributed species, extending from Borneo, the Philippines, New 

Guinea to Australia. In Sabah and Sarawak common and widespread in the lowlands. Also 


Taxonomy. Braid (I.c.) split the former A. excelsa into a group of species poorly differentiated 

in leaf shape and size. Examination of numerous specimens throughout Malesia and 

308


A 

IE 

E 

Fig. 1. Alphitonia excelsa. A, flowering leafy twig; B, flower; C, infructescence; D, dehisced 

mericarps of a fruit; E, seed almost completely covered with a thin reddish brown aril CA & B from 

SAN A 364, C-E from Schirarend 5.) 

309


l~ 

o 

Fig. 2. Colubrina beccariana. A, leafy twig; B, flower; C, fruit; D, dehisced endocarpids of a fruit. 

CA from Sin clair et al. 8946, B from SAN 54539, C from SAN 15374, D from Kostermans 21228.) 

310


Australia supports the conclusion that there is only one highly variable and widely 

distributed species. 

Ecology. Fairly common element of lowland secondary forests or of depauperate, disturbed 

primary forests throughout the area. Maiden (I.e.) reports that the trees grow well on poor 

sandy soils. 

Uses. Braid (I.e.) states that the species is used as fodder plant. He also records its 

medicinal properties and notes that it is valuable for tanning. 

2. COLUBRINA Rich. ex Brongn. 

(Latin, eolubrinus = like a snake; perhaps from the French name bois eouleuvre, 

the snake wood or serpent tree, C ferruginosa, from Martinique Island) 

Ann. Sci. Nat. 10 (1827) 61; Ridley I.c. 465; Backer & BakhuizenJ I.e. 84; Johnston, Brittonia 23 

(1971) 2; Latiff I.e. 298; Whitmore, Tantra & Sutisna I.e. 293. 

Deciduous or evergreen trees and shrubs, rarely scandent. Stipules minute, early deciduous. 

Leaves alternate to rarely opposite, chartaceous to subcoriaceous; margin entire, serrate or 

crenate; venation pinnate or acrodromous from the base. Inflorescence an axillary cyme or 

small thyrse, sessile or shortly peduncled, mainly few-flowered. Flowers usually 5-merous, 

bisexual, perigynous; hypanthium hemispherical; sepals 5, triangular, rather densely pubescent 

outside, conspicuously keeled within, deciduous; petals 5, clawed; stamens 5, about as long 

as petals, dehiscing introrsely; disc massive, fleshy; ovary semi-inferior, 3 (-4)-locular, 

styles 3-lobed to trifid, stigmas 3. Fruit a subglobose, 3-lobed capsule; mesocarp thin, dry 

or rather fleshy; endocarp cartilaginous to woody, splitting into 3 ventrally dehiseent en doearpids. 

Seeds broadly obovate, shiny, rarely with a minute basal aril, surface smooth or 

pitted; testa coriaceous to bony; endosperm fleshy, thick. 

Distribution. An almost pantropical genus of about 24 species, mainly centered in the 

Neotropics. Two species (C asiatiea and C beeeariana) are found in Sabah and Sarawak. 

Uses. The very hard and heavy wood of several neotropical species, known as "West 

Ironwood" (C arboreseens, C el/iptiea) is of some local importance. 

Key to Colubrina species 

Small to medium-sized tree. Leaves subcoriaceous, oblong-elliptic to oblong-obovate, 3- 

veined from the acute base ............................................................................. C. beccariana 

Small coastal shrub. Leaves chartaceous, broadly ovate to cordate, pinnately veined ............ . 

C. asiatica (L.) Brongn. 

I.e. 62; Masamune I.c. 434. Basionym: Ceanothus asiatieus L., Sp. PI. (1753) 196. 

Sprawling shrub. Leaves ovate to cordate, very thin, pinnately veined, margin dentate. 

Flowers in few-flowered thyrses. Fruit a globose capsule with explosive dehiscence. 

311


India, China, throughout Malesia. In Sabah and Sarawak scattered in coastal area, and 

collected from Pulau Tiga FR (FDBNB 38625) and Timbun Mata FR (FDBNB 44652) 

in Sabah, and Telok Pinang, 1st Div. in Sarawak (s. 41941). 

Colubrina beccariana Warb. Fig. 2. 


Bot. Jabrb. Syst. 13 (1891) 367; Ridley I.e. 465; Johnston I.e. 44; Latiff I.e. 298; Whitmore, Tantra & 

Sutisna I.e. 293. Type: Warburg 20187, Dutch New Guinea (holotype B, destroyed; isotype A). 

Synonym: C. anomala King, J. As. Soc. Beng. 45 (1896) 377. 

Small to medium-sized tree to 25 m high, bole up to 20 m, diameter to 50 cm, sometimes 

with weak buttresses. Bark smooth or slightly dimpled, very brittle, reddish brown to dark 

brown; inner bark light red, pale yellow near cambium. Wood very hard, sapwood pale 

yellow. Young branches, axillary buds, petioles, major leaf venation, peduncles, pedicels 

and gynoecium densely rusty pubescent. Stipules subulate, to 4 mm long, early deciduous. 

Leaves alternate, subcoriaceous, oblong-elliptic to oblong-obovate, 10-22 x 3.5-8.5 cm; 

base acute, margin slightly revolute, entire to finely serrulate, individual teeth minute, 

appressed, bearing a black, glandular apex, apex acuminate, acumen 1-2 cm long and 2-3 

mm wide, apically rounded; venation pinnate, slightly impressed above, more or less 

prominent below; lateral veins 4-6 pairs, basal ones extending into the upper half of the 

lamina; intercostal veins rectangular to the midrib, strongly percurrent between 

superadjacent lateral veins; petioles 6-15 mm long. Inflorescences axillary fascicles or 

shortly peduncled cymes; pedicels 1-3 mm long. Flowers 5(-6)-merous; hypanthium 

shallow patelliform; sepals triangular to acuminate, distinctly keeled within; petals obovate, 

cucullate; disc fleshy, slightly grooved; ovary immersed in the fleshy disc, densely brownish 

pubescent, 2-3-locular; style more or less deeply bi- to tri-fid, stigmas 3, slightly capitate. 

Fruits globose, 10-16 mm in diameter, glabrous; exocarp flaky, black when dry; mesocarp 

thin; endocarp woody, splitting into 3 explosively dehiscing endocarpids when ripe; 

pedicels to 2 cm long. Seeds 8-10 mm across, testa glossy, reddish brown. 

Vernacular names. Sabah----obar-obar (Dusun), odok-odok (Kadazan), udok-udok (Malay). 

Distribution. Peninsular Malaysia and Borneo (Sabah and Kalimantan). In Sabah, collections 

have been made from Kinabatangan, Lamag, and Sandakan (Kabili-Sepilok FR) districts. 

Ecology. A typical lowland species, mainly found in undisturbed primary rainforest. 

3. RHAMNUS L. 

(after the ancient Greek plant name rhamnos) 

Sp. PI. (1753) 193; Masamune I.e. 434; Anderson I.e. 290; Latiff I.e. 298. Synonym: Oreorhamnus 

RidI., J. Fed. Malay. st. Mus. 10 (1920) 131. 

Deciduous or evergreen trees and shrubs, rarely climbing, often thorny. Stipules small, 

caducous. Leaves alternate or rarely opposite, glabrous to rather densely pubescent, 

312


membranous to coriaceous, venation pinnate; margin entire to toothed. Inflorescences 

axillary cymes, sessile or peduncled, rarely reduced to a solitary flower. Flowers perigynous, 

bisexual or unisexual by abortion; hypanthium campanulate to urceolate; sepals 4-5, 

deltoid, valvate in bud; petals 4-5 or rarely wanting, minute, usually shortly clawed, 

concave to hooded; stamens 4-5, sterile and rudimentary or completely wanting in female 

flower; disc nectariferous, lining the floral tube; ovary superior, 2-3 (-4)-10cular, each 

locule with one solitary, erect, anatropous ovule; stigma 3-lobed or stigmata 2-3. Fruit a 

globular to obovoidal drupe with 2-3(-4) endocarpids; mesocarp fleshy to coriaceous, 

often with tanniferous idioblasts or mucilage cavities; endocarp cartilaginous to woody; 

endocarpids abaxially convex, adaxially slightly angular to almost flat, dehiscent or 

indehiscent. Seeds narrow ellipsoidal to obovoidal, rarely suborbicular; testa membranous; 

endosperm fleshy, scanty; embryo large. 

Distribution. The genus extends over nearly the whole area of the family, being abundant 

in Eastern Asia and Southwestern North America, rather scanty in Europe and Africa, and 

absent from Madagascar, Australia and Polynesia; one species in Sabah and Sarawak. 

Ecology. Primarily distributed in the temperate and subtropical regions of both hemispheres, 

the genus extends into tropical highlands and is found even in semiarid to arid 

places. Of significance is the obvious avoidance of hot tropical lowlands. 

Uses. Several species are grown as ornamentals or hedge-plants, other furnish wood of 

local importance or drugs (e.g., R. cathartica). 

Rhamnus borneensis Steenis 


Fig. 3. 

1. Bot. 72 (1934) 6; Masamune I.e. 434; Anderson I.e. 290; Latiff I.e. 298. Type: Clemens 27876, 

British North Borneo, Mt. Kinabalu (L). Synonyms: R. laneifolia Steenis I.e. 7, Anderson I.e. 290; 

R. borneensis Steenis var. borneensis Latiff I.e. 299. 

Small tree to 15 m high; crown small; bole straight, diameter to 20 cm. Bark rugulose; 

inner bark yellow, fibrous, without latex. Sapwood silky white, heartwood dark brown. 

Young twigs, axillary buds, stipules, petioles, pedicels, flower buds and lower surface of the 

leaves densely yellowish pubescent, indument composed of stellate hairs; branches soon 

glabrescent, with smooth, red to red-brown bark, without lenticels. Leaves lanceolateelliptic 

to obovate, chartaceous to subcoriaceous, shiny and glabrous above, densely whitish 

to yellowish pubescent below, 7.5-14 x 3-5.5 cm; base acute, margin slightly revolute, 

entire to finely serrulate, apex rounded or acute to acuminate, acumen blunt or broadly 

triangular; lateral veins 7-11 pairs, straight to slightly curved, parallel, ex-medially 

anastomosing with each forming well-developed marginalloopes. Inflorescences axillary 

fascicles or shortly peduncled, few-flowered cymes. Flowers 4-5 mm in diameter; pedicels 

4-7 mm long; hypanthium hemispheric to campanulate; sepals deltoid to acute triangular, 

1.5-2 mm long and wide, distinctly keeled within; petals cucullate, clawed, distinctly 2- 

lobed, 1.5-2 mm long; stamens 1-1.5 mm long; disc thin, lining the hypanthium, glabrous; 

ovary very small, conical to globose, 3-locular, glabrous; style undivided, cylindric, stigma 

313


slightly 3-partite, inconspicously capItate. Fruits red, globose to obovoid, with 2-3 endocarpids; 

mesocarp coriaceous; endocarp stony, endocarpids lenticular. Seeds elliptic to 

cylindric. 

Distribution. Sumatra, Peninsular Malaysia and Borneo. Uncommon; in Sabah known 

from a few collections (e.g., Clemens 27876 and SAN 56276 from Mt. Kinabalu area), and 

in Sarawak scattered and confined to montane forest (e.g., Nooteboom & Chai 1996 from 

Mt. Murut; S. 35822 and S. 38139 from Mt. Mulu; and S. 47355 from Mt. Berumput). No 

record from Brunei and Kalimantan. 

Ecology. A species of submontane to montane ridge and mossy forests to 2700 m. 

4. ZIZIPHUS Mill. 

(from the Persian plant name zizuf) 

Gard. Diet. Abr. ed. 4 (1754); Ridley I.c. 461; Merrill, PEB (1929) 177; Masamune I.c. 435; Backer 

& BakhuizenJ I.c. 81; Anderson I.c. 290; Corner I.c. 610; Latiff I.c. 299; Whitmore, Tantra & 

Sutisna I.c. 293. 

Deciduous or evergreen, small to medium-sized trees, erect or straggling shrubs or to 30 m 

high woody climbers, often armed. Stipules small, deciduous or often transformed into 

straight or recurved spines. Leaves alternate, usually distinctly 3-veined from the base, 

rarely pinnately veined, often slightly asymmetric; margin entire or serrate to crenate. 

Inflorescences axillary fascicles or sessile or peduncled, umbel-like cymes, rarely terminal 

or axillary thyrses. Flowers perigynous, bisexual, yellowish to greenish; hypanthium 

shallow patelliform to hemispherical; sepals 5, keeled inside, early deciduous; petals 5, 

cucullate, clawed, rarely absent; disc nectariferous, fleshy, mainly flat, 5- to lO-lobed, 

lining the hypanthium; ovary 2-3 (-4)-locular, styles 2-4, stigma 2-3(-4), very small, 

slightly capitate. Fruits drupaceous, fleshy or almost dry, exclusively one-stoned; mesocarp 

fleshy to leathery; endocarp bony to woody, surface smooth to furrowed. Seeds ellipsoid, 

nearly pIano-convex, raphe lateral, testa thin, membranous; endosperm scanty, fleshy; 

embryo straight. 

Distribution. An almost pantropical genus of about 100 species centered in tropical 

America and in Southeastern Asia. In Sabah and Sarawak, 8 species, which are mostly 

thorny straggling shrubs or woody climbers of the lowlands. 

Ecology. The genus is widely distributed in tropical and subtropical regions of the world, 

extending locally into temperate or even semi-arid to arid zones. 

Uses. Two species, Z. jujuba Mill. and Z. mauritiana Lam. are widely cultivated for their 

edible fruits. The wood of some other species is of local importance, e.g., for housebuilding, 

furnitures, cabinet-making, and for agricultural tools. 

314


! " 

//" _." 

---. - .. -- 

5mm 

c 

I~ 

Fig. 3. Rhamnus bomeensis. A, flowering leafy twig; B, flower with 2 sepals, 2 petals and 2 stamens 

removed; C, twig with mature fruits. (All from SAN 56276.) 

315


/'/: 

'--:'


Key to Ziziphus species 

l. Small to medium-sized unarmed tree. Leaves to 30 cm 10ng ................. Z. angustifolius 

Erect or straggling shrub or woody climber with stipulary spines. Leaves smaller. ....... 2 

2. Leaves less than la cm long ...... . 

Leaves 10-17 cm long .............. . . .... 7 

3. Fruits ellipsoid, with flattened base and acuminate apex, to 4 cm long .......................... . 

Z. borneensis Merr. 

I.e. (1929) 178; Masamune I.e. 435; Whitmore, Tantra & Sutisna I.e. 294. 

Straggling shrub or woody climber with recurved stipulary spines, plant often 

densely ferruginous-tomentose. Leaves elliptic to oblong-elliptic, chartaceous to 

subcoriaceous; base often distinctly asymmetric, margin entire to minutely dentate, 

apex acuminate. Flowers in axillary fascicles. Fruits a very large ellipsoid drupe. 

Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan). 

Fruits globose to obovoid, to 1 cm in diameter.. ................ . . ..... .4 

4. Leaves very thin, venation pattern of the lower surface dark coloured and typically 

distinct. Style undivided and glabrous ................................................. . 

Z. cumingiana Merr .. 

Phi1ipp. 1. Sci. 1, Suppl. 3 (1906) 206. 

Erect or straggling shrub with one recurved stipulary spine per node. Leaves ovate, 

often distinctly asymmetric, very thin; margin finely dentate. Flowers in fewflowered 

axillary cymes, very small. Drupes glabrous. Borneo (Sabah) and the 

Philippines. 

Leaves chartaceous to coriaceous, venation pattern never distinctly coloured. Styles 2(- 

3)-lobed, densely pubescent. ......................................................................................... 5 

5. Leaves with distinct arcs of veins diverging from the lateral veins ................................ . 

Z. horsfieldii Miq. 

Fl. Ned. Ind.1, 1 (1855) 643; Masamune I.e. 435; Latiff I.e. 300; Whitmore, Tantra & 


Straggling shrub with one short stipulary spine per node. Leaves ovate to elliptic, 

very thin, often somewhat asymmetric; base obtuse, margin minutely dentate, apex 

shortly acuminate. Flowers in axillary, dichotomously branched cymes. Drupes 

globose, densely brownish pubescent. Sumatra, Peninsular Malaysia, Java, Borneo 

(Sabah, Sarawak, Brunei, Kalimantan). 

Leaves without such lateral veins ............................................. . . .......... 6 

6. Leaves bearing a distinct intra-marginal vein ................. . 

Z. havilandii Rid!. 

Kew Bull. (1931) 495; Masamune I.e. 435; Whitmore, Tantra & Sutisna I.e. 294. 

Straggling shrub with one recurved stipulary spine per node, plant more or less 

densely yellowish pubescent, glabrescent. Leaves ovate, often asymmetric, very 

thin; base obtuse, margin minutely dentate. Flowers in dichotomously branched, 

317


axillary cymes. Drupes globose to obovoid, densely yellowish to brownish 

pubescent. Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan). 

Leaves without any intramarginal vein .............. . 

Z. suluensis Merr. 

Philipp. J. Sci. 30 (1926) 408, I.c. (1929) 177; Masamune l.c. 435; Whitmore, Tantra & 

Sutisna l.c. 294. 

Straggling shrub with one recurved stipulary spine per node. Leaves elliptic, subcoriaceous; 

base obtuse to rounded, often somewhat asymmetric, margin minutely 

dentate, apex acuminate. Flowers in small, axillary, few-flowered cymes. Drupes 

globose to obovoid, densely brownish pubescent. Sumatra, Borneo, Philippines and 

the Moluccas. 

7. Leaves chartaceous to subcoriaceous, venation pattern typically prominent on the lower 

surface. Bark of twigs cream-coloured, densely lenticellate .................. . 

Z. crebrivenosa C.B. Rob. 

Philipp. J. Sci. 3 (1908) 201; Whitmore, Tantra & Sutisna I.c. 294. 

Woody climber with one recurved stipulary spine per node and densely lenticellate 

branches. Leaves obovate to elliptic, often somewhat asymmetric, chartaceous; 

base obtuse to rounded, margin faintly denticulate, apex acute to obtusely 

acuminate and mucronate. Flowers in axillary cymes. Drupes globose, to 3 cm 

diameter, densely lenticellate. Borneo (Sabah, Sarawak, Kalimantan), Philippines 

and Celebes. 

Leaves coriaceous, venation pattern very delicate, never prominent. Bark of twigs 

brownish, not lenticellate ........................... . 

Z. calophylla Wall. 

in Roxb., Fl. Ind. 2 (1824) 366; Merrilll.c. (1926) 364, I.c. (1929) 177; Rid1ey, Kew Rull. 

(1931) 494; Masamune I.c. 435; Latiff I.c. 300; Whitmore, Tantra & Sutisna I.c. 294. 

Sturdy straggling shrub, with short, solitary or paired stipulary spines. Leaves 

coriaceous, elliptic to oblong-elliptic, base usually narrowed, margin minutely 

dentate, apex acuminate. Flowers in axillary or terminal cymes. Drupes globose, 

densely brownish tomentose. Sumatra, Peninsular Malaysia and Borneo (Sabah, 

Brunei, Kalimantan). 

Ziziphus angustifolius (Miq.) Hatusima ex Steenis 

(Latin, angustifolius = having narrow leaves) 

Fig. 4. 

Nova Guinea, Botany 3 (1960) 13; Hatusima, Cat. Hort. Bot. (1957) 242; Merrill & Perry, J. Am. 

Arb. 20 (1939) 337; Anderson I.c. 290; Latiff l.c. 299; Whitmore, Tantra & Sutisna I.c. 293. 

Basionym: Solenostigma angustifolium Miq., Fl. Ned. Ind., Suppl. (1861) 412. Type: Miquel HB 

758 and HB 2501, Sumatra West Coast (syntypes L). Synonyms: Celtis angustifolia (Miq.) P1anch. 

in DC., Prod. 17 (1873) 186; C. grewioides Warb., Bot. Jahrb. 13 (1891) 287; Ziziphus inermis 

Merr., Govt. Lab. Pub!. Phi1ipp. 35 (1906) 37; Z.forbesii Bakerf, J. Bot. 61, Suppl. (1923) 10; Z. 

grewioides (Warb.) Perry ex Steenis, Blumea 7 (1954) 595. 

Small to medium-sized, unarmed tree to 30 m high and 40 cm diameter; crown narrow 

conical; bole straight, rarely crooked, up to 20 m high, sometimes with buttresses of about 1 

318


m high and 1-1. 5 m out. Bark not fissured, not peeling, with densely spaced lenticels, 

greyish to brownish, about 0.2 mm thick; inner bark yellow to brownish, blaze strawcoloured, 

cambium orange. Sapwood hard, light to greyish brown, to 15 cm thick; 

heartwood hard and heavy, dark brown. Young twigs, axillary buds, stipules, petioles, 

peduncles, pedicels and flower buds rather densely yellowish to brownish pubescent; older 

branches glabresc~nt, greyish to brownish, lenticellate. Stipules minute, narrow lanceolate, 

not transformed into spines, 1-2 mm long, early deciduous. Leaves alternate, chartaceous 

to subcoriaceous, symmetric, oblong-elliptic to oblong-ovate, 12-20 x 3-7 cm; base obtuse 

to acute, margin slightly revolute, finely serrate, apex acute to acuminate, acumen about 1- 

2 cm long, 0.5 cm wide, apically distinctly mucronate; venation 3-veined from the base; 

intercostal veins rectangular to the midrib, strongly parallel between the midrib and the 

lateral veins, outer intercostal veins anastomosing to form a slightly looped marginal vein. 

Inflorescences axillary, peduncled, branched cymes, to 3 cm long. Flowers 4-6 mm in 

diameter, 5-merous, greenish to yellowish; pedicels 2-4 mm long; hypanthium patelliform; 

sepals triangular, keeled within; petals clawed, cucullate; stamens about 1.5 mm long, 

filaments pale greenish, anthers green; disc prominent, glandulous, glabrous, yellow to 

ochre; ovary immersed in the disc, 2-3-locular, style 2-3-lobed, increasingly diverging 

during anthesis, stigmas distinctly papillous. Fruits globose to obovoid, often I-seeded, 

10-20 mm diameter; mesocarp comparatively thin; endocarp woody. 

Vernacular names. Sabah-kayu labu (Kadazan), pasil-pasil (Bajau). 

Distribution. India, Burma, Thailand, Sumatra, Peninsular Malaysia, Borneo (throughout), 

Philippines, Celebes, Moluccas, New Guinea, and Solomon Island. In Sabah recorded from 

Kinabatangan, Tenom, Tongod, Lahad Datu, and Sandakan districts; and in Sarawak from 

various localities in the 1st, 4th and 7th Divisions. Also occurs in Brunei and Kalimantan. 

Ecology. Lowland mixed dipterocarp forest to c. 600 m. 

319


RHIZOPHORACEAE 

L. Madani & K.M. Wong 




Merrill, EB (1921) 420; Masamune, EPB (1942) 515; Browne, FTSB (1955) 296; Ding Hou, FM 1, 

5 (1958) 429; Burgess, TBS (1966) 431; Ashton, MNDTS 2 (1988) 342; Juncosa & Tomlinson, Ann. 

Missouri Bot. Gard. 75 (1988) 1278; Kochummen, TFM 4 (1989) 302; Whitmore, Tantra & Sutisna, 

eLK 2, 1 (1990) 295. 

Trees or shrubs, the mangrove species often with characteristic root formations (branching 

stilt-roots in Rhizophora, knee pneumatophore-roots in Bruguiera). Bark invaginating 

finely and regularly into the wood in the inland genera (Carallia, Gynotroches, Pellacalyx). 

Twigs solid or (in Gynotroches and Pellacalyx) hollow; branch nodes swollen. Leaves 

opposite and decussate, entire or toothed, pinnately veined; stipules interpetiolar, caducous, 

with colleters on the inner side. Inflorescences usually axillary, cymose or a fascicle of 

flowers or flowers solitary. Flowers bisexual or (in Gynotroches) unisexual (the plants 

dioecious), radially symmetrical; calyx-lobes 4-16, valvate; petals 4-16, free, fleshy, 2-jid 

to fimbriate or fringed with apiculate appendages or lacerate or lobed (rarely entire); 

stamens free or (in Pellacalyx) inserted on the calyx-tube, 1-3 times the number of petals 

or (in Kandefia) many; anthers 4-celled or (in Kandefia) many-celled, dorsifixed, splitting 

lengthwise or (in Rhizophora) opening by a ventral valve; ovary superior or semi-inferior 

or inferior, 2-12-celled (cells often incompletely or not separated at floral maturity and 

then apparently l-celled), style simple, stigma capitate or lobed, ovules 2-many per cell, 

pendulous, placentation axile; disc usually present, fleshy, annular, entire to lobed. Fruit a 

berry, calyx persistent. Seeds I-many per fruit, pendulous, endospermous; embryo with 

laminar cotyledons; germination epigeal or in the mangrove species viviparous (while the 

seed still enclosed in the fruit and the fruit still attached to the tree). 

Distribution. 15 genera with c. 145 species, pantropical. In Sabah and Sarawak 7 genera 

with 23 species. 

Ecology. The mangrove species occur on coastlines with warm currents, and many also 

grow in brackish water conditions within the estuaries or along the lowest stretches of 

rivers. The inland species are mostly of moist (primary or secondary) tropical forests. 

Uses. The mangrove species have been exploited for poles used in scaffolding and the 

construction of large fishing platforms and traps. The bark of almost all mangrove species 

(in Bruguiera, Ceriops and Rhizophora) are used for tanning. The mangrove species are 

also important in providing poles for the local charcoal industry, the calorific value of the 

wood being excellent. In the inland forests, Caralfia species grow to timber size and are 

exploited for logs. 

321


Taxonomy. Rhizophoraceae have been traditionally placed in the Myrtales and sometimes 

with the Cornales. This was based on the incorrect assumption that the Rhizophoraceae 

typically have an inferior ovary, whereas in fact this is a rare and derived condition. 

Sometimes, also based on a single character of the presence of interpetiolar stipules, the 

Rhizophoraceae has been said to be related to the Rubiaceae and Cunoniaceae. Dahlgren 

(1988) (Ann. Missouri Bot. Gard. 75: 1259), after considering a number of characters, 

places the Rhizophoraceae in the Celastrales, together with Celastraceae and Elaeocarpaceae. 

Cronquist (1981) (An Integrated System of Classification of Flowering Plants) separates 

the Rhizophoraceae sensu stricto in its own order, the Rhizophorales. 

Although a number of authors have treated the genera Anisophyllea and Combretocarpus 

as part of the Rhizophoraceae, it is now well established that these two genera should be 

separately classified in a family, the Anisophylleaceae (this volume). Within the Rhizophoraceae 

s.s. Juncosa & Tomlinson I.c. consider three tribes, Rhizophoreae (Bruguiera, Ceriops, 

Kandelia, Rhizophora), Gynotrocheae (Carallia, Gynotroches, Pellacalyx, Crossostylis) 

and Macarisieae (which includes seven genera not in our area). 

Some Rhizophoraceae can be confused with the Rubiaceae at a glance, as both typically 

have opposite leaves with interpetiolar stipules. The Rubiaceae have fused corolla members 

and have a corolla-tube, and the stamens are as many as the corolla-lobes. In the 

Rhizophoraceae, the corolla members (petals) are free, and frequently the stamens are at 

least twice the number of petals. 

Key to genera 

1. Plants of mangrove forests. Bark not invaginating into wood. Seeds germinating while 

still in the fruits and the fruits still attached to the tree (viviparous) ............................ 2 

Plants of inland forests. Bark invaginating finely and regularly into wood. Seeds 

germinating only after the fruits have fallen off the tree ............................................... 5 

2. Leaf apex pointed .. ................................. 3 

Leaf apex blunt or rounded ............ . .................... .4 

3. Plants developing buttresses and knee pneumatophores (breathing roots). Calyx-lobes 

8-16, lanceolate-linear. ............................................................................. 1. Bruguiera 

Plants developing conspicuous branching stilt-roots. Calyx-lobes 4, broadly ovate 

............................................................................................................... 7. Rhizophora 

4. Plants developing only short buttresses, trunk not usually thickened at base. Leaves 

broadly elIiptic to obovate, length at most twice the width. Hypocotyl (first elongating 

axis) in the germinating fruit with sharp longitudinal ridges ......................... 3. Ceriops 

Plants typically without buttresses, trunk developing a conically thickened base. Leaves 

narrowly elIiptic-oblong, length at least 2.5-3-times the width. Hypocotyl in the 

germinating fruit smooth ............................................................................ 5. Kandelia 

5. Twigs solid. Leaf-margin distinctly toothed or crenate. Inflorescence cymose, with a 

distinct main stalk. ....................................................................................... 2. CaraIlia 

322

RHIZOPHORACEAE (MADANI & WON G) 

Twigs hollow. Leaf-margin faintly toothed to entire. Inflorescence a fascicle of flowers 

without any distinct main stalk. ................................................................................... 6 

6. Stipules with overlapping margins. Ovary superior (calyx at the base of the fruit) 

............................................................................................................. 4. Gynotroches 

Stipules not overlapping, flat to slightly spreading. Ovary inferior (calyx at the apex of 

the fruit) .................................................................................................... 6. Pellacalyx 

1. BRUGUIERALam. 

(J.O. Bruguieres, 1750-1798, who worked for Lamarck's Encyclopaedia Methodica) 

beus, lenggadai (Malay) 

Tab. Enc. Meth. (1797) t. 397; Merrilll.e. (1921) 421; Masamune I.e. 515; Browne I.e. 299; Ding 

Hou I.e. 457; Burgess I.e. 431; Ashton I.e. 349; Kochummen I.e. 309; Whitmore, Tantra & Sutisna 

I.e. 295. 

Medium-sized trees with buttresses and knee pneumatophores, sometimes with aerial roots 

when young. Leaves usually coriaceous, entire, glabrous, finely black-dotted on the lower 

side; stipules lanceolate. Inflorescence a 2-5-flowered cyme or reduced to a solitary 

flower. Flowers bisexual, articulate at the base with the stalk; calyx thick, 8-15-lobed, 

lobes lanceolate-linear, acute; petals 2-lobed, falling off early; disc cup-shaped and attached to 

the calyx-tube; stamens twice the number of petals and paired; ovary inferior, 2-4-celled, 

ovules 2 per cell, style filiform, stigma obscurely 2-4-lobed. Fruits included in and adnate 

to the calyx-tube, usually l-celled, 1(-2)-seeded; germination viviparous; hypocotyl rounded 

or obscurely ribbed, blunt. 

Distribution. 6 species, tropical east Africa to Asia, throughout Malesia to Australia and 

Polynesia. In Sabah and Sarawak, 4 species. 

Ecology. In mangroves, usually occurring behind Rhizophora stands that are inundated. 

Forming pure stands in some sites. 

Uses. Not suitable for poles and for construction. The wood, however, is used for making 

charcoal. 

Key to Bruguiera species 

1. Flowers solitary. Bark in mature trees usually fissured-flaky ................................... 2 

Flowers several in a cyme. Bark usually smooth or lenticellate ..................................... 3 

2. Flower-stalks 10-12 mm long; tips of petal acute, each with 3 or 4 distinct bristles 

........................................................................................................ 2. B. gymnorrhiza 

Flower-stalks 6-8 mm long; tips of petals obtuse, each with 1-3 bristles ...... 4. B. sexangula 

323


3. Flowers small, the calyx 3-4 mm long, smooth; stalks 2-3 mm long. Fruit with 

reflexed calyx-lobes .............................................................................. 1. B. cylindrica 

Flowers larger, the calyx 6-10 mm long, ribbed; stalks 5-8 mm long. Fruit with erect 

calyx-lobes ............................................................................................ 3. B. parviflora 

1. Bruguiera cylindrica (L.) Blume 

(Latin, cylindricus = cylindric; the hypocotyl) 

En. PI. Jav. 1 (1827) 93; Browne I.e. 300; Ding Hou I.e. 467; Burgess I.e. 432; Ashton I.e. 353; 

Koehummen I.e. 310; Whitmore, Tantra & Sutisna I.e. 295. Basionym: Rhizophora cylindriea L., 

Sp. PI. (1753) 443. Type: Kari-kandel Rheede, Hort. Mal. 6 (1686) 59, t. 33. Synonyms: R. 

earyophylloides Burm.f, Fl. Ind. (1768) 109; R. eeratophylloides Gmel., Syst. Veg. (1796) 749; B. 

earyophylloides (Burm.f) Blume I.e. (1827) 93; B. malabariea Am., Ann. Mag. Nat. Hist. 1 (1838) 

369; Kanilia earyophylloides (Burm.f ) Blume, Mus. Bot. Lugd. Bat. 1 (1849) 141. 

Tree to 25 m tall, 45 cm diameter; buttresses to 1 m high. Bark grey, with many large 

prominent lenticels; inner bark pink. Sapwood yellowish brown. Leaves elliptic, 7-17 x 2- 

8 cm, thinly coriaceous; base cuneate, apex acute; lateral veins c. 7 pairs, distinct to 

obscure on both sides; stalk 1-4.5 cm long; stipules 2.5-3.5 cm long. Flowers in 3- 

jlowered cymes; stalks 2-3 mm long; calyx-tube smooth, 3-4 mm long, lobes 8, as long as 

the tube; petals 4-5 mm long, the lobes 113 the length of the petal, apex obtuse, each with 

1-3 bristles to 1.5 mm long, outer margin fringed with white hairs. Fruits with calyx-tube 

1-1.2 cm long, the calyx-lobes rejlexed; hypocotyl cylindric, to 15 cm long, 0.5 cm thick. 

Vernacular names. Sabah-beus (Malay). Sarawak--berus ngayong (Lundu Malay), berus 

puteh (Malay), ngayong (Lundu Malay). 

Distribution. SE Asia, throughout Malesia to N Queensland. In Sabah recorded from Pulau 

Gaya and Pulau Tiga in the west coast; in Sa.rawak recorded at Lundu and the Rejang delta. 


Ecology. Often gregarious on newly formed sandy clays and behind Avicennia on the 

seaface; it is succeeded by other species on better drained sites. 

Uses. The timber is mainly used for firewood and is not durable in contact with soil, hence 

it is not favoured as a pole in house construction. 

2. Bruguiera gymnorrhiza (L.) Lam. 

(Greek, gymnos = naked, rhiza = root; the exposed germinating hypocotyl) 

Eneyel. Meth. Bot. 4 (1798) 696; Browne I.e. 301; Ding Hou I.e. 461; Burgess I.e. 431; Ashton I.e. 

350; Koehurnmen I.e. 310; Whitmore, Tantra & Sutisna I.e. 296. Basionym: Rhizophora gymnorrhiza L., 

Sp. PI. (1753) 433. Type: Kandel Rheede, Hort. Mal. 6 (1686) 57, t. 31 & 32. Synonyms: R. palun 

DC., Prod. 3 (1828) 33; B. rheedii Blume I.e. (1827) 92; R. tinetoria Blaneo, Fl. Filip. (1837) 394; B. 

eapensis Blume I.e. (1849) 137; B. wightii Blume I.e. (1849) 138; B. rnmphii Blume I.e. (1849) 138; 

324

RHIZOPHORACEAE (MADANI & WONG) 

B. eylindriea sensu Hance (non Blume), 1. Bot. 18 (1879) 10; B. eonjugata sensu Merr. (non R. 

eonjugata L.), Philip. 1. Sc. 9 (1914) Bot. 118, I.e. (1921) 421, Masamune I.e. 515. 

Tree to 35 m tall, 45 cm diameter; often with buttresses to 1 m high. Bark dark grey to 

brown to black, deeply fissured and roughly flaky, lenticels conspicuous only when young; 

inner bark yellowish pink. Sapwood pale yellowish brown. Leaves elliptic-oblong, 8-20 x 

4-7 cm, thinly coriaceous; base cuneate or rarely obtuse, apex acute; lateral veins 9-10 

pairs, indistinct on both sides; stalk 2-4.5 cm long; stipules 3-4 cm long. Flowers solitary; 

stalks 10-12 mm long; calyx-tube distinctly ribbed at upper part, 8-12 mm long, lobes 10- 

16, 17-22 mm long; petals 13-15 mm long, the lobes l/rl/2 the length of the petals, apex 

acute, each with 3 or 4 bristles of 2-3 mm long, outer margin fringed with white silky 

hairs. Fruits with calyx-tube to 2.5 cm long, the calyx-lobes ascending, not reflexed; 

hypocotyl slightly ribbed, to 25 cm long, 2 cm thick. 

Vernacular names. Sabah--putut (Malay). Sarawak-berus kurong, berus merah, kurong 

(Malay). 

Distribution. Tropical east Africa, S and SE Asia to the Ryukyus, throughout Malesia to 

Australia, Micronesia and Polynesia. In Sabah and Sarawak all districts with a coastline. 


Ecology. One of the largest trees of the mangrove, found at the inland margins. 

Uses. The wood is hard, difficult to work and not ornamental, although it is easily 

impregnated with preservative. It can be used for rail sleepers, and is locally used in 

charcoal making and for firewood, and the poles are sometimes used in house construction. 

The bark has been used for the manufacture of cutch but is said to be inferior to that from 

Rhizophora. 

3. Bruguiera parviflora (Roxb.) Wight & Am. ex Griff 

(Latin, parvus = little,flores = flowers) 

Fig. 1. 

Trans. Med. Phys. Soc. Ca1c. 8 (1836) 10; Merril1l.e. (1921) 421; Masamune I.e. 515; Browne I.e. 

301; Ding Hou I.e. 464; Burgess I.e. 431; Ashton I.e. 353; Kochummen I.e. 310; Whitmore, Tantra & 

Sutisna I.e. 296. Basionym: Rhizophora parviflora Roxb., Fl. Ind. ed. Carey 2 (1832) 461. Type: 

Roxburgh no. 1246 (K). Synonyms: R. eylindriea sensu Roxb. (non L.), Hort. Beng. (1814) 36; 

Kanilia parviflora (Roxb.) B1ume I.e. (1849) 140; B. ritehiei Merr., Publ. Govt. Lab. Phi1ip. 6 

(1904) 11. 

Tree to 30 m tall, 45 cm diameter. Bark pale grey mottled, pale brown, smooth with small 

obscure lenticels; inner bark pinkish red. Sapwood pale brown. Leaves elliptic-oblong, 8- 

15 x 2.5-5 cm, thinly coriaceous; base cuneate, apex acute; lateral veins 9-10 pairs, slightly 

distinct on both sides; stalk 0.6-1.2 cm long; stipules 3-6 cm long. Flowers 2-5 in a 

cyme; stalks 5-8 mm long; calyx-tube distinctly ridged to the base, 6-10 mm long, lobes 

10-12, 2-3 mm long; petals 2-3 mm long, the lobes 1/2 the length of the petals, apex 

obtuse and reflexed, each with 1-3 bristles to 1.3 mm long, outer margin fringed with 

white silky hairs. Fruits with calyx-tube to 4 cm long, the calyx-lobes ascending, not 

reflexed; hypocotyl angular, to 6 cm long, 1.5 cm thick. 

325


Vernacular names. Sabah--beus (Malay), langarai (Idahan, Suluk), lanjing lanjing 

(Kcdayan), lenggadai (Malay). Sarawak--berus lenggadai, lenggadai (Malay). 

Distribution. Sri Lanka, SE Asia, throughout Malesia, New Britain. In Sabah and Sarawak 

common in mangroves. 

Ecology. Oftcn mixed with Rhizophora and occasional along the margins of mangrove 

channels. 

Uses. The wood lasts only a year in contact with soil in the tropics. A minor source of 

charcoal viCod and firewood, unpopular as a pole. 

4. Bruguiera sexangula (Lour.) Poir 

(Latin, sex = six, angulus = angle; the ridged hypocotyl) 

Encycl. Supp. 4 (1816) 262; Merrilll.e. (1921) 422; Masamune I.e. 515; Ding Hou I.c. 463; Burgess 

I.e. 431; Ashton I.e. 351; Kochummen I.e. 312; Whitmore, Tantra & Sutisna I.e. 296. Basionym: 

Rhizophora sexangula Lour., Fl. Coch. (1790) 297. Type: Loureiro, S.n. (BM). Synonyms: B. 

angularis Spreng., Syst. Veg. 2 (1825) 602; B. eylindriea sensu Blume (non R. eylindriea L.) I.e. 

(1827) 93; R. polyandra Blanco, Fl. Filip. (1837) 396; R. plieata Blanco I.e. 398; B. eriopetala 

Wight & Am. ex Am., Ann. Mag. Nat. Hist. 1 (1838) 368, Brovme I.e. 301; B. australis A. Cunn. ex 

Am. I.e. 368; R. australis (A. Cunn. ex. Am.) Steud., Nomencl. ed. 2,2 (1841) 449; R. eriopetala 

Steud. I. e. 449; B. parietosa Griff., Notu!. 4 (1854) 670; B. lO-angulata Griff. I. e. 669; B. oxyphylla 

Miq., Sumatra (1861) 324; B. malabariea sensu F.-Vii!. (non Am.), Nov. App. (1880) 79. 

Trce to 40 m tall, 60 cm diameter; buttresses to 1 m high, stilt-roots sometimes developing. 

Bark pale grey-brown, conspicuously lenticel/ate; inner bark pale brown. Sapwood 

yellowish brown. Leaves elliptic-oblong to oblanceolate, 8-14 x 3-6 cm, thinly coriaceous; 

base cuneate, apex acute; lateral veins 7-11 pairs, slightly distinct to obscure on both sides; 

stalk 0.7-2.5 cm long; stipules 3-6 cm long. Flowers solitary; stalks 6-8 mm long; calyxtube 

distinctly ridged to the base, 10-12 mm long, lobes 10-12, 18-20 mm long; petals c. 

15 mm long, the lobes 1/2 the length of the petals, apex obtuse and reflexed, each with ]-3 

bristles, outer margin fringed with white silky hairs. Fruits with calyx-tube 15-18 mm 

long, the calyx-lobes ascending, not reflexed; hypocotyl angular, to 6-8 cm long, l.5 cm 

thick. 

Vernacular names. Sabah-mata buaya (Malay). Sarawak--berus putut, putut (Malay). 

Distribution. Sri Lanka and SE Asia to New Britain and throughout Malesia. In Sabah and 

Sarawak common in mangroves. Also in Brunei and Kalimantan. 

Ecology. Scattered towards the inland margins of mangroves; on wetter soils together with 

B. cylindrica and B. parviflora. 

Uses. Mainly for charcoal making and firewood, and also in local house construction. 

326


Fig. 1. Bruguiera parviflora. Fruiting leafy twig. (From SAN 75544.) 

327


2. CARALLIA Roxb., nom. cons. 

(from karalli, an Indian plant name) 

PI. Corom. 3 (1811) 8; Merrilll.e. (1921) 421; Masamune I.e. 515; Browne I.e. 305; Ding Hou I.e. 

481; Ashton I.e. 291; Koehummen I.e. 312; Whitmore, Tantra & Sutisna I.e. 296. Synonyms: 

Symmetria Blume, Bijdr. (1826-27) 1130; Sagitfipetalum Merr., Philip. J. Se. 3 (1908) Bot. 247. 

Small to medium-sized trees, sometimes with small buttresses or stilt-roots. Bark smooth 

and hoop-marked, becoming roughened by minute longitudinal fissures and transverse 

cracks, sometimes scaly, grey-brown or pinkish brown; inner bark invaginating finely and 

regularly into the wood. Sapwood with conspicuous radiating lines. Stipules lanceolate. 

Leaves often black-dotted on lower side, margin entire or toothed. Inflorescence a 

condensed or lax cyme, or flowers in pairs or solitary. Flowers sessile or stalked, calyxlobes 

5-8, deltoid, acute to acuminate; petals 5-8, clawed; stamens twice the number of 

petals, usually free; disc angular, fleshy; ovary inferior, 5-8-celled with 2 ovules per cell, 

or I-celled with IO-12 ovules, styles filiform or slightly conical. Fruits small, ellipsoidobovoid, 

pulpy, l-celled, usually I-seeded. Seeds ellipsoid or reniform, endospermous; 

germination epigeal. 

Distribution. c. 11 species, Madagascar, India, Sri Lanka, SE Asia and Malesia to N 

Australia. 8 species in Sabah and Sarawak. 

Ecology. Mostly lowland forests, sometimes swamps, to hills and ridges to c. 2000 m. 

Uses. The timber of some species are suitable for making tool-handles and inferior construction 

such as picture-frames and panelling. It is also sometimes used for firewood and charcoal. 

Taxonomy. Four species enumerated here are not named but merely indicated by numbers; 

their precise naming will depend on the collection of better flowering material and, in one 

or two cases, will need to be based on a study of wider scope. 

Key to Carallia species 

1. Leaves without conspicuous dark gland-dots all over the blade ............................... 2 

Leaves with conspicuous dark gland-dots all over the blade ......................................... 5 

2. Leaves not longer than 4 cm; lateral veins 4-6 pairs, intercostal veins inconspicuous 

on both sides. Flowers solitary in each unbranched inflorescence ......... 8. Carallia sp. 5 

Leaves typically longer than 4 cm; lateral veins 6-11 pairs, intercostal veins 

conspicuous and often prominent on both sides. Flowers several to many on clearly 

branched cymes ........................................................................................................... 3 

3. Leaves stiffly leathery, margin coarsely saw-toothed; intercostal veins very prominent 

on upper side ....................................................................................... 6. Carallia sp. 3 

Leaves chartaceous to thinly leathery, margin entire to subentire or faintly toothed 

towards apex; intercostal veins only slightly prominent on upper side ......................... .4 

328

RHIZOPHORACEAE (MADANl & WONG) 

4. Leaf-margin faintly toothed to entire; lateral veins looping towards the margin forming 

only one clear submarginal vein. Flowers sessile, on an apparently simple axis 

(actually a cyme with branches reduced), peduncle nil or very short and 

inconspicuous ......................................................................................... 3. C. coriifolia 

Leaf-margin always slightly but distinctly toothed; lateral veins forming 2-several 

series of angular loops between midrib and margin. Flowers short-stalked, in a 

distinctly branched cyme, peduncle 1-1. 8 cm long, distinct... .............. 1. C. borneensis 

5. Leaf lateral veins forming 2-several series of angular loops between midrib and 

margin; intercostal veins typically net-like; margin typically with distinct fine teeth 

(rarely shortly toothed to subentire) ...................................................... '4. Carallia sp. 1 

Leaf lateral veins typically looping to form only one distinct submarginal vein; 

intercostal veins more or less parallel to the lateral veins; margin either shortly toothed 

or entire ....................................................................................................................... 6 

6. Leaf-margin shortly toothed to sub entire. Inflorescence peduncle 1-2.5 cm long. 

Flowers at least shortly stalked .............................................................. 2. C. brachiata 

Leaf-margin entire. Inflorescence peduncle to only 1 cm long at most. Flowers sessile 

.7 

7. Leaves elliptic to slightly obovate, to 7-8 cm long at most, chartaceous to thinly 

leathery, veins inconspicuous to slightly prominent on upper side ........ 5. Carallia sp. 2 

Leaves broad-obovate, larger, stiffly leathery, veins prominent on upper side ................. 

... 7. Carallia sp. 4 

1. Carallia borneensis Oliver 


Fig. 2. 

in Hooker, Ic. Pl. (1896) t. 2459; Merrilll.e. (1921) 421; Masamune I.e. 515; Ding Hou I.e. 484; 

Burgess I.e. 432; Ashton I.e. 356; Whitmore, Tantra & Sutisna I.e. 296. Type: Creagh, s.n., British 

North Borneo (K). Synonyms: Sagittipetalum mindanaensis Merr., Philip. J. Sc. 3 (1908) Bot. 247; 

S. palawanense E1mer, Leafl. Phi1ip. Bot. 5 (1913) 1830; Carallia mindanaensis (Merr.) Merr., En. 

Phi1ip. 3 (1923) 146. . 

Small or medium-sized tree to 25 m tall. Bark finely fissured, pale to brownish, inner bark 

yellowish brown. Sapwood yellowish brown. Leaves broadly elliptic to obovate, 5.5-10 x 

2.5-7 cm, chartaceous, without conspicuous dark gland-dots on the blade; base cuneate, 

margin distinctly densely short-toothed, apex acute to acuminate to cuspidate; lateral veins 

6~ pairs, slightly raised on lower side, obscure or distinct above, forming 2-several series 

of angular loops between midrib and margin; intercostal veins indistinct to slightly distinct 

on both sides; stalk 5-8 mm long; stipules 1-1.5 cm long. Inflorescences to 3.5 cm long, 

(di-)trichotomously branched; peduncle 1-1.8 cm long. Flowers 4-5 mm long, shortly 

stalked; calyx-lobes 6, c. 4 mm long, shortly white-hairy on inner side; petals sagittate, to 5 

mm long; filaments filiform, 3-5 mm long; disc cup-shaped, divided to halfway into 

deltoid lobes; ovary I-celled with 12 ovules, style filiform, stigma capitate. Fruits oblongellipsoid, 

8-10 x 4-6 mm, I-seeded. Seeds oblong-ellipsoid, 2-3 mm long. 

Vernacular name. Sabah--kemuning hutan (Dusun). 

329


Distribution. Borneo, Philippines, New Guinea. In Sabah and Sarawak, common throughout. 


Ecology. Primary mixed dipterocarp forest, lowlands to about 1000 m, and secondary forest 

on sandy soils. 

2. Carallia brachiata (Lour.) Merr. 

(Latin, brachiatus = having decussate branches; the inflorescence) 

Philip. J. Sc. 15 (1919) 249; Masamune I.e. 515; Burgess I.e. 432; Ashton I.e. 356; Kochummen I.e. 

313; Whitmore, Tantra & Sutisna I.e. 296. Basionym: Diatoma braehiata Lour., Fl. Coch. (1790) 

296. Type: Loureiro, s.n., "Habitat in sylvis Cochinchinae" (Merrill, Comm. Lour., 1935, 281, states 

that this specimen cannot be found in the BM). Synonyms: Symmetria obovata Blume I.e. (1826-27) 

1131; C. eelebiea Blume I.e. (1849) 131, MerrillI.e. (1921) 421; C. eonfinis Blume I.e. (1849) 129, 

Merrilll.e. (1921) 421; C. euspidata Blume I.e. (1849) 129, MerrillI.e.(1921) 421; C. lueida Roxb., 

Corom. Pl. 3 (1811) 8, Merrilll.e. (1921) 421; C. multiflora Blume I.e. (1849) 131; c. timorensis 

Blume I.e. (1849) 128; C. eerisopsifolia Miq., Analecta Pt. 3 (1852) 8; C. floribunda Miq., F1. Ind. 

Bat. 1, 1 (1858) 1088; C. ealycina Benth., J. Linn. Soc. 3 (1859) 75; Stalagmites lamponga Miq., 

Sumatra (1861) 496; Gareinia eymulosa Miq., Ann. Mus. Bot. Lugd. Bat. 1 (1864) 208; C. arguta 

Koord. & Valeton, Bijdr. Booms. Java 4 (1896) 301; C. seorteehinii King, J. As. Soc. Beng. 66, 2 

(1897) 319; C. spinulosa Ridl., J. Str. Br. R. As. Soc. 82 (1920) 184; C. euprea Ridl., Kew Bull. 

(1938) 282; C. viridifolia Ridl. I.e. (1938) 282. 

Medium-sized to large tree to 35 m tall, 70 cm diameter, sometimes buttressed and sometimes 

with stilt-roots. Bark smooth or scaly-Ienticellate, brownish; inner bark yellowish 

brown. Sapwood pale yellowish to brownish. Leaves elliptic to obovate, 5-12.5 x 2-7.5 

cm, chartaceous, with conspicuous dark gland-dots on the lower side when dry; base 

cuneate to obtuse, margin shortly toothed to sub entire, apex acuminate to cuspidate; lateral 

veins 8-12 pairs, faint to slightly raised on both sides; intercostal veins more or less 

parallel to lateral veins, indistinct on both sides; stalk 1.5-2.5 cm long; stipules 0.8-1.2 

cm long. Inflorescence a cyme to 2.5-4 cm long, (di-)trichotomously branched, peduncle 

1-2.5 cm long. Flowers 2.5-3 mm long, shortly stalked; calyx-lobes (4-)6-8, 1-1. 5 mm 

long, glabruus on inner side; petals suborbicular, to 1.5-2 mm long; filaments filiform, 2-3 

mm long; disc cup-shaped, divided to halfway into deltoid lobes; ovary 5-8-celled with 2 

ovules per cell, style filiform, stigma lobed. Fruits globose, to 7 mm across, I-seeded. 

Seeds globose, c. 2 mm across. 

Vernacular names. Sarawak---rabong, radipah (Melanau Matu and Oya). 

Distribution. Widely distributed from Madagascar, Sri Lanka, India, SE Asia and Malesia 

to N Australia. In Sabah and Sarawak common throughout. Also in Brunei and Kalimantan. 

Ecology. Primary and secondary forests, lowlands to 1800 m; on infertile organic soils in 

mixed dipterocarp forest, heath forest, and on the margins of freshwater swamp forests. 

Uses. The wood is good for furniture and other interior finishings such as parquet flooring 

and panelling, as it is hard and has an attractive oak-like figure; it is, however, difficult to 

season. 

330


Fig. 2. Carallia bomeensis. Flowering leafy twig. (From SAN 103017.) 

331

TREE FLORA OF SABAHAND SARAWAKVOL.1 (1995) 

3. Carallia coriifolia Ridl. 

(Latin, coriaceus = leathery, -folius = -leaved) 

I.e. (1938) 283; Ding Hou I.e. 483; Ashton I.e. 357; Whitmore, Tantra & Sutisna I.e. 296. Type: 

Haviland 1797, Sarawak, Kuching (holotype K; isotypes L, SAR). 

Small tree to 10 m tall. Bark flaky, dark grey; inner bark yellowish. Sapwood yellowish. 

Leaves ovate to oblong or elliptic, 7-17 x 3-6 cm, chartaceous, without conspicuous dark 

gland-dots on the lower side when dry; base cuneate or rarely rounded, margin faintly 

toothed to entire, apex acuminate; lateral veins 8-12 pairs, slightly raised on both sides; 

intercostal veins slightly prominent on upper side, visible on lower side; stalk 0.5-1 cm 

long; stipules 1.5-2.5 cm long. Inflorescence a cyme to 1.5 cm long with alternate 

branches reduced, peduncle nil or inconspicuous. Flowers 4-5 mm long, sessile; calyxlobes 

5-6, 1-1.5 mm long, glabrous on inner side; petals sagittate, 2.5-3 mm long; 

filaments filiform, 2-2.5 mm long; disc cup-shaped, divided to halfway into deltoid lobes; 

ovary 1-celled with 10-12 ovules, style filiform, stigma capitate. Fruits ellipsoid, c. 12 mm 

long, 7 mm wide, I-seeded. Seeds oblong-ellipsoid, 2-2.5 mm long. 

Distribution. Endemic to Borneo. In Sabah recorded from the Penampang and Beaufort 

districts, in Sarawak in Kuching and Serian districts. Also in Kalimantan. 

Ecology. Mixed dipterocarp forest to 1000 m. 

4. Carallia sp. 1 

Small tree to 15 m tall, 30 cm diameter. Bark smooth, dark grey-brown; inner bark brown. 

Sapwood pale yellowish. Leaves broadly elliptic to obovate, 6-7 x 4-8 cm, chartaceous to 

thinly coriaceous, with conspicuous dark gland-dots on the lower side when dry; base 

cuneate, margin finely toothed, apex acuminate to cuspidate; lateral veins 6-12 pairs, 

raised on both sides, forming 2-several series of angular loops between midrib and 

margin; intercostal veins net-like, indistinct on upper side, slightly distinct on lower side; 

stalk 0.5-1 cm long; stipules 1-2 cm long. Inflorescence a dense cyme to 2.5 cm long. 

Flowers 2-2.5 mm long; calyx-lobes 5-6, c. 1.5 mm long, glabrous on inner side; petals 

sagittate, to 2 mm long; filaments filiform, 2-2.5 mm long; disc cup-shaped, divided to 

halfway into rounded lobes; ovary 5-celled with 2 ovules per cell, style filiform, stigma 

capitate. Fruits globose, to 6 mm across, I-seeded. Seeds oblong-ellipsoid, 2-2.5 mm long. 

Vernacular names. Sabah---meransi (Brunei Malay), merawai, tikolod, yulu tambang 

(Dusun). 

Distribution. Known so far from Sabah, where it is apparently common (all districts) and 

often confused with C. borneensis, and apparently also present in Sarawak (4th Division at 

Niah National Park, and 7th Division at UIu Belaga). Also in Brunei (Niga NN 47, Belait). 

Sabah collections include SAN 124068, SAN 85908, Nooteboom 1306 (from Tambunan), 

SAN 63504 (Sandakan), and SAN 89461 (Ranau). The Sarawak collections, which have 

entire leaf margins, are S. 43802 (UIu Belaga) and S. 40118 (Niah). 

332


Ecology. Lowland secondary forest and along rivers, and mixed dipterocarp forest to c. 600 

m. 

5. CaraIIia sp. 2 

Small tree to 25 m tall, 30 cm diameter. Bark slightly fissured, brown; inner bark brown. 

Sapwood pale pinkish brown. Leaves oblong-elliptic to slightly ob ovate, 2-7 x 2-4 cm, 

chartaceous to thinly coriaceous, with conspicuous dark gland-dots on the lower side when 

dry; base cuneate to obtuse, margin entire, apex obtuse, slightly cuspidate; lateral veins 6- 

10 pairs, raised on both sides; intercostal veins more or less parallel to lateral veins, 

obscure above, slightly raised below; stalk 0.5-1 cm long; stipules 0.5-0.7 cm long. 

Inflorescence a cyme to 1.4 cm long, peduncle to 1 cm long. Flowers known only as young 

buds, sessile. Fruits not known. 

Vernacular name. Sarawak--rabong (lban). 

Distribution. Recorded only from Sarawak: S 39097 (Niah, Ulu Sungai Sah) and S 43023 

(Limbang, Ulu Medamit). 

Ecology. Lowland mixed dipterocarp forest on sandy clay. 


Small tree to 10 m tall, 15 cm diameter. Leaves broadly elliptic to obovate, 5-9 x 4-5 cm, 

stiffly coriaceous, without conspicuous dark gland-dots on the lower side when dry; base 

cuneate, margin coarsely saw-toothed, apex acute; lateral veins 8-10 pairs, raised on both 

sides, forming 2-several series of angular loops between midrib and margin; intercostal 

veins prominent on both sides; stalk 0.5-0.8 cm long; stipules 0.5-1 cm long. Inflorescence a 

cyme, to 2 cm long. Flowers not known. Fruits not known. 

Distribution. Recorded only from Sarawak at the Nyabau Catchment area, Bintulu, Miri 

(S 24594). 

Ecology. Lowland mixed dipterocarp forest on yellow red sandy humuIt ultisols, at about 

150m. 


Shrub or small tree to 6 m tall, 5 cm diameter. Leaves broadly ob ovate, 8-11 x 4-7 cm, 

stiffly coriaceous, with conspicuous dark gland-dots on the lower side when dry; base 

cuneate, margin entire, apex obtuse and very slightly cuspidate to rounded; lateral veins 8- 

10 pairs, raised on both sides; intercostal veins prominent on both sides; stalk 0.7-1 cm 

long; stipules c. 1 cm long. Inflorescence a cyme to 1.8 cm long, peduncle to 1 cm long. 

Flowers 3-4 mm long, sessile; calyx-lobes 5, c. 2 mm long, glabrous on inner side; petals 

sagittate, to 2.5 mm long; filaments filiform, to 2 mm long; disc cup-shaped, divided to 

333


halfway into deltoid lobes; ovary 5-celled with 2 ovules per cell, style filiform, stigma 

capitate. Fruits not known. 

Distribution. Known only from Bukit Tawai, Telupid in Sabah (SAN 39321), on ultramafic 

soil. 

Ecology. Secondary forest. 

8. Carallia sp. 5 

Shrub or small tree 1-2 m tall. Leaves oblong-elliptic to obovate, to 2.7 x 2.4 cm, thinly 

coriaceous, without conspicuous dark gland-dots on the lower side when dry; base cuneate, 

margin faintly toothed, apex acute; lateral veins 4-6 pairs, slightly raised on both sides; 

intercostal veins inconspicuous on upper side, slightly prominent on lower side; stalk c. 0.5 

cm long; stipules 0.5-1 cm long. Inflorescences to 1.2 cm long, with a solitary flower. 

Flowers (buds) with 4 calyx-lobes. Fruits (young) ellipsoid, c. 4 mm long. 

Distribution. Recorded only from Sarawak at the summit of Batu Lawi in the Kelabit 

Highlands area (Nooteboom & Chai 2259). 

Ecology. Forest on sandstone at about 2000 m. 

3. CERIOPS Am. 

(Greek, ceras = horn, ops = obscure; alluding to the fruit 

from which the hypocotyl has yet to elongate) 

Arm. Mag. Nat. Hist. 1 (1838) 363; Merrilll.e. (1921) 420; Masamune I.e. 516; Browne I.e. 302; 

Ding Hou I.e. 468; Burgess I.e. 432; Ashton I.e. 357; Kochummen I.e. 315; Whitmore, Tantra & 


Small or medium-sized trees with stilt-roots to 1 m high; pneumatophores prominent. 

Leaves broadly elliptic to obovate, coriaceous, sometimes black-dotted on lower side; 

margin entire; stipules lanceolate. Inflorescence a condensed cyme, shortly stalked or 

subsessile, (2-)4-many-flowered. Flowers with calyx deeply 5-6-lobed; petals 5-6, each 

embracing 2 stamens; ovary semi-inferior, 3-celled, ovules 2 per cell, style simple, stigma 

simple or obscurely 2-3-lobed. Fruits ovoid, almost entirely superior. Seed germination 

viviparous; hypocotyl prominently ridged or grooved. 

Distribution. 2 species, tropical east Africa to Malesia, N Australia and the Pacific. Both 


Ecology. At the mouths of estuaries and bays, and behind the sea beach. 

Uses. The bark is a source of tannin and a dye for the batik industry, and is used in the 

manufacture of an alcoholic drink. The wood is used for firewood and charcoal. 

334


Key to Ceriops species 

Petals fringed with many long narrow processes at the apex; anthers as long as or longer 

than the filament. Fruit with erect to ascending calyx-lobes; hypocotyl 9-15 cm long when 

fully extended at fruit-fall ............................................................................. 1. C. decandra 

Petals with 3 short appendages at the apex; anthers less than half the length of the filament. 

Fruit with reflexed calyx-lobes; hypocotyl 15-35 cm long when fully extended at fruit 

.2. C. tagal 

1. Ceriops decandra (Griff.) Ding HOll 

(Greek, deca = ten, -andrus = male; with ten stamens in the flower) 

I.e. 471; Burgess I.e. 432; Ashton I.e. 357; Kochununen I.e. 315; Whitmore, Tantra & Sutisna I.e. 

296. Basionym: Bruguiera deeandra Griff. I.e. (1836) 10. Type: GrifJith, s.n., Burma (K). 

Synonyms: C. roxburghiana Am. I.e. 364, Merrilll.e. (1921) 420, Masamune I.e. 516, Browne I.e. 

303; C. zippeliana Blume I.e. (1849) 143; Rhizophora glomerulata Zipp. ex Blume I.e. pro syn. R. 

deeandra Roxb. ex Griff. I.e. (1854) 663. 

Shrub to small tree to 15 m tall, 30 cm diameter. Bark smooth to papery-flaky to 

lenticellate, pale brown; inner bark red. Sapwood pale yellowish brown. Leaves obovate to 

elliptic-oblong, 4.5-10 x 2.5-6 cm; base cuneate, margin slightly recurved when dry, apex 

obtuse, rounded or notched; lateral veins 9-10 pairs, obscure, joining near margin to form 

a submarginal vein; stalk 1-2.5 cm long; stipules 1.5-2.5 cm long. Flower petals white, 

fringed with many long narrow processes at apex; stamens 1 mm long, anthers ovoid, more 

than half the length of the filaments; style c. 1 mm long, stigma capitate. Fruits ovoidconical, 

c. 1.8 cm long, calyx-lobes erect to ascending; hypocotyl club-shaped, 9-15 cm 

long, sharply ridged. 

Vernacular names. Sabah-tengar (Brunei Malay). Sarawak---bakau lali, tengar tikus 

(Malay). 

Distribution. India, Indo-China, Peninsular Malaysia, Banka, Java, Borneo, Philippines, 

Celebes, Moluccas and New Guinea. In Sabah recorded from Kudat (Balambangan Island 

also), Sandakan and Lahad Datu; in Sarawak from Selabat to the Rejang delta. Also in 

Kalimantan 

Ecology. Edges of mangrove swamps and along tidal creeks. 

2. Ceriops tagal (Pers.) c.B. Rob. 

(a Philippine plant name) 

Fig. 3. 

Philip.1. Sc. 3 (1908) 306; MerrillI.e. (1921) 420; Masamune I.e. 516; Browne I.e. 302; Ding Hou 

I.e. 469; Burgess I.e. 432; Ashton I.e. 360; Kochununen I.e. 316; Whitmore, Tantra & Sutisna I.e. 

297. Basionym: Rhizophora tagal Pers., Mem. Soc. Linn. Paris 3 (1824) 138. Type: Perrottet, 

Philippines, Zamboanga (No specimen seen by Merrill, see En. Phi1ip. FI. PI. 3, 1923, 144). 

Synonyms: Rhizophora timorensis DC., Prod. 3 (1828) 32; C. eandolleana Am. I.e. 364; c. 

paueijlora Benth. in Hooker /, Lond. 1. Bot. 2 (1843) 218; C. fosteniana Blume I.e. (1849) 143; C. 

335


boviniana Tul., Ann. Se. Nat. 4,6 (1856) 112; C. lucida Miq., Sumatra (1861) 325; c. timoriensis 

Domin, Bibl. Bot. 8 (1928) 444. 

Small to medium-sized tree, to 25 m, 45 cm diameter, with small stilt-roots. Bark smooth 

to flaky or dippled, grey-brown; inner bark pale yellowish white to pinkish. Sapwood 

yellowish white to pale brown. Leaves obovate to obovate-oblong or elliptic-oblong, 5-12 x 

2-7 cm; base cuneate, margin slightly recurved when dry, apex obtuse to slightly notched; 

lateral veins 9-12 pairs, faint, joining at margin to form a submarginal vein. Flower petals 

white, with 3 club-shaped appendages at the apex; stamens 3-5 mm long, anthers 

sagittate, less than half the length of the filaments; style c. 2 mm long, simple. Fruits 

ovoid, 1.5-2.5 cm long, calyx-lobes re curved; hypocotyl club-shaped, 15-35 cm long, 

sharply ridged. 

Vernacular names. Sabah--tagal (Suluk), tangal (Suluk), tanug (Bajau) tengar (Brunei 

Malay), tenug (Ilanun). Sarawak-tengar samak (Malay). 

Distribution. East Africa, Madagascar, Seychelles, Sri Lanka, India, Indo-China, Taiwan, 

throughout Malesia to Micronesia and Australia. In Sabah, recorded from the southwest 

part around Sipitang to Kota Kinabalu, and the Sandakan and Lahad Datu mangroves. In 

Sarawak, recorded from the Sarawak and Rejang deltas. Also in Brunei and Kalimantan. 

Ecology. In brackish water environments, on relatively well-drained sites within the reach 

of tides in the inner fringes of mangroves. It is much more common than the preceeding 

species. 

Uses. A most durable mangrove wood. The bark is used for tanning fishing lines, nets and 

sails. The trunk is used in house-building. It is also used for charcoal and known as an 

excellent firewood. 

4. GYNOTROCHES Blume 

(Greek, gune = woman, trochos = wheel; alluding to the shape of the stigma) 

Bijdr. (1825) 218; Merrilll.e. (1921) 420; Masamune I.e. 516; Ding Hou I.e. 488; Ashton I.e. 362; 

Koehummen I.e. 317; Whitmore, Tantra & Sutisna I.e. 297. 

Small trees; young branches hollow. Leaves with entire to faintly toothed margin; stipules 

lanceolate, margins overlapping. Flowers very small, solitary on short stalks or fascicled 

in dense axillary clusters, stalks jointed; calyx deeply 4-5-lobed; petals 4-5, obovate to 

elliptic; stamens 8-10, free; ovary superior, slightly ridged, 4-6-celled, ovules 3-8 per cell, 

style simple, stigma a 4-8-lobed discoid structure or slender with spreading lobes. Fruit a 

berry, globose to oblong, few- to many-seeded. 

Distribution. One species, from Upper Tenasserim, throughout Malesia, to the Western 

Pacific. 

Taxonomy. Although the wider concept of Ding Hou I.c. is followed here, there is some 

suggestion (Juncosa & Tobe, Ann. Missouri Bot. Gard. 75 (1988) 1410) that the material 

336


u~ 

A 

li 

3cm 

:,. 

q, H'l 

Fig. 3. Ceriops tagal. A, flowering leafy twig; B, germinating fruit. (From SAN 84104.) 

337


20m[ t 

Fig. 4. Gynotroches axillaris. Fruiting leafy twig. (From SAN 55769.) 

338


from Sarawak may represent at least two ecotypes which probably constitute distinct 

species. 

Gynotroches axillaris Blume 

(Latin, axillaris = axillary; the flowers) 

Fig. 4. 

I.e. (1825) 219; Merrilll.e. (1921) 420; Masamune I.e. 516; Ding Hou I.e. 488; Ashton I.e. 362; 

Koehummen I.e. 317; Whitmore, Tantra & Sutisna I.e. 297. Type: Blume, s.n., Java (L). Synonyms: 

Dryptopetalum eoriaeeum Am. I.e. 373; G. dryptopetalum Blume I.e. (1849) 127; G. micrantha 

Blume I.e. (1849) 127, Masamune I.e. 517; G. retieulata A. Gray in Wilkes, Bot. U.S. Expl. Exped. 

1 (1854) 607; G. parvifolia Merr., Publ. Gov. Lab. Philip. 35 (1905) 46; G. puberula Merr., Philip. 

J. Se., Bot. 10 (1915) 333; G. laneeolata Merr., Philip. J. Se., Bot. 11 (1916) 21. 

Small or medium-sized tree, to 35 m tall, 70 cm diameter. Bark smooth, often hoopmarked, 

grey to blackish; inner bark yellowish to reddish brown, fibrous, finely and 

regularly invaginating into the wood. Sapwood yellowish brown to pale reddish brown. 

Leaves ovate to elliptic-oblong or lanceolate, 5-15 x 3-7.5 cm, usually coriaceous, 

glabrous or (rarely) pubescent on the midrib and lateral veins below; base cuneate, apex 

acute to acuminate; midrib flat on upper side, raised on lower side; lateral veins 8-12 pairs, 

flat on upper side, raised on lower side, forming distinct loops just behind the margin; 

intercostal veins net-like, flat on upper side, distinctly prominent on lower side; stalks 0.5- 

1. 5 cm long; stipules to 1. 5 cm long. Flowers greenish white, 1-16 in each leaf axil, 

bisexual but sometimes male by abortion; calyx-lobes with tufts of hairs at apex; petals 

clawed, divided into appendages towards the apex; stamens 1-2 mm long; disc cup-shaped 

or nearly flat, 8-1O-10bed; ovary ovoid; style to 2 mm long. Fruits usually globose, c. 3 mm 

across, green ripening red to shiny black, persistent calyx-lobes erect or reflexed. 

Vernacular names. Sabah--bulu bulu (Malay), kandis batu (Malay), kupi kupi (Brunei 

Malay), pahau pahau (Murut). Sarawak-kelalud (Melanau), sawar bubu (Iban). Bruneikerakas 

payau (Kedayan), kerakas payoh (Dusun), sawar bubu (Jban). 

Distribution. Burma, Thailand, Malesia except the Lesser Sunda Islands, to Melanesia and 

Micronesia. In Sabah and Sarawak, in all districts. Also recorded for Brunei and 

Kalimantan. 

Ecology. Lowlands to about 2200 m, in marshy places especially along riv.ers, and also 

well-drained sandy sites. Occasionally also in secondary forests. 

5. KANDELIA Wight & Am. 

(from a Malabar (Indian) plant name) 

Prod. (1834) 310; Merrilll.e. (1921) 421; Masamune I.e. 517; Ding Hou I.e. 473; Ashton I.e. 363; 

Koehummen I.e. 319. . 

Small trees. Leaves with linear stipules. Inflorescence a peduncled cyme bearing 4-9 

flowers. Flowers with calyx deeply 5(-6)-lobed; petals 5(-6), bi/obed, with a long bristle 

339


~~ 

:tt 

!\ 

:~:I 

: :~! 

';" 

B 

Fig. 5. Kandelia candel. A, nowering leafy twig; B, germinating fruit. CA from S. 30640, B from SAN 

102906.) 

340


between lobes, each lobe laciniate to fimbriate; stamens many, free, of unequal length, 

exserted, anthers many-celled; ovary I-celled, ovules 6, apically attached, style simple. 

Fruit ovoid. Seed solitary; germination viviparous; hypocotyl cylindric to spindle-shaped. 

Distribution. One species; India, Indo-China to S China and western Malesia (NE 

Sumatra, Peninsular Malaysia, Nand W Borneo). 

Kandelia candel (L.) Druce 

(a Malabar plant name) 

Fig. 5. 

Rep. Bot. Exch. Club. Br. IsI. 3 (1914) 420; Merrilll.e. (1921) 421; Masamune I.e 517; Ding Hou 

I.e. 473; Ashton I.e. 363; Kochum.men I.e. 319. Basionym: Rhizophora eandel 1., Sp. PI. (1753) 443. 

Shrub or small tree, to 10 m tall; trunk with swollen conical base but no buttresses. Bark 

flaky, lenticellate, greyish to reddish brown; inner bark thin, pink. Sapwood pale yellowish 

brown. Leaves narrowly oblong-elliptic to ooovate-oblong, 6-13 x 2-6 cm, coriaceous; 

base cuneate, margin frequently recurved, apex acute-blunt to rounded; lateral veins 6-7 

pairs, inconspicuous on both sides; stalks 1-1. 5 cm long; stipules 0.5-1. 5 cm long. 

Flowers c. 2 cm long, white; calyx 5-6-lobed, reflexed after anthesis; petals c. 14 mm 

long; stamens with small narrow anthers on slender filaments; style filiform, stigmas 3. 

Fruits ovoid, 1-2.5 cm long, I-seeded; hypocotyl to 40 cm long, terete, smooth. 

Vernacular names. Sabah--berus berus (Malay), beus (Bajau, Malay), linggayong, 

linggayong laut (Brunei Malay). Sarawak-bakau aleh aleh (Malay). 

Distribution. India, Burma, Thailand, Indo-China, China, Ryukyus, S Japan, NE Sumatra, 

Peninsular Malaysia, Wand N Borneo. In Sabah, recorded so far from the Papar and 

Sandakan areas; in Sarawak from the Kuching, Sajingkat and Kelepu mangroves. Also in 

Brunei and W Kalimantan 

Ecology. Scattered along the banks of brackish water (tidal) rivers and channels in 

mangrove areas. 

6. PELLACAL YX Korth. 

(Greek, pella = hide, kalux = calyx; the hair-covered calyx) 

merbuloh (lban, Malay; Sarawak), sawar (sambar) bubu (lban; Sarawak) 

Tijd. Nat. Gesh. Phys. 3 (1836) 20; Merrilll.e. (1921) 422; Masamune I.e. 517; Ding Hou I.e. 490; 

Ashton I.e. 363; Kochummen I.e. 320; Whitmore, Tantra & Sutisna I.e. 297. Synonyms: Plaesiantha 

Hook. f in Benth. & Hooker j, Gen. PI. 1 (1865) 861; Craterianthus Valeton ex Heyne, Nutt. PI. 

Indon. 4 (1917) 196. 

Small to medium-sized trees; young branches hollow. Bark invaginating finely and 

regularly into the wood. Leaves oblong to ovate; sparsely to densely pubescent with stellate 

341


\4'1 

Fig. 6. Pellacalyx axillaris. Fruiting leafy twig. (From SAN 75981.) 

342


or simple hairs (rarely a mixture of the two), sometimes glabrescent to glabrous; margin 

entire or obscurely to distinctly serrulate; stipules with margins not overlapping. Flowers 

fascicled, 2-8 in a: leaf axil, or on short dense inflorescences; calyx tubular, lobes (3-)4- 

5(-6), tube hairy inside at the lower part; petals 4-5, inserted on the margin of the calyxtube, 

densely short-hairy outside; stamens twice as many as petals, inserted on the margin 

of the calyx-tube, filaments connate at the base or free, in 1 or (rarely) 2 series, unequal, 

anthers sub orbicular; ovary inferior, 9-1O(-12)-celled, ovules 8-25 per cell, style columnar 

and usually hairy, stigma discoid or capitate, obscurely 8-1O-lobed. Fruit a berry, subglobose. 

Seeds few to many. 

Distribution. 7 or 8 species; Burma and S China to Sumatra, Borneo, Philippines, and 

Celebes. In Sabah and Sarawak, 4 species. 

Ecology. In Sabah and Sarawak in mixed dipterocarp forest to 1300 m. 

Key to Pellacalyx species 

1. Leaf lateral veins 5-7 pairs. Calyx 4-merous, the outside covered in very minute hairs 

not easily visible to the unaided eye (appearing glabrous) ............................. 3. P. lobbii 

Leaf lateral veins 9-12 pairs. Calyx 5-merous, the outside conspicuously hairy ............ 2 

2. Lower leaf surface with a mixture of simple hairs and stellate hairs, both types 

similarly abundant. Inside of calyx-tube with scanty short straight hairs at the base. 

Stamens in two series ..................................................................... 4. P. symphiodiscus 

Lower leaf surface with predominantly stellate scales (mixed with a few simple hairs) 

or predominantly simple hairs (mixed with a few stellate scales). Inside of calyx-tube 

with either a band of woolly hairs and scanty short straight hairs below the band, or a 

zone of villous hairs at the base. Stamens in one series ................................................ 3 

3. Lower leaf surface with predominantly stellate scales mixed with a few simple hairs. 

Calyx-lobes reflexed at floral maturity; inside of calyx-tube with a band of woolly hairs 

and scanty short straight hairs below the band. Petal tips fringed with linear segments 

attenuating into fine processes .................................................................. 1. P. axillaris 

Lower leaf surface with predominantly simple hairs mixed with a few stellate scales. 

Calyx-lobes ascending at floral maturity; inside of calyx-tube with a zone of villous 

hairs near the base. Petal tips with very fine hair-like processes ............... 2. P. cristatus 

1. Pellacalyx axillaris Korth. Fig. 6. 

(Latin, axillaris = axillary; the inflorescences) 

I.e. 20, t. 2; Ding Hou I.e. 493; Ashton I.e. 364; Kochummen I.e. 320; Whitmore, Tantra & Sutisna 

I.e. 297. Type: Korthals, s.n., Sumatra (L). 

Small to medium-sized tree, to 25 m tall, 60 cm diameter; buttresses if present to 60 cm 

high. Bark pale reddish brown, finely cracked to rough-fissured; inner bark pinkish yellow. 

Sapwood yellow. Leaves oblong to obovate-oblong, 8-17 x 3.5-7 cm, lower side densely 

or sparsely covered with stellate scales mixed with a few simple hairs; base broadly cuneate 

to rounded, margin entire or toothed (with a tuft of short hairs on each tooth), apex acute to 

343


acuminate; lateral veins 9-12 pairs, flat on upper side, raised on lower side; stalk 5-8 mm 

long; stipule 1-1.5 cm long. Flower calyx conspicuously hairy outside, the tube 5-6 mm 

long, inside with a few short hairs at the base and a band of woolly hairs above these, 

lobes 5, 2-2.5 mm long, rejlexed at anthesis; petals 1.5-2.5 mm long, the apices fringed 

with linear segments attenuating into fine processes; stamens in one series; style 1.5-2 mm 

long. Fruits ovoid or subglobose, c. 10 mm across. 

Vernacular name. Sarawak--danguh (Bidayuh). 

Distribution. Sumatra, Peninsular Malaysia, Borneo and Mindanao (Philippines). In Sabah 

recorded from Kudat, Beaufort and Telupid. In Sarawak recorded from the Padawan area 

(1st Div.) and Sg. Beria in the Kapit District (7th Div.). Also in Brunei and Kalimantan. 

Ecology. Common where it occurs in lowland primary mixed dipterocarp forest and secondary 

forest, usually in valleys and moist sites. 

2. Pellacalyx cristatus Hems!. 

(Latin, cristatus = crested; the persistent calyx crowning the fruit) 

in Hookerf, le. PI. 16 (1886) sub t. 1546; Masamune I.e. 517; Ding Hou I.e. 493; Ashton I.e. 364; 

Whitrnore, Tantra & Sutisna I.e. 297. Type: Beeeari PB 1258, Sarawak (G, K). 

Small tree to 10 m tall, 10 cm diameter. Bark yellowish brown, smooth; inner bark pale 

brown. Sapwood pale yellow. Leaves narrowly oblong to ovate-oblong, 10-17 x 3-5 cm, 

lower side densely or sparsely covered with simple hairs mixed with a few stetlate scales, or 

glabrescent; base obtuse to rounded, margin toothed (with a tuft of short hairs on each 

tooth), apex acuminate; lateral veins 9-12 pairs, flat on upper side, raised on lower side; 

stalk 7-10 mm long; stipule 1-1.5 cm long. Flower calyx conspicuously hairy outside, the 

tube 9-10 mm long, inside with a few short hairs at the base and a zone of villous hairs 

above these, lobes 5, 1.5-2 mm long, ascending at anthesis; petals 2.5-3.5 mm long, the 

apices fringed with fine hair-like processes; stamens in one series; style 2.5-3 mm long. 

Fruits subglobose, to 10 mm across. 

Distribution. Endemic to Borneo. In Sabah, recorded from Nabawan northwards to Mt. 

Kinabalu; in Sarawak from the Kuching area, Belaga and Mt. Api, Mulu National Park. 

Also in Kalimantan. 

Ecology. Hills and mountain ridges to c. 900 m. 

3. Pellacalyx lobbii (Hook./) Schimp. 

(Thomas Lobb, 1820-1894, plant collector) 

in EngI. & Prantl, Pfl. Fam. 3,7 (1893) 54; Masamune I.e. 517; Ding Hou I.e. 491; Ashton I.e. 365; 

Whitmore, Tantra & Sutisna I.e. 297. Basionym: Plaesiantha lobbii Hook. f in Benth. & Hooker f, 

Gen. PI. 1 (1865) 145. Type: Lobb, s.n., Brunei (K). 

Medium-sized tree, to 45 m tall, 65 cm diameter; stilt-roots at times present. Bark medium 

brown, cracking-fissured; inner bark pale brown. Sapwood yellow. Leaves obovate to 

obovate-oblong, 7-17 x 2-6 cm, lower side sparsely covered with stellate hairs; base 

344


cuneate, margin entire or faintly toothed (glabrous on each tooth), apex acute to abruptly 

acuminate; lateral veins 5-7 pairs, flat on upper side, raised on lower side; stalk 5-8 mm 

long; stipule 0.5-1 cm long. Flower calyx very minutely hairy outside (appearing glabrous 

to the unaided eye), the tube 6-7 mm long, inside with a few short hairs at the base, lobes 

4,2.5-3 mm long, ascending at anthesis; petals 1.5-2 mm long, the apices finely serrulate; 

stamens in one series; style 1-1. 5 mm long. Fruits subglobose, to 8 mm across. 

Vernacular names. Sarawak-paserujan gunugo, rasu (Iban). 

Distribution. Sumatra and Borneo. In Sabah and Sarawak, in most districts. Also in 


Ecology. Primary and secondary forests. 

4. PelIacalyx symphiodiscus Stapf 

(Greek, sumphuein = fused, discus = disc; of fused parts forming the floral disc) 

Kew Bull. (1898) 224; Merrilll.c. (1921) 422; Masamune I.c. 517; Ding Hou I.c. 492; Ashton I.c. 

365; Whitmore, Tantra & Sutisna I.c. 297. Type: Haviland 2206, Sarawak (K). 

Small to medium-sized tree, to 25 m tall, 40 cm diameter. Bark pale brown, smooth; inner 

bark pale brown. Sapwood yellow. Leaves ovate-oblong to obovate-oblong, 9-19 x 3.5-7 

cm, lower side densely or sparsely covered with a mixture of stellate hairs and simple hairs 

(both types similarly abundant); base acute or rounded, margin toothed (with a tuft of short 

hairs on each tooth), apex acute to shortly acuminate; lateral veins 9-12 pairs, flat on 

upper side, raised on lower side; stalk 6-10 mm long; stipule 0.8-1.2 cm long. Flower 

calyx conspicuously hairy outside, the tube 4-5 mm long, inside with a few short hairs at 

the base, lobes 5, 1.5-2 mm long, reflexed at anthesis; petals 1.5-2.5 mm long, the apices 

fringed with fine hair-like processes; stamens in two series; style 1.5-2 mm long. Fruits 

sub globose, to 9 mm across. 

Distribution. Endemic to Borneo. In Sabah, known only from the Penampang and Lamag 

districts; in Sarawak from the Kuching, Selang and Pengkalan Ampat areas. Also in 


Ecology. Mixed dipterocarp forest to c. 650 m. 

7. RHIZOPHORAL. 

(Greek, rhiza = root, pherein = bearing; the stilt-roots) 

bakau (Malay) 

Gen. PI. ed. 5 (1754) 202; Merrilll.c. (1921) 420; Masamune I.c. 517; Browne I.c. 296; Ding Hou 

I.c. 448; Burgess I.c. 433; Tomlinson & Womersley, Contrib. Herb. Austr. 19 (1976) 1; Ashton I.c. 

366; Kochummen I.c. 321; Whitmore, Tantra & Sutisna I.c. 298. Synonyms: Mangium Rumph. ex 

Scop., Introd. Hist. Nat. (1777) 218. 

345


Fig. 7. Rhizophora apiculata. A, flowering leafy twig; B, genninating fruit with long hypocotyl. (A 


346


Small to medium-sized trees with conspicuous branching stilt-roots but no pneumatophores. 

Leaves succulent, entire, apex acute, lower side usually black-dotted; stipules 

lanceolate, conspicuous. Inflorescence a cyme, 2-3-branched. Flowers bisexual; calyx 

leathery, deeply 4-lobed, lobes broadly ovate, ascending but reflexed in the fruit; petals 4, 

falling off early; stamens 8-12, filaments very short or hardly distinct; ovary semi-inferior, 

2-celled, ovules 2 per cell, style obscure or to 6 cm long, stigma simple or somewhat 2- 

lobed. Fruits ovoid, usually I -seeded; germination viviparous. 

Distribution. 7 species, mainly along tropical coasts and throughout Malesia. In Sabah, 3 

species and in Sarawak, 2. 

Ecology. Mangrove swamps and tidal parts of rivers; the most widespread mangrove 

genus. 

Uses. The poles are used in fishing platforms, traps and house frames. The wood is durable 

as piling timber and is also used in concrete foundations for buildings. Excellent as 

firewood and for making charcoal. The bark is also used for tanning. 

Key to Rhizophora species 

I. Inflorescences 2-flowered, shorter than the leaf-stalks, mostly in the axils of leaf-scars 

on the branches just below the current cluster of leaves. Petals glabrous save for the 

occasional presence of a few scattered short-hairs .................................... 1. R apiculata 

Inflorescences 2-16-flowered, longer than the leaf-stalks, in leafaxils within the 

current cluster ofleaves. Petals long-hairy on the margins and inner side .................... 2 

2. Leaves typically 11-18 x 5-10.5 cm. Style obscure or very short, not exceeding 1.5 

mm ...................................................................................................... 2. R mucronata 

Leaves typically 6-12 x 3-6 cm. Style filiform, 4-6 mm long ..................... 3. R stylosa 

1. Rhizophora apiculata Blume Fig. 7. 

(Latin, apiculatus = with a short abrupt point; the leaf apex) 

I.e. (1827) 91; Ding Hou I.e. 452; Burgess I.e. 433; Ashton I.e. 366; Kochummen I.e. 322; Whitrnore, 

Tantra & Sutisna I.e. 298. Type: Pee-Kandel Rheede, Hort. Mal. 6 (1686) 61, t. 34. Synonyms: R. 

eandelaria DC., Prod. 3 (1828) 32, Merrilll.e. (1921) 420, Masamune I.e. 517; R. conjugata Am. 

(non 1.) I.e. 363, Browne I.e. 298. 

Medium-sized tree to 30 m tall, 75 cm diameter. Bark pinkish grey to grey, smooth to 

shallowly fissured. Sapwood pale brown. Leaves narrowly elliptic to sublanceolate, 7-16 x 

3-7 cm, coriaceous; base cuneate, apex acute to pointed, with a short mucro; stalk 1. 5-3 

cm long; stipules 4-8 cm long. Inflorescences 2-flowered, shorter than the petioles, 

peduncle 0.5-1.; cm long, mostly in the axils of leaf-scars on the branches just behind the 

current cluster of leaves. Flowers sessile, calyx-lobes 10-14 x 6-8 mm; petals 8-11 x 1.5- 

2 mm, glabrous except for the occasional presence of a few scattered short-hairs; stamens 

347

· TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995) 

10-12, sessile; style 0.5-1 mm long. Fruits obpyriform (inve!1:ed pear-shaped), 2-2.5 cm 

long, hypocotyl to 33 cm long. 

Vernacular names. Sabah-bangkita (Malay). Sarawak-bakau minyak (Malay). 

Distribution. Tropical SE Asia, Sri Lanka, throughout Malesia to Micronesia and New 

Britain, the Solomons and New Hebrides. In Sabah, the largest stands occur between the 

Bay of Brunei and Klias in the SW, but generally common in mangroves. In Sarawak, with 

good stands at the Sarawak River Delta, the Lower Rejang and generally elsewhere. Also in 


Ecology. Dominant species of the mangroves, forming pure stands on soft recent muddy 

substrates, immediately behind the river banks or up to them, in the lower parts of major 

river deltas. 

2. Rhizophora mucronata Lam. 

(Latin, mucronatus = with an abrupt short point; the leaf apex) 

Encycl. 6 (1804) 189; Merrill/.c. (1921) 421; Masamune I.c. 517; Browne I.c. 298; Ding Hou I.c. 

453; Burgess I.c. 433; Ashton I.c. 367; Kochumrnen I.c. 322; Whitmore, Tantra & Sutisna I.c. 298. 

Type: Commerson, S.n., isle de France (Mauritius) (P). Synonyms: R. mangle Roxb. (non L.), Hort. 

Beng. (1814) 36; R. macrorrhiza Griff. I.c. (1836) 8; R. lingissima Blanco, Fl. Filip. (1837) 398. 

Medium-sized tree to 30 m tall, 210 cm diameter. Bark grey to black, roughly gridcracked. 

Sapwood pale brown. Leaves broadly elliptic to oblong, 11-18 x 5-10.5 cm, 

coriaceous; base cuneate, apex acute to slightly obtuse, with a short mucro; stalk 2.5-5.5 

cm long; stipules 5.5-8.5 cm long. Inflorescences 2-5(-12)-jlowered, longer than the 

petioles, peduncle 2.5-5.5 cm long, in leafaxils within the current cluster of leaves. 

Flowers with stalks 4-8 mm long, calyx-lobes 13-15 x 5-7 mm; petals 8-10 x 2-3 mm, 

long-hairy on the margins and inner side; stamens 8, subsessile; style 0.5-1.5 mm long. 

Fruits narrowly ovoid, 5-7 cm long, hypocotyl to 62 cm long. 

Vernacular name. Sabah and Sarawak-bakau kurap (Malay). 

Distribution. E Africa through S and SE Asia to Queensland, Melanesia and Micronesia, 

also in Tonga. In Sabah and Sarawak, common in mangrove areas. Also in Brunei and 

Kalimantan. 

Ecology. Usually in small stands or scattered groups along the lower parts of tidal rivers, 

both on clay-rich and sandy deposits. 

3. Rhizophora stylosa Griff. 

(Latin, stylosus = with a conspicuous style; the flower) 

I.c. (1854) 665; Ding Hou I.c. 456; Kochumrnen I.c. 323. Type: Griffith, s.n., Malacca, Pulo Bissar 

(K). Synonym: R. mucronata var. stylosa Schimp., Bot. Mitt. Trap. 3 (1891) 92. 

348


Small tree to 15 m tall, 25 cm diameter. Bark dark brown to black, fissured. Sapwood pale 

~'ello\\"ish brown. Leaves elliptic to sublanceolate, 6.5-12 x 3--6 cm, coriaceous; base 

cuneate, apex acute to pointed, with a short mucro; stalk 1.5-2.5 cm long; stipules 4-6 cm 

long. Inflorescences 4-8(-J6)-jlowered, longer than the petioles, peduncle 2.5-5 cm long, 

in leafaxils within the current cluster of leaves. Flowers with stalks 0.5-1 mm long, 

calyx-lobes 9-12 x 3-5 mm; petals 7-8 x 2.5-3.5 mm, long-hairy on the margins and 

inner side; stamens 8, filaments short but distinct; style 4--6 mm long. Fruits ovoid to 

obpyriform, 2.5-4 cm long, hypocotyl to 54 cm long. 

Distribution. Taiwan, Peninsular Malaysia, Java, Borneo, Philippines, Celebes, Moluccas, 

New Guinea to Melanesia, New Britain and Queensland. In Sabah, recorded only from the 

Lahad Datu area (SAN 26145) and Pulau Balambangan in the north (SAN 85589). Not 

recorded for Sarawak, Brunei and Kalimantan. 

Ecology. Exclusively along sandy shores and coral terraces facing open sea. 

349


RUTACEAE 

David T. Jones 

South Florida Natural Resources Center, 

Everglades National Park, Homestead, Florida, U.S.A. 

Hooker J, Fl. Brit. Ind. 1 (1875) 484; Kurz, For. Fl. Brit. Bunna 1 (1877) 178; Merrill, EB (1921) 

313, PEB (1929) 113; Ridley, FMP 1 (1922) 340; Craib, Fl. Siam Enum. 1 (1931) 215; Masamune, 

EPB (1942) 357; Browne, FTSB (1955) 315; Backer & BakhuizenJ, FJ 2 (1965) 94; Swingle, Citrus 

Industry 1 (1967) 190; Stone, TFM 1 (1972) 367; Anderson, CLTS (1980) 307; Perry, MPESA 

(1980) 361; Corner, WSTM 2 (1988) 656; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 304; Ng, 

MFFSS 2 (1992)493. 

Aromatic trees, shrubs, or woody climbers, or seldom herbs; twigs and branches sometimes 

armed with spines or thorns; tissues usually dotted with lysigenous oil-glands containing 

volatile essential oils. Leaves pinnate or trifoliolate or unifoliolate, less commonly simple; 

opposite, alternate or spirally arranged, rarely whorled, often conspicuously pelluciddotted; 

petioles sometimes winged, usually articulated with the blade, without stipules. 

Inflorescences terminal or axillary panicles, cymes or racemes, or flowers seldom solitary, 

rarely epiphyllous. Flowers bisexual or (in dioecious plants) unisexual, regular or rarely 

somewhat irregular; sepals 4-5, rarely 2-3, usually imbricate, free or fused basally to form 

a cup-like calyx; petals 4-5, rarely 2-3 or 6, alternate with the sepals, imbricate or valvate, 

free or fused basally, rarely united to form a tube; stamens 4-5 or 8-10 or more (to 60), 

equal or unequal, sometimes intermixed with staminodes, filaments free or fused basally, 

rarely united to form a tube; anthers 2-celled, often gland-tipped; disc surrounding base of 

ovary, intrastaminal, usually annular, nectariferous, sometimes modified into a gynophore, 

rarely lacking; ovary superior, carpels 4-5, rarely 1-3 or 6 or more, completely to partially 

fused or free except for the united styles, ovules 1-2 per locule, rarely more, placentation 

axile, stigma small or broad, sometimes lobed. Fruit a capsule, follicle, samara, drupe, 

berry or hesperidium (Citrus), dry or fleshy; hesperidia composed of jUice-jilled pulpvesicles, 

with or without oil-droplets; pericarp leathery to woody or fleshy, usually 

glandular. Seeds usually 1 per carpel, ovoid to ellipsoid or oblong, with or without endosperm, 

sometimes winged, sometimes hairy; testa firm or membranous; monoembryonic or 

polyembryonic; embryo straight or curved; cotyledons white or green, flat or folded. 

Distribution. A cosmopolitan family compnsmg approximately 160 genera and 1650 

species, distributed largely in tropical and subtropical parts of the world, with good 

representation in arid areas of the warm temperate zone (Australia and S Africa). In Sabah 

and Sarawak., 23 genera with about 75 species, of which 17 genera and 43 species are 

native trees and shrubs. The remaining taxa are scandent shrubs (15 species), or are 

cultivated (c. 17 species). The genera Fortunella (2-3 species), Limonia (1 species), and 

Triphasia Cl species) are represented only by introduced species found under cultivation 

and are excluded from this treatment. 

351


Ecology. Native species are found in a wide range of natural habitats occurring in the 

lowlands, hills, and mountains, along coastal areas, and on offshore islands. These habitats 

include primary and secondary forests (flat or undulating lands, ridges, hillsides), forest 

margins, open places and disturbed sites, mangrove forest (Merope), heath forest, swamp 

(freshwater, peat) forest, streamsides and riverbanks, and beach forest (Severinia). Some 

species show affinities for specific soil types such as ultramafic (Lunasia) and limestone 

(some Clausena, Glycosmis, Murraya and Micromelum). The majority of species are found 

below 1300 m elevation, and a few ascend to higher altitudes (Citrus and Tetractomia to 

1800 m, and Melicope to 2400 m). Thirteen tree and shrub taxa are endemic to Sabah 

and/or Sarawak, four of which are found in each of the genera Glycosmis and Melicope, 

followed by Maclurodendron (2 species), Clausena, Monanthocitrus, and Pleiospermium. 

Abundance varies among species, ranging from widespread and common to infrequent and 

rare. Merrillia is very rare in Sabah, and Burkillanthus is endangered, if not already 

extinct, in Sarawak. 

The typically small size and drab colour (white or cream to greenish or yellowish) of the 

flowers of the Rutoideae (Melicope, Tetractomia, Zanthoxylum) and Toddalioideae 

(Acronychia, Maclurodendron), and the larger, often fragrant flowers of the Aurantioideae 

(Citrus, Murraya, Pleiospermium, etc.), suggests insect pollination. The watery nectar 

secreted through stomata in the outer portions of the floral disc serves as both an attractant 

and food source. Fruit and seed dispersal is by birds (Melicope, Zanthoxylum), wind 

(Tetractomia), water (Merope), and fruit- and seed-eating animals (Citrus, Clausena, 

Glycosmis, Micromelum, Severinia, and most other Aurantioideae). 

Uses. The family includes the wild and cultivated citrus fruits of the subfamily 

Aurantioideae (chiefly from Citrus and Fortunella) which serve as a source of food and 

flavourings, essential oils, and medicines. A number of other genera provide lesser known 

edible fruits (Clausena, Limonia, and Triphasia), flavours (Murraya), and traditional 

medicines and poisons (Glycosmis, Merope, Micromelum, Murraya, Paramignya, Severinia). 

Although few of the species reach timber size, the wood of some is valued for its special 

qualities such as grain, colour, or hardness (Citrus, Merrillia, Micromelum, and Murraya). 

A number of species are prized as ornamentals or hedge-plants due to their attractive 

foliage and fragrant flowers (Citrus, Fortunella, Glycosmis, Luvunga, Murraya, and 

Triphasia). Certain species of Pleiospermium and Severinia have been suggested as a 

source of rootstock material for Citrus breeding and improvement programmes. 

More than half the species of the non-aurantioid genera (subfamilies Rutoideae and 

Toddalioideae) attain timber size, yet only Acronychia (1 species), Maclurodendron (1 

species) and Melicope (1 species) have reportedly been used for construction purposes. All 

of the genera (except Maclurodendron), including Toddalia, are said to have minor 

medicinal importance. Lesser-known uses are as a source of resin (Melicope), food and 

flavouring (Acronychia and Melicope), and fish poison (Acronychia). 

Taxonomy. The Rutaceae is generally accepted as closely allied to the Meliaceae, 

Simaroubaceae, and in some characters to other families included in the order Sapindales, 

e.g., the Anacardiaceae, Burseraceae, and Sapindaceae. From the Meliaceae and Simaroubaceae 

(its closest allies), the family differs by its glandular-punctate leaves and the presence of 

secretory cavities containing aromatic esthereal oils in the parenchymatous and pericarp 

352

RUTACEAE (JONES) 

tissues. In the most detailed treatment of the family, Engler (in Engler & Prantl, Pfl. Fam. 

19 a (1931) 187-359) recognised 7 subfamilies. Airy-Shaw (in Willis, Dict. FI. PI. & Ferns 

8th ed. (1973) 1014), however, raised the subfamily Flindersioideae and Rhabdodendroideae to 

family status, thus reducing the number of subfamilies to 5. Of these 5 or 7 subfamilies, 

only the Aurantioideae, Rutoideae and Toddalioideae are represented by native species in 

Sabah and Sarawak. More recently, Da Silva et al. (PI. Syst. EvoI. 161 (1988) 97-134), on 

the basis of phytochemical, morphological and geographical evidence, proposed the 

inclusion of the subfamily Toddalioideae in the Rutoideae, and the reorganisation of tribes 

and genera in the subfamily Aurantioideae. 

Key to subfamilies 

l. Carpels usually 3-5, usually joined only by the styles, separated in fruit. Fruits follicular, 

woody, dehiscing along the upper suture ............................................... subfam. Rutoideae 

(Genera: Lunasia, Melicope, Tetractomia, Zanthoxylum) 

Carpels 2-20, joined completely or nearly so. Fruit a drupe or berry, dry or fleshy, 

indehiscent. ............................................................................................................................ .2 

2. Carpels 2-5, rarely 1, rarely completely joined. Fruit a single, several-Ioculed drupe, or 

2-4 druplets joined at the base. Seeds with endosperm, cotyledons obscure ................. . 

...................................................... ......................................... .. subfam. Toddalioideae 

(Genera: Acronychia, Maclurodendron, Toddalia) 

Carpels usually 4-20, completely joined. Fruit a dry or fleshy berry or hesperidium, 

with or without pulp-vesicles. Seeds lacking endosperm, cotyledons well-developed 

........................... subfam. Aurantioideae 

(Genera: Burkillanthus, Citrus, Clausena, Fortunella, 

Glycosmis, Limonia, Luvunga, Merope, Merrillia, 

Micromelum, Monanthocitrus, Murraya, Paramignya, 

Pleiospermium, Severinia, Triphasia). 

Key to native tree and shrub genera 

l. Twigs, branchlets, and older branches unarmed. Leaves opposite ................................. 2 

Twigs and branchlets and/or older branches armed with spines or prickles, or 

ullClfmed. Leaves alternate or spirally arranged ............................................................ 5 

2. Leaves trifoliolate and/or unifoliolate, rarely bifoliolate; petioles 1-30 cm long, not 

swollen apically .......................................................................................... 8. Melicope 

Leaves strictly unifoliolate; petioles not longer tIlan 5 cm, generally swollen 

apically ........................................................................................................................ 3 

3. Leaves generally leathery. Fruit a group of 1-4 dehiscent follicles ....... 16. Tetractomia 

Leaves thinly leathery. Fruit a 4-loculate, indehiscent drupe ........................................ 4 

4. Younger branchlets glabrous to finely hairy. Leaflets usually elliptic; lateral veins and 

reticulations conspicuous below. Flowers bisexuaL ................................ 1. Acronychia 

353


Younger branchlets brown- to rusty-hairy. Leaflets usually obovate to oblanceolate; 

lateral veins and reticulations inconspicuous below. Flowers unisexuaL ...................... . 

..... 7. Maclurodendron 

5. Twigs and branchlets unarmed (or with spine-like paraphylls in some Severinia) ........ 6 

Twigs and branchlets and/or older branches armed with spines or prickles (or unarmed 

or nearly so in some Pleiospermium) .......................................................................... 12 

6. Leaves simple; blades often rigid when older, not articulated at the base ...................... 7 

Leaves pinnate and/or unifoliolate; leaflet blades not as above, articulated at the base .......... 8 

7. Leaf-margins sinuate or coarsely dentate. Petioles swollen apically. Fruit a group of 

ribbed follicles .............................................................................................. 6. Lunasia 

Leaf-margins subentire to obscurely crenulate. Petioles not as above. Fruit a fleshy 

berry ......................................................................................................... 15. Severinia 

8. Young branch-tips densely rusty-hairy. Leaves pinnate and/or unifoliolate, often leathery 

Glycosmis 

Young branch-tips glabrous or hairy (if hairy, not as above). Leaves pinnate, thinly 

leathery ........................................................................................................................ 9 

9. Leaf-rachis narrowly winged. Leaflets alternate to subopposite. Fruit a large berry, 7 

cm in diameter or more; pericarp thick, leathery ....................................... l0. Merrillia 

Leaf-rachis wingless. Leaflets alternate. Fruit a small berry, 1-2 cm in diameter or 

less; pericarp thin ...................................................................................................... 10 

10. Leaflets generally 3-7 (or more in some of the cultivated species); bases not obliquely 

asymmetric (or asymmetric in some of the cultivated species). Inflorescence an axillary 

panicle (or terminal and/or corymbose in some of the cultivated species) ... 13. Murraya 

Leaflets generally 5-15 (or more in some Clausena); base obliquely asymmetric. 

Inflorescence a terminal panicle or corymb ................................................................ 11 

11. Leaflets 9-15. Inflorescence a flat-topped corymb. Petals valvate in bud .... 11. Micromelum 

Leaflets 5-31. Inflorescence a panicle. Petals imbricate in bud .................... 4. Clausena 

12. Trees or scandent to erect shrubs. Twigs and branchlets and/or older branches armed 

with numerous thick spines or small (sometimes hollow or recurved) prickles. Leaves 

pinnate. Fruit of 1-4 follicles ............................................................. 17. Zanthoxylum 

Trees or erect shrubs. Twigs and branchlets and/or older branches armed with straight, 

terete, axillary spines, or unarmed or nearly so. Leaves 2-3-foliolate and/or 

unifoliolate, or simple. Fruit a fleshy berry or hesperidium ......................................... 13 

13. Leaves 2-3-foliolate (or unifoliolate in juvenile growth), rather stiff when older 

........................................................................................................... 2. Burkillanthus 

Leaves strictly unifoliolate or simple .......................................................................... 14 

14. Shrubs or small trees. Petioles wingless. Fruits lacking pulp-vesic1es ......................... 15 

Shrubs or small to large trees. Petioles narrowly to broadly winged, or rarely winglcss. 

Fruits containing pulp-vesicles ................................................................................... 16 

354

RUT ACEAE (IONES) 

15. Leaves oblong-ovate, thick, fleshy; lateral veins inconspicuous. Petiole articulated with 

the blade. Mangrove swamp plants ................................................................ 9. Merope 

Leaves oblong-oblanceolate, thin, not fleshy; lateral veins conspicuous. Petiole not 

articulated with the blade. Inland lowland forest plants .................. 12. Monanthocitrus 

16. Spines solitary or paired, generally only on older branches, or branches unarmed or 

nearly so. Fmits 1.5-2.5 cm in diameter... ....................................... 14. Pleiospermium 

Spines solitary, predominantly on younger branches and twigs, older branches often 

unarmed. Fmits mostly 4 cm in diameter or larger .......................................... 3. Citrus 

Key to native genera 

l. Carpels 3-4, seldom 2, often joined by the styles but separated in fmit. Fmits follicular, 

leathery or woody, dehiscing along the upper suture; endocarp sometimes detaching ... 2 

Carpels 2-18, incompletely or completely united. Fmit an indehiscent, dry or fleshy 

dmpe or berry .............................................................................................................. 5 

2. Flowers 4-5-merous, unisexual or bisexual. Branches armed with spines or prickles, or 

unarmed ...................................................................................................................... 3 

Flowers 3-4-merous, unisexual (or bisexual in some Melicope). Branches unarmed ... .4 

3. Ovary 4-carpellate. Follicles boat-shaped; exocarp leathery, not glandular. Seeds winged, 

testa parchment-like. Flowers bisexual. Branches unarmed .................. 16. Tetractomia 

Ovary 2-4-carpellate. Follicles subglobose; exocarp conspicuously glandular. Seeds unwinged, 

testa black, glossy. Flowers unisexual. Branches armed with spines or prickles 

.......................................................................................................... 17. Zanthoxylum 

4. Flowers 3-merous, unisexual. Follicles flattened laterally, ribbed, with a short beak. 

Leaves simple, alternate ................................................................................ 6. Lunasia 

Flowers 4-merous, unisexual or bisexual. Follicles subglobose, not ribbed, without 

beak. Leaves trifoliolate and/or unifoliolate, opposite .................................. 8. Melicope 

5. Carpels rarely completely united; ovary usually shallowly lobed. Fmit a dmpe ............ 6 

Carpels always completely united; ovary not lobed. Fmit a berry or hesperidium ......... 8 

6. Scandent shmbs, armed with.prickles. Leaves trifoliolate, alternate or spiral. Flowers 

unisexual. ............. . 

Toddalia Juss. 

Gen. PI. 3 (1789) 371; Hooker J l.c. (1875) 497; Kurz I.c. (1877) 183; Craib I.c. 220; 

Backer & BakhuizenJ I.c 101; Perry l.c. 368. 

1 species; Africa, Madagascar, India, Sri Lanka, S China, Indonesia, Borneo 

(Sabah and Sarawak), and the Philippines. 

Climbers with prickly branchlets. Leaves trifoliolate; leaflets with prickly midrib 

and toothed margin. Inflorescence an axillary or terminal panicle. Flowers 4-5- 

merous, unisexual; sepals small; petals to 5 mm long, oblong; stamens 4-5, female 

flowers with staminodes; ovary 4-5-carpellate, on short stalk. Fmit a small, round 

dmpe. Seeds smooth, angular; embryo ~urved, embedded in fleshy endosperm. 

355


Erect shrubs or trees, unarmed. Leaves unifoliolate, opposite. Flowers unisexual or 

bisexual ...................................................................................................................... 7 

7. Flowers unisexual; buds globose; petals imbricate; filaments glabrous; ovaries and 

fruits glabrous ................................................................................ 7. Maclurodendron 

Flowers bisexual; buds cylindrical; petals valvate; filaments ciliate at base; ovaries and 

fruits hairy or sparsely so ........................................................................ 1. Acronychia 

8. Twigs and branchlets unarmed. Ovary 2-S-carpellate (or rarely 6-carpellate in 

Merrillia); ovules 1-2 per locule (or 8-10 in Merrillia). Fruits rather small (or large in 

Merrillia), without pulp-vesicles, dry or slightly juicy (or mucilaginous in Merrillia) 

.............. 9 

Twigs and branchlets (especially in young plants) and/or older branches armed (or 

unarmed or with spine-like paraphylls in some Severinia; unarmed or nearly so in 

some Pleiospermium). Ovary 2-S-carpellate (or 6-I8-carpellate in Citrus), ovules 2 

per locule or more (or rarely 1). Fruits small to large, with or without pulp-vesicles, 

usually juicy or mucilaginous ..................................................................................... 13 

9. Inflorescence a terminal, flat-topped corymb. Petals valvate in bud. Cotyledons thin, 

folded ................................................................................................. 11. Micromelum 

Inflorescence a terminal or axillary panicle, raceme or cyme, or 1-2-flowered. Petals 

imbricate in bud. Cotyledons thick, flat... ................................................................... lO 

10. Inflorescence I-2-flowered. Petals c. 2.S cm long. Ovary S-6-carpellate. Fruits 9 cm or 

more in diameter; pericarp thick, leathery ................................................. 10. Merrillia 

Inflorescence a many-flowered panicle, raceme or cyme. Petals rarely exceeding 2 cm 

long. Ovary 2-S-carpellate. Fruits less than 2.S cm in diameter; pericarp thin, 

glandular .................................................................................................................. 11 

Il. Style persistent, thick, shorter than the ovary. Locules 1-2-ovulate. Young growth 

densely reddish hairy ................................................................................ 5. Glycosmis 

Style deciduous or persistent, shorter than the ovary or as long or longer. Locules 2- 

ovulate (rarely I-ovulate). Young growth not as above ............................................... 12 

12. Petals 10-21 mm long or more. Style as long as or longer than the ovary, stigma 

capitate. Floral buds cylindrical or oblong ................................................. 13. Murraya 

Petals 3-6 mm long. Style shorter than the ovary, stigma flattened. Floral buds globose 

or ovoid ...................................................................................................... 4. Clausena 

13. Stamens twice as many as the petals. Fruits 1-3 cm in diameter; locules containing 

mucilage or scant flesh, pulp-vesicles lacking. Erect or scandent shrubs or trees ........ 14 

Stamens 2-4 times as many as the petals or more. Fruits variable in size, to 20 cm in 

diameter; locules containing juice-filled pulp-vesicles. Erect shrubs or trees .............. 17 

14. Scandent shrubs armed with solitary, recurved or straight spines. Leaves trifoliolate 

and/or unifoliolate ..................................................................................................... IS 

Erect shrubs or small trees armed with solitary or paired, straight spines. Leaves 

unifoliolate or simple ................................................................................................. 16 

3S6

RUT ACEAE (JONES) 

IS. Leaves trifoliolate (or unifoliolate on juvenile stems). Petioles 7-30 cm long, rarely 

Luvunga Buch.-Ham. in Wall. ex Wight & Am. 

Procir. 1 (1834) 90; Hooker! I.e. (1875) 508; Kurz I.e. (1877) 193; Merrilll.e. (1921) 315, 

I.e (1929) 115; Ridley I.e. (1922) 354; Craib I.e. 235; Masamune I.e. 360; Backer & 

Bakhuizen! I.e. 105; Burkill, EPMP (1966) 1396; Swingle I.e. (1967) 265; Anderson I.e. 

308; Perry I.e. 371; Stone, Proc. Ac. Nat. Sc. Phil. 137 (1985) 221. 

c. 12 species; India, Sri Lanka, Burma, Indo-China, Malesia; 7-8 species in Sabah 


Woody climbers with hook-like or (in juvenile plants) straight axillary spines. 

Leaves trifoliolate or (in juveniles) unifoliolate. Leaflets coriaceous; margin entire; 

petiolules pulvinate. Inflorescence an axillary panicle or raceme. Flowers 4-Smerous; 

calyx cup-like, slightly lobed; petals oblong-linear; stamens usually 8 or 

10; ovary 2-4-carpellate, on short stalk, style thick. Fruit a glandular berry, thickwalled. 

Seeds 1-3 per fruit. 

Leaves unifoliolate. Petioles to 2.S cm long, usually shorter ................. . 

Paramignya Wight 

Ill. Ind. Bot. 1 (1840) 108, 110; Hooker! I.e. (1875) 509; Kurz I.e. (1877) 193; Ridley I.e. 

(1922) 355; Burkill, Gard. Bull. S.S. 5 (1931) 213, I.e. (1966) 1691; Craib I.e. 235; 

Swingle I.e. (1967)270; Stone I.e. (1972) 384; Perry I.e. 371; Corner I.e. 669. 

c. 12 species; India, Sri Lanka, Burma, S China, Indo-China, Thailand, Malesia, 

Australia; 3-4 species in Sabah and Sarawak. 

Woody climbers or (rarely) shrubs, with hook-like or short straight axillary spines. 

Twigs zigzag. Leaves unifoliolate, margins entire; petioles short, pulvinate. 

Inflorescences axillary, often solitary. Flowers 4-S-merous, often large; calyx cuplike, 

lobed; petals lanceolate-oblong; stamens 8 or 10; ovary 3-S-carpellate, 

stalked. Fruits round or lobed, glandular, resinous. Seeds 1 to several per fruit, 

ovoid or flattened. 

16. Ovary ovoid, glabrous. Fruits ovoid to ellipsoid, angular, green when ripe. Seeds 

flattened. Leaves unifoliolate ....................................................................... 10. Merope 

Ovary bottle-shaped, sparsely hairy. Fruits obovoid to pyriform, yellow-orange when 

ripe. Seeds thick. Leaves simple ..................................................... 12. Monanthocitrus 

17. Stamens 4-8 times as many as the petals. Pulp-vesicles tapered at both ends (fusiform), 

on slender stalks ............................................................................................ 13. Citrus 

Stamens twice as many as the petals. Pulp-vesicles cylindrical or conical or irregular in 

shape, stalkless or nearly so ....................................................................................... 18 

18. Ovary S-carpellate, locules 22-26-ovulate. Fruits 8-11 cm in diameter, obovoid to oblong 

........................................................................................................... 2. Burkillanthus 

Ovary 2- or 4-S-carpellate, locules 1-2-ovulate. Fruits to 2.5 cm in diameter, usually 

less, round or nearly so .............................................................................................. 19 

19. Disc small, annular to cup-like. Fruits 15-25 mm in diameter, pericarp rough-glandular. 

Petioles winged or narrowly so, articulated with the leaf-blades ....... 14. Pleiospermium 

Disc cup-like, enclosing the base of the ovary. Fruits 6-1S mm in diameter, pericarp 

smooth. Petioles wingless, not articulated with the leaf-blades ................. 15. Severinia 

357


1. ACRONYCHIA 1. R. Forst. & G. Forst. 

(Greek, akron = tip, onychos = claw; the claw-like tip ofthe petals) 

Char. Gen. PI. ed. 1 (1775) 27; Hooker f I.c. (1875) 498; Kurz I.c. (1877) 183; King, J. As. Soc. 

Beng. 62,2 (1893) 214; Rid1ey I.c. (1922) 347; MerrillI.c. (1929) 113; Craib I.c. 221; Masamune 

I.c. 357; Backer & Bakhuizenf I.c. 101; BurkillI.c. (1966) 40; Stone I.c. (1972) 371; Hart1ey, J. 

Am. Arb. 55 (1974) 469; Anderson I.c. 307; Perry I.c. 361; Whitmore, Tantra & Sutisna I.c. 304. 

Shrubs or trees or rarely climbers. Branches unarmed. Leaves opposite, trifoliolate or 

unifoliolate, petioles wingless; leaflets pinnately veined, margin entire, articulated at the 

base. Inflorescences axillary, paniculate, subcorymbose, or 1- to few-flowered. Flowers 

bisexual, 4-merous; sepals 4, free or fused at base, imbricate; petals 4, free, valvate, pelluciddotted, 

white or cream to greenish or yellowish, becoming reflexed; stamens 8, unequal, 

alternately long and short, filaments flattened and tapering to a sharp apex, usually densely 

ciliate at the base, becoming reflexed, anthers ovoid to ellipsoid, basifixed; disc shallowly 

8-10bed; ovary 4-carpellate, with septicidal fissures or without, ovules 2 per locule, style 

twisted, stigma shallowly 4-10bed. Fruit a 4-loculate drupe, with septicidal fissures or 

without; epicarp semi-fleshy, spongy or woody when dry, with or without evident mesocarp; 

endocarp cartilaginous to parchment-like. Seeds 1-2 per locule, narrowly ellipsoid, slightly 

bent; testa dull to shiny, smoothish to finely rugose; endosperm fleshy; embryo straight. 

Distribution. 42 species; from India to SW China, and throughout Malesia to E Australia 

and New Caledonia. The greatest number are found in Australia, Irian Jaya (Indonesia) and 

Papua New Guinea, including several rare and endemic species. One species, A. pedunculata, 

occurs in Sabah and Sarawak. 

Ecology. Found in primary and secondary rain forests, from the lowlands and hills to cloud 

forests and subalpine habitats above 3000 m and in coastal scrub. 

Acronychia pedunculata (L.) Miq. Fig. 1. 

(Latin, pedunculatus = slender-stalked; the inflorescence) 

Fl. Ned. Ind., SuppI. (1861) 532; MerrillI.c. (1929) 113; Craib I.c. 221; Masamune I.c. 357; Stone 

I.c. (1972) 371; Hart1ey I.c. 549; Perry I.c. 361; Whitmore, Tantra & Sutisna I.c. 305. Basionym: 

lambolifera pedunculata L., Sp. PI. 1 (1753) 349. Type: lambolifera L., Fl. ZeyI. (1747) 139. 

Synonyms: Acronychia laurifolia Blume, Cat. (1823) 63; A. arborea Blume, Bijdr. (1825) 244; 

Melicope conferta Blanco, FI. Filip. ed. 2 (1845) 205; Acronychia apiculata Miq. I.c. (1861) 532. 

Shrub or small to large tree to 30 m tall and 50 cm in diameter; bole to 23 m tall. Younger 

branchlets glabrous to finely hairy. Leaves unifoliolate; leaflet usually elliptic or ellipticoblong 

to obovate or oblanceolate, 3.5-24.5 x 2-8.5 cm, thinly leathery; base cuneate or 

occasionally rounded, apex acuminate or occaSionally blunt or rounded; lateral veins 7-14 

pairs; reticulation~ conspicuous above and below; petioles 0.5-5 cm long. Inflorescences 

few- to many-flowered, 2-25 cm long, axes glabrous or nearly so, pedicels 2-12 mm long. 

Flowers 4-12 mm long; sepals triangular or rounded, minute; petals linear or oblanceolate, 

glabrous to sparsely hairy; disc glabrous; ovary hairy, rarely only at apex; style hairy at 

base. Fruits to 15 mm in diameter, usually subglobose to ellipsoid or conical, more or less 

358


2 cm [ 

B 

',m [ 

A 

c 

5mm 

Fig. 1. Acronychia pedunculata. A, fruting leafy twig; B, part of inflorescence; C, flower with one 

petal removed. (From SAN 85676.) 

359


1

RUTACEAE (lONES) 

4-lobed or longitudinally ribbed, usually sparsely hairy, base with a ring of dense hairs, 

apex rounded to shortly tipped; epicarp when dry to 3 mm thick; mesocarp woody or nearly 

so; endocarp cartilaginous. Seeds reddish black to black, 3-7 mm long. 

Distribution. India, Sri Lanka, eastward to Taiwan and throughout Malesia to Papua New 

Guinea. In Sabah, frequent in primary and secondary forests, often on hill-sides, from near 

sea-level to 1500 m. In Sarawak, collected only twice (Baram and Lundu Districts). Also 

occurs in Brunei and Kalimantan. 

Uses. The wood, roots, bark and leaves are used medicinally to treat rheumatism, scabies 

and colic, and as pain reliever (China, Indo-China, Indonesia). The roots are used as a fish 

poison (Indo-China). The wood is rarely used in construction and for making charcoal, and 

its young leaves are eaten as a condiment (Java). - 

2. BURKILLANTHUS Swingle 

(I. H. Burkill, 1870-1965, one time director of the Singapore Botanic Gardens) 

J. Am. Arb. 20 (1939) 255, I.c (1967) 294; Stone I.c. (1972) 373; Whitmore, Tantra & Sutisna l.c. 

305. 

Trees; crown broad, dense. Branches armed, drooping in older trees. Leaves alternate, 1- 

3-foliolate; petioles narrowly winged to nearly wingless, pulvinoid at base; leaflets thinly 

leathery, becoming stiff when older, glabrous, articulated at the base. Inflorescences 

axillary, few-flowered clusters, pedicels short. Flowers bisexual, large, 5-merous; sepals 5, 

nearly free to the base, with large oil-glands; petals 5, gland-dotted distally; stamens 10, 

free, slender, anthers small, oblong; disc cylindrical; ovary 5-carpellate, obovoid, sparsely 

hairy, glandular on upper half, fluted with 5 grooves; ovules 22-26 per locule in 2 rows, 

style slender, glabrous, stigma globose. Fruit a large, obovoid to oblong berry; exocarp 

thick, leathery, light green becoming pale yellow, roughened with numerous oil-glands; 

endocarp thin, woody; pulp-vesicles sessile, cylindric, tips acute, pale yellow. Seeds numerous, 

large, cream with brown cap, embedded in mucilage; monoembryonic. 

Distribution. 1 species, restricted to N Sumatra, Peninsular Malaysia and Sarawak. Uncommon, 

and possibly endangered. 

Ecology. Found as solitary trees or small populations of several trees, on stream banks, 

slopes, and ridge tops in primary and secondary forest, on rocky and sandy soils, from 5 to 

200 m. 

Burkillanthus malaccensis (Ridl.) Swingle 

(of Malacca) 

Fig. 2. 

I.c. (1939) 257, I.c. (1967) 295; Whitmore I.c. 373; Stone I.c. (1972) 373, Fed. Mus. J. 23 (1978) 

114; Whitmore, Tantra & Sutisna I.c. 305. Basionym: Citrus malaccensis Ridl. I.c. (1922) 359. 

Type: Goodenough 1273, Malacca, Nya1as (ho1otype SING; isotypes K, US). 

361


Small tree to 15 m tall and 40 cm in diameter; bole to 4 m tall, fluted. Branches generally 

armed with stout, straight, usually paired spines. Leaves unifoliolate (on juvenile growth) 

or 2-3-foliolate; terminal leaflets lanceolate or elliptic-lanceolate, 12.5-27 x 4.5-11.5 cm; 

base broadly cuneate, margins subentire to slightly coarse-crenate, apex acute or acuminate; 

lateral veins 12-18 pairs, deeply sunken above; petioles 4-5 cm long; petiolules 2-2.5 mm 

long, pulvinoid; lateral leaflets to 8 x 4 cm, petioles 1-4 cm long, otherwise similar to 

terminal leaflet. Inflorescences 1-4-flowered, pedicels 5-7 mm long. Flower 5-6 cm 

across; sepals lanceolate-acuminate; petals 2-2.5 cm long, subspathulate; ovary 6.5-7 mm 

long. Fruits 10-8 x 8-11 cm; exocarp c. 1 cm thick; endocarp 2-3 mm thick, pulp-vesicles 

1.5-3 cm long. Seeds broadly obovoid, tapering to an acute base, 2.2-2.7 cm long and c. 8 

mm thick; testa thin, slightly wrinkled; cotyledons pale buff. 

Vernacular name. Sarawak-limau hantu (Malay). 

Distribution. Throughout the range of the genus. In Sarawak very uncommon, collected 

once in 1961 (S. 15848) from primary riparian forest in the Labang FR, Bintulu. 

3. CITRUS L. 

(Latin name for the citron tree) 

Sp. PI. 1 (1753) 401; Hooker! I.e. (1875) 514; Kurz I.e. (1877) 195; Merrilll.e. (1921) 315; Ridley 

I.e. (1922) 358; Burkilll.e. (1931) 220, I.e. (1966) 568; Craib I.e. 236; Masamune I.e. 357; Santiago, 

Bull. Mal. Div. Agr. 111 (1962) 1; Backer & Bakhuizen! I.e. 107; Swingle I.e. (1967) 358; Stone 

I.e. (1972) 374, I.e. (1985) 226; Perry I.e. 361; Corner I.e. 659; Whitmore, Tantra & Sutisna I.e. 305; 

PROSEA2 (1991) 119, 325. 

Shrubs or small to large trees. Branches generally armed with axillary, solitary spines, 

older branches often unarmed. Young twigs angular, flattened, becoming terete. Leaves 

alternate, simple or unifoliolate, blades thinly leathery to leathery, glabrous or short-hairy 

below; margins entire to serrate or crenulate; petioles often winged or emarginate, usually 

articulated with the blade. Inflorescences axillary, short corymbose racemes or cymes, or 

solitary. Flowers bisexual or male only, 4-5-merous, fragrant or not; calyx cup-like; sepals 

4-5; petals 4-8, commonly 5, oblong-linear, thick; stamens about 4-8 times the number of 

petals, free or fused at the base and in bundles; disc annular; ovary subglobose, (6-)8-18- 

carpellate, usually glabrous, ovules (1-)4-8(-12) per locule in 2 rows, style cylindric, 

stigma globose-capitate. Fruit a fleshy hesperidium; pericarp leathery, outer layer (exocarp or 

flavedo) densely glandular and pigmented, middle layer (mesocarp or albedo) white, dry, 

inner layer (endocarp) membranous, thin, flesh of stalked fusiform, pulp-vesicles filled with 

watery, sweet or bitter juice, with or without distinct oil-droplets. Seeds few to many, 

angular-obovoid, often flattened, pale; mono- or polyembryonic; embryo white or green. 

Distribution. 17 species; from India, Sri Lanka and Burma to Japan, China, Taiwan and 

Indo-China, and throughout Malesia eastward to the Pacific Islands. Two species, C. 

halimii and C. macroptera, are native to Sabah, the latter species also occurs in Sarawak. 

In addition, a number of species have long been introduced and are cultivated in Sabah and 

Sarawak. Numerous hybrids and variants of these species .exist, many having unknown 

origins which makes classification difficult. 

362


Stone (I.e. 1985) reported a collection of Citrus grandis from Ulu Segama, Sabah (SAN 

75565) and suggested that it might be indigenous there, although he admitted that the 

species could have been introduced into that area by early gold prospectors. On this basis 

and until more specimens are available, this species is excluded from the present treatment. 

Uses. Citrus fruits are eaten fresh, canned, preserved or candied. The juice is extracted and 

made into drinks or concentrates. Pectin and essential oils are made from the rind, while 

waste fruit-pulp is processed into cattle-feed. Citric acid is manufactured from certain 

species. The leaves, flowers, and fruits of some species are sources of expensive essential 

oils (oil of neroli, bergamot oil) used in perfumery. Various plant parts (leaves, flowers, 

fruits, and seeds) are used to flavour foods and often enter into native medicinal remedies. 

Some species are reported to show insecticidal and germicidal activity. F1avonoids from the 

inner cortex are said to have anti-tumour properties. Some species are utilised as rootstock 

for the cultivation of commercially important citrus because of their resistance to certain 

diseases. Citrus is less commonly used as a source of wood for cabinetry. In Malaysia, the 

plants have reportedly been used for occult purposes. 

Taxonomy. The taxonomic delimitation of species, subspecies, and forms of Citrus is 

complicated by the peculiar reproductive behaviour found in the genus including 

hybridisation, polyembryony (asexual seed production), the spontaneous production of 

autotetraploids, and mutations. Two widely opposing systems of classification exist. That of 

Tanaka (Revisio Aurantiacearum - IX, Jap. Soc. Prom. Sci., Ueno, Tokyo, 1954) proposes 145 

species (increased to 157 a few. years later), while that of Swingle (l.c.1967) recognises only 

16 species. This difference reflects a lack of agreement on what constitutes a species and 

whether supposed hybrids should be assigned the rank of species. Swingle, whose system is 

followed in this treatment, did not consider valid many of Tanaka's species which were 

clearly horticultural forms, and mostly the result of interspecific hybridisation. Several 

other systems of classification put forward by various authors are intermediate. Recent 

studies employing numerical taxonomy, chemosystematics, and other experimental methods 

have attempted to elucidate relationships among taxa and clarify the "species" problem in 

Citrus. Several such studies support the notion that the genus centres around three basic 

species or "biotypes", viz. C. grandis, C. medica, and C. reticulata. 

Key to Citrus species 

1. Petioles narrowly winged or wingless, or if broadly winged, never as broad as the leaf 

blades. Flowers large, usually 2.5-4.5 cm wide, fragrant. Stamens free or united in 

bundles ........................................................................................................................ 2 

Petioles broadly winged, often as broad as the leaf-blades. Flowers small, usualiy less 

than 2 cm wide, not fragrant. Stamens free .................................................................. 9 

2. Petioles wingless, not articulated with the leaf-blade. Flowers perfect or staminate. 

Fruits usually oblong, pericarp thick. Cultivated ........................................................... .. 

C. medica L. 

Sp. PI. 2 (1753)782. Synonym: C. aurantium var. medica Wight & Arn. I.e. 98. 

Probably native to NE India and upper Burma. Cultivated widely in the tropics 

and subtropics for its fruits. Vernacular names: citron, limau susu. 

363


Petioles winged, often narrowly so, articulated with the leaf-blade. Flowers all perfect 

(except in C. limon). Fruits ovoid or globose or pear-shaped, pericarp thin (except in C. 

grandis) ....................................................................................................................... 3 

3. Petioles very narrowly winged. Flowers perfect or staminate; stamens usually more 

than 4 times the number of petals. Fruits ovoid. Cultivated ..................... . 

C. limon (L.) Burm. f 

Fl. Ind. (1768) rl3. Basionym: Citrus medica var. limon L. I.e. (1753) 782. Synonym: 

Citrus limonum Risso, Ann. Mus. Hist. Nat. Paris 20 (1813) 20l. 

Probably native to S Asia and now widely cultivated in the tropics and subtropics 

for its fruits. Vernacular names: lemon, limau mata kerbau. 

Petioles narrowly to broadly winged. Flowers usually perfect; stamens usually 4 times 

the number of petals. Fruits globose or pear-shaped, sometimes slightly flattened ....... .4 

4. Fruits with loose pericarp, easily detached. Seeds small, plump. Embryo green ............ 5 

Fruits with adherent pericarp, not easily detached. Seeds various. Embryo not green .... 6 

5. Leaves somewhat rhomboid, tips acute. Fruits small to medium-sized, sometimes 

flattened. Cultivated ...................................................... . 

C. reticulata Blanco 

Fl. Filip. (1837) 610. Synonyms: Citrus nobilis Andrews, Bot. Repos. 9 (1809) 608, non 

Lour. (1790); Citrus delieiosa Tenore, Ind. Sem. Hort. Neap. (1840) 9 .. 

Native to SE Asia. Widely grown for its edible fruits. Vernacular names: mandarin, 

limau manis. 

Leaves elliptic, tips rounded. Fruits small, globose. Cultivated ................. . 

C. microcarpa Bunge 

Mem. Ac. Imp. Sc. st. Petersb. 2 (1833) 84. Synonyms: Citrus mandurensis Lour., F1. 

Cochineh. 2 (1790) 467; Citrus mitis Blaneo I.e. (1837) 610. 

Originated in China as a natural hybrid between Citrus reticulata var. austera 

Swingle (sour mandarin) and a species of Fortunella (kumquat); currently classified as 

xCitrofortunella microcarpa (Bunge) Wijnands, Baileya 22 (1984) 134. Widely 

grown in SE Asia for its acid juice. Vernacular names: calamondin, limau kesturi. 

6. Petioles broadly winged. Fruits large to very large, usually 10-20 cm in diameter, 

round to pear-shaped. Seeds large, rough, monoembryonic. Cultivated ..... 

C. grandis (L.) Osbeck 

Dagb. Ostind. Resa (1757) 98. Basionym: Citrus aurantium var. grandis L. I.e. (1753) 783. 

Synonyms: Citrus deeumana L., Syst. ed. 12 (1767) 508; Citrus maxima (Bunn. f) Merr., 

Interp. Rumph. Herb. Amb. (1919) 296. 

Native to Malesia; widely cultivated in SE Asia for its edible fruits. Vernacular 

names: pummelo, limau besar. 

Petioles narrowly winged to nearly wingless. Fruits small to medium-sized, generally 

4-9 cm in diameter. Seeds various, usually polyembryonic (except in C. halimii) ......... 7 

7. Fruits medium-sized, 5-9 cm in diameter; pulp orange. Cultivated ............... . 

364


C. sincnsis (L.) Osbeck 

Rcisc Ostind. China (1765) 250. Basionym: Citrus aurantium var. sinensis L. I.e. (1753) 

782. 

Probably native to the region between China and Vietnam. Cultivated widely in 

the subtropics and tropics for its edible fruits. Vernacular names: sweet orange, 

limau manis. 

Fruits small, 4-7 cm in diameter; pulp greenish to yellowish. Cultivated or wild ......... 8 

8. Leaves small, mostly 5-7.5 cm long. Stamens 20-25. Fruits globose to ovoid, 

sometimes with apical papilla; pericarp thin, smooth. Cultivated .................................. . 

C. aurantifolia (Christm. & Panz.) Swingle 

1. Wash. Ac. Se. 3 (1913) 465. Basionym: Limonia aurantifolia Christm. & Panz., Pfl. 

Syst. 1 (1777) 618. Synonym: Citrus acida Roxb., Fl. Ind. ed. 2, 3 (1832) 390. 

Native to N India and Burma, or N Malesia. Cultivated throughout the tropics and 

warm subtropics for its fruits. Vernacular names: lime; limau nipis. 

Leaves large, 8-15 cm long or more. Stamens 18-20. Fruits subglobose to slightly 

pyriform; pericarp thick, smooth to bumpy. Hills and mountains ............... 1. C. haIimii 

9. Petiolar wings with subentire margins. Fruits large, 8-15 cm in diameter; pericarp 

smooth ............................................................................................... 2. C. macroptera 

Petiolar wings with serrate-crenulate margins. Fruits small, 5-7 cm in diameter; 

pericarp irregularly bumpy. Cultivated ................................................... . 

C. hystrix DC. 

Cat. PI. Hort. Bot. Monsp. (1813) 97. Synonym: Citrus torosa Blanco I.e. (1832) 609; 

Citrus papeda Miq., Fl. Ned. Ind. 1 (1859) 530. 

Of unknown origin but widely naturalised in Malesia, Sri Lanka and Burma. 

Cultivated for its fruits. Vernacular names: Mauritius papeda, limau purut. 

1. Citrus halimii Stone 

(Sultan Abdul Halim Nu'azzam Shah Ibni Almarhum Sultan Badlishah, Sultan of Kedah 

and former King of Malaysia) 

Biotropica 5 (1973) 102, I.e. (1978) 114; Stone & Jones, Proc. Ac. Nat. Sc. PhiI. 140 (1988) 267; Ng 

I.e. 495; Whitmore, Tantra & Sutisna I.e. 305; PROSEA 2 (1991) 119. Type: Stone 9550, Peninsular 

Malaysia, Pahang, G. Nuang (holotypc KLU; isotypes KEP, L, SING, US). 

Medium-sized or large tree to 25 m tall and 60 cm in diameter; trunk straight, cylindric; 

crown meagre, of few ascending branches. Spines on youngest branches of saplings straight 

to 2.5 cm long, absent on adult plants. Leaves unifoliolate, elliptiC or slightly ovate, 8-15 x 

1 )-7,. cm (larger on "igO!"0US young shoots), thinly le:lther),; margin 5ubentire vr 

subcrenulate, apex acute to shortly acuminate and emarginate; lateral veins 7-11 pairs; 

petioles usually 1-2 cm long, very narrowly winged or wingless, articulated with the blade. 

Inflorescences solitary, pedicels 1-3.5 mm long. Flowers 5-merous; sepals deltoid; petals 

ovate-elliptic, white; stamens 18-20, free or in 2-3 bundles; disc thin; ovary (6-)9-10- 

365


carpel/ate, ovules 1-3(-5) per locule, style columnar, stigma flat, aIlgled. Fruits subglobose to 

slightly pyriform, 5-7 cm in diameter, green ripening glossy deep yellow, smooth or bumpy; 

pericarp c. 6 mm thick, glandular; pulp-vesicles pale greenish to yellowish, juicy, acrid oi/ 

droplets absent. Seeds numerous, to 2.1 cm long, testa veiny-reticulate; monoembryonic; 

cotyledons white. 

Vernacular name. Sabah-limau hutan (Malay). 

Distribution. S Thailand, Peninsular Malaysia, and Borneo. In Sabah, uncommon (Kota 

Marudu, Ranau and Tambunan districts), occurring as solitary trees on slopes and ridges in 

primary submontane forest, at 900-1800 m. Not reported from Sarawak, Brunei and 

Kalimantan. 

Uses. The fruits are eaten as a condiment (rinds?), and are said to be thirst-quenching 

(Peninsular Malaysia). The flesh is said to be acidic and pleasant to taste. 

2. Citrus macroptera Montr. Fig. 3. 

(Greek, makros = large, pteron = wing; the leaf-stalk) 

Mem. Ac. R. (Imp.) Lyon Sc. 10 (1860) 187; BurkillI.e. (1931) 220, I.e. (1966) 576; Santiago I.e. 

131; Swingle I.e. (1967) 395; Stone I.e. (1972) 374; Corner I.e. 661; PROSEA 2 (1991) 326. Type: 

Montrouzier, s.n., New Caledonia, ne Art (LY). Synonyms: Citrus papllana F.M. Bail., Contr. Ft. 

Brit. N. Guin. (1903) 1; c. aurantillm subsp. saponaeea Safford, Contr. US. Nat. Herb. 9 (1905) 

226; C. hystrix (non DC.) Ridl. I.e. (1922) 358. 

var. macroptera 

Small or medium-sized tree to 20 m tall and 50(-100) cm diameter, generally much 

smaller; crown broad, dense. Branches and twigs armed with straight spines to 1.5 cm 

long, longer on juvenile growth. Leaves unifoliolate, to 30 cm long, blade ovate-lanceolate, 

5-15 x 3-6.5 cm; base rounded, margin shallowly crenate, apex acuminate; petioles broadly 

winged, with wing as large as the blade, obovate to nearly spathulate, margin subentire to 

crenate in the upper half, articulated with the blade. Inflorescence a few-flowered cluster. 

Flowers l.3-2 cm in diameter; buds subglobose; sepals triangular, small; petals oblong, 

concave; stamens c. 20, free; ovary 10-12-carpel/ate, slightly hairy, style thick, stigma 

depressed. Fruits globose to pyriform, sometimes depressed, 8-15 cm in diameter or 

larger, smooth, green turning pale yellow; pericarp to 2.5 cm thick, segments 10-12, pulpvesicles 

greenish yellow, juice scant, bitter, acrid oil-droplets present. Seeds 1-2 per 

segment, flattened, to c. 2 cm long; monoembryonic. 

Vernacular names. Sabah and Sarawak-limau hantu, limau hutan (Malay). 

Distribution. The species and the variety are distributed in Indo-China, Malesia, New 

Caledonia, and Polynesia. Three varieties are known to occur, only one of which (var. 

macroptera) is in Sabah and Sarawak. Found occasionally in SW Sabah (Pinangah, 

Kinabatangan, Lahad Datu, and Tawau districts) as solitary trees along the banks of 

streams and on forested ridges and hills in primary forest, at 150-600 m. In Sarawak, 

366


',m [ 

A 

2cm 

Fig. 3. Citrus macroptera var. macroptera. A, leafy twig; B, fruit. (From SAN 71072.) 

367


Fig. 4. Clausena excavata. A, flowering leafy twig; B, part of inflorescence (one flower with two 

petals removed); C, part ofinfructescence. (A & B from SAN 89309, C from SAN 56188.) 

368


known only from a single collection made from lowland riverine forest (Taujik, s.n., Kg. 

Bedup, Serian; KLU). 

Uses. The fruits are reported to be edible (Philippines, Indonesia, Sabah), and are said to 

cause impotence (Indonesia). The leaves are eaten as a vegetable and used medicinally 

(Indonesia). Sticks are made from its wood (Papua New Guinea). 

4. CLAUSENA Bunn.f 

(Clausen, a botanist known to Burmann) 

F1. Ind. (1768) 243; Hooker f I.c. (1875) 503; Kurz l.c. (1877) 187; Merrilll.c. (1921) 314, I.c. 

(1929) ll5; Ridley I.c. (1922) 352; Craib I.c. 231; Masamune I.c. 358; Backer & Bakhuizen.j I.c. 

103; Burkilll.c. (1966) 584; Swingle I.c. (1967) 209; Stone l.c. (1972) 375; Anderson I.c. 307; Perry 

I.c. 363; Corner I.c. 662; Ng I.c. 495; Whitmore, Tantra & Sutisna I.c. 305; PROSEA 2 (1991) 141. 

Shrubs or small trees. Branches unanned; branchlets short-hairy. Leaves alternate, pinnate, 

rachis sometimes winged; leaflets alternate, usually 5-9(-31), often with asymmetric base, 

articulated with the petiolules. Inflorescences lax panicles or racemes, terminal or axillary. 

Flowers bisexual, small, 4-5-merous; buds small, subglobose or ovoid; sepals 4-5, calyx 

cup-like, lobed; petals elliptic or oval, free, imbricate; stamens (7-)8 or 10, free, unequal, 

in 2 whorls, the outer opposite the sepals and longer, filaments inflated or flattened at base, 

often narrowed at apex, glabrous, anthers ovate or elliptic, dorsifixed; disc annular, 

glabrous; ovary 2-5-carpellate, sometimes obscurely lobed, usually hairy, glandular, seated 

on a glabrous, hourglass-shaped gynophore, ovules (1-)2 per locule, style short, stout, 

deciduous or persistent, usually narrowed where it joins the ovary, stigma flattened, 

obscure. Fruit an ovoid or oblong berry. Seed usually 1 per fruit; cotyledons green. 

Distribution. c. 29 species; from tropical Africa to India, S China, Taiwan, Indo-China, 

throughout Malesia and N Australia. Two species, C. calciphila and C. excavata, are in 

Sabah, the latter occurring also in Sarawak, Brunei and Kalimantan. A third species, C. 

lansium, is sparingly cultivated. 

Key to Clausena species 

1. Leaves 15-31-foliolate. Flowers mostIy4-merous. Stamens 8 .................. 2. C. excavata 

Leaves 5-11-foliolate. Flowers 5-merous. Stamens 10 .................................................. 2 

2. Leaves 5-7-foliolate. Stigma flat. Fruits oblong-ellipsoid, to l.5 cm long ..... 1. C. calciphila 

Leaves 9-11-foliolate. Stigma 5-lobed. Fruits ovoid-globose, to 2.5 cm long. Cultivated ...... . 

C. lansium (Lous) Skeels 

USDA Bur. PI. Ind. Bull. 168 (1909) 31. Basionym: Quinaria lansium Lour., F1. Cochinch. 

(1790) 272. Synonyms: Cookia wampi Blanco I.c. 358; Clausena wampi (Blanco) Oliv., 1. 

Lilill. Soc. Bot. 5, Suppl. 2 (1861) 34. 

Native to S China and Indo-China. Widely cultivated in the tropics and subtropic 

for its edible fruits. Vernacular names: wampee, wampi. 

369


1. Clausena calciphila Stone 

(Latin, calx = chalk, Greek, phi/os = loving; referring to its preference for limestone habitats) 

I.c. (1978) Ill. Type: Erwin & Paul S. 27430, Sarawak, Kuching, Bukit Pa'it (SAR). 

Small tree to 10 m tall. Leaves to 60 cm long, commonly 5-7-foliolate; leaflets ellipticovate, 

the lowest ones 7-13 x 4-7 cm, the lateral and terminal ones larger, 15-25 x 12-16 

cm; base rounded, obliquely asymmetric, margin entire to subentire, apex acuminate; 

lateral veins 5-8(-10) pairs, together with midrib raised below; petioles 10-12 cm long, 

rachis glabrescent, wingless,· petiolules 5-9 mm long. Inflorescence a loose, terminal 

panicle, 20 cm long or more during flowering, to 40 cm long in fruit; buds subglobose, c. 2 

mm long. Flowers 5-merous; sepals 5, broadly triangular, minute, margin ciliate; petals 5, 

elliptic, c. 3 mm long, glabrous, margin hyaline; stamens 10, alternately long and short, 

filaments narrowly elliptic, anthers oblong-ovoid, connectives I-glandular; disc annular, 

yellowish; ovary 5-carpellate, globose, glabrous, papillate-glandular, style shortly cylindrical. 

Fruits oblong-ellipSOid, 1.5-1.6 cm long, gland-dotted. 

Distribution. Endemic to Sarawak. Uncommon, known from only three localities in 

Kuching (Bt. Pa'it, Bt. Bra'ang) and Baram (Bt. Mentagai) districts; on rocky limestone 

slopes, at 150-300 ill. 

2. Clausena excavata Burm. f Fig. 4. 

(Latin, excavatus = to make hollow; the bases of the filaments) 

I.c. 243; HookerJ I.c. (1875) 504; Kurz I.c. (1877) 188; Merrilll.c. (1921) 314, l.c. (1929) 115; 314; 

Ridley I.c. (1922) 352; Craib I.c. 231; Masamune I.c. 358; Backer & BakhuizenJ l.c. 104; Burkill 

I.c. (1966) 585; Swingle I.c. (1967) 212; Stone I.c. (1972) 375; Anderson l.c. 308; Perry l.c. 363; 

Corner I.c. 662; Whitmore, Tantra & Sutisna I.c. 306. Type: Burmann 29, India, Dauhon Kongeere 

(L). Synonyms: Clausena javanensis Raeusch. ex DC., Prodr. 1 (1824) 538; Murraya burmanni 

Spreng., Syst. Veg. 2 (1825) 315; Amyris graveolens Buch.-Ham. ex Steud., Nom. Bot. ed. 2, 1 

(1840) 81. 

var. excavata 

Small tree to 15 m tall and 20 cm diameter. Branchlets softly hairy. Leaves 20-50 cm long, 

rachis slender, cylindrical, not winged; leaflets 15-31, oblong-ovate to lanceolate or 

slightly crescent-shaped, 2-9 x l.5-4 cm, thinly leathery, glabrous above, sparsely hairy 

below; base rounded, obliquely asymmetric, margins toothed, apex tapered or acuminate; 

petiolules short. Inflorescence a much-branched panicle, terminal, 10-30 cm long, pyramidshaped, 

branches hairy. Flowers 4(-5)-merous, buds round; calyx minute, hairy; petals 

oval, 3.5-5 mm long, glabrous, yellowish or greenish; stamens 8, filaments inflated and 

concave at base; ovary ovoid to ellipsoid, hairy or hirsute, slightly lobed, style cylindrical, 

persistent, not narrowed where it joins the ovary. Fruits broadly ellipsoid, 1-l.8 cm long, 

smooth, glabrous; peel green ripening pink, translucent. Seed 1 per fruit, oblong. 

Distribution. India, Burma, S China, Taiwan, Indo-China, and throughout Malesia. Two 

varieties exist with only the widespread var. excavata in Sabah and Sarawak. Widespread 

370


and common in Sabah and parts of Sarawak (Kuching and Baram districts). Also occurs in 


Ecology. In forests and open places in the lowlands and hills, especially disturbed areas 

around villages and forest margin, and on limestone, from sea-level to 1500 m. 

Uses. The leaves are reported to have insecticidal properties, and the roots, stems, leaves 

and flowers are used to treat paralysis, stomach disorders, intestinal worms, and fever 

(Taiwan, Indo-China, Peninsular Malaysia, Indonesia). Its wood has been used to make 

axe-handles (Java). 

5. GLYCOSMIS COrrea 

(Greek, glukus = sweet, osme = scent; sweet-smelling plant) 

Ann. Mus. Hist. Nat. Paris 6 (1805) 384; HookerJ I.c. (1875) 499; Kurz I.c. (1877) 184; Merrilll.c. 

(1921) 314, I.c. (1929) 113; Ridley I.c. (1922) 348; Craib I.c. 222; Narayanaswami, Rec. Bot. Surv. 

Ind. (1941) 1; Masamune I.c. 359; Backer & BakhuizenJ I.c. 102; Burkilll.c. (1966) 1103; Swingle 

l.c. (1967) 206; Stone I.c. (1972) 380, I.c. (1978) 75, l.c. (1985) 1; Anderson l.c. 308; Corner l.c. 

665; Whitmore, Tantra & Sutisna I.c. 306. 

Shrubs or trees. Branches unarmed. Shoot tips and young inflorescences densely hairy, the 

hairs reddish. Leaves alternate or opposite, pinnate or unifoliolate, rarely simple, petioles 

wingless; leaflets 1-5(-15), alternate or sometimes opposite, chartaceous or leathery; 

margin entire or obscurely toothed, articulated at the base. Inflorescences axillary or 

pseudoterminal racemes or cymose panicles. Flowers bisexual, 4-5-merous; sepals fused at 

base, imbricate, glandular; petals ovate to elliptic, imbricate, greenish to whitish; stamens 8 

or 10, usually unequal, filaments abruptly narrowed at the apex, anthers small, ovate to 

elliptic, generally with a terminal gland, ojten with 1 or several glands on the connective; 

disc annular, with small lobes; ovary 2-5-carpellate, club- to flask-shaped to nearly conical, 

often raised on a stout gynophore, ovules 1-2 per locule, style short, thick, persistent, 

stigma slightly broadened, faintly lobed. Fruits globose to oblong or ellipsoid berries, dry 

or fleshy, outer wall glandular, white to pink or purplish. Seeds 1-2(-3) per fruit, round to 

pIano-convex, thin-walled, cotyledons green, fleshy. 

Distribution. c. 40 species; India, Sri Lanka, SE Asia, S China, Taiwan, Malesia, and 

Australia. Seven species occur in Sabah and Sarawak, two of which (Glycosmis chlorosperma 

and G. macrantha) are widespread and common. Glycosmis parviflora is occasionally 

cultivated in Sabah and possibly also in Sarawak. 

Taxonomy. Although readily distinguished from its closest relatives (Clausena and 

Murraya), the chaotic state of its constituent species (65 names have been published to 

date) has persisted due to lack of any critical taxonomic and nomenclatural review. Stone 

(I.c. 1978 and 1985), whose interpretation of the genus is followed here, admits that 

Glycosmis is still imperfectly known and several taxonomic problems remain to be solved. 

371


Key to Glycosmis species 

1. Leaves predominantly pinnate, never uniformly unifoliolate. Ovary smooth ................. 2 

Leaves predominantly unifoliolate, or some of them 2-3-foliolate. Ovary papillateglandular. 

............................................................................................. 3. G. lanceolata 

2. Ovary glabrous, 4-5-carpellate .................................................................................... 3 

Ovary densely hairy, 2-3-carpellate ................................................... 6. G. sapindoides 

3. Inflorescences pseudoterminal. ............ . ................................. 4 

Inflorescences axillary ................................... ........................... 7 

4. Flowers large, 7-11 mm long; petals glabrous ............................................................. 5 

Flowers medium-sized to small, rarely over 6 mm long; petals hairy, at least below ..... 6 

5. Leaves alternate; leaflets not over 15 cm long, lateral veins 5-9, petiolules to 7 mm 

long .................................................................................................... 5. G. macrantha 

Leaves sometimes opposite, leaflets over 15 cm long, lateral veins 10-14, petiolules 

10-20 mm long ....................................................................................... 7. G. superba 

6. Lateral veins not prominently raised below. Sepals narrowly triangular, c. 2.5 mm 

long. Fruits subellipsoid, c. 7.5 mm long ........................................... 4. G. longisepala 

Lateral veins prominently raised below. Sepals deltoid-ovoid, 1-1.5 mm long. Fruits 

globose, over 10 mm 10ng ............................................................... 1. G. chlorosperma 

7. Leaflets predominantly entire. Inflorescences corymbose. Fruits ellipsoid, usually 

purplish to black. ............................................................................... 2. G. cyanocarpa 

Leaflets sometimes obscurely toothed. Inflorescences narrow, somewhat elongate. 

Fruits globose, white to yellowish or pinkish. Cultivated ............................................... . 

G. parviflora (Sims) Little 

Phytologia 2 (1948) 463. Basionym: Limonia parvijlora Sims, Bot. Mag. (1823) t. 2416. 

Synonym: Glycosmis citrifolia (Willd.) Lindl., TrailS. Hort. Soc. Lond. 6 (1826) 72. 

Native to S China and Indo-China. Introduced to horticulture and botanical gardens in 

various parts of the World. 

1. Glycosmis chlorosperma (Blume) Spreng. 

(Greek, khloros = green, sperma = seed) 

Syst. Veg. cd. 16,4 (1827) 162; Narayanaswami l.c. 40; Masamune I.c. 359; Backer & BakhuizenJ 

I.c. 102; Swing1e I.c. (1967) 207; Stone I.c. (1972) 382, l.c. (1978) 93, I.c. (1985) 2; Anderson I.c. 

308; Corner I.c. 666; Whitmore, Tantra & Sutisna l.c. 306. Basionym: Cookia chlorosperma Blume. 

I.c. (1825) l35. Type: Blume, s.n., Java, Mt. Sa1ak (L). Synonyms: Glycosmis monticola Ridl., J. 

Str. Br. R. As. Soc. 75 (1917) 14; G. malayana Ridl.l.c. (1917) 12. 

Shrub or small tree. Leaves pinnate, rarely unifoliolate, opposite or alternate, glabrous, 

~vith or without axillary lanceolate paraphylls to 2.5 cm long; leaflets (1-)3-5(-7), ovate to 

372


oblong-lanceolate, or narrowly so, 5-20(-30) x 2-9(-14) cm, leathery; base acute, margin 

entire, apex acuminate; lateral veins 5-11 pairs, rather distant, strongly inarching near the 

margin, depressed above, conspicuously raised below; petiolules 3-5 mm long. Inflorescences 

pseudoterminal cymose-panicles, 2-16 cm long, branches rusty hairy, buds small, 

minutely hairy. Flowers at tips of short branchlets, 5-merous; sepals ovate, to 2 mm long, 

margin hyaline, ciliate; petals whitish, obovate-oblong, to 6 mm long, rusty hairy below, 

margin membranous; stamens 10, alternating long and short, anthers ovate-cordate, 

sometimes minutely gland-tipped; disc ring-like, yellow; ovary (4-)5-carpellate, glabrous, 

flask-shaped, on a thick gynophore, style short, equalling the ovary in length. Fruits 

globose, 10 -12.5 mm long. 


1. Leaflets ovate, 5-7 cm long, lateral veins 5 pairs. Inflorescences 2-3 cm long .............. . 

var. bidiensis Stone 

I.c. (1985) 3. Type: 1. & MS. Clemens 20663, Sarawak, Kuching, Bidi Cave (holotype K). 

Small tree. Leaves 3-5-foliolate; leaflets ovate, 5-7 x 2-2.3 cm; lateral veins 5 

pairs. Inflorescences 2-3 cm long. Flowers small; buds 2-3 mm long; anthers 

glandular; disc distinctly constricted, 0.8-0.9 mm wide; ovary 4-5-carpellate. 

Endemic to Sarawak, where it is known only from the type locality, on limestone. 

Leaflets oblong-lanceolate to elliptic, 5-30 cm long, lateral veins 9-11 pairs. Inflorescences 

longer than 5 cm ................... . 

2. Leaflets oblong-lanceolate, mostly less than 15 cm long. Inflorescences 7.5-11 cm 

long ..................................... . 

var. chlorosperma 

Shrub or small tree to 10 m tall and 15 cm diameter. Leaves 3-6-foliolate; leaflets 

oblong-lanceolate, 5-15(-20) x 2-8.5 cm; base sometimes slightly asymmetric; 

lateral veins 9-10 pairs. Inflorescences 7.5-11 cm long. Flowers small; sepals c. 

1 mm long; petals c. 2.5 mm long; ovary 5-carpellate, style almost lacking. Vernacular 

name: segera (Malay). Throughout the range of the species. Widespread in Sabah 

and Sarawak, in mixed dipterocarp forest on fertile clay loam soils, especially on 

basic volcanic substrates, occasionally on limestone, from near sea-level to 800 m. 

Leaflets oblong-elliptic, 15-30 cm long. Inflorescences generally larger, to 16 cm long ......... . 

var. elmeri (Merr.) Tanaka 

Med. Rijksherb. 69 (1931) 3; Stone l.c. (1978) 94, l.c. (1985) 3. Basionym: Glycosmis 

elmeri Merr., Philip. J. Sc. 30 (1926) Bot. 400, Merrilll.c. (1929) 113, Masamune I.c. 359. 

Type: Ramos & Edano 44150, Philippines, Sulu Archipelago, Tawitawi (UC). 

Shrub or small tree to 10 m tall and 20 cm diameter. Leaves 3-4-foliolate; leaflets 

oblong-elliptic, 15-20(-30) x 6-9(-14) cm; base broadly acute; lateral veins 9-11 

pairs. Inflorescences 6-16 cm long, side branches to 5 cm long. Flowers sessile, 

buds brownish. Philippines (Sulu Archipelago), Borneo. Common in Sabah and 

Sarawak, occurring in lowland and hill forest, usually on ridges arid slopes, and 

occasionally on limestone, from near sea-level to 1280 m. 

373


Distribution. S Burma, S Thailand, Peninsular Malaysia, Sumatra, Java, Borneo (Sabah, 

Sarawak, Kalimantan) and S Philippines. A variable species with seven recognised 

varieties, three of which occur in Sabah and Sarawak. 

2. Glycosmis cyanocarpa (Blume) Spreng. 

(Greek, kuanos = dark blue, karpos = fruit) 

I.c. (1827) 161; Kurz l.c. (1877) 184; Masamune l.c. 359; Backer & BakhuizenJ I.c. lO2; Swing1e 

l.c. (1967) 207; Stone l.c. (1978) 97, I.c. (1985) 5; Whitmore, Tantra & Sutisna I.c 306. Basionym: 

Cookia cyanocarpa Blume I.c. (1825) 136. Type: Blume, s.n., Java, Mt. Salak (L). Synonyms: 

Glycosmis longifolia Tanaka, Bull. Soc. Bot. Fr. 75 (1928) 709; G. cymosa (Kurz) Narayanaswami 

I.c.26 

var. platypbylla (Merr.) Stone 

I.c. (1978) 98, l.c (1985) 6. Basionym: Glycosmis platyphylla Merr., Philip. J. Sc. 12 (1917) Bot. 

273. Type: Wenze11611, Philippines, Leyte (US). Synonym: Glycosmis clemensii Tanaka I.c. (1931) 

4. 

Shrub or small to medium-sized tree to 15 m tall and 30 cm diameter. Leaves 3-4-foliolate, 

alternate, 11-20 cm long; leaflets (1-)3-9, elliptic or lanceolate, 4-30 x 1-10 cm, glabrous 

or rusty-scurfy,' sometimes only on midrib, below; base acute, margin usually entire or 

sometimes finely toothed, apex caudate-acuminate; lateral veins 5-10(-20) pairs, sometimes 

strongly raised below, inarching near the margin; petiolules 2-4 mm long. Inflorescences 

predominantly axillary corymbs, 1-12 cm long, open-branched, branches spreading, hairy, 

with linear, leaf-like bracts. Flowers 4-5-merous; buds glabrous, c. 5 mm long; sepals 

broad, rounded, c. l.3 mm long, hairy, especially towards apex, margin rusty-ciliate; petals 

oblong to oblong-obovate, 3.5-4 mm long, hairy, especially towards apex; stamens 10, 

nearly equal to unequal, filaments broad, narrowed at apex, anthers ovate, glandularapiculate; 

disc elevated, minutely lobed or glandular; ovary 4-5-carpellate, ellipsoid, 

constricted at base and in the middle, glabrous, not glandular, seated on a well-developed 

gynophore, style short, stigma broad. Fruits oblong or ellipsoid, 1-l.5 cm long, purplish. 

Seed 1 per fruit. 

Distribution. India, Nepal, SW China, Burma, Indo-China, and throughout W Malesia 

(Sumatra, Peninsular Malaysia, Java, Borneo and the Philippines). A variable species 

comprising 10 varieties, one of which (var. platyphylla) occurs in Borneo, Philippines, and 

the Talaud Islands. In Sabah and Sarawak (Lundu district), var. platyphylla occurs on flat 

and undulating land in primary and secondary lowland forest, from near sea-level to 200 

m; also in Kalimantan. 

3. Glycosmis lanceolata (Blume) Spreng. 

(Latin, lanceolatus = spear-shaped; the leaves) 

Cat. Hart. Bog. (1866) 208; Narayanaswami I.c. 61; Stone I.c. (1985) 10; Whitmore, Tantra & 

Sutisna I.c. 306. Basionym: Sclerostylis lanceolata Blume l.c. (1825) 134. Type: Blume, s.n., 

374


Batavia (L). Synonyms: Glyeosmis simplieifolia Spreng. I.e. (1827) 162; G. montana Pierre, Fl. For. 

Cochinch. 4 (1893) 285; G. greenei var. simplex Stone I.e. (1978) 90. 

Shrub or small to medium-sized tree to 18 m tall and 25 cm diameter. Branches with shiny 

and cracked bark. Leaves 1-5-foliolate, alternate; leaflets, oblong-ovate or oblanceolate, 

8.5-17 x 3-8 cm; base rounded, margin entire, apex shortly acuminate or emarginate; 

lateral veins 8-12 pairs; petiolules 3-4 mm long, 7-9 mm long in unifoliolate leaves. 

Inflorescences short, axillary cymose-panicles, 1-3 cm long, branches short, rusty scurfyhairy. 

Flowers small, 5-merous; sepals 1-1.5 mm long, glabrous below; petals longer, 

oblong-obovate, rusty scurfY-hairy below, margins hyaline; stamens 10, alternating long 

and short, anthers oblong-cordate, connectives obscurely glandular; ovary (2-)3- 

carpellate, ellipsoid-ovoid, glabrous, distinctly papillate glandular, seated on a welldeveloped 

gynophore, tapering into style, stigma 3-10bed. Fruits globose. 

Distribution. Indo-China, Hainan, Sumatra, Java, Borneo, S Philippines, and the Lesser 

Sunda Islands. Widespread in Sabah on ridges and slopes in lowland forest, occasionally on 

limestone, from near sea-level to 240 ID. In Sarawak, known from only one locality in 

Kuching district (Stone 13739, Padawan, Bt. Pa'it). Also known from Kalimantan. 

4. Glycosmis Iongisepala Stone 

(Latin, longus = long, sepalum = sepal) 

l.e (1985) 11; Whitmore, Tantra & Sutisna I.e. 306. Type: J. & MS. Clemens 20001, Sarawak, Mt. 

Pueh (holotype K). 

Small tree. Leaves 5-7-foliolate, to 30 cm long; leaflets ovate-lanceolate or elliptic, 7-16 x 

2.8-6 cm; base acute, margin entire, apex acuminate; lateral veins 6-8 pairs; petiolules 3-8 

mm long. Inflorescences pseudoterminal cymose-panicles, elongate or pyramid-shaped, to 

13 cm long. Flowers 5-merous; sepals narrowly triangular, c. 2.5 mm long, surfaces hairy, 

margin ciliate; petals c. 3 mm long, hairy below; stamens 10, anthers c. 1 mm long, 

glandular-apiculate, connective glandular dorsally; ovary (4-)5-carpellate, to 2 mm long, 

glabrous, glandular, seated on a well-developed gynophore. Fruits sub ellipsoid, c. 7.5 mm 

long. Seeds 2 per fruit. 

Distribution. Endemic to Sarawak; known only from the type locality, in forest, at 1000- 

1400 m. 

5. Glycosmis macrantha Merr. 

(Greek, makros = large, anthos = flower) 

Fig. 5. 

I.e. (1929) 114; Masamune I.e. 360; Swingle I.e. (1967) 207; Stone I.e. (1978) 92, I.e. (1985) 11; 

Whitmore, Tantra & Sutisna I.e. 306. Type: Elmer 21456, British North Borneo, Tawau (holotype 

VC; isotypes MO, VS). Synonym: Glyeosmis oliveri Stapf ex Ridl., Kew Bull. (1930) 80, Masamune 

I.e. 360. 

Small to medium-sized tree to 20 m tall and 20 cm diameter; bole to 12 m tall. Leaves 5-9- 

foliolatc, 17-30 cm long, alternate; leaflets oblong to elliptic, 7-14 x 3-6 cm, glabrous, 

375


chartaceous; base acute, apex acuminate, acumen 1-2 cm long; lateral veins 5-9 pairs, 

strongly raised below; petiolules 4-7 mm long. Inflorescences terminal or axillary cymes. 

Flowerslarge, 4-5-merous, whitish, on short stalks or nearly sessile; sepals orbicular, 2.5- 

3 mm long, sparsely hairy or glabrous; petals 5, oblong, glabrous, c. 7 x 3 mm; stamens 10, 

equal, filaments thick, c. 4 mm long, anther connectives dorsally glandular; ovary 5- 

carpellate, glabrous, seated on a well-developed gynophore, style cylindrical. Fruits ovoid, 

c. 1 cm long. 

Distribution. Endemic to Borneo. In Sabah, widespread and abundant in primary and 

secondary forest, predominantly on hillsides and ridges, and often in disturbed habitats, to 

1400 m. In Sarawak, uncommon and rarely collected. Also in Kalimantan. 

6. Glycosmis sapindoides Lindl. in Wall. ex Oliver 

(resembling the genus Sapindus) 

J. Linn. Soc. Bot. 5, Supp!. 2 (1861) 38; Hooker! I.e. (1875) 501; King I.e. 217; Ridley I.e. (1922) 

351; eraib I.e. 225; Narayanaswami I.e. 55; Backer & Bakhuizen! I.e. 102; Swingle I.e. (1967) 207; 

Stone I.e. (1972) 381, I.e. (1985) 18; Whitmore, Tantra & Sutisna I.e. 307. Type: Wallieh Cat. 6376, 

Penang (lectotype K). Synonyms: Glyeosmis cyanoearpa var. sapindoides (Lind!.) Kurz, J. Bot. 14 

(1876) 34; G. elata Rid!., J. Fed. Ma!. St. Mus. 10 (1920) 130; G. maerophylla (Blume) Miq., F1. 

Ind. Bat. 1, 2 (1859) 522. 

var. sapindoides 

Shrub or small tree to 10 m tall and 10 cm diameter. YouYJg branches yellow, shiny. Leaves 

pinnate, alternate; leaflets (3-)5-9, ob lanceolate, 14-22 x 4-8 cm; base acute, margin 

entire, apex acuminate; lateral veins 9-11 pairs, prominently raised below; petiolules to 3 

mm long. Inflorescences axillary cymose-panicles, 1.5-2(-7.5) cm long; buds rusty hairy. 

Flowers small, 5-merous, in clusters at tips of short branchlets; sepals broadly ovate, to 0.5 

mm long, hairy below, margin ciliolate; petals oblong-elliptic, c. 2.5 mm long, hairy below; 

stamens 10, alternating long and short, filaments slender, c. 1.5 mm long, glabrous, anthers 

crescent-shaped, gland-tipped; ovary (2-)3-carpellate, ellipsoid, densely nJsty hairy, stigma 

broad. Fruits ellipsoid. 

Distribution. Mainland SE Asia and W Malesia, to Papua New.Guinea and Australia. Of 

the three known varieties, only the commonest (var. sapindoides) occurs in Sabah. This 

variety occurs in the western and central parts of the range of the species, as far east as 

Papua New Guinea. In Sabah, known only from Kota Kinabalu district (Pulau Sipanggar) 

where it is found on ridge tops and hillsides in primary forest, to 200 m. Also in 

Kalimantan. 

7. Glycosmis superba Stone 

(Latin, superb us = splendid; the large, attractive leaves) 

I.e. (1978) 95, I.e. (1985) 20; Anderson I.e. 308. Whitmore, Tantra & Sutisna I.e. 307. Type: 

Anderson, Tan & Wright S. 26059, Sarawak, Ulu Sg. Sekaloh (SAR). 

376


2mm [ 

c 

>om r 

A 

Fig. 5. Glycosmis macrantha. A, fruiting leafy twig; B, part of inflorescence; C, flowers. CA, from 

SAN 57109, B & C from SAN 44669.) 

377


'J,,/ vi' ,"~~ -~ 1 

\ ... "" 3 cm 

), 

"f,f'/" 

Fig. 6. Lunasia amara. Fruiting leafy twig, (From SAN 92007.) 

378

RUT ACEAE (lONES) 

Shrub or small tree to 10 m tall and 10 cm diameter. Leaves large, opposite, 3-6-foliolate, 

to 33 cm long; leaflets elliptic-oblanceolate, 20-31 x 8-12 cm, leathery; base cuneate, 

margin obscurely wavy to subentire, apex acuminate-caudate; lateral veins 10-14 pairs; 

petiolules ]-2 cm long. Inflorescences pseudoterminal, pyramid-shaped, densely cymosepaniculate, 

subglabrous, 33-45 mm long. Flowers 5-merous; sepals broadly orbicularovate, 

c. 3 mm long, glabrous below, margin hyaline; petals at least 6 mm long, glabrous, 

glandular; stamens 10, anther connectives 4-6-glandular, apex glandular-apiculate; disc 5- 

lobulate; ovary 5-carpellate, cylindrical, glabrous, stigma 5-lobed, style glandular. 

Distribution. Endemic to Sarawak (Kuching, Bintulu and Miri districts) and Brunei. 

Ecology. Found on ridges and undulating ground in primary mixed dipterocarp forest and 

secondary forest on sandy humult ultisols, from near sea-level to 100 m. 

6. LUNASIA Blanco 

(Tagalog, lunas; a native name for L. amara) 

Fl. Filip. ed. I (1837) 783; Merrilll.e. (1929) 113; Masamune I.e. 360; Backer & BakhuizenJ I.e. 

99; Hartley, J. Am. Arb. 48 (1967) 460; Anderson I.e. 308; Perry I.e. 366. 

Erect shrubs or small trees; dioecious. Branches unarmed. All vegetative and reproductive 

parts with grey to reddish brown scale-like and/or star-shaped hairs. Leaves alternate, 

simple; leaf blades pinnately veined, leathery, becoming stiff when older; petioles wingless, 

swollen at apex. Inflorescences axillary panicles, flowers in small globose-clusters, 3-6 

mm in diameter. Flowers unisexual, 3-merous, fragrant; sepals 3, free, valvate; petals 3, 

free, valvate, white to greenish or yellowish; stamens 3, opposite the sepals, rudimentary in 

female flowers, anthers dorsifixed; ovary 3-carpellate, carpels fused at base, rudimentary in 

male flowers, ovules 1 per locule; styles 3. Fruits of 1-3 dehiscent follicles, the undeveloped 

follicles persistent in fruit; pericarp dry at maturity; endocarp cartilaginous, discharged 

when the follicle dehisces. Seeds 1 per locule; endosperm lacking; cotyledons oily. 

Distribution. One species; E Java, Borneo, Philippines, Celebes, Papua New Guinea and 

Australia (Queensland). 

Ecology. Occurs in well-drained, primary and secondary rain forests, thickets, and garden 

regrowth, from near sea-level to 900 m. 

Lunasia amara Blanco Fig. 6. 

(Latin, amarus = bitter; referring to the alkaloid content) 

I.e. 783; Merrill I.e. (1929) 113; M!lsamune I.e. 360; Backer & Bakhuizen J I.e. 99; Hartley I.e. 

(1967) 464; Anderson I.e. 308; Perry I.e. 366. Type: Eseritor BS 20776, Philippines, Luzon Island 

(neotype A). Synonyms: Piloearpus amara (Blanco) Blanco I.e. (1845) 540; Lunasia retieulata 

Elmer, Leafl. Philip. Bot. 4 (1912) 1511; L. gigantifolia Merr., Philip. J. Sc. 21 (1922) Bot. 519. 

379


var. amara 

Shrub or small tree to 15 m tall (usually smaller), 30 cm diameter, sparsely branched. 

Leaves crowded toward the bran ch let tips; blades oblanceolate to obovate or elliptic, 6-60 

x (up to) 18 cm; base cuneate to narrowly rounded or cordate, margin subentire to sinuate 

or coarsely dentate, apex rounded to acuminate; lateral veins 9-35 pairs, strongly raised 

below; petioles 1.5-15 cm long. Male inflorescences to 28 cm long and 8 cm wide; sepals 

ovate, minute; petals obovatc-acuminate, c. 1 mm long; staminal filaments glabrous. Female 

inflorescences to 25 cm long and 2 cm wide; sepals broady ovate, 1-1. 5 mm long; petals 

ovate-acuminate, c. 2 mm long; staminodes 3; styles fused at base, stigmas flattened and 

spreading over the tops of the carpels. Fruits obovate follicles, 6-15 x 5-10 mm, flattened 

laterally, ribbed on the sides, with a beak to 5 mm long, densely hairy. Se£ds nearly 

obovoid; testa dark brown to reddish, papery. 

Distribution. One variety (var. amara) with the same range of distribution as that of the 

genus and species, occurs in Sabah and Sarawak. The other known variety, babuyanica 

(Merr.) Rartley, is endemic to the Babuyan Islands in the Philippines. In Sabah, var. amara 

is widespread and found in a variety of habitats, including offshore islands, ridges and 

slopes in primary forest, and forest on ultramafic soils, from near sea-level to 900 m. In 

Sarawak, frequent on limestone cliffs and slopes at low elevations (Miri and Baram 

districts). 

Uses. Bark, leaves, and seeds have been used medicinally to treat skin diseases, swollen 

limbs, snake bite, and stomach ailments (Philippines, Indonesia). A number of alkaloids 

are reported from the plant. Lunasin and lunacrin, two alkaloids extracted from the bark, 

were shown to have negative effects on the voluntary and smooth muscles, blood vessels, 

and hearts of laboratory animals (Wirth, J. Am. Pharm. Assoc. 20 (1931) 1254), causing 

death due to the simultaneous cessation of the respiratory and circulatory systems. 

7. MACLURODENDRON T.G. Hartley 

(Floyd A. McClure, 1897-1970, American botanist and plant explorer) 

Gard. Bull. Sing. 35 (1982) l. 

Small to medium-sized trees; dioecious. Branches unarmed. Young branchlets hairy, 

trichomes brownish to rust-coloured. Leaves opposite, unifoliolate; blade pinnately veined, 

margin entire; petioles wingless, swollen at the apex, articulated with the blades. Inflorescences 

axillary, paniculate or racemose. Flowers unisexual, 4-merous, round in bud; sepals 

4, triangular, fused at base, valvate; petals 4, ovate, free, imbricate, white or greenish or 

yellowish; stamens 8, unequal, alternately long and short, filaments glabrous, nearly linear, 

curved inward, anther ovoid to ellipsoid, dorsifixed, pollen lacking in female flowers; disc 

irregularly 8-lobed; ovary 4-carpellate, shallowly 4-lobed, glabrous, rudimentary in male 

flowers, 2 ovules per locule, style straight, stigma capitate, 4-lobed. Fruit a 4-loculate 

drupe, glabrous; exocarp somewhat leathery, glandular; mesocarp spongy, thin when dry; 

endocarp parchment-like, shiny. Seeds ovoid to kidney-shaped, dark brown to black, shiny, 

380


1-2 per locule; outer testa usually spongy, inner testa bony; endosperm fleshy; embryo 

straight or bent. 

Distribution. 6 species; Thailand, Sumatra, Peninsular Malaysia, Borneo, the Philippines, 

Vietnam, and Hainan Island. One species (M porteri) commonly occur in Sabah and 

Sarawak, while M parviflorum is endemic to Sarawak and M pubescens to Sabah. 

Ecology. Mainly in well-drained primary rain forests and occasionally in secondary forests 

and heath forests, from near sea-level to 1500 m. 

Taxonomy. Several species were previously described under Acronychia (e.g., A. porteri). 

The two genera are similar in a number of characters including the opposite leaves, 4- 

merous flowers, 2-ovulate locules, and drupaceous fruits. However, Maclurodendron differs 

in its exclusively unifoliolate leaves (sometimes trifoliolate in Acronychia), unisexual 

flowers, imbricate petals, glabrous staminal filaments, and glabrous ovaries and fruits, 

never with septicidal fissures. 

Key to Maclurodendron species 

1. Fully developed leaf-blades hairy below, especially on themidrib and lateral veins 

...... 3. M. pubescens 

Fully developed leaf-blades glabrous or nearly so ......................................................... 2 

2. Inflorescences 1-3 cm long. Petals c. 1.5 mm long. Seed surface irregularly roughened 

......................................................................................................... 1. M. parviflorum 

Inflorescences 2-15 cm long. Petals 2-2.5 mm long. Seed surface minutely reticulate 

................................................................................................................. 2. M. porteri 

1. MacIurodendron parviflorum T. G. Hartley 

(Latin, parvus = small,jlos = flower) 

l.c. (1'.182) 14. Type: Anderson S. 25426, Sarawak, Kuching district (holotype L; isotype SAR). 

Small tree to 5 m tall. Branchlets glabrous or nearly so. Leaves elliptic or ob ovate to 

oblanceolate, 9-16 x 3-9 cm, thinly leathery, drying pale green, glabrous or nearly so; 

base narrow, apex acuminate or occasionally rounded; lateral veins 7-11 pairs; intercostal 

veins and reticulations faint; petioles 1-2 cm long. Inflorescences 1-3 cm long; peduncle 

axis and branches sparsely hairy to glabrous, pedicels sparsely hairy, 1-3 mm long. 

Flowers 1-1.5 mm wide in bud; sepals nearly 1 mm long, sparsely hairy; petals c. 1.5 mm 

long, glabrous. Fruits c. 7 mm in diameter, nearly round to ovoid, shallowly 4-10bed, acute 

at the apex. Seeds c. 5 mm long, surface irregularly roughened; outer testa not spongy. 

Distribution. Endemic to Sarawak, where it is known from only four collections in 

Kuching district. 

Ecology. In primary heath (kerangas) forest and secondary forest on podzolised soils, at 

Iow elevations. 

381


2. Maclurodendron porteri (Hook./) T.G. Hartley 

(G. Porter, first Curator of Waterfall Garden, Penang) 

Fig. 7. 

I.e. (1982) 8. Basionym: Aeronyehia porteri Hook.! (1875) 498; Anderson I.e. 307. Type: Maingay 

Kew Distr. No. 280, Penang (lectotype K; iso1ectotypes BM, GH, L). Synonyms: lambolifera porteri 

(Hook.f) Kuntze, Rev. Gen. PI. 1 (1891) 102; Melieope unifoliolata Merr., Philip. J. Sc. l3 (1918) 

Bot. 74, I.e. (1921) 314, Masamune I.e. 360. 

Small to medium-sized tree to 25 m tall and 40 cm diameter; bole to 12 m tall. Branchlets 

glabrous or nearly so. Leaves obovate to ob lanceolate or elliptic, 5-24 x 2-10 cm, thinly 

leathery, drying pale brown to dark brown, glabrous or nearly so; base narrow or acute, 

apex abruptly acuminate or occasionally rounded or acute; lateral veins 6-11 pairs; 

intercostal veins and reticulations faint; petioles 0.7-5 cm long Inflorescences 2-15 cm 

long; peduncle axis and branches sparsely hairy to glabrous, pedicels nearly glabrous, 1-7 

mm long. Flowers 1.5-2 mm wide in bud; sepals 0.5-1 mm long, glabrous to hairy; petals 

2-2.5 mm long, glabrous to hairy. Fruits 6-11 mm in diameter, nearly round to ovoid, often 

shallowly 4-lobed, often tipped. Seeds 4-8 mm long, surface minutely reticulate, outer 

testa spongy. 

Vernacular name. Sarawak-rawang (Malay). 

Distribution. Burma, Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, Borneo, 

and the Philippines. Widespread in Sabah and Sarawak. Also in Brunei and Kalimantan. 

Ecology. Locally on hillsides and ridges in primary mixed dipterocarp forest and occasionally 

in secondary forest and heath (kerangas) forest (in Sarawak), on low nutrient organic soils, 

from near sea-level to 1400 m. 

Uses. The timber, while not durable, has been used for building purposes (Peninsular 

Malaysia). 

3. Maclurodendron pubescens T. G. Hartley 

(Latin, pubescens = hairy; the leaves and floral parts) 

I.e. (1982) 1l. Type: Patriek SAN 26359, Sabah, Sandakan district (holotype L; isotype K). 

Small to medium-sized tree to 30 m tall and 35 cm diameter; bole to 18 m tall. Branchlets 

hairy, becoming glabrous. Leaves obovate to oblanceolate or elliptic, 11-23 x 5-11 cm, 

thinly leathery, drying pale greenish brown to brown, pubescent below especially on the 

midrib and veins, glabrous above; base narrow, apex acuminate to rounded; lateral veins 8- 

11 pairs; intercostal veins and reticulations faint; petioles 1.5-5 cm long. Inflorescences 3- 

11 cm long; peduncle axis and branches sparsely to densely hairy; pedicels hairy, 3-5 mm 

long. Flowers 1.5-2 mm wide in bud; sepals c. 7 mm long, hairy; petals c. 2 mm long, 

hairy below, glabrous to sparsely hairy above. Fruits 10-12 mm in diameter, nearly round 

to ovoid, shallowly 4-lobed, acute at the apex. Seeds 5-7 mm long, surface minutely 

reticulate; outer testa spongy. 

382


A If 

I 

2 ern 11 B 

Fig. 7. Maclurodendron porteri. A, flowering leafy twig; B, part of inflorescence; C, fruits. (A & B 

from SAN 34008, C from SAN 55734.) 

383


Distribution. Endemic to Sabah, where it is found in Sandakan (especially common in 

Leila and Sepilok-Kabili FR), Keningau, and Lamag districts. 

Ecology. On ridges and hillsides in primary forest, from near sea-level to 140 m. 

8. MELICOPE J.R. Forst. & G. Forst. 

(Greek, meli = honey, kope = a cutting; 

the emarginate lobes of the nectar -secreting disc) 

serang (Malay) 

Char. Gen. PI. (1776) 55; Hooker f I.e. (1875) 491; Kurz I.e. (1877) 181; King I.e. 212; ATlderson 

I.e. 308; Hartley, Gard. Bull. Sing. 34 (1981) 91, Sandakania 4 (1994) 47; Hartley & Stone, Taxon 

38 (1989) 199. 

Shrubs or small to tall trees; dioecious in unisexual plants. Branches unarmed. Leaves 

opposite or whorled, trifoliolate and/or unifoliolate, petioles wingless; leaflets leathery or 

thinly so, margin mostly entire, articulated at the base. Inflorescences cymes or densely 

flowered panicles or solitary, axillary or on branchlets below leaves, rarely terminal or on 

stems. Flowers small, bisexual or unisexual, 4-merous; sepals 4, fused at base; petals 4, 

free, valvate or imbricate, white or cream to greenish or yellowish, rarely pink; stamens 4 

or 8, rarely 4-8, free, rudimentary in female flowers; disc cushion- to ring-like or cupular; 

ovary 4-carpellate, carpels fused completely or only at base, rudimentary in male flowers, 

ovules (1-)2 per locule, styles united, rarely stylar branches divergent, stigma small to 

capitate or peltate, often lobed, or 4-branched. Fruits of J-4 basally fusedfoUicles, erect or 

spreading at maturity, grading to a 4-locular capsule; endocarp cartilaginous, remaining 

attached in the dehisced follicle, fused to or separate from epicarp. Seeds 1-2 per locule, 

not expelled from mature fruit at dehiscence; testa thick, hard, covered by a shiny, black 

epidermis; endosperm abundant; embryo straight or nearly so; cotyledons flat. 

Distribution. About 230 species ranging from Madagascar to India, S China, throughout 

Malesia, Polynesia, the Hawaiian Islands, Australia and New Zealand. 14 species in Sabah 


Taxonomy. In his revision of the genus Tetradium Lour., Hartley (l.c. 1981) redefined 

Melicope and suggested transferring to it many of the species placed in the genus Euodia J. 

R. Forst. & G. Forst. As characterised by Hartley, Melicope and Euodia are largely 

differentiated on the basis of their seeds (smooth, shiny and remaining attached in the 

dehisced follicle in Melicope, and dull, rough and discharged from the follicle in Euodia), 

floral characteristics (bisexual or unisexual flowers with 4 or 8 stamens in Melicope, and 

bisexual flowers with 4 stamens in Euodia), and geographical ranges (Euodia has a 

narrower distribution, occurring in New Guinea, NE Australia, and east to Samoa and 

Hawaii). As a consequence of this new generic delimination, there are no representatives of 

the genus Euodia in Sabah or Sarawak. 

384


Key to Melicope species 

I. Stamens 8. Seeds attached to dehiscent fruit by a partially detached strip of pericarp 

and/or raphe. Leaves trifoliolate and/or unifoliolate ......................................................... 2 

Stamens 4. Seeds attached to dehiscent fruit by a funiculus. Leaves mostly trifoliolate, 

or (in M subunifoliolata) sometimes uniformly unifoliolate ...................................... .4 

2. Sepals shorter than 1 mm. Follicles 2-5 mm long. Leaves mostly trifoliolate ............. . 

...... . 14. M. triphylla 

Sepals 1-2 mm long. Follicles 10-13 mm long. Leaves trifoliolate and/or unifoliolate ....... 3 

3. Terminal bud\glabrous. Petiolules 1-2 mm long ....................................... 9. M. jugosa 

Terminal bud hairy. Petiolules 3-15 mm long ........................................... 12. M. sororia 

4. Flowers bisexual; stigma not lobed or wavy. Inflorescences axillary and/or on 

branchlets below leaves .............................................................................................. 5 

Flowers usually unisexual; stigma lobed or wavy. Inflorescences axillary ................... 6 

5. Leaflet-margin entire. Pedicels 4-8 mm long. Petals densely hairy above. Follicles 4- 

6 mm long .......................................................................................... 2. M. bonwickii 

Leaflets-margin entire or (in cultivated forms) lobed or wavy. Pedicels less than 3 mm 

long. Petals glabrous. Follicles 2-3 mm long ....................................... 5. M. denhamii 

6. Trichomes mostly simple ........................ . ......... 7 

Trichomes mostly in bundles or star-shaped ........... ... 13 

7. Leaflet-base cordate to rounded; petiolules lacking ............................... l0. M. latifolia 

Leaflet-base rounded or acute to narrowly tapered; petiolules lacking or up to 20 mm 

long. 

8. Petals 1.3-1.8 mm long. Leaflets l.5-5.5 cm wide ....................................................... 9 

Petals 1.5-3 mm long. Leaflets 2.5-21 cm wide ............................................................. 10 

9. Leaflets glabrous or midrib sparsely hairy below. Follicles 7-9 mm long; epicarp 

subfleshy, becoming glabrous ............................................................ 3. M. clemensiae 

Leaflets glabrous above; midrib hairy to velvety below. Follicles 10-10.5 mm long; 

epicarp dry, hairy ..................................................................... 13. M. subunifoliolata 

10 Petals silky-hairy above. Raphe of seeds contorted ............................. : ... .4. M. confusa 

Petals glabrous or sparsely hairy to hairy above, especially on lower half. Raphe of seeds 

not contorted ........................................................................................................... 11 

11. Terminal leaflets obovate to broadly obovate, 7.5-16.5 cm long; apex abruptly 

acuminate; lateral veins 9-15 pairs. Follicles 3-4 mm long ........................ 6. M. glabra 

Terminal leaflets elliptic to obovate, 3.5-46 cm long; apex usually acuminate; lateral 

vei ns 8-24 pairs. Follicles 3-12 mm long ................................................................. 12 

385


2. Melicope bonwickii (F. Mue11.) T.G. Hartley 

(l Bonwick, student of Australian geography) 

I.e. (1994) 56. Basionym: Euodia bonwiekii F. Mue1l., Fragm. 5 (1865) 56. Type: Dallaehy, s.n., 

Australia, Queensland, Rockingham Bay (holotype MEL). Synonyms: Euodia speciosa Rchb. f & 

Zoll. ex Teijsm. & Binnen., Nat. Tijd. Ned. Ind. 29 (1867) 255; E. villamilii Merr., Philip. J. Sc. 9 

(1914) Bot. 296. 

Tree to 40 m tall. Young branchlets glabrous or rarely sparsely hairy; terminal bud 

appressed hairy. Trichomes simple. Leaves opposite, trifoliolate, petioles 2-14.5 cm long; 

leaflets elliptic to obovate, 10-30 x 5-15 cm, glabrous or sparsely hairy on veins below; 

base acute or narrowly tapered and sometimes asymmetric, apex acuminate; lateral veins 

14-24 pairs, sometimes raised above, sometimes with aXillary cavities below; petiolules 

lacking or to 4 mm long. Inflorescences on branchlets below leaves, rarely axillary, 

glabrous to sparsely hairy, 3.5-10 cm long; pedicels 4-8 mm long. Flowers bisexual; sepals 

nearly round, to 2 mm long, glabrous to sparsely hairy below; petals pin.k or rarely white, 

ovate to elliptic or elliptic-oblong, 4-5.5 mm long, densely hairy above; stamens 4, 

filaments glabrous; disc and ovary hairy to densely hairy, style glabrous, stigma capitate. 

Fruits of nearly round to obovoid follicles, 4-6 mm long; epicarp dry, sparsely hairy to 

almost glabrous; endocarp glabrous. Seeds nearly round to ellipsoid or nearly hemispherical, 

3-4.5 mm long; funiculus 1-3 mm long. 

Distribution. Java, Borneo, Philippines, and east to Papua New Guinea and Australia. In 

Sabah, in primary and secondary forest, from near sea-level to 750 m; also occurs in 

Kalimantan. 

Uses. The bark is reported to be used in the treatment ofleech bites (Indonesia). 

3. Melicope clemensiae T. G. Hartley 

(Mary S. Clemens, 1873-1968, plant collector ofthe Malesian region) 

I.e. (1994) 64. Type: Aban Gibot SAN 60767, Sabah, Mt. Kinabalu Park (holotype L; isotype SAN). 

Shrub or tree to 15 m tall. Young branchlets and terminal buds glabrous, or nearly so, to 

velvety. Trichomes mostly simple. Leaves opposite, trifoliolate (rarely unifoliolate); petioles 

1.5-8 cm long; leaflets elliptic to obovate or oblanceolate, 5-16 x 2-5.5 cm, glabrous or 

midrib sparsely hairy below; base acute to narrowly tapered and often asymmetric, apex 

acuminate; lateral veins 11-18 pairs, raised above; petiolules 1.5-12 mm long. Inflorescences 

axillary, laxly flowered, 2.5-16 cm long, glabrous to somewhat velvety; pedicels 2-5 mm 

long. Flowers unisexual; sepals round to ovate-triangular, less than 1 mm long, hairy or 

glabrous below; petals ovate to elliptic, c. 1.5 mm long, glabrous; stamens 4, filaments 

glabrous; disc and ovary nearly glabrous to hairy, style glabrous, stigma weakly 4-lobed. 

Fruits of round to obovoid follicles, 7-9 mm long; epicarp subfleshy, becoming glabrous; 

endocarp glabrous. Seeds round, ovoid or ellipsoid, slightly compressed, 5-7 mm long; 

funiculus 1.5-2 mm long. 

Distribution. Endemic to Sabah and Sarawak; in primary and secondary forest, on ridges 

and slopes, sometimes on limestone, from 650 to 1800 m. 

388


4. Melicope confusa (Merr.) Liu 

(Latin, confusio = disorder; referring to this taxon's long confusion with Euodia glabra) 

Ill. Native Introd. Lign. PI. Taiwan 2 (1962) 876; Hartley I.e. (1994) 59. Basionym: Euodia eon/usa 

Merr., Philip. J. Se. 20 (1922) Bot. 391. Type: Ramos BS 15055, Philippines, Luzon, Laguna 

Province (A). 

Tree to 30 m tall. Young branchlets glabrous to sparsely hairy; terminal buds hairy to 

velvety. Trichomes simple. Leaves opposite, trifoliolate; petioles 2.5-18.5 cm long; leaflets 

elliptic to ob ovate or ovate, 12-35 x 5-8 cm, glabrous to hairy below; base acute to 

narrowly tapered, rarely asymmetric, apex acuminate; lateral veins 11-24 pairs, raised to 

depressed above; petiolules 3-15 mm long. Inflorescences axillary, 6-30 cm long, peduncle 

glabrous to hairy, pedicels 1-3 mm long. Flowers unisexual; sepals nearly round or ovate, 

to 1 mm long, hairy below; petals elliptic, 2.5-3 mm long, silky hairy above; stamens 4, 

filaments sparsely hairy towards base or glabrous; disc and ovary hairy, style hairy at least 

basally, stigma weakly 4-lobed. Fruits of round to ellipsoid follicles, 4.5-6 mm long, fused 

basally and erector diverging; epicarp subjleshy, sparsely hairy or glabrate; endocarp 

glabrous. Seeds round to ovoid or ellipsoid, 3.5-4 mm long; funiculus 1-1.5 mm long, 

raphe contorted. 

Distribution. Borneo, Philippines, Celebes and Moluccas. In Sabah, found mainly in primary 

and secondary forest, from near sea-level to 90 m; no record from Sarawak. 

Uses. The bark is used medicinally for treating enlargement of the spleen (Philippines); a 

decoction of the root or leafy shoot with liquor is ingested to treat hives (Taiwan). 

5. Melicope denhamii (Seemen) T.G. Hartley 

(H. M. Denham, the 19th century British sea captain) 

I.e. (1994) 57. Basionym: Pierasma denhamii Seemen, Fl. Vit. (1865) 33. Type: MeGillivray 46, 

New Hebrides, Aneitum (holotype BM). Synonyms: Euodia tenuistyla Stapf, Trans. Linn. Soc. Bot. 

4 (1894) 137, Merrilll.e. (1921) 314, Masamune I.e. 359; E. ridleyi Hoehr., Icon Bog. 2 (1905) 151; 

E. schullei var. ridleyi (Hoehr.) Lauterb., Bot. Jahrb. 55 (1918) 230; E. suaveolens var. ridleyi 

(Hoehr.) Bakhuizenj, Blumea 6 (1950) 365. 

Shrub or tree to 25 m tall. Young branchlets and terminal buds glabrous to densely hairy 

or velvety. Trichomes simple, star-shaped or in bundles. Leaves opposite, trifoliolate 

(occasionally unifoliolate); petioles 1-16 cm long; leaflets elliptic, 10-19 cm long, sparsely 

hairy to hairy on midrib and veins below; base cuneate to narrowly tapered, apex 

acuminate; lateral veins 15-22 pairs, raised or depressed above; petiolules lacking or up to 

1 cm long. Inflorescences axillary and/or on branchlets below leaves, 2-7 cm long, hairy, 

pedicels 0.2-2.3 mm long. Flowers bisexual; sepals ovate to triangular, to 0.8 mm long, 

glabrous to hairy below; petals ovate-elliptic or elliptic, 1.2-2.2 mm long, glabrous; stamens 

4, filaments glabrous; disc glabrous; ovary hairy, style glabrous to sparsely hairy, stigma 

capitate. Fruits of nearly round follicles, 2-3 mm long; epicarp dryish, glabrous to 

sparsely hairy; endocarp glabrous. Seeds nearly round or hemispherical, 1.5-2.5 mm 

long; funiculus to 1.5 mm long. 

389


rounned or abruptly acuminate, lateral veins 7-11 pairs, sometimes raised above; petiolules 

1-2 mm long; leaflets of unifoliolate leaves elliptic or rarely elliptic-obovate, to 13.5 x 6 

cm, base rounded or rarely acute, lateral veins 9-13 pairs; petiolules lacking, or similar to 

that of trifoliolate leaves. Inflorescences axillary, glabrous to sparsely hairy, 1-2 cm long; 

pedicels to 3.5 mm long. Flowers unisexual; sepals nearly round or ovate, to 2 mm long, 

sometimes ciliolate; petals elliptic, to 4 mm long; stamens 8; stigma capitate, weakly 4- 

lobed. Fruits glabrous, follicles ellipsoid, c. 10 mm long; epicarp dry. Seeds ellipsoid, c. 8 

mm long. 

Distribution. Endemic to Sabah, in forest from 2250 to 2400 m. 

la. Melicope latifolia (DC.) T.G. Hartley 

(Latin, latus = broad, folium = leaf) 

I.c. (1994) 72. Basionym: Euodia latifolia DC.l.c. 724, Merrilll.c. (1921) 314, Masamune l.c. 358, 

Anderson I.c. 308. Type: Doleschall 335, Molueeas, Ambon (neotype W). 

Shrub or tree to 30 m tall. Young branchlets and terminal buds glabrous to softly hairy. 

Trichomes simple. Leaves opposite, trifoliolate (occasionally unifoliolate); petioles 4-29 

cm long; leaflets elliptic to ovate or obovate, 8-37 x 3-19 cm, glabrous to hairy above and 

below; base cordate to rounded, sometimes asymmetric, apex acuminate; lateral veins 14- 

31 pairs, raised to slightly depressed above; petiolules lacking. Inflorescences axillary, 

often densely flowered, 5-24 cm long, glabrous to softly hairy; pedicels lacking or to 3 mm 

long. Flowers unisexual, sometimes bisexual; sepals ovate to triangular or rounded, to 1.5 

mm long, softly hairy below; petals ovate to elliptic, 2-4 mm long, glabrous to hairy below; 

stamens 4, filaments glabrous; disc more or less glabrous; ovary glabrous to hairy, style 

glabrous, stigma 4-lobed or wavy. Fruits of elliptic follicles, 3.5-4.5 mm long, sometimes 

fused up to full length; epicarp dry, glabrous to sparsely hairy; endocarp glabrous. Seeds 

round to ellipsoid or hemispherical, 2-3 mm long; funiculus 0.5-1.5 mm long. 

Distribution. Peninsular Malaysia, Java, Borneo, Phillipines, Papua New Guinea and east 

to Samoa. In Sabah, in primary and secondary forest and open places, from near sea-level 

to 600 m. No record from Sarawak. 

Uses. The leaves have been used for treating fever and cramps (Peninsular Malaysia, 

Indonesia). Resin collected from the trunk has been used as a varnish and adhesive 

(Indonesia). 

11. Melicope lunu-ankenda (Gaertn.) T.G. Hartley 

(the Sri Lankan name for this species) 

I.c. (1994) 61. Basionym: Fagara lunu-ankenda Gaertn., Fruet. Sem. PI. 1 (1788) 334. Type: 

Koenig, s.n., Ceylon (holotype L). Synonyms: Euodia aromatica Blume l.c. (1825) 246; Zanthoxylon 

aromaticum (Blume) Miq. I.c. (1859) 670; Ampacus aromatica (Blume) Kuntze I.c. 98; Zanthoxylum 

roxburghianum Cham., Linnaea 5 (1830); E. roxburghiana (Cham.) Benth., F1. Hongk. (1861) 59; 

Ampacus roxburghiana (Cham.) Kuntze I.c. 98; E. lunu-ankenda (Gaertn.) Merr., Philip. J. Se. 7 

392


(1912) Bot. 378, Merrilll.e. (1921) 314, Anderson I.e. 308; E. arborea Elmer, Leafl. Philip. Bot. 8 

(1915) 2806; E. malayana Rid!. I.e. (1922) 342, Masamune I.e. 358, Anderson I.e. 308; E. punetata 

MeIT., J. Str. Br. R. As. Soc. 86 (1922) 315, I.e. (1929) 113, Masamune I.e. 358, Anderson I.e. 308; 

E. triphylla var. pubeseens Rid!. I.e. (1930) 77, Masamune I.e. 359; E. eoncinna Rid!. I.e (1930) 78, 

Masamune I.e. 358; E. obtusifolia Rid!. I.e. (1930) 78, Masamune I.e. 358. 

Shrub or tree to 30 m tall. Young branchlets glabrous to velvety; terminal buds sparsely 

hairy to velvety. Trichomes mostly simple. Leaves opposite, trifoliolate (occasionally 

unifoliolate); petioles 1.S-1S cm long; leaflets elliptic to obovate, 3.S-23 x 2.S-7 cm, 

glabrous or nearly so; base rounded to narrowly tapered and often asymmetric, apex acute 

to acuminate, rarely rounded or emarginate; lateral veins 8-17 pairs, sometimes raised 

above; petiolules O.S-IS mm long. Inflorescences axillary, 2-32 cm long with main 

branches ascending, glabrous to velvety; pedicels 0.3-3 mm long. Flowers unisexual, 

rarely bisexual; sepals round to ovate-triangular, 0.S-1.2 mm long, glabrous to hairy below; 

petals ovate to elliptic, 1.S-3 mm long, glabrous to sparsely hairy below, hairy above on 

lower half or glabrous; stamens 4, filaments hairy at base or glabrous; disc glabrous to 

densely hairy or velvety; ovary nearly. glabrous to densely hairy, style glabrous to hairy, 

stigma weakly 4-lobed. Fruits of ellipsoid to obovoid follicles, 6-10 mm long (9-12 mm 

long in montane plants); epicarp sub fleshy, glabrous; endocarp glabrous. Seeds round to 

ovoid or ellipsoid, sometimes compressed, 3-6 mm long; funiculus 0.S-3 mm long. 

Distribution. Himalayas, Sri Lanka, Java, Borneo, SW Philippines, and Celebes. In Sabah 

and Sarawak, common in primary and secondary well-drained or swampy forest and 

montane shrubbery, from near sea-level to 2200 m. Also known from Brunei and 

Kalimantan. 

Taxonomy. Hartley (l.c. 1994) recognises lowland and montane variants of this species. 

Plants from montane habitats have larger follicles and more leathery leaflets, usually with 

rounded to abruptly tipped apices. Because the two intergrade, they are treated as a single 

taxon. 

Uses. The roots are reportedly used to treat colds and rheumatism (Taiwan), and the leaves 

and flowers for menstrual disorders and fever (Peninsular Malaysia). Its timber is weak but 

used in construction. The leaves are eaten as a condiment and have been used to flavour 

food. 

12. Melicope sororia T.G. Hartley 

(Latin, sororius = sisterly; referring to its close relationship to Melicope jugosa) 

I.e. (1994) 53. Type: 1. & M.S. Clemens 29477, British North Borneo, Mt. Kinabalu, Tenompok 

(holotype NY; isotypes A, B, BO, L, UC). 

Shrub or small tree to S m tall. Young branchlets glabrous, rather corky; terminal bud 

hairy. Trichomes mostly simple. Leaves opposite, trifoliolate and/or unifoliolate, rarely 

bi/oliolate, glabrous, petioles 1-10 cm long; leaflets of trifoliolate leaves elliptic to obovate, 

6.S-14 x 3-6 cm, base acute to narrowly tapered and somewhat asymmetric, margin entire 

or less commonly toothed toward apex, apex acuminate or rarely rounded, lateral veins 7- 

393


11 pairs, sometimes raised above, petiolules 3-15 mm long; leaflets of unifoliolate leaves to 

16 x 8 cm, petiolule to 1 cm long, otherwise similar to leaflets of trifoliolate leaves. 

Inflorescences axillary, glabrous, 2.5-10 cm long; pedicels to 3.5 mm long, sparsely hairy. 

Flowers unisexual; sepals nearly round or ovate or elliptic, to 1.5 mm long, sometimes 

ciliolate; petals elliptic, to 2.5 mm long; stamens 8; stigma capitate, weakly 4-lobed. Fruits 

glabrous, follicles ellipsoid, 10-13 mm long; epicarp dry. Seeds ellipsoid, 9-10 mm long. 

Distribution. Endemic to Sabah, occurring in forest from 1500 to 2400 m.- 

13. Melicope subunifoliolata (Stapt) T. G.Hartley 

(Latin, sub = nearly, un us = one, folium = leaf; the almost unifoliolate leaves) 

I.e. (1994) 66. Basionym: Euodia subunifoliolata Stapf I.e. 138, MerrillI.e. (1921) 314, Masamune 

l.e. 359. Type: Haviland 1193, British North Borneo, Mt. Kinabalu (holotype K). 

Shrub or tree to 15 m tall. Young branch lets hairy to velvety; terminal buds velvety. 

Trichomes mostly simple. Leaves opposite, trifoliolate and/or unifoliolate, glabrous above, 

hairy to velvety on midrib (sparsely hairy on veins) below; petioles 1-9 cm long; leaflets of 

trifoliolate leaves elliptic to obovate, 5-15.5 x 1.5-5.5 cm, base acute to narrowly tapered, 

apex acuminate, lateral veins 10-20 pairs, sometimes raised above, petiolules 0.5-10 mm 

long; leaflets ofunifoliolate leaves elliptic-obovate, to 8.5 x 3.5 cm, base acute, lateral veins 

11-14 pairs, otherwise similar to leaflets of trifoliolate leaves. Inflorescences axillary, 

laxly flowered, 3-13 cm long, hairy to velvety; pedicels 1-2.5 mm long. Flowers 

unisexual; sepals ovate to triangular, 0.7-1.5 mm long, hairy below; petals elliptic to ovate, 

1.3-1.8 mm long, glabrous; stamens 4, filaments glabrous; ovary and top of disc hairy, 

style hairy at base, stigma weakly 4-lobed. Fruits of ellipsoid follicles, 10-10.5 mm long; 

epicarp dry, hairy; endocarp glabrous. Seeds round to ovoid, sometimes compressed, 5-6.3 

mm long; funiculus 0.5-1.5 mm long. 

Distribution. Endemic to Sabah, occurring in primary montane forest, often on ridges and 

slopes, from 1200 to 2600 m, and sometimes descending to 180 m. 

14. Melicope tripbyIIa (Lam.) Merr. 

(Greek, treis = three, phyllon = leaf; having trifoliolate leaves) 

I.e. (1912) 375; Anderson I.e. 309; Hartley I.e. (1994) 54. Basionym: Fagara triphylla Lam., Eneyel. 

2 (1788) 375. Type: Sonnerat, s.n., Philippines (holotype P). Synonyms: Euodia triphylla (Lam.) 

DC. I.e. 724; Zanthoxylum triphyllum (Lam.) G. Don, Gen. Hist. 1 (1831) 804; Ampaeus triphylla 

(Lam.) Kuntze I.e. 98. 

Shrub or tree to 15 m tall. Young branchlets glabrous to hairy, sometimes glaucous; terminal 

bud glabrous to velvety hairy. Tricl).omes, if present, simple. Leaves opposite, mostly 

trifoliolate, petioles 3-11 cm long; leaflets elliptic to oblanceolate, 8-25 x 2.5-6 cm, 

glabrous; base acute to narrowly tapered and sometimes asymmetric, apex acuminate; 

lateral veins 7-19(-25) pairs, sometimes raised above; petiolules 2-15 mm long. Inflorescences 

axillary and/or on branchlets below leaves, glabrous to hairy. 1-11 cm long, 

394


pedicels lacking or to 4 mm long. Flowers unisexual; sepals ovate-triangular, to 0.8 mm 

long, apex sometimes sparsely hairy; petals lanceolate to ovate or elliptic, to 3.5 mm long; 

stamens 8, filaments sparsely hairy or glabrous; disc hairy or glabrous; ovary glabrous, 

stigma capitate, 4-lobed. Fruits glabrous, follicles ellipsoid or nearly round, 2.5-5 mm 

long; epicarp sub fleshy. Seeds nearly round to ellipsoid, 2.4-4.5 mm long. 

Distribution. Taiwan and Ryukyu Islands, south to Borneo, and east to Papua New Guinea. 

In Sabah and Sarawak, in primary and secondary forest, from near sea-level to 1300 m. 

Also in Kalimantan. 

9. MEROPE Roem. 

(Merope of Greek mythology, one of the Heliades) 

Syn. Hesper. 1 (1846) 44; Backer & Bakhuizenj I.e. 106; Swingle I.e. (1967) 258; Stone I.e. (1972) 

382, I.e. (1978) 115. 

Shrubs or small trees. Branches armed with axillary, solitary or paired spines. Leaves 

alternate, unifoliolate, leathery; petioles short, stout, wingless, articulated with the blade. 

Inflorescences 1 (-2)-flowered, rarely few-flowered, axillary. Flowers bisexual, 4-5-merous; 

calyx cup-like, 5-lobed; petals 5(-6), free, imbricate in bud; stamens 10, free, equal; ovary 

ovoid, 3(-4)-carpellate, on a tall gynophore, ovules 2-4 per locule, style short, thick, stigma 

flat. Fruit an ovoid, angular berry, with 3-4-jlattened sides; pericarp thick, glandular, strongly 

aromatic when crushed, filled with sticky clear fluid. Seeds few to many per frUit, oblong, 

flattened, overlapping. 

Distribution. One species, ranging from Burma to Indo-China and throughout Malesia. 

Found in Sabah and Sarawak. 

Ecology. A widely scattered, highly specialised genus which thrives in the saline soil of 

tidal forests and mangrove swamps, often on the banks of streams. The tides aid in the 

dispersal of the buoyant fruits. 

Merope angulata (Willd.) Swingle 

(Latin, angulatus = angled; the fruit) 

Fig. 9. 

J. Wash. Ac. Sc. 5 (1915) 420, I.e. (1967) 258; Backer & Bakhuizenj I.e. 106; Stone I.c. (1972) 

382, I.e. (1978) 115. Basionym: Citrus angulata Willd., Sp. PI. ed. 4, 3 (1800) 1426. Type: 

Limonellus angulosus Rumph., Herb. Amboin. 2 (1741) 110. Synonyms: Sclerostylis spinosa Blume 

I.e. (1825) 134; Paramignya longispina Hook.j I.e. (1875) 511; P. angulata (Willd.) Kurz, J. As. 

Soc. Beng. 44 (1875) 135. 

Erect shrub or small tree to 3 m tall and 10 cm diameter, often with multiple stems arising 

from a root crown. Branches sparse, with paired, rarely solitary, stout spines, to 5 cm long 

on juvenile stems. Leaves oblong-ovate to ob ovate, 4.5-16 x 2-6 cm, thick, glabrous; base 

rounded, margin subentire to faintly notched, apex acute or shortly acuminate; midrib 

395


prominent below; lateral veins and reticulations inconspicuous; petioles 5-7 mm long. 

Inflorescences J (-2)-flowered, glabrous; pedicels short. Flowers small, fragrant, buds 

ovoid; sepals triangular, c. I mm long, glabrous; petals oblong-lanceolate, 7-9 mm long, 

glabrous, white; filaments glabrous, white, anthers linear-oblong, yellow; ovary slender, 

glabrous. Fruits ovoid to ellipsoid, 3-4.5 cm long, 3-4-sided in cross-section, apex bluntly 

acuminate, green when ripe. Seeds 2-3 cm long, somewhat reniform, tapering to a blunt 

point; testa rough; cotyledons green. 

Vernacular names. Sabah and Sarawak-limau buaya, limau laut (Malay). 

Distribution. Throughout the range of the genus. In Sabah and Sarawak, occurs in rather 

small but dense populations in inland areas of tidal swamps and mangrove forest, often on 

the edges of streams and on riverbanks, and usually in association with Nipa palm (Nypa 

fruticans). 

Uses. The fruits are used for the treatment of stomach disorders and colic (Peninsular 

Malaysia). 

10. MERRILLIA Swingle 

(E. D. Merrill, 1876-1954, American botanist) 

I.c. (1918) 337, I.c. (1967) 240; Rid1ey I.c. (1922) 353; Craib l.c. 231; Burkill l.c. (1966) 1481; 

Stone I.c. (1972) 382; Corner I.c. 667; Stone & Jones I.c. (1988) 268. 

Trees; crown rather bushy; trunk and main branches covered with pale fissured bark; 

branches unarmed. Leaves alternate, pinnate; leaflets 5-J 3, lower ones small, higher ones 

increasingly larger, terminal leaflet largest, thinly leathery, articulated at the base; rachis 

flattened, narrowly winged. Inflorescences axillary, 1-2-flowered. Flowers bisexual, large, 

slightly irregular, 5-merous; calyx cup-like, sepals 5, small; petals 5, free, long, imbricate; 

stamens 10, free, unequal; ovary 5(-6)-carpellate, bottle-shaped, on a well-developed 

gynophore, ovules 8-10 per locule, style long, slender, stigma capitate. Fruit a subglobose 

to oblong or ellipsoid berry; pericarp thick, resinous, 5-chambered, chambers with cartilaginous 

walls and filled with clear mucilage. Seeds numerous, flattened, densely scaly; 

cotyledons pIano-convex. 

Distribution. One species; S Thailand, Sumatra, Peninsular Malaysia, and Borneo (Sabah). 

Ecology. Scattered as solitary trees on stream banks and slopes of hills and ridges, in moist 

primary and secondary forest, to 400 m. 

Taxonomy. This anomalous genus contains a number of metabolites in common with 

Murraya sect. Murraya (Kong et al., Biochem. Syst. Eco!. 16 (1988) 47). One of these 

metabolites, yuehchukene, is of particular interest because of its anti-implantation effect in 

rats. Swingle (l.c. 1967) suggested a possible origin of the genus from a Murraya-like 

ancestor. Morphologically, there are a number of similarities between Merrillia and 

396

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Murraya sect. Murraya. Both have yellowish stem- and root-bark, and some species of sect. 

Murraya possess a winged leaf-rachis and densely villous seeds, characteristics found also in 

Merrillia caloxylon. The two differ markedly, however, in the size and shape of their 

flowers and fruits. 

Merrillia caloxylon (Rid!.) Swingle 

(Greek, kalos = beautiful, xylon= wood) 

Fig. 10. 

I.e. (1918) 338, I.e. (1967) 241; Ridley I.e. (1922) 354; Craib I.e. 231; Burkill I.e. (1966) 1481; 

. Stone I.e. (1972) 383; Corner I.e. 667; Stone & Jones I.e. 68; Ng I.e. 495. Basionym: Murraya 

ealoxylon Ridl., J. Str. Br. R. As. Soc. 50 (1908) 113. Type: Wray & Robinson 5548, Upper Perak, 

Kenering (holotype BM; isotypes SING, US). 

Small to medium-sized tree to 20 m tall and 25 cm diameter; bole short. Leaves to 20 cm 

long; leaflets usually 7-9, alternate to subopposite, lowest leaflets stipule-like, larger 

laterals elliptic, 7.5-9 x 4 cm; base triangular and slightly asymmetric, margin wavy to 

slightly toothed, apex acuminate; petiolules nearly lacking. Inflorescences usually 1- 

flowered. Flowers pendulous, trumpet-shaped; sepals triangular or ovate, to 2.5 mm long; 

petals oblanceolate, gradually tapering to a narrow base, 2.5-5.5 cm long, greenish-white; 

ovary and style hairy. Fruits c. 11 x 9 cm or larger; pericarp c. 13 mm thick, leathery, 

warty, greenish becoming yellow. Seeds 8-10 per chamber, lens-shaped, c. 13 x 3 mm, 

scales membranous, flattened, slightly fimbriate. 

Distribution. Throughout the range of the genus. In Sabah, collected only once from the 

bank of a stream in secondary forest at 20 m (Junaidi Payne, s.n., Sandakan, Ulu Gum 

Gum; SAN). 

Uses. In Peninsular Malaysia, the durable, handsome wood, which is yellow with dark 

brown streaks, has been used to make walking sticks, smoking pipes, parang handles and 

sheaths, and other small objects. 

11. MICROMELUM Blume 

(Greek, mikros = small, melon = apple; the shape of the fruits) 

I.e. (1825) 137; Hooker f I.e. (1875) 501; Kurz I.e. (1875) 136; King I.e. 218; Rid1ey I.e. (1922) 351; 

Merrilll.e. (1929) 114; Craib I.e. 227; Masamune I.e. 360; Backer & Bakhuizen.j I.e. 103; Burkill 

I.e. (1966) 1492; Swing1e I.e. (1967) 197; Stone I.e. (1972) 383; Anderson I.e. 309; Perry I.e. 366; 

Corner I.e. 667; Whitmore, Tantra & Sutisna I.e. 307. 

Shrubs or trees; branches unarmed. Leaves alternate, pinnate, rachis wingless; leaflets (1-) 

7-15(-23), laterals alternate, subchartaceous to thinly leathery, usually obliquely asymmetric 

and articulated at the base. Inflorescences terminal, flat-topped paniculate 

corymbs, often very large, flowers numerous. Flowers bisexual, 5-merous; calyx cup-like, 

398


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c 

8 t------l 

2cm 

Fig. 10. Merrillia calaxylan. A, leafy twig; B, flowering shoot; C, fruit. (All from fresh material from 

a cultivated tree at the Forest Research Institute Malaysia; Non-Dipterocarp Arboretum D13 - Tree 

No. 213.) 

399


sepals triangular, small; petals oblong-linear, free, valvate; stamens 10, alternately long and 

short, filaments linear, glabrous, anthers ovate, sub-basifixed; disc annular; ovary 2-6- 

carpellate, glabrous to densely hairy, radial walls often twisted, one oil-gland over each 

locule, ovules 2 per locule, style slender, constricted at the base, detaching from ovary, 

stigma flattened to subglobose. Fruit a sub globose or oblong berry, dryish; pericarp thin, 

glandular, glabrate to hirsute. Seeds ellipsoid; cotyledons thin, folded. 

Distribution. About 10 specit:s; from W Pakistan, India and Sri Lanka to S China, and 

south through Indo-China and Malesia to Australia, New Caledonia and S Pacific. One 

species, M minutum, in Sabah and Sarawak. 

Micromelum minutum (G. Forst.) Wight & Am. 

(Latin, minutus = small; the flowers) 

Fig. 11. 

I.c. (1834) 93; Craib l.c. 227; Masamune l.c. 360; Backer & BakhuizenJ l.c. 103; Burkilll.c. (1966) 

1493; Swingle I.c. (1967) 203; Stone I.c. (1972) 383; Anderson I.c. 309; Perry l.c. 366; Corner l.c. 

668; Whitmore, Tantra & Sutisna l.c. 307; Ng I.c. 496. Basionym: Limonia minuta G. Forst., Prodr. 

(1786) 33. Type: Forster, s.n., Friendly Islands (BM). Synonyms: Micromelum pubescens Blume 

I.c. (1825) 138, Merrill I.c. (1929) 114, Masamune I.c. 361, Anderson l.c. 309; M. glabrescens 

Benth. in Hooker, Lond. J. Bot. 2 (1843) 212. 

var. minutum 

Shrub or small to medium-sized tree to 20 m tall and 15 cm diameter. Young branchlets 

densely short-hairy, the trichomes greyish. Leaves to 30 cm long or more; leaflets 9-15, 

ovate-lanceolate, 3-12 x 1.5-6 cm, thinly leathery, sometimes drying almost black, subglabrous 

to shortly hairy on veins below; base obtuse and asymmetric, margin entire or 

wavy to shallowly dentate-crenate, apex attenuate-acuminate; lateral veins 5-8 pairs; 

petiolules to 5 mm long. Inflorescences 13-20 cm long, hairy; bracts short deltoid-linear; 

pedicels to 5 mm long. Flowers small; sepals shortly hairy, sometimes glabrate; petals to 

5-8 mm long, spreading, hairy below, pale green or yellowish-white; staminal filaments to 

9 mm long, narrowed at apex, white; ovary cylindric, 5-carpellate, c. 1.5 mm long, hairy 

becoming glabrous, style glabrous or sparingly hairy, stigma flattened to subcapitate. Fruits 

ellipsoid-oblong, to 1 cm long, glabrate, yellow or red when ripe. Seeds with green, wrinkled 

cotyledons. 

Distribution. Sri Lanka, Indo-China, Peninsular Malaysia, Borneo, Philippines, Papua 

New Guinea, NE Australia and the Pacific. A variable species with four varieties, the most 

widespread of which, var. minutum, occurs in Sabah and Sarawak. The var. minutum is 

scattered in Sabah and Sarawak, in primary and secondary forest on low undulating 

ground, disturbed open sites, and forest margins, and frequently on limestone outcrops and 

sandy coasts of the mainland and islands, from near sea-level to 600 m. 

Uses. The roots and leaves are used in treating scabies, intermittant fever, headache, and 

menstrual disorders (lndo-China, Peninsular Malaysia, Philippines). Its light, durable 

timber is used for construction purposes (Peninsular Malaysia). 

400


',m [ 

Fig. 11 Micromelum minutum var. minutum. Fruting leafy twig. (From SAN 93274.) 

401

TREE FLORA OF SABAH AND SARAWAK VOL. I (1995) 

12. MONANTHOCITRUS Tanaka 

(Greek, monos = single, anthos = flower) 

J. Am. Arb. 9 (1928) 138; Swing1e I.e. (1967) 252; Stone I.e. (1985) 216; Stone & lones I.e. (1988) 

268. 

Erect shrubs to small trees. Branches slender, armed, spines axillary and paired or single, 

unequal in length, rarely unarmed. Leaves alternate, simple; petioles short, wingless, not 

articulated with the blade. Inflorescences axillary, usually solitary. Flowers small, bisexual, 5- 

merous, buds ovoid or ellipsoid; calyx cup-like, slightly lobed, sepals 5, imbricate; petals 5, 

ovate, imbricate; stamens 10, equal, free, anthers oblong-linear; disc annular; ovary 3-5- 

carpellate, globose to oblong, glabrous to sp?f-sely hairy, ovules 2-10 per locule, style stout, 

stigma 3-5-lobed. Fruit a globose to ovoid 0'[ pyriform berry; pericarp somewhat leathery, 

thin, glandular. Seeds 6-10 per fruit, tightly packed in the locules and embedded in 

mucilage or scant pulp, flattened to concave or pIano-convex, with or without thin 

membranous margin; testa sometimes spotted. 

Distribution. 4 species; Borneo, Irian Jaya, and Papua New Guinea. One species, M. 

oblanceolata, in Sabah. 

Taxonomy. Proposed by Tanaka in 1928, the genus remained monotypic until 1985, when 

Stone described a new species, M. bispinosa, and transferred one other from Wenzelia 

Merr., a related genus occurring in the Philippines, Irian Jaya, Papua New Guinea, and 

Fiji. The two genera were originally distinguished chiefly on the character of the seeds 

(margin laciniate-membranous and testa spotted in Monanthocitrus, and margin thinmembranous 

and testa not spotted in Wenzelia). These differences do not hold up between 

the genera as here understood, and their distinctness now remains somewhat problematical. 

The size of the flowers (generally smaller in Monanthocitrus) and nature of the spines 

(generally paired and subequal in Monanthocitrus) may be more significant characters, but 

conclusive analysis will depend upon the availability of new material. 

Monanthocitrus oblanceolata Stone & lones 

(Latin, ob = reversed, lancea = spear; the leaf shape) 

Fig. 12. 

I.e. 268. Type: Abu Bakar SAN 36203, Sabah, Sandakan, Labuk Road (ho1otype SAN; isotypes K, 

L). 

Shrub or small tree to 5 m tall. Bark smooth, dark brown. Spines paired, 5-40 mm long or 

more. LeafY stems zig-zag. Leaves oblong-oblanceolate or narrowly subelliptic, 4-20 x 

1.5-7 cm, thinly leathery; base subcuneate or obtuse or subcordate, margin entire, apex 

acuminate-caudate, tip to 26 mm long; midrib below prominent, sparsely hairy; lateral 

veins clearly visible, 8-10 pairs; reticulations obscure; petioles 2.5-7 mm long. Inflorescences 

1(-2)-flowered; pedicels to 5 mm long. Flowers with calyx c. 2 mm wide, sepals ovatetriangular, 

c. 1 mm long, margin minutely ciliate; petals subelliptic, c. 5 mm long, 

glabrous, whitish; staminal filaments c. 1.5 mm long, anthers with a minutely glandularapiculate 

apex; ovary narrowly bottle-shaped, 3-4-carpellate, sparsely hairy, style cylindrical, 

402


4 cm 

A 

Fig. 12. Monanthocitrus oblanceolata. A, fruting leafy twig; B, detail of shoot with three spines. CA 


403


lcm 

c 

>om [ 

A 

B 

Fig. 13. Murraya paniculata. A, fruiting leafy twig; B, inflorescence; C, flower. (From SAN 50299.) 

404


stigma minutely lobed. Fruits obovoid or pyriform, to 5 x 2.8 cm, beaked; pericarp yelloworange, 

smooth, on pedicels to 1 cm long. Seeds 8-10 per fruit, embedded in whitish, 

sweet-tasting flesh; testa thin; cotyledons pIano-convex, thick, green, obscurely glandular. 

Distribution. Endemic to Sabah and rather uncommon in Kota Belud, Beluran and 

Sandakan districts, south to Lahad Datu and Tawau. 

Ecology. Found as solitary trees or small populations in the understorey of lowland forests, 

usually occurring on slopes and ridges of low hills, and sometimes bordering streams on 

flat land, to 600 m. 

Taxonomy. The species superficially resembles Merope angulata. However, the more 

conspicuous leaf venation, smaller flowers, obovoid fruits, and different habitat conditions 

readily distinguish this from Merope angulata which is associated with mangrove swamps. 

The two species are very similar in size and habit. 

13. MURRA Y A Koenig ex L. 

(1. A. Murray, 1740-l791, student of Linnaeus) 

Mant. 2 (1771) 554,563; Hooker! I.c.(1875) 502; Kurz I.c. (1877) 190; King I.c. 219; Ridley l.c. 

(1922) 353; Craib l.c. 230; Browne I.c. 315; Backer & Bakhuizen! l.c. 103; Burkilll.c. (1966) 

1531; Swingle l.c. (1967) 231; Stone I.c. (1972) 384; Perry I.c. 367; Corner I.c. 668. 

Shrubs or trees, deciduous or evergreen. Branches unarmed. Bark usually smooth, often 

pale. Leaves alternate, pinnate or rarely unifoliolate, rachis wingless; leaflets usually 3-9 

(to as many as 2S), articulated at the base. Inflorescences axillary or terminal panicles, 

cymes or corymbs. Flowers bisexual, medium-sized to large (over I cm long), S-merous, 

buds cylindrical or oblong; sepals ovate or lanceolate, fused at base; petals linear to 

oblanceolate, imbricate; stamens 10, free, nearly equal or alternately long and short, 

filaments slightly flattened, glabrous, anthers ovate to elliptic, nearly basifixed; disc annular to 

cylindric; ovary 2-S-carpellate, ovoid to ellipsoid, glabrous or sometimes finely hairy, on a 

gynophore, ovules (1-)2 per locule, style slender, not persistent, stigma capitate. Fruit an 

ovoid to round berry, pulp mucilaginous, peel thin, glandular. Seeds 1 to several per fruit, 

smooth or hairy; testa thin; cotyledons green, pIano-convex. 

Distribution. About IS species; from India to S China, Indo-China and Taiwan, and 

throughout Malesia, eastward to NE Australia and New Caledonia. One native species 

(Murraya paniculata) occurs in Sabah. Two additional species, M exotica and M koenigii, 

are occasionally cultivated in Sabah and Sarawak. 

Taxonomy. A study by Kong et al. (Biochem. Syst. Eco!. 14 (1986) 491) on the distribution 

of the alkaloids yuehchukene and girinimbine in the roots of eight species of Murraya has 

revealed a division within the genus between species producing one or the other of these 

alkaloids. This division is supported by morphological differences which have long been 

recognised. However, the most recent classification of the genus (Swingle I.c. 1967) does 

40S ...


not adequately reflect this information. One group (section Murraya) is characterised by 

species containing yuehchukene (no girinimbine) and possessing yellowish stems and roots, 

larger petals (1-2 cm long), and red, ovoid to ellipsoid fruits. Plants in the second group 

(section Bergera) contain girinimbine (no yuehchukene) and possess brown stems and 

roots, smaller petals (4-7 mm long), and purplish-black, globose to ellipsoid fruits. Of the 

species occurring in Sabah and Sarawak, M paniculata and M exotica represent section 

Murraya, andM. koenigii represents section Bergera. 

Key to Murraya species 

1. Leaf stalks hairy. Inflorescences many-flowered corymbs. Fruits blackish when ripe. 

Cultivated ..................................................................................................................... . 

M. koenigii (L.) Spreng. 

I.c. (1825) 315. Basionym: Bergera koenigii L. l.c. 563. Synonym: Murrayafoetidissima 

Teijsm. & Binn., Nat. Tijd. Ned. Ind. 27 (1864) 41; Chalcas koenigii (L.) Kurz l.c. (1875) 

132. 

India, Sri Lanka, Burma, Indo-China, Hainan and S China. Commonly cultivated 

in tropical countries, including in Sabah and Sarawak, for its leaves (Curry leaves; 

daun kari). 

Leaf stalks glabrate. Inflorescences few-flowered panicles or cymes. Fruits red when 

...................................... 2 

2. Leaflets obovate, less than 4 cm long, apex somewhat obtuse. Inflorescences terminal. 

Fruits elliptic to subglobose. Cultivated ......................................................... . 

M. exotica L. 

I.c. (1771) 563. Synonym: Murraya paniculata var. exotica (L.) Huang, Act. Phytotax. 

Sinica 8 (1959) 100. 

Native to China and Taiwan (7). Commonly cultivated throughtout the tropics. 

Leaflets ovate or elliptic to rhomboid, 3-7 cm long, apex acuminate. Inflorescences 

axillary. Fruits ovoid with acuminoid apex. Wild species ........................... M. paniculata 

Murraya paniculata (L.) Jack Fig. 13. 

(Latin, paniculatus = tufted; the inflorescence) 

Mal. Misc. 1 (1820) 31; Ridley I.c. (1922) 353; Craib l.c. 230; Browne I.c. 315; Backer & Bakhuizen 

f I.c. 103; Burkilll.c. (1966) 1531; Swinglel.c. (1967) 232; Stone I.c. (1972) 384; Perry I.c. 367; 

Corner I.c. 669; Ng I.c. 496. Basionym: Chalcas paniculata L., Mant. 1 (1767) 68. Type: "India" 

(LlNN); based partly on Camunium Rumph., Herb. Amboin. 6. Synonyms: Limonia lucida G. Forst. 

I.c. (1786) 33; Murraya sumatrana Roxb., Fl. Ind. ed. 2,2 (1832) 375; M. odorata Blanco I.c. (1845) 

256. 

var. paniculata 

Shrub or small to medium-sized tree to 20 m tall, 25 cm diameter. Bark thin, pale to 

whitish. Young shoots, twigs, sepals, petals, and ovary glabrous to slightly hairy. Leaves 

,.. 

406

pinnate, to 17 cm long; leaflets 3-7, rarely unifoliolate, ovate or ovate-elliptic to rhomboid, 

3-7 x 2-3.5 cm, chartaceous or thinly leathery, glossy and darker above, glabrous; base 

cuneate to rounded, margin entire or faintly crenulate, apex acuminate; lateral veins 5-8 

pairs; petiolules 2-6 mm long. Inflorescences axillary panicles or cymes, few-flowered; 

peduncle 1-4 cm long; pedicels 2-9 mm long. Flowers large; sepals (4-)5, narrowly 

deltoid, c. 1 mm long, glandular; petals (4-)5, elliptic to oblong-obovate, to 15-21 mm 

long or more, white; stamens alternately long and short, the longer to 12 mm long, 

filaments dilated below; ovary 2-carpellate, glandular, style columnar, nearly 1 cm long, 

stigma enlarged, bilobed. Fruit an ovoid berry, c. 12 mm long, apex acuminoid; peel shiny 

red, gland-dotted, glabrous. Seeds densely hairy; cotyledons thick, fleshy. 

Vernacular name. Sabah-kemuning (Malay). 

Distribution. Throughout the range of the genus. Of the four known varieties, only var. 

Shrubs or small trees. Branches armed with axillary, solitary or paired spines, or unarmed 

or nearly so. Young twigs angular, becoming terete. Leaves alternate, 1-3-foliolate; petioles 

winged or wingless; leaflets thinly leathery, glabrous, margin entire, articulated at the base. 

Inflorescences axillary or pseudoterminal panicles or racemes, often few-flowered, hairy. 

Flowers small, bisexual, 4-5-merous, fragrant, buds cylindric; sepals triangular to linear- 

407 

1. Wash. Ae. Se. 6 (1916) 426, I.c. (1939) 258, I.c. (1967) 290; Merrilll.c. (1929) 115; Masamune 

I.e. 361; Baeker&BakhuizenJ I.c. 104; Andersonl.c. 309. 

14. PLEIOSPERMIUM (Engl.) Swingle 

(Greek, pleio = few, sperma = seed) 

Uses. The leaves, fruits and bark ofbothM. paniculata and M exotica have been used for 

treating venereal disease, intestinal worms, skin disorders, and dysentery, among others 

(China, Philippines, Peninsular Malaysia, Indonesia). Their heavy, yellow wood has been 

used to make walking sticks, kris handles, and various small objects (Peninsular Malaysia, 

Thailand). A face powder has been made from its bark and roots (Burma), and the fragrant 

flowers are used in cosmetics (Java). The leaves contain an unidentified, water-soluble 

toxin that has been shown to kill eggs and nymphs of the Asian citrus blackfly, 

Aleurocanthus woglumi Ashby, with topical exposure (Dowell, Pan-Pacific Entomol. 65 

(1989) 163). 

Taxonomy. M exotica has usually been placed in synonymy with this species or as a 

variety of it. In this treatment, M paniculata is restricted to the wild form found locally, 

with comparatively larger leaflets and flowers. M exotica, locally found only in cultivation, 

differs in the shape of its leaflets (ob ovate to subelliptic) and fruits (ellipsoid to subglobose), 

in having terminal inflorescences, and in its more restricted natural geographical range 

(coastal areas of Hong Kong and China). 

panicuiata occurs in Sabah, where it is occasionally found in lowland and hill forests, 

usually on rocky soils or limestone, from near sea-level to 600 m. 

RUT ACEAE (lONES)

2cm 

,= [ 1I A 

Fig. 14. Pleiospermium longisepalum. A, fruting leafy twig; B, flowering twig; C, mature flower with 

bud. (From SAN 90784.) 

408 

'mm[ I c 


r----""l

lanceolate, glabrous to finely hairy; petals linear-oblong, glabrous or finely hairy below, 

white to greenish, imbricate in bud; stamens 8 or 10, free, alternately long and short, 

filaments glabrous, anthers oblong to linear-oblong; disc annular to shortly cup-like; ovary 

4-5-carpellate, cylindric or ovate, glabrous or hairy, on a short gynophore or nearly sessile, 

ovules 2 per locule, style slender or stout, stigma subcapitate. Fruit a globose to oblong 

berry, 1.5-2.5 cm in diameter; pericarp rough-glandular; pulp-vesicles slender, to 1 cm 

long; juice oily-resinous. Seeds ovoid, flattened, to 1 cm long; testa smooth or wrinkled; 

monoembryonic; cotyledons pIano-convex, thick, green. 

Distribution. 6 species; S India, Sri Lanka, Vietnam, Sumatra, Java, and Borneo. Two 

species, P. latialatum and P. longisepalum, are found in Sabah. The former species also 

occurs in Sarawak. 

Key to Pleiospermium species 

Distribution. Endemic to Borneo. Widespread and common in Sabah and Sarawak, 

occurring in the lowlands and hills and along streams and rivers in primary mixed 

dipterocarp and secondary forests on fertile clay-rich loam soils, and (in Sarawak) on 

limestone slopes and ridges, from near sea-level to 300 m. 

409 

Vernacular names. Sarawak-limo antu (preferred common name), limo to' (Kayan), limo 

bali (Kenyah), para bileh (Berawan). 

Shrub or small to medium-sized tree to 25 m tall, 30 cm diameter; bole to 15 m tall; trunk, 

bole, and main branches armed with paired spines to 3 cm long, generally unarmed on 

upper branches. Leaves unifoliolate, oblong-elliptic or lanceolate, 9-15 x 3.5-7 cm; base 

rounded or cuneate, apex acute-caudate, acumen 4-6 mm long and emarginate; lateral 

veins 10-13 pairs; petioles 1.5-2.5 cm long, broadly winged, obcordate, top 1-2 cm wide, 

narrower at base and pulvinoid. Inflorescences axillary clusters or pseudoterminal 

panicles. Flowers with sepals triangular, (2-)4-6 x 2-4 mm, glabrous, perSistent and 

spreading at right angles below the fruit; petals glabrous; ovary hairy. Fruits globose, 1.5- 

2.5 cm in diameter; pericarp greenish yellow, with numerous raised oil-glands; pulpvesicles 

slender, conical, 3--4.5 mm long, tips acute. Seeds 2-3 per fruit. 

I.e. (1939) 261, I.e. (1967) 293; Masamune I.e. 361; Anderson I.e. 309. Type: Elmer 21542, British 

North Borneo, Tawau (holotype AA; isotypes BO, K, L, P, PNH, US). 

1. Pleiospermium latialatum Swingle 

(Latin, latus = broad, ala = wing; the petiole) 

Petioles narrowly winged, 3-5 mm wide. Sepals 9-12 mm long, linear ......... 2. P. longisepalum 

Petioles broadly winged, 10-20 mm wide. Sepals 4-6 mm long, triangular ........ 1. P. latialatum 

RUTACEAE (JONES)

Perry I.e. 37l. 

Shrubs or small trees. Branches spreading, armed with stout axillary spines, or unarmed. 

Leaves alternate, simple, often rigid, conspicuously parallel-veined; petioles short, wingless, 

not articulated with the blade. Inflorescences densely flowered, axillary or terminal 

panicles or racemes, or few-flowered clusters. Flowers bisexual, small, 3-S( -7)-merous; 

calyx cup-like, sepals imbricate, glabrous or hairy; petals va!vate, glabrous or nearly so; 

stamens S-1S, free, filaments dilated or flattened, anthers small, disc cup-like, enclosing 

base of ovary; ovary subglobose or oblong, 1-S-carpellate, on a short gynophore, ovules 1 

per locule, style short,stigma ovoid. Fruit a small berry, round, juicy or semi-dry; pericarp 

. dotted with oil-glands, smooth; pulp-vesicles irregular in size and shape, lacking stalks. 

Seeds small, ovoid, thick; testa thin; monoembryonic. 

Distribution. 6 species; from S China, Taiwan, and lndo-China, south to Java, N Borneo, 

the Philippines, Moluccas, and Irian Jaya. Two species (s. disticha and S. panieulata) 

occur in Sabah. 

410 

----- - 

Ind. Sem. Bort. Neap. 3 (1840); Masamune I.e. 357; Swingle I.e. (1967) 283; Stone I.e. (1978) 116; 

Ecology. Usually found on the slopes of hills and adjacent to streams in primary and 

secondary inland forests, and on high ground near the sea, to 1300 m. 

Distribution. Endemic to Sabah where it is not uncommon in the foothills and low 

mountainous areas of Kota Marudu, Ranau, and Tambunan districts, and occasionally on 

off-shore islands (pulau Banggi = Banguey Is., Pulau Bohayan). 

base. Inflorescences axillary clusters or pseudoterminal panicles. Flowers with sepals 

linear, 9-12 x 2-3 mm, chartaceous, apex rounded or emarginate, persistent and reflexed 

below the fruit. Fruits globose, tipped by the short persistent style, 2-2.S cm in diameter; 

pericarp l.S-2 mm thick, pimply from numerous raised oil-glands, ripening pale brownish 

yellow, pulp-vesicles 4-6 mm long, tapering to a blunt point. Seeds 1-2 per fruit, c. 1 cm 

long. 

1.S cm long, narrowly winged (wing 3-5 mm wide at top), tapering to a wingless, pulvinoid 

unarmed or nearly so. Leaves unifoliolate, elliptic, 7.S-11.S x 3-S.S cm; base rounded or 

cuneate, apex acuminate, acumen rounded or retuse; lateral veins 8-10 pairs; petioles 1- 

Shrub or small tree to IS m tall, 40 cm diameter; trunk sometimes fluted. Branches 

Fig. 14. 

I.e. (1939) 259, I.e. (1967) 293. Type: Castro & Melegrito 1348, British North Borneo, Banguey Is. 

(Severinus, 6th Archbishop of Rome, 640 A.D.) 

15. SEVERINIA Tenore 

Vernacular names. Sabah-limau hutan, limau limau (Malay). 

2. Pleiospermium Iongisepalum Swingle 


(Latin, longus = long, sepalum = sepal) 

(holotype NA; isotypes K, US).


Taxonomy. The genus has been confused with Atalantia Corr. which resembles in it leaf 

characiers (shape, size, and venation) and habit. All species of Severinia have at one time 

been placed in Atalantia. Atalantia differs, however, in having leaf-blades articulated with 

the petioles, larger flowers, filaments more or less fused, well-formed pulp-vesicles, and 

small, orange-like fruits. Its 11 species are distributed from India to W Malesia and is 

represented in Peninsular Malaysia by two species. The relationship of Severinia to 

Atalantia needs to be examined more critically. 

Key to Severinia species 

Leaves ovate-lanceolate, apex acute-acuminate. Inflorescences axillary or terminal, 

racemose or paniculate. Ovary 2-carpellate ............................................................ 1. S. disticha 

Leaves oblong-ovate, apex rounded. Inflorescence terminal, paniculate. Ovary 4-carpellate 

S. paniculata 

1. Severinia disticha (Blanco) Swingle Fig. 15. 

(Latin, distichus = of two rows; the leaf arrangement). 

J. Wash. Ae. Se. 28 (1938) 533, I.e. (1967) 287; Masamune I.e. 357; Stone I.e. (1978) 116. 

Basionym: Limonia distieha Blaneo I.e. (1837) 356. Type: Merrill, Sp. BIaneoanae No. 594, 

Philippines, Luzon (L). Synonyms: Limonia eorymbosa Blaneo I.e. (1845) 251; AtaIantia nitida 

Oliv. I.e. 24; A. distieha (Blaneo) MelT., Bull. Gov. Lab. Philip. 27 (1905) 28, Masamune I.e. 357. 

Shrub or small tree to 10 m tall, 15 cm diameter. Branches unarmed, occasionally with 

paired spine-like paraphylls. Leaves 3-8(-11) x (1.5-)2.5-4 cm or more, ovate or ovatelanceolate, 

wavy in dried speCimens; base cuneate to obtuse, margin subentire, sometimes 

faintly crenulate, apex acute or acuminate, acumen usually blunt, often emarginate; lateral 

veins parallel, straight, numerous, 15-20 pairs, prominent below; petioles 3-9 mm long, 

flattened above, glabrous to hairy. Inflorescences many-flowered, axillary or (rarely) 

terminal panicles or racemes, 3-9 cm long; pedicles 3-5 mm long. Flowers 5-merous; 

sepals 5, broadly rounded, margins ciliate; petals 5, oblong, minutely tipped, to 7 mm long, 

white; stamens 10, alternately long and short, filaments to 5 mm long, flattened, narrowed 

at apex and base, anther:s ovate, apiculate-glandular; ovary 2-carpellate, oblong, glabrous 

to sparsely hairy, style cylindrical, stout, stigma slightly lobed. Fruits subglobose, to 1.5 cm 

wide, blackish-purple when ripe; pulp greenish, juicy. Seeds 1-2 per fruit, greenish white. 

Distribution. Borneo (Sabah), E Java, Philippines, Moluccas, Flores, and Irian Jaya. In 

Sabah, not uncommon in the understorey of beach strand vegetation, and in primary and 

secondary forest on hills and ridges near the coast, frequently on offshore islands, usually 

on sandy or rocky soil, rarely on limestone, from near sea-level to 90 m. 

Uses. The ripe, juicy fruits are rather sweet and eaten fresh (Sabah, Indonesia). The roots 

are said to be used medicinally, and its fruits made into glue (Philippines). The timber has 

been used in construction (Indonesia, Philippines). 

411


2. Severinia paniculata (Warb.) Swingle 

(Latin, paniculatus = tufted; the inflorescence) 

I.c. (1938) 533, l.c. (1967) 288; Stone l.c. (1978) 116. Basionym: Atalantia paniculata Warb., Bot. 

Jahrb. 13 (1891) 340. Type: Warburg 20132, Mo1uccas, Ceram Laut (B). Synonyms: Atalantia 

maritima Merr., I.c. (1914) 293; A. disticha (B1anco) Merr. var. paniculata (Warb.) Tanaka I.c. 

(1928) 141. 

Shrub or small tree. Branches unarmed. Leaves oblong-ovate, 9-12 x 3-5 cm, or larger; 

base gradually tapered, margin obscurely crenu1ate, apex rounded or emarginate; lateral 

veins numerous, parallel, more or less prominent below; petioles c. 8 mm long, terete, 

hairy. Inflorescences terminal panicles, 4-6 cm long or more, hairy; pedice1s 2-4 mm 

long. Flowers 5-merous; sepals 5, c. 1 mm long, tips blunt, margins ciliate; petals 5, 4-7 

mm long, lanceolate, tips rounded, subg1abrous; stamens 10, alternately long and short, 

filaments 4-5 mm long, glabrous, anthers broadly cordate; ovary 4-carpellate, style 

deciduous. Fruits globose, to 12 mm in diameter. Seeds 2-4 per fruit. 

Distribution. Borneo (Sabah), S Philippines, Mo1uccas, and Sumbawa (the Lesser Sunda 

Is.). In Sabah, less widespread than the related species S. disticha, and found in many of 

the same habitats, particularly beach forest and offshore islands, at sea level. 

Taxonomy. This species is very closely related to S. disticha and Tanaka (I.c. 1928) 

considered it to be a variety. Many of their morphological characters appear to overlap. 

Swing1e (I.c. 1967) distinguished it from S. disticha based upon its larger leaves, smaller, 

panicu1ate inflorescences, and ovaries with four carpels instead of two. A re-evaluation of 

the taxonomic status of this species (and most other Severinia) would be useful. 

16. TETRACTOMIA Hook.! 

(Greek, tetra = four, tome = a separation; the four carpels which split in fruit) 

I.c. (1875) 490; King l.c. 211; Merrill I.c. (1921) 314; Rid1ey l.c. (1922) 345; Craib I.c. 215; 

Masamune l.c. 361; Burkilll.c. (1966) 2182; Stone l.c. (1972) 385; Hart1ey, J. Am. Arb. 60 (1979) 

127; Anderson l.c. 309; Whitmore, Tantra & Sutisna l.c. 307. Synonym: Terminthodia Ridl., J. Fed. 

Mal. St. Mus. 6 (1915) 14l. 

Shrubs or small to large trees. Branches unarmed. Leaves opposite, unifoliolate, margins 

entire, pinnate1y veined; petioles wingless, articulated with the blade. Inflorescences 

axillary panicles, sometimes 1- to few-flowered. Flowers bisexual, 4-merous; sepals 4, 

fused at base, va1vate or partially imbricate; petals 4, free, usually va1vate, triangular, 

spreading and becoming recurved; stamens 4, free, opposite the sepals, filaments flattened, 

tapered, glabrous, elongating after anthesis, anthers ellipsoid, dorsifixed; staminodes 4, 

alternating with the stamens, anthers minute, without pollen; disc broad, glabrous, rounded 

to 4-ang1ed; ovary 4-carpellate, carpe.ls erect, free or fused at the base, joined near the apex 

by a single style, 10cu1es 2-ovu1ate; style straight, composed of 4 elements twisted together, 

stigma capitate. Fruits of 1-4 erect follicles, undeveloped carpels persistent in fruit, 

follicles boat-shaped, free or fused at the base; exocarp and mesocarp leathery; endocarp 

412


3 cm 

Fig. 15. Severinia disticha. Fruiting leafy twig. (From SAN 36104.) 

413


cartilaginous, separating from the mesocarp. Seeds 1-2 per follicle, Winged; testa papery; 

endosperm fleshy; embryo straight. 

Distribution. 6 species; S Thailand, Sumatra, Peninsular Malaysia, Borneo, Philippines, 

Celebes, Papua New Guinea, and the Solomon Islands. The widely distributed T. 

tetrandrum occurs in Sabah and Sarawak. The remaining species are localised endemics in 

other parts of the range. 

Ecology. Lowland to montane forests, peat swamp forests, heath forests, and subalpine 

habitats, from near sea-level to 3300 m. 

Tetractomia tetrandrum (Roxb.) Merr. Fig. 16. 

(Greek, tetra = four, andros = male; the number of stamens) 

J. Str. Br. R. As. Soc. 76 (1917) 87, I.e. (1921) 314; Rid1ey I.e. (1922) 346; Craib I.e. 215; 

Masarnune I.e. 361; Burkill I.e. (1966) 2182; Stone I.e. (1972) 385; Hart1ey I.e. (1979) 132; 

Anderson I.e. 309; Whitrnore, Tantra & Sutisna I.e. 308. Basionym: Melieope tetrandra Roxb., Hort. 

Bengal. (1814) 88. Type: Roxburgh, Ieones 1411, Penang (K). Synonyms: Tetraetomia beeearii 

Hook. f I.e. (1875) 489, Merrill I.e. (1921) 314, Masamune I.e. 361, Anderson I.e. 309; 

Terminthodia viridiflora Ridl. I.e. (1915) 141; Tetraetomia obovata Merr. I.e. (1917) 86, I.e. (1921) 

314, Masamune I.e. 361; Tetraetomia holttumi Ridl., J. Bot. 62 (1924) 295; Tetraetomia latifolia 

Ridl. I.e. (1930) 79, Masamune I.e. 361, Anderson I.e. 309; Tetraetomia montana Ridl. I.e. (1930) 

79, Masamune I.e. 361, Anderson I.e. 309; Tetraetomia parviflora Ridl. I.e. (1930) 78, Masamune 

I.e. 361, Anderson I.e. 309. 

Shrub or small to large tree to 30 m tall, 50 cm diameter; bole to 18 m tall. Branchlets 

glabrous. Leaves sometimes clustered at branch ends, blades glabrous, leathery, obovate 

to ob lanceolate or elliptic, (2.5-)7.5-22(-32) x 1-14 cm; base rounded to attenuate, apex 

emarginate or rounded to acuminate or mucronate; lateral veins 4-11 pairs; petioles 0.3-5 

cm long, swollen at both ends. Inflorescences 1- to many-flowered, (0.8-)4-16 cm long, 

glabrous to sparsely hairy; pedicels 0.5-4 mm long. Flowers 2.5-10 mm wide; sepals 

rounded to triangular, 0.4-l.5 mm long, glabrous to sparsely hairy, ciliolate when young; 

petals green to yellowish, ovate-triangular, 1-4.5 mm long, usually glabrous; carpels free 

at the base, glabrous or sparsely hairy, style glabrous. Fruits with free follicles, 4-11 mm 

long, glabrous or nearly so. 

Vernacular names. Sabah and Sarawak-jampang,jampang rusa, medang rawang (preferred 

common name), rawang mata, rawang paya (Malay). 

Distribution. Throughout the range of the genus. In Sabah and Sarawak widespread; also 


Ecology. Common in secondary forests and open places on infertile organic soils, and also 

mixed dipterocarp forest on humult ultisols, lowland peat swamp and heath forests, from 

near sea-level to 1900 m. 

Taxonomy. An extremely variable species in habit, leaf size and shape, and inflorescence, 

flower and fruit size. In Sabah and Sarawak, this variation is characterised by a widespread, 

414


c 

1 >om 

lcm 

D 

,t 

n ~! 

~~ 

1 cm I 11'111 U 

gl 

.~ 

~u' 

A 

J "m 

Fig. 16. Tetractomia tetrandrum. A, fruiting leafy twig; B, fiuit; C, part of inflorescence; D, detail of 

c1usterofflowers. CA & B from SAN 65010, C & D from SAN 22689.) 

415


generalised form and several specialised forms or races which reflect the distribution of 

certain habitats, i.e., lowland peat swamp forest, heath forest, lower montane forest, and 

lowland dipterocarp forest. Hartley (l. c. 1979) describes these races and provides the names 

of relevant previously described taxa assigned to these entities. 

17 . ZANTHOXYLUM L. 

(Greek, xanthos = yellow, xulon = wood) 

I.c. (1753) 270; Hooker J I.c. (1875) 492; Kurz I.c. (1877) 180; King I.c. 213; Ridley I.c. (1922) 346; 

Craib l.c. 218; Backer & BakhuizenJ l.c. 96; BurkillI.c. (1966) 2326; Hartley, J. Am. Arb. 47 

(1966) 171, 51 (1970) 423; Stone I.c. (1972) 386; Anderson I.c. 309; Perry I.c. 369; Corner l.c. 670; 

Whitmore, Tantra & Sutisna I.c. 308. Synonym: Fagara L., Syst. ed. 10 (1759) 897. 

Scan dent or erect shrubs or trees; dioecious or rarely monoecious, evergreen or deciduous. 

Twigs and branches armed with spines or prickles. Leaves alternate, unifoliolate, trifoliolate 

or pinnate with up to 15 pairs of leaflets, with or without a terminal leaflet, leaflets 

articulated at the base, leaf rachis sometimes winged. Inflorescences terminal or axillary 

racemes, panicles or cymes. Flowers small, unisexual, rarely bisexual; sepals 4-5; petals 

4-5; or perianth segments 6-8 and undifferentiated; stamens 4-6, opposite the sepals, 

rudimentary in female flowers; disc flat or cushion-like; ovary 1-5-carpellate, rudimentary 

in male flowers, carpels free or fused at base, ovules 2 per locule, styles fused to divergent, 

stigma capitate. Fruits of 1-5 follicles, free or basally fused; outer wall glandular, red to 

black; endocarp cartilaginous. Seeds ovoid to round, 1 per follicle, often hanging from the 

dehisced follicle at maturity; testa black or reddish, glossy; endosperm white, fleshy. 

Distribution. A large genus comprising approximately 200 or more species of mainly 

pantropical distribution, with a few representatives in temperate eastern Asian and North 

America. In Sabah and Sarawak, 4 species occur, two of which are trees or shrubs, and the 

remaining two being scandent shrubs. 

Ecology. Most of the species generally grow in rain forests and thickets at low and medium 

elevations. The attractive seeds are dispersed by birds, accounting for the often wide and 

sometimes discontinuous distribution of some of the species. 

Uses. Various species are sources of spices and condiments, and local medicines. The 

attractive wood of some is used for cabinetry and other fine work. 

Key to Zanthoxylum species 

1. Erect shrubs or trees. Branchlets armed with straight prickles. Ovary 2-3-carpellate ... 2 

Scandent or suberect shrubs. Branchlets and/or leaf axes armed with recurved prickles 

or unarmed. Ovary 4-carpellate .................................................................................... 3 

2. Inflorescences paniculate. Leaflets 8-18 cm long. Carpels 3 .......... 2. Z. myriacanthum 

Inflorescences cymose. Leaflets 1-8 cm long. Carpels 2 ........................ 1. Z. aviccnnac 

416


3. Lateral leaflets subopposite to alternate. Midribs ofleaflets not armed ..... 

Z. scandens Blume 

I.c. (1825) 249; Backer & BakhuizenJ I.c. 96; Hartley I.c. (1966) 177; Whitmore, Tantra 

& Sutisna l.c. 308. 

India, China, Taiwan, Ryukyu Islands, Sumatra, Java, Borneo; uncommon in 

Sabah and Sarawak, known only from montane forest. 

Climbing (occasionally suberect) shrub, dioecious. Leaves pinnate; leaflets 2-12 

pairs, with margins finely glandular toothed. Inflorescences axillary, or axillary 

and terminal panicles. Flowers 4-merous; petals occasionally purple-margined; 

ovary 4-carpellate. Follicles in groups of 1-4,4-5 mm in diameter. 

Lateral leaflets opposite. Midribs ofleaflet usually armed ............................................. . 

Z. nitidum (Roxb.) DC. 

I.c. (1824) 727; Ridley I.c. (1922) 1347; Burkilll.c. (1966) 2327; Hartley I.c. (1966) 180, 

I.c. (1970) 423; Stone l.c. (1972) 386; Whitmore, Tantra & Sutisna I.c. 308. 

India, Thailand, Vietnam, Malesia, Solomon Is., Australia. In Sabah (Ranau and 

Pinangah districts), in forest to 1100 m. 

Climbing (occasionally suberect) shrub, dioecious or (rarely) monoecious. Leaves 

pinnate, leaflets 2-4 pairs, with margin entire to glandular toothed. Inflorescences 

terminal and/or axillary, racemes or panicles. Flowers 4-merous; ovary 4- 

carpellate. Follicles in groups of 1-4, 5-7 mm in diameter. 

1. Zanthoxylum avicennae (Lam.) DC. 

(Ibn Sina, or Avicenna, 980-1037, Arabian medical writer and philosopher) 

I.c. (1824) 726; Hartley I.c. (1966) 190, I.c. (1970) 423; Perry I.c. 370. Basionym: Fagara avicennae 

Lam. I.c. 445. Type: d'/ncarville 179, China, Kwangtung (P). Synonyms: Zanthoxylum diversifolium 

Warh.l.c. 339; Z. iwahigense Elmer, Leaf}. Philip. Bot. 5 (1913) 1833. 

Scandent or erect shrub or small tree to 15 m tall; dioecious, evergreen. Smaller branches 

and twigs with straight or recurved prickles. Leaves pinnate, to 30 cm long, axis often 

narrowly winged; leaflets 2-11 pairs, sub opposite, ovate to elliptic-lanceolate, 1-8 x 1-3 

cm, glabrous, thinly leathery; base rounded to cuneate, margin nearly entire to finely 

glandular toothed, apex rounded to acuminate; lateral veins 4-11 pairs; petiolules 2-5 mm 

long. Inflorescences terminal (occasionally axillary), flat-topped cymes, to 20 cm long, 

axis glabrous. Flowers unisexual, 5-merous; sepals 5, triangular or rounded, under 1 mm 

long; petals 5, elliptic, 1-2.5 mm long, white to greenish; stamens 5, rudimentary in female 

flowers; disc flat or cushion-like; ovary 2-carpellate, rudimentary in male flowers, styles 

and stigmas united, peltate. Fruits of /-2 nearly roundfollicles, c. 4.5 mm in diameter, the 

undeveloped carpels (if any) persistent in fruit. 

Distribution. China, Vietnam, Thailand, Java, Borneo, Philippines, Celebes, Lesser Sunda 

Islands and the Moluccas. In Sabah, reported only from Ranau district. 

Ecology. Found locally on flat sites and hillsides in primary and secondary forest, 

occasionally on ultramafic soil, to 1500 m. 

Uses. The stems and bark are used medicinally as a tonic and for treating snake bite (Indo 

China, Philippines). 

417

• / J •••• ~ 

, ••••• _ 


2 cm [ 

4cm 

A 

B 

lcm 

""r" 

'." ........... ~ ( 

.. 

......-... : /. '". •.:: .'. . .' .... 

.... 

. 

~ /". - 

- " 

." 

c 

Fig. 17. Zanthoxylum myriacanthum. A, flowering leafy twig; B, detail of lower leaflet surface; C, 

detail of upper leaflet surface; D, part of infructescence; E, dehisced fruit. (A-C from SAN 87166, D 

& E from SAN 116158.) 

418


2. Zanthoxylum myriacanthum Wall. ex Hook.! 

(Greek, myrios = numberless, akantha = thorn) 

Fig. 17. 

I.c. (1875) 496; King I.c. 214; Rid1ey I.e. (1922) 347; Burkilll.c. (1966) 2327; Hart1ey I.c. (1966) 

185; Stone I.e. (1972) 386; Anderson I.e. 309; Perry I.e. 371; Corner I.e. 671. Whitmore, Tantra, 

Sutisna I.e. 308. Type: Porter (Wallieh Cat. No. 1214), Malacca (K). Synonyms: Fagara 

myriacantha (Wall. ex Hook./) Engl. in Eng1er & Prant11.c. (1896) 118. 

Small to large tree to 30 m high, 35 cm diameter; dioecious, evergreen; bole to 15 m tall. 

Stems with thick spines to 3 cm long; smaller branches and twigs with straight, hollow 

prickles usually housing ants. Leaves pinnate, to 60 cm long; leaflets 4-11 pairs, opposite 

or subbpposite, elliptic, 8-18 x 3-8 cm, glabrous or shortly hairy below, leathery; base 

blunt to nearly cordate and slightly asymmetric, margin finely glandular toothed, apex 

acuminate; lateral veins 8-18 pairs; petiolules to 5 mm long. Inflorescences terminal and 

axillary panicles, 15-25 cm long, axes mostly glabrous. Flowers unisexual, 5-merous; 

sepals 5, triangular, minute; petals 5, elliptic, 1.5-2.5 mm long, white or yellowish, 

sometimes purplish; stamens 5; disc flat; ovary 3 (-4)-carpellate, rudimentary in male 

flowers, styles and stigmas united, peltate. Fruits of ]-3 nearly roundfoIlicles, 3-6 mm in 

diameter, the undeveloped carpels (if any) persistent in fruit. 

Distribution. E India, N Vietnam, SW China, Sumatra, Peninsular Malaysia, Borneo, 

Philippines. In Sabah, occurs scattered in Ranau, Tambunan and Keningau districts. In 

Sarawak, collected once (Omar, s.n., Ulu Lawas). 

Ecology. Locally found in primary and secondary forest in the hills and low mountains, to 

1200 m. 

Uses. The smoke of burning seeds is said to be inhaled for treating ulcerated syphilitic nose 

(Peninsular Malaysia). 

419


SIMAROUBACEAE 

Julius Kulip & K.M. Wong 




Merrill, EB (1921) 315; Rid1ey, FMP 1 (1922) 360; Masamune, EPB (1942) 361; Nooteboom, FM 1, 

6 (1962) 193, FM 1, 6 (1972) 968, Blumea 11 (1962) 509; Keng, OFMSP (1969) 178; Kochummen, 

TFM 2 (1972) 345; Cockburn, TS 1 (1976) 217; Anderson, CLTS (1980) 322; Corner, WSTM 2 

(1988) 696; Whitmore, Tantra & Sutisna, CLK 2,1 (1990) 329. 

Trees or shrubs (some sprawling), usually containing very bitter substances. Hairs mostly 

simple and unicellular, sometimes with a glandular head. Leaves spirally arranged, simple 

or pinnate, sometimes (in Ailanthus, Brucea and Soulamea) with pitted, concave, or flattish 

glands on the lower surface; stipules usually absent except in Allantospermum, Jrvingia and 

Picrasma. Inflorescences usually compound, axillary, rarely terminal; plants monoecious 

or dioecious. Flowers usually small, regular, unisexual or bisexual; sepals 3-5, almost 

always partly united; petals 3-5, free; stamens inserted at the base of the disc, sometimes 

arranged in two whorls, with the inner whorl alternate with the petals and the outer whorl 

(if present) opposite the petals, anthers 2-celled, opening lengthwise; disc intrastaminal, 

sometimes rather inconspicuous; ovary superior, 2-5-lobed, with 1-5 chambers, or with 

free carpels; ovule one in each carpel, anatropous, placentation axile. Fruits usually not 

splitting, often drupe-like, sometimes a samara or in A llantospermum septicidally splitting 

into 5 valves. Seeds with scant or no endosperm. 

Distribution. Some 30 genera and 200 species, distributed in the tropics and subtropics and 

some in temperate Asia. In Sabah and Sarawak, represented by 9 genera with 11 species of 

which only one (Harrisonia) includes scrambling shrubs. 

Ecology. Simaroubaceae species are found mostly in lowland forest. Quassia indica shows 

preference for temporarily inundated areas while Eurycoma longifolia has a distinct 

preference for acidic, leached, well-drained soils. Soulamea amara occurs in the Barringtoniaformation 

of the coastal vegetation and prefers calcareous or rocky beaches. Pollination is 

probably by insects as the flowers are often reported to be fragrant. 

Uses. Ailanthus, Allantospermum and Jrvingia are the only genera reaching timber size. All 

the bitter-tasting genera are used locally in the preparation of traditional medicines, 

especially as tonics, antidysenterics and antihelminthics, and the roots of Eurycoma 

longifolia has been reported to contain biologically active compounds useful as anti-malaria 

drugs (Chan et al., Planta Medica 52 (1986) 105). 

Taxonomy. The Simaroubaceae are closely related to the Burseraceae, Meliaceae and 

Rutaceae. While the genera are quite distinctly recognised, the limits of these families 

overlap somewhat. Forman (Kew Bull. 19 (1965) 517) placed Jrvingia in the Irvingiaceae 

421


andAllanthospermum in the Ixonanthaceae but Nooteboom (I.e. (1972) 970) concluded that 

the morphological, phytochemical and palynological evidence favoured including these 

genera in the Simaroubaceae. 

Key to genera 

1. Sprawling or scrambling spiny shrubs .......................................................... .. 

Harrisonia R. Brown ex A. Juss. 

Mem. Mus. Hist. Nat. Paris 12 (1825) 517; Masamune I.e. 362; Nooteboom, FM 1,6 

(1962) 207 . 

. 3-4 species in the Old World Tropics. 1 species, H. perforata (Blanco) Merr., 

occurs in Sabah. Scandent prickly shrub; branches with thorns; leaf-stalk winged. 

Lowlands to 400 m, also on Banggi island. Vernacular names: Sabah--bogua 

(Dusun Banggi), kait-kait (Tenom Murut), kukualang (Malay). 

Shrubs or trees, unarmed ....................................... .. .. ..................... 2 

2. 

Leaflets margin toothed ..................................... 

.. ............. 3. Brucea 

Leaves or leaflets margin entire .... .. .. ............ 3 

3. Leaves simple 

Leaves pinnate . 

.. .4 

..7 

4. Leaves spirally arranged, densely clustered at shoot tips, obovate, stalks 3-8 cm long 

.................................................................................................................. 8. Soulamea 

Leaves spiral to alternate and well-spaced along the shoots, elliptic, stalks 0.5-2 cm 

long ............................................................................................................................. 5 

5. Leaves with pitted glands. Stipules absent.. ..................................... 7. Quassia (in part) 

Leaves without such glands. Stipules either conspicuous conical structures or present 

only in leaf buds on young shoots and falling off early ................................................. 6 

• 

6. Twigs with annular stipule-scars, not swollen at leaf insertion. Leaf-stalks 10-20 mm 

long .............................................................................................................. 5. Irvingia 

Twigs without annular stipule-scars, swollen at points of leaf insertion. Leaf-stalks 5- 

44 mm long .................................................................................... 2. Allantospermum 

7. Leaflets sessile .. .. ........... .4. Eurycoma 

Leaflets distinctly stalked .... ........................................................................ 8 

8. Leaflets very unequal at base. Fruits winged ................ . ..1. Ailanthus 

Leaflets symmetric at base. Fruit a drupe ............. . ........................................ 9 

9. Veins sunken on both upper and lower leaflet surfaces; stipules absent.. ....... 

....... ...... ................ .......... ... ...................... ... ....... ................. ............ 7. Quassia (in part) 

Veins prominent on both upper and lower leaflet surfaces; stipules present, suborbicular 

..................................................................................................... 6. Picrasma 

422

SIMAROUBACEAE (KULIP & WONG) 

1. AILANTHUS Desf., nom. cons. 

(from the Amboinese plant name aylanto) 

Mem. Phys. Math. Ac. R. Se. Paris (1786) 270, t. 8; Nooteboom, FM 1,6 (1962) 215; Koehummen 

I.e. 346; Whitmore, Tantra & Sutisna l.c. 329. 

Dioecious trees with straight bole but without buttresses. Bark pale grey, smooth or 

lenticellate; inner bark yellowish brown. Sapwood pale white. Twigs thick, with large leafscars. 

Leaves more or less tufted at the ends of twigs, pinnate, to 60 cm long, without a 

terminal leaflet; leaflets 1O-l3 pairs, 7.5-12.5 x 2.5-5 cm, hairy below, with scattered 

glands in the forks of the veins on the underside; base very unequal, margins entire, apex 

pointed; the leaves eventually drooping with bowed stalk. Flowers 5-6-merous; calyx small, 

5-6-lobed, closed in bud and later irregularly splitting (often 2-lobed) to the base, rarely 

cupular; petals 5-6, induplicate-valvate in bud, concave; stamens 10, in male flowers 

inserted below the outer margin of the disc, in female flowers either of subnormal size (but 

without pollen), or vestigial, or absent, anthers opening laterally or externally, the 2 cells 

free in their lower half; ovary 2-5-carpellate, carpels free, flat, in the male flower vestigial 

or absent; styles 2-5, free or united; ovule 1 in each carpel. Fruit a samara, elliptic or 

oblong-lanceolate. Seeds flat, orbicular or obovate or somewhat triangular, without endosperm. 

Distribution. 5 species in tropical and subtropical SE Asia from Turkestan and India to 

China, through Malesia to Solomon Islands, Queenslands and northern New South Wales 

in Australia. In Sabah and Sarawak, 2 species are known. 

Ecology. In Sabah and Sarawak, both species are uncommon and found mainly in lowland 

forests below 1000 m, in valleys, along streams, and open places. 

Key to Ailanthus species 

Leaflets with lower surface glabrous and with a pair of large glands .......... 1. A. integrifolia 

Leaflets with lower surface hairy or sparsely hairy and with scattered small glands ............. . 

...................................................................................................................... 2. A. triphysa 

1. Ailanthus integrifolia Lam. 

(Latin, integer = entire,folia = leaves or leaflets; the leaflet margins) 

Diet. 3, 2 (1792) 417; Merrill, Interpr. Rumph. (1917) 299; Nooteboom, FM 1, 6 (1962) 218; 

Koehummen I.c. 346; Coekbum I.e. 219; Whitmore, Tantra & Sutisna, I.e. 229. Type: Rumphius 

Herb. Amb. 3:205, t. 132. Synonyms: A. blancoi Merr., Sp. Blanc. (1918) 205; A. peekelii Mekh., 

Notizbl. Berl. Dahl. 10 1930) 893; Dysoxylum dasyphyllum Miq., Ann. Mus. Bot. Lugd. Bat. 4 

(1868) 19. 

Tree to 55 m tall, 65 cm diameter. Bark smooth, light brown or grey. Sapwood white, 

yellow, pale brown or creamish, very soft; heartwood absent. Leaves 30-200 cm long, 

423


stalks 5-20 cm long; leaflets 2-9 pairs, 3.5-14 x 3.3-6.2 cm, glabrous on both surfaces; 

base very oblique or sickle-shaped, apex blunt -acute; stalks 0.5-1. 5 cm long; lateral veins 

6-13 pairs; glands on lower surface large, black, flat, oblong, 0.5-5.0 mm diameter, 

mostly paired near the base. Inflorescences to c. 40 cm long, glabrous, pedicels to c. 15 

mm long. Flowers with calyx more or less pubescent, closed in bud, rupturing and toothed 

irregularly, rarely cupular, 1-4 mm high, rarely caducous; petals puberulous, acute or 

bluntish, to c. 9 x 3 mm; filaments with many long spreading hairs to glabrous, usually 

thickened downwards, c. 0.5 mm in female flowers, to 4 mm long in male flowers; anthers 

c. 1 mm in female flowers, to 2.5 mm long in male flowers; ovary 5, usually densely 

puberulous; styles 5, connate at the base, including the long, stellately spreading stigmas, to 

c. 6 mm long. Fruits of (1-)3-5 samaras, each somewhat elliptic, 11-22 x 2.5-5 cm, the 

vein reticulations distinct on the outside, pale green; stalk 2.5-5 cm long. Seeds flat; testa 

thin; cotyledons 2. 

Distribution. Malesia: all islands, except Java and the Lesser Sunda Islands, and 

Melanesia (Bismarcks and Solomons). In Sabah, the species has been recorded from the 

Sandakan and Beaufort districts. In Sarawak, once collected at Suai, Miri, 4th Div. (s. 

39203). Also in Kalimantan. 

Ecology. In primary rain forest or rarely secondary forest, very rare. 

Uses. In New Guinea and the Bismarcks, the timber is made into planks for house 

construction (Nooteboom l. c.). 

2. Ailanthus triphysa (Dennst.) Alston Fig. 1. 

(Greek, tri = 3-partite, phusis = in character; the calyx-tube) 

Handb. Fl. Ceyl. 6, Suppl. (1931) 41; Nooteboom, FM 1,6 (1962) 219; Koehummen I.e. 346; 

Coekbum I.e. 219; Whitmore, Tantra & Sutisna I.e. 329. Basionym: Adenanthera triphysa Dennst., 

Sehluss. Hort. Mal. (1818) 32. Type: Dennst., Sehluss. Hart. Mal. (1818) t. 32. Synonyms: A. 

philippinensis Merr., Publ. Gov. Lab Phi lip. 35 (1906) 25; Hebanga abliqua Radlk., Philip. J. Se. 6 

(1911) Bot. 366. 

Tree to 60 m tall and 50 cm diameter. Bark pale brown, smooth to lenticellate; inner bark 

yellowish brown, mottled. Sapwood white. Twigs thick, reddish brown hairy. Leaves to 30 

cm long, rachis slightly pubescent; leaflets almost sickle-shaped, c. 18 pairs, 7.5-11.7 x 3- 

4 cm, membranous, upper surface glabrous, lower surface densely hairy; base distinctly 

unequal (one side sharply acute, the other side rounded), margin wavy, apex pointed; 

lateral veins 8-20 pairs, more or less sunken above; glands small, scattered over the whole 

surface especially on the midrib and veins. Inflorescences many-flowered, more or less 

pubescent, c. 20-60 cm long; bracts small, ovate to triangular, falling early; pedicels to c. 4 

mm. Flowers with calyx pubescent, less than 1 mm high, the triangular acute lobes as long 

as the tube or a little longer; petals glabrous or nearly so, 3-5 x 1-1.5 mm; filaments 

twisted-folded in bud, filiform or sometimes attenuating from the base to the top; anthers c. 

1.2 mm long, 1 mm wide in male flowers, smaller in female flowers. Fruits of 1-3(-4) 

samaras, each somewhat obovate, 4.5-8 x 1.5-2.5 cm, ripening red; stalk 0.8-2 cm long. 

424


Fig. 1. Ailanthus triphysa. A, leafy twig; B, infructescence. CA from SAN 59251, B from SAN 49456.) 

425


B 

I,,· 

A 

Fig. 2. Allantospermum bomeense subsp. bomeense. A, fruiting leafy twig; B, infructescence. (All 

from S. 15016.) 

426


Distribution. India, Sri Lanka, Burma, Thailand, Vietnam, through Malesia (except 

Sumatra, Peninsular Malaysia, Lesser Sunda Is., and New Guinea) to Queenslands and the 

northern parts of New South Wales. In Sabah uncommon, recorded from the interior parts 

at Sook, Keningau and Tenom, and in the east coasts at Lahad Datu. Not yet recorded in 

Sarawak. Also in Kalimantan. 

Ecology. In primary lowland and hill forests. 

Uses. The resin is tapped and used as incense and traditional medicine in India. In Indo 

China the bark is burned as incense. The bark and the leaves are used for making a tonic, 

especially for post-childbirth. They also possess febrifuge properties and are used for 

treating dyspeptic complaints. The wood is used for making wooden shoes in Luzon (the 

Philippines), for fishing floats, catamarans, sword-handles, and spear-sheath in India, and 

for tea-boxes in Sri Lanka (Nooteboom l.c.). 

2. ALLANTOSPERMUM Fonnan 

(Greek, ala = wing, sperma = seed; the winged seeds) 

Kew Bull. 19 (1965) 516; Nooteboom I.e. (1972) 968; Kochummen I.e. 347; Cockburn I.e. 217; 

Whitmore, Tantra & Sutisna I.e. 329. 

Trees. Twigs without annular stipular scars, swollen at pOints of leaf insertion. Leaves 

simple, entire. Inflorescence a panicle. Flower bisexual, 5-merous; sepals 5; petals 5; 

stamens 10, free, anthers versatile; disc lO-lobed, intrastaminal; ovary superior, shallowly 

5-lobed, stigma with a tiny papillose head. Fruit a capsule, broadly ellipsoidal, twisted 

after splitting into 5 valves along the septa, leaving a central columella. Seeds cylindrical, 

ellipsoid, shiny and waxy. 

Distribution. 2 species, 1 in Malesia (Peninsular Malaysia and Borneo) and the other in 

Madagascar (A. multicaule (Capuron) Noot.). 

AIlantospermum borneense Forman 


Fig. 2. 

I.e. 517, t. 1; Nooteboom I.e. 1972) 972; Kochummen I.e. 347; Cockburn I.e. 217; Whitmore, Tantra 

& Sutisna I.e. 329. Type: Galau S. 15262, Sarawak, 1st Div., Semengoh Forest Reserve (ho1otype K; 

isotypes L, SAN, SAR). 

Tree, to 90 m tall and 60 cm diameter; bole fluted and often crooked; buttresses sharp and 

spreading, to 3 m high. Bark pale brown with grey patches, smooth or with distant, 

adherent, large, thinnish scales, minutely lenticellate; inner bark pink, mottled white. 

Sapwood yellowish brown. Twigs brown, slender, more or less zig-zag. Leaves elliptic to 

oblong, 6.5-15 x 1.7-3.3 cm, glossy above, dull beneath; base cuneate to broadly rounded, 

apex blunt-acuminate; midrib and veins prominent on both surfaces; lateral veins 5-10 

pairs, intermediate veins extending about half-way to the margin; stalks blackish, channelled 

427


/~::'jl 

/7 A~'~~// 

~ ./ 

,"_.;r ,". _'..__-- 

/~7 ~ .. !\, 

'i" " & 

I ,.. 

Fig. 3. Bruceajavanica. Fruiting leafY twig. (From Zainuddin 1713.) 

428


above, 0.5-4.4 cm long. Inflorescences (J -)3.3-5.9 cm long, bearing the scars of early 

caducous bracts. Flowers with a stalk 7-9 cm long; sepals white, boat-shaped, 3-4 x 2 mm, 

rounded at the apex, reflexed at anthesis; petals white, elliptic to ob ovate, 4-5 x 2.5-3 mm, 

membranous, reflexed at anthesis, caducous; stamens to 6 mm long, anthers c. 1 mm long; 

disc c. 1.5 mm diameter and 0.5 mm thick; ovary 5-lobed, c. 1.5 x 2 mm, style filiform, 3- 

4 mm long, purple, stigma knob-like. Fruits broadly ellipsoid, 5-lobed, 2.8-4.2 x 1-3.5 

cm, apical beak 2-10 mm long. Seeds cylindrical, often slightly curved, 2-2.5 x 4-6 mm. 


LeafY branches conspicuously zig-zag. Infloresences 3.3-5.9 cm long, laxly branched. 

Fruits 2-3.5 cm wide, apical beak 2-4 mm long .................. . 

subsp. borneense 

Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan). In primary mixed 

dipterocarp forest on sandy humult ultisols; in Sabah also on ultramafic soils. Apparently 

gregrariously flowering and fruiting at several year intervals. 

LeafY branches only slightly zig-zag. Inflorescences 1-2 mm long, condensed. Fruits about 

1.1 cm wide, apical beak 6-10 mm long ....................................... . 

subsp. rostratrum Noot. 

I.e. (1972) 972; Cockburn I.e. 217; Whitrnore, Tantra & Sutisna I.e. 330. Type: Agama SAN 

36068, Sabah, Lahad Datu, Pu1au Sakar Cholotype L; isotypes K, SAN). 

Known only from Sabah (Sandakan and Lahad Datu areas). 

Vernacular names. Sarawak--nyalin (lban). Brunei--kayu tulang (Malay), tulang (Iban). 

3. BRUCEA l.F. Mill., nom. cons. 

(J. Bruce, 1730-1794, a Scottish scholar and explorer) 

Icon. (1779) t. 25; Merrill I.e. (1921) 316; Ridley I.e. 361; Masamune I.e. 361; Corner I.e. 697; 

Nooteboom, FM 1, 6 (1962) 209; Kochummen I.e. 348; Cockburn I.e. 219; Anderson I.e. 322. 

Shrubs or small trees. Leaves pinnate, with a terminal leaflet; leaflets with toothed 

margins; stipules none. Flowers unisexual, in axillary inflorescences; sepals 4, united at 

base; petals 4, free; disc thick with 4 lobes; stamens 4, with short filaments, vestigial or 

absent in female flowers; ovary 4-carpellate, carpels free, styles free or united at base; 

ovules 1 in each carpel, attached above the middle. Fruit a drupe, hardly fleshy, with a 

stone. Seeds with very thin endosperm. . 

Distribution. 6 species in tropical Africa and Asia, including 2 in Malesia. In Sabah and 

Sarawak one species. 

Brucea javanica (L.) Merr. 


Fig. 3. 

J. Am. Arb. 9 (1928) 3; Kochummen I.e. 348; Cockburn I.e. 219; Nooteboom, FM 1, 6 (1962) 210; 

Anderson I.e. 322. Basionym: Rhusjavaniea L., Sp. PI. (1753) 265. Type: Osbeek, s.n., Java CL). 

429


.~, 

4 cm 

c 

I 

:f.1: 

~J-li 

2mm 

D 

n"';~ Od," 

Fig. 4. Ewycoma longifolia. A, flowering leafy twig; B, lower side of leaf; C, flower bud; D, open 

flower; E, infructescence. (A-D from SAN 88096, E from SAN 73999.) 

430


Synonyms: Lussa radja Rumph., Herb. Amb. (Auet.) 7 (1755) 27, t. 15; Brueea sumatrana Roxb., 

Hort. Beng. (1814) 12; Brueea sumatrensis Spreng., PI. Min. Cogn. 2 (1815) 90; Brueea amarissima 

Desv. ex Gomes, Mem. Aead. Se. Lisb. n.s. 4, 1 (1872) 30. 

Shrub or small tree to 5 m high. Leaves 20-40 cm long; leaflets ovate to oblong-lanceolate, 

3-15,4.5-11 x 1.5-4 cm, sparsely hairy above, more or less pubescent below, sometimes 

glabrous; stalks 2-5 mm, the terminal one much longer. Flowers greenish white to 

greenish red or purple. Fruits 1-4 together, 4-5 mm long. 

Vernacular names. Sabah---kuinin (Dusun/Kadazan Tambunan), mara (Maga), pail-pait 

(DusunlKadazan Kinabatangan), payas (DusunlKadazan Ranau), tongkat ali (papar Malay; in 

common with Eurycoma). Sarawak-jaloot (Murut). 

Distribution. From Sri Lanka and the Deccan Peninsula through SE Asia to S China and S 

Formosa, throughout Malesia and N Australia. In Sabah, it is found all over the state. In 

Sarawak, it has been recorded from the 1st, 2nd, 4th and 7th Div. Also in Brunei. 

Ecology. A common, light-tolerant plant, preferring open sites and secondary forest and 

thickets, forest edges and ridges, even occurring in sunny places in sandy dunes and on 

limestone rock. Flowering and fruiting throughout the year. 

Uses. The roots and fruits contain bitter principles which possess medicinal value and used 

as concoctions in the treatment of dysentery, diarrhoea and fever. 

4. EURYCOMA Jack 

(Greek, eurus = broad, kome = tuft or crust; the leaves crowded at the ends of branches) 

,Mal. Mise. 2 (1822) 45; Meuill I.e. (1921) 316; Ridley I.e. 361; Burkill, EPMP (1935) 984; 

Masamune I.e. 361; Corner I.e. 698; Nooteboom, FM I, 6 (1962) 203; Koehummen I.e. 349; 

Coekbum I.e. 219; Anderson I.e. 323. 

Trees, treelets or shrubs. Twigs stout with large leaf-scars. Leaves pinnate, with terminal 

leaflet, crowded at branch ends; leaflets sessile, opposite or subopposite; base slightly 

oblique, attached to the rachis with a prominent joint; lateral veins inconspicuous above 

and below. Inflorescence a downturned axillary panicle; plants monoecious or dioecious. 

Flowers unisexual, female always with large sterile stamens, males always with a pistillode; 

calyx small, 5-6-lobed; petals 5-6; stamens 5-6, on the sepals alternating with 5-6 

small staminodes, stamens and staminodes sometimes united with the base of petals; disc 

inconspicuous; carpels 5-6, each with 1 ovule, free, the styles slightly united, stigma peltate, 5- 

6-lobed. Fruit a nut, to 5 per flower, each on a short stalk c. 3 mm long, ellipsoid or ovoid. 

Seeds without endosperm. 

Distribution. 3 species in tropical SE Asia, Sumatra, Borneo and S Philippines. In Sabah 

and Sarawak, only 1 species. 

431


4cm 

'mm [ 

4cm 

D 

E 

Fig. 5. Irvingia malayana. A, flowering leafy twig; B, flower; C, flower with petals and stamens 

removed; D, twig with young fruit; E, fibrous mesocarp of fruit. (A-C from SAN 43118, D from SAN 

63884, E from SAN 74993.) 

432


Eurycoma iongifolia Jack 

(Latin, longus = long,folium = leaves) 

Fig. 4. 

I.e. 45; Ridley I.e. 362; MerrillI.e. (1921) 316, PEB (1929) 116; Masamune I.e. 361; BurkillI.e. 

984; Corner I.e. 604; Nooteboom, FM 1, 6 (1962) 205; Kochummen I.e. 349; Cockbum I.e. 219; 

Anderson I.e. 323. Type: Jack s.n., Sumatra (ho1otype K). Synonyms: E. merguensis Planch. in 

Hooker, Lond. J. Bot. 5 (1846) 584; Pieroxylon siamense Warb., Fedde Rep. 16 (1919) 256; Manoles 

asiatiea Gagn., Bull. Soc. Bot. FT. 98 (1951) 207. 

Spindly unbranched tree or shrub, to 8 m tall and 15 cm diameter, or with a few upright 

branches, each crowned by an umbrella-like rosette of leaves. Bark greyish brown, smooth. 

Leaves to 100 cm long; leaflets lanceolate to obovate-lanceolate, rarely oblong, 5-20 x 

1.5-6 cm; base oblique, apex blunt to slightly acuminate; midrib raised on both surfaces; 

lateral veins inconspicuous above and sunken below. Flowers reddish; petals hairy on both 

sides, c. 4.5-5.5 x 1.5-2.5 mm; styles rather long, stigma c. 1 mm above the ovaries. Fruits 

10-17(-20) x 5-12 mm. 

Vernacular names. Sabah--ionadiandau, nuad-mandau (Runggus), tombuid (Dusun/Kadazan 

Tambunan), tongkat ali (Malay), tongkat langit (Kuala Penyu Malay). Sarawak-bedara 

(Semantan Malay), sengkanyat (Iban), sengkayap (Iban), tongkat ali (Santubong & Miri), 

tungkat ali (Lundu). 

Distribution. Lower Burma, Thailand, Laos, Cambodia, Indo-China; Malesia: Sumatra, 

Peninsular Malaysia, and Borneo. Common throughout both Sabah and Sarawak; also in 

Brunei and Kalimantan 

Ecology. Abundant on well-drained sandy soils below 1200 m, in primary and secondary 

mixed dipterocarp, heath and submontane forests. 

Uses. In Sabah the roots are mixed with other medicinal plants, e.g., Cinnamomum species, 

and used to prepare a health tonic. In Brunei, the bark is used as a blood coagulant in 

complication during childbirth. The young leaves can be eaten raw to cure stomach-aches. 

In Peninsular Malaysia, Chan et al. (Planta Medica 52 (1986) 105) reported that methanolextracts 

of roots contained biologically active compounds showing a strong anti plasmodial 

activity against a multi-drug resistant Kl strain of Plasmodium falciparum from Thailand. 

The putative aphrodisiac properties of the roots have, as yet, been substantiated by rigorous 

experiment. 

5. IRVINGIA Hookf 

(E.G.lrving, 1816-1855, Scottish botanist) 

pauh kijang 

Trans. Linn. Soc. 23 (1860) 167; Rid1ey I.e. 363; Corner I.e. 699; Nooteboom, FM 1, 6 (1962) 223; 

Kochummen I.e. 350; Cockbum I.e. 221; Anderson I.e. 323; Whitmore, Tantra & Sutisna I.e. 330. 

Large trees; buttresses steep, to 6 m high. Bark fawn, smooth with distant loose scales, 

minutely lenticellate; inner bark mottled, cream-yellow. Sapwood orange-brown. Twigs 

433


with stipules forming a narrow, conical cap surrounding the terminal buds, soon falling, 

leaving conspicuous annular scars. Leaves simple, glabrous, entire. Inflorescences axillary 

and terminalpanicies. Flowers (4-)5-merous, bisexual; sepals connate (united) at the base; 

petals overlapping in buds; stamens twice as many as petals, inserted beneath the large, 

cushion-shaped, intrastaminal disc; ovary 2-chambered, conical or somewhat flattened, 

sessile; style 1, stigma inconspicuous; ovules solitary. Fruit a drupe, large, 1-2-seeded, 

resembling a mango. 

Distribution. 3 species in tropical Africa and 1 species in tropical SE Asia and W Malesia. 

Ecology. Frequent in lowland forests. 

Uses. The fruit of all species is edible, but usually only the seeds are eaten. 

Irvingia malayana Oliv. ex AW. Benn. 

(of Malaya) 

Fig. 5. 

in Hooker [, Fl. Brit. Ind. 1 (1875) 522; Rid1ey I.e. 364; Corner I.e. 699; Nooteboom, FM 1, 6 

(1962) 223; Kochummen I.e. 350; Cockburn I.e. 221; Burgess TBS (1966) 455; Anderson I.e. 323; 

Whitmore, Tantra & Sutisna I.e. 330. Type: Maingay 298, Malacca (holotype K). Synonyms: 

Irvingia oliveri Pierre, Fl. For. Coch. 4 (1892) t. 263 B; Irvingella malayana van Tiegh., Ann. Se. 

Nat. 9, 1 (1905) 276; Irvingella oliveri (Pierre) van Tiegh. and Irvingella harmandiana van Tiegh. 

I.e. 279; Irvingia harmandiana (van Tiegh.) Pierre ex Lecomte, Fl. Gen. I.-C. 1 (1911) 701; Irvingia 

longipedieellata Gagnep., Fl. Gen. I.-C. Suppl. 1 (1946) 670. 

Medium-sized to large tree reaching 50 m tall and 50 cm diameter, with big limbs; 

buttresses steep, plank-like and spreading, to 3 m high. Bark greyish to whitish, scaly to 

flaky, sometimes smooth, minutely lenticellate. Leaves elliptic-oblong to lanceolate, 8-20 x 

2.5-9 cm, upper surface shiny, lower surface slightly glaucous especially when fresh; base 

often rounded, apex usually pointed; midrib raised above; lateral veins 10-16 pairs, looping 

and joining at margin, prominent on both surfaces; stipule-cap 3-4 cm long. Flowers 

greenish white or yellowish, small. Fruits ellipsoid, c. 6 x 4 cm, slightly glaucous. Seedling 

with first two leaves opposite; germination epigeal. 

Vernacular names. Sabah--mengkudu (DusunlKadazan TuaranIRanau; doubtful, as this 

name normally refers to Morinda in the Rubiaceae), pauh kijang (Malay), selangan tandok 

(Malay), tenghilan (DusunlKadazan Tuaran). Sarawak-patok entilit (Iban). 

Distribution. Thailand, Indo-China, and Malesia (Sumatra, Peninsular Malaysia, Borneo 

and Bawean). Widespread in Sabah and Sarawak. Also in Brunei and Kalimantan. 

Ecology. Scattered in mixed dipterocarp forest on clay-rich soils, to 300 m. 

Timber. Burgess (i.c. 455) summarises the timber properties of this species. Pauh kijang 

produces a very strong and springy timber, hard to saw and work, due to the high density. 

It takes a very fine finish, and requires very little filling, and turns very well. In Sandakan, 

furniture of this timber glued with synthetic resin glues has tended to fail at the glue-line. 

434


Uses. The yellow wood is too hard to work with and not very durable. In Peninsular 

Malaysia, it has been used for making kris-handles and handles of tapping knives 

(Nooteboom, I.c.). The seeds contain a creamy yellow, nice-smelling fat known as "dika" 

fat in Europe, used for making soap, wax, and candles. The seeds can also be eaten 

(Nooteboom,l.c.). 

6. PICRASMA Blume 

(Greek, pikros = bitter, osme = smell or taste; the bark and other parts) 

Bijdr. 5 (1825) 247; Ridley I.e. 361; Burkill I.e. 1723; Nooteboom, FM 1, 6 (1962) 212; 

Kochummen I.e. 351; Anderson I.e. 323; Whitmore, Tantra & Sutisna I.e. 330. 

Trees or shrubs. Leaves pinnate, with terminal leaflet, stalk base and rachis nodes usually 

swollen; leaflets opposite or subopposite, entire, veins prominent on both upper and lower 

surfaces; stipules present, suborbicular and falling off early. Inflorescences axillary, longpeduncled, 

compound-cyme, unisexual (plants monoecious or dioecious). Flowers 4-5- 

merous, female usually twice as large as male; sepals small, free to united half way up; 

petals persistent in female, much longer than the sepals; stamens 4-5; disc thick; carpels 

up to 7, free, each with 1 ovule, vestigial or absent in male; styles united except at base, 

sometimes 1 or 2, free; stalks jointed in the lower half. Fruit 1-4, drupe-like; exocarp thin, 

fleshy, wrinkled when dry; endocarp hard. Seeds without endosperm. 

Distribution. About 8 species, 6 in tropical America, 2 species in Asia including 1 in 

Malesia. 

Uses. Picrasma species contain alkaloids, the source of quassia chips used in insecticide. 

Picrasma javanica Blume 


Fig. 6. 

I.e. 248; Ridley I.e. 361; BurkillI.e. 1723; Nooteboom, FM 1, 6 (1962) 213; Kochummen I.e. 351; 

Anderson I.e. 323; Whitrnore, Tantra & Sutisna I.e. 330. Type: Blume, s.n., Java (holotype L). 

Synonyms: P. nepalensis A.W. Benn., PI. Jav. Rar. (1844) 201; P. andamaniea Kurz ex A.W. Benn. 

in Hooker f I.e. (1875) 520; P. philippinensis Elmer, Leafl. Philip. Bot. 5 (1913) 1837. 

Tree to 24 m tall and 25 cm diameter; bole fluted. Bark dark, smooth, brittle; inner bark 

dull yellow. Sapwood with clearly visible vessels. Leaves with 5-7 leaflets, stalk 2-6 cm 

long; leaflets entire, 4-20 x 1-10 cm; base wedge-shaped, margin wavy or wrinkled, apex 

acuminate; lateral veins 3-8 pairs, petiolules to 7 mm long; stipules leafY, nearly rounded, 

7-25 x 5-20 mm, usually falling off early leaving a large scar. Inflorescences to 20 cm 

long. Flowers 4-merous, white to yellow or green; sepals glabrous to puberulous, triangular 

to ovate, c. 1 mm; petals ovate-oblong or oblong, often acute-acuminate to mucronate, 

glabrous, or sparsely hairy, with a conspicuous midrib; stamens usually longer than petals 

in male flowers, shorter than petals in female flowers, filaments gradually thinner towards 

the top, hairy at base, 0.5-2 mm long in female flowers and 1-5 mm in male flowers, 

anthers 1-2 mm long in male and to 1 mm and empty in female flowers; ovary 4-lobed, 4- 

435


carpellate, glabrous to hairy, styles 1-1. 5 mm long, stigmas e. 2 mm. Fruits green to red or 

blue, ovoid to depressed globlose, 9-10 x 7-12 mm. Seeds with a broad hilum; testa rather 

thick and hard. 

Vernacular names. Sabah-balimbing, panguban (DusunlKadazan Tambunan). Sarawakkayu 

pahit (Malay). 

Distribution. Tropical SE Asia (from Sikkim, Assam, Burma and Tonkin southward to 

Malesia). In Borneo, recorded in Sarawak only at about 400 m on the Gunung (Mt.) Api 

limestone, at Mulu, and in Sabah on the west coasts. 

Ecology. Uncommon, usually scattered in rainforests from near sea-level to 1500 m. 

Uses. The bark contains quassin, which gives its bitter taste. In Java, the leaves have been 

applied for treating sores (Nooteboom I.e.). The trunk is too small for timber and the wood 

is not durable (Burkilll. c.). 

7. QUASSIA L. 

(named after a slave in Surinam who reported 

the medicinal properties of the wood to Dalberg, friend of Linnaeus) 

Sp. Pl. ed. 2 (1762) 553, l.c. (1763) 1679; Merrilll.c. (1921) 315 (as Samadera); Ridley I.c. 363; 

Burkilll.c. 1945; Masamune·l.c. 362 (asSamadera); Nooteboom, FM 1, 6(1962) 198, Blumea 11 

(1982) 514; Kochummen l.c. 352; Cockburn l.c. 217. 

Trees or shrubs. Leaves pinnate or simple, with pitted glands on the upper surface along 

the margin and especially at the apex; stipules and scars absent. Inflorescence a simple or 

branched raceme, a panicle, or an umbel. Flowers 4-6-merous, unisexual or bisexual, or 

polygamous; petals imbricate or contorted in bud, longer than the calyx, sometimes very 

long; stamens hairy, adaxial scale with a shorter or longer free apex; disc cylindrical or 

subglobose; carpels free, more or less puberulous; style 1, with a terminal, inconspicuous 

stigma. Fruits 1-6 per flower, drupaeeous or woody, often compressed laterally, with a 

narrow, unilateral, sharp-edged thinner part in the apical half. Seeds with a thin testa, 

without endosperm. 

Distribution. Pantropical, e. 25 species in tropical and subtropical America, 5-10 species 

in Africa, 2 species in lower Burma and Cambodia (one of which is found almost 

throughout Malesia), 1 species endemic to Borneo and Sumatra, and 2 species in Queenslands. 

Ecology. In lowland rain forests. 

Key to Quassia species 

Leaves pinnately compound. Flowers in panicles. Fruits 1-5 from each flower, drupaceous, 

slightly flattened, ellipsoid ......................................................................... 1. Q. borneensis 

436

437 

Fig. 6. Picrasma javanica. A, flowering leafy twig; B, stipules; C, male flower; D, female flower; E, 

fruits. (A from de Wilde & de Wilde-DuyjJes 14843, B-'-E after FM 1,6 (1962) 213, fig. 15.) 

'ml 'mm [ i ],mm 

D B E 

SIMAROUBACEAE (KULIP & WONG)

""~ 

438 


Fig. 7. Quassia indica. A, fruiting leafy twig; B, inflorescence. CA from S. 42968, B from S. 16407.)


Leaves simple. Flowers in umbel-like clusters. Fruits 1-4 from each flower, strongly 

flattened, with straight inner and semicircular outer margin .............................. 2. Q. indica 

1. Quassia borneensis Noot. 


FM 1,6 (1962) 203, Blumea 11 (1962) 518; Cockbum I.e. 219; Anderson I.e. 323; Whitmore, Tantra 

& Sutisna I.e. 330. Type: Meijer SAN 20499, Sabah (holotype L; isotypes K, SAN). 

Tree to 25 m tall and 25 cm diameter; buttresses low. Bark pale yellow to greyish brown, 

densely fissured-corky. Sapwood white. Leaves pinnate, spirally arranged; leaflets 2-4 

pairs, elliptic to obovate-oblong, 8-12 x 4-4.5 cm, glabrous, upper surface shiny, lower 

surface dull; with small pitted glands along the margins and in the acumen on the upper 

surface; base cuneate, apex shortly rounded to acuminate; lateral veins sunken on both 

surfaces, obscure, ending in a marginal vein; stalk c. 5 cm long, rachis terete, petiolules 1- 

1.5 cm long, articulated at the base. Inflorescences puberulous all over, shorter than the 

leaves. Flowers (male) 4-5-merous, pedicels to 7 mm long; calyx c. 1 mm high; petals 

contorted or imbricate in bud, glabrous, elliptic to ovate-oblong, 3-4 x 2 mm; stamens 

slightly shorter than the petals, anthers oblong; disc c. 0.5 mm high, at the base c. 2 mm 

wide and at the apex c. 1 mm wide, the upper half distinct from the lower half and folded 

around the barren ovaries. Fruits 1-5 in each flower, drupaceous, prune-shaped, dark 

purple-red when ripe, slightly flattened-ellipsoid, with a faint dorsal and ventral ridge, 2-3 

x 1.5 cm; pericarp thin but hard. Seeds with a thin testa; cotyledons large, green, planoconvex. 

Vernacular names. Sabah-mamungal (Malay), pait-pait (Malay). Sarawak-medang pahit 

(Malay). 

Distribution. Malesia: Sumatra (Indragiri), Borneo (Sabah and Sarawak). Uncommon. 

Ecology. Primary mixed dipterocarp forest on humult ultisols; also, rarely, in peat swamp 

and kerangas (heath) forests. 

2. Quassia indica (Gaertn.) Noot. 

(of the Indies) 

Fig. 7. 

FM 1,6 (1962) 199, Blumea 11 (1962) 517; Kochummen I.e. 352; Cockbum I.e. 217; Anderson I.e. 

323; Whitmore, Tantra & Sutisna I.e. 330. Basionym: Samadera indiea Gaertn., Fruct. 2 (1791) 352, 

t. 156, f. 3. Type: Gaertner, Fruet. 2 (1791) 352, t. 156, f 3. Synonyms: Manungala pendula 

Blanco, Fl. Filip. (1837) 306; Samadera brevipetala Scheff., Nat. Tijd. Ned. Ind. 32 (1871) 410. 

Tree or shrub, to 20 m and 8 cm diameter. Bark brownish green, smooth; inner bark 

pinkish. Sapwood pale yellow; cambium yellow. Branchlets with small pith, with several 

stiff persistent scales at the base of each shoot. Leaves simple, elliptic-oblong to lanceolate, 

12-13 x 4-12 cm; base acute or sometimes rounded, or subcordate, apex blunt or 

acuminate or sometimes rounded; midrib, lateral and intercostal veins prominent on both 

439


'om [ 

D 

lmm[~ ....;.. · 

~ 

lmm 

Fig. 8. Soulamea amara. A, fruiting leafy twig; B, fruit; C, flower; D, longitudinal section of flower 

with pistil removed. (After FM 1, 6 (1962) 222, fig. 21.) 

440


surfaces; stalks 1-2.5 cm long. Inflorescences jointed at the lower half, 0.5-2.5 cm, 

growing during anthesis; bracts minute. Flowers to 20 or more; calyx 4-lobed, 2-3 mm 

long, lobes about as long as or longer than the tube, puberulous outside; petals 4, free, 

dorsally puberulous, obtuse, usually narrowed to the base, creamy green to violet, to 3 x I 

cm; filaments puberulous, hairy except toward the apex, to 2.5 cm long, inserted at the base 

of the disc, anthers lanceolate to oblong, c. 4 x 2 mm; styles to 2 mm long. Fruits 1-4 

together, flattened, with straight inner and semicircular outer margin, which is sharp and 

thinner in the upper half, the apex more or less overtopping the subapical stylar scar, 4-9 x 

2.5 cm. 

Vernacular names. Sabah--kacang-kacang (Malay), kelapahit (Sook Murut). Sarawakmanuggal 

(Iban). 

Distribution. Madagascar, Sri Lanka, S Concan, Malabar, Lower Burma (Martaban, 

Tenasserim), Andamans, and Cochinchina, through Malesia to the Bismarcks and 

Solomons. In Sabah, it is found mainly in the east coast (Sandakan & Lahad Datu 

districts), with only a few records from the west coast (Sipitang district). In Sarawak, it is 

found throughout the state. 

Ecology. Locally abundant in tidal swamp forests below 150 m, sometimes in localities 

which are periodically inundated by fresh or salt water, for example on the edge of 

mangroves. Occasional in freshwater swamp forest; also occurs in mixed dipterocarp forest. 

Uses. In Sarawak, the wood is used for making knife-handles. The seeds are given as an 

emetic and purgative, and sometimes in bilious fevers (Nooteboom I.c.). 

8. SOULAMEA Lam. 

(soelamoe, a Ternatean name for the plant) 

Diet. Enc. Meth. 1 (1783) 449; Masamune I.e. 362; Nooteboom, FM 1,6 (1962) 221; Coekburn I.e. 

217. 

Shrubs or small trees. Leaves simple, obovate, spirally arranged, densely clustered at 

shoot tips, sometimes with few glands underneath. Flowers in axillary racemes or narrow 

thyrses, 3(-4-5)-merous, bisexual; floral parts persistent; sepals more or less connate at the 

base, slightly imbricate in bud; petals longer than sepals; stamens twice as many as petals, 

in 2 distinct rows, inserted under the lower outer margin of the disc, anthers versatile; disc 

3 (-4-5)-lobed, each lobe forked; ovary (l-)2-3-carpellate, styles horizontally adnate to 

their carpels, stigmas small; ovules sessile. Fruits dry, (1-)2(-3)-celled, indehiscent, flattened, 

distinctly winged, more or less emarginate, rarely flattened, ovoid, acute. Seeds attached 

adaxially nearly halfway down; testa thin; cotyledons pIano-convex. 

Distribution. 9 species. One species is endemic to the Seychelles, 6 species occur in New 

Caledonia, and one species in Fiji. One species is widely distributed in Malesia and Polynesia, 

and it occurs in Sabah and Sarawak. 

441


Soulamea amara Lam. Fig. 8. 

(Latin, amarus = bitter; the taste of the tissues) 

l.c. (1783) 449; Mique1, Fl. Ind. Bat. 1,2 (1859) 129;.Masamune I.c. 362; Nooteboom, FM 1, 6 

(1962) 221; Cockburn I.c. 217. Type: Rumphius Herb. Amb. t. 415 (L). Synonyms: Rex amaroris 

Rumph., Herb. Amb. 2 (1743) 129, t. 41; Cardiocarpus amarus Reinw., Syll. Ratisb. 2 (1826) 14; 

Cardiophora hindsii Benth. & Hook.f, Lond. J. Bot. 2 (1843) 216. 

Shrub or small tree to 5(-15) m tall; young shoots and buds rusty tomentose. Leaves 

crowded at the apex of the branchlets, on dropping leaving large scars; blade obovateoblong, 

10-35 x 4-12 cm; base cuneate, apex blunt but sometimes mucronate; midrib and 

veins hairy below; midrib slightly immersed or inconspicuous above, strongly prominent 

beneath; lateral veins straight, parallel, ending in an intramarginal looped vein, sulcate, 

slightly prominent or inconspicuous above; intercostal veins finely dense-reticulate beneath; 

stalks pithy, shrunken at the base when dry, sometimes also at apex, hairy, 3-8 cm long. 

Inflorescences erect, shorter than the leaves, 3-l2 cm long. Flowers c. 2 mm long; pedicels 

to 5 mm long; sepals puberulous, erect, appressed, 0.5-1.0 mm long; petals concave, 

spreading, finally reflexed, sparsely hairy to glabrous, accrescent, to 2.5 x 1 mm; stamens 

with glabrous filaments to 1 mm long, anthers c. 0.75 mm iong; ovary 2-3-carpellate, never 

with more than 2 carpels fertile, carpels connate except at the top. Fruits obcordate, to 2 x 

2.5 cm, strongly emarginate; pericarp hard and corky; wings often nearly touching near the 

inward curved style-bases. Seeds round, 0.5-1 cm across. 

Distribution. From Borneo eastwards to Micronesia (West and East Carolines and 

Marshalls) and Melanesia (New Britain, Solomons, New Hebrides); in Malesia: Borneo 

(Sabah, Sarawak, Karimata Is.), Moluccas, and New Guinea. 

Ecology. A typical constituent of the Barringtonia-formation of the beach vegetation, but 

much rarer than most of the species belonging to that formation, though locally common on 

the sandy beaches and behind coral reefs. 

Uses. None known in Sabah and Sarawak. The roots and the fruits of this very bitter plant 

have been used to treat cholera, pleurisy, and other fevers; a beverage prepared from 

powdered leaves is taken against colic and cough, and the fruits have been used to induce 

vomitting in treating snake bites (Nooteboom l.c.). 

442

SONNERA TIACEAE 

Othman Bojo 

Universiti Malaysia Sarawak, 

Kota Samarahan, Malaysia 

MerrilI, EB (1921) 418; Holthuis & Lam, Blumea 1, 5 (1942) 216; Backer & van Steenis, FM 1,4 

(1951) 280, FM 1, 5 (1958) 557, FM 1,6 (1972) 973; Whitmore, TFM 1 (1973) 442; Cockburn, TS 

1 (1976) 223; Anderson, CLTS (1980) 323; Ashton, MNDTS 2 (1988) 374; Whitmore, Tantra & 

Sutisna, CLK2, 1 (1990) 331. 

443 

Taxonomy. Previous authors such as Ridley (FMP 1 (1922) 819), Watson (MFR 6 (1928) 

50), Browne (FTSB (1955) 248), Hsuan Keng (OFMSP (1978) 155), Corner (WSTM 1 

(1988) 470) and Brummitt (Vasc. ·PI. Fam. & Gen.-Dicot. (1992) 607) included Duabanga 

Uses. The soft pale wood of the Sonneratiaceae is not durable and its only known use is as 

firewood. However, timber of Duabanga is occasionally used for boat-building. Fruits of 

some species (S caseolaris and S ovata) are edible, and the leaves and pneumatophores of 

S caseolaris are reported to have medicinal value. 

Ecology. Whereas Sonneratia is a mangrove genus found mainly on sandy and muddy tidal 

flats, estuaries, brackish streams and coral terraces at tide level, Duabanga is a component 

of lowland and hill forests, and being light-demanding, is frequently found in secondary 

forests, forest-edges and river-banks. The flowers produce a mild sour to musty odour, 

expand at sunset, last for only one night and are pollinated by nectarivorous bats. The 

fruits/seeds are dispersed either by water (Sonneratia) or by wind (Duabanga). 

Distribution. 2 genera with 7 species distributed in the Old World Tropics from East 

Africa to the Pacific islands. In Sabah and Sarawak, 2 genera with 4 species. 

Trees. Bark cream to grey, smooth or shallowly irregularly flaky; inner bark soft, brown. 

Sapwood soft, pale yellow. Leaves simple,. opposite, biseriate, entire, coriaceous, shortstalked, 

without stipules. Inflorescences terminal corymbs or 1-3-flowered fascicles. 

Flowers bisexual, pedicelled, rather large, regular; sepals persistent, thickly leathery, 

connate at base to form a tube or cup with 4-8 triangular lobes, segments valvate in bud, 

often reddish inside; petals absent or as many as sepals, broad and wrinkled or very narrow 

and smooth, alternating with the sepals; stamens 12 to many, inserted on the sepals, 1- 

many seriate, inflexed in bud, filamens filiform to subulate; anthers 2-celled, kidneyshaped 

or oblong, opening lengthwise; ovary superior, sessile and with a broad base, 

enclosed by the calyx-base until it expands, 4-20-celled, septa thin; style 1, long, stout, 

stigma 1, capitate or slightly lobed; ovules anatropous many; placentation axile. Fruits 

many-seeded berries (in Sonneratia) or dehiscent capsules (in Duabanga), subtended by 

the persistent calyx-tube. Seeds small, without endosperm; embryo straight. 


$. 

'-- 

"'-,r 

lcm '\\' C 

o 

Fig. 1. Duabanga moluccana. A, flowering leafy twig; B, flower with some sepals and petals 

removed; C, petal; D, longitudinal section of flower with sepals and petals removed; E, fruit. (A-D 

from UNIMAS/OB 82, E from Hansen 207.) 

444 


~ ,">.~'

SONNERATIACEAE (OTHMAN) 

and Sonneratia in the Lythraceae. This contention seems to be supported by anatomical 

(Metcalfe & Chalk, Anat. Dicot. 1 (1964) 6), embryological (Venkateswarlu, Proc. Indian 

Acad. Sci. Ser. B, 5 (1937) 206), and palynological evidence (Erdtman, Poll. Morph. & PI. 

Tax. 1 (1952) 412; Thanikaimoni & Jayaweera, Tran. Sect. Sci. & Techn., Inst. Pranc. 

Pondichery 5, 2 (1966) 1; Muller, Pollen et Spores 2 (1969) 223; Germeraad, Hopping & 

Muller, Rev. Palaeobot. & Palyn. 6 (1968) 189). However, accounts by Backer & van 

Steenis (I.c.), Cockburn (l.c.), Ashton (l.c.), Mabberley (PB (1987) 546), and van Steenis 

(CheckI. Gen. Names Males. Bot., Sperm. (1987) 115) include the two genera in the 

Sonneratiaceae, a view which is adopted here. For anatomical information on the family, 

the work of Reinders-Gouwentak (FM 1, 4 (1951) 513) should be consulted . 

. Key to genera 

Monopodial trees of inland forests, without breathing roots; branches drooping. Leaves 

coriaceous, glaucous below, with many prominent lateral veins and intra-marginal veins, 

base heart-shaped. Flowers in terminal corymbs; petals broad, wrinkled. Fruits 4-8-valved 

capsules. Seeds tailed at each end by the protracted testa .................................. 1. Duabanga 

Sympodial trees of mangroves and sea-coasts, with breathing roots; branches not drooping. 

Leaves succulent, not glaucous, venation obscure; base not heart-shaped. Flowers in 

clusters of 1-3 at the ends of the branchlets; petals absent or very narrow. Fruits 

indehiscent berries. Seeds not tailed ................................................................ 2. Sonneratia 

1. DUABANGA Buch.-Ham. 

(from duyabangga, an Indian vernacular name) 

Trans. Linn. Soc. Lond. 17 (1893) 177; Rid1ey, FMP 1 (1922) 824 (under Lythraceae); Burkill, 

EPMP (1935) 868; Backer & van Steenis I.e. 288; Jayaweera, 1. Am. Arb. 48 (1967) 89; Ashton I.e. 

375; Kochumrnen & Wyatt-Smith, MFR 17 (1964) 271; Cockburn I.e. 223; Corner, WSTM (1988) 

476 (under Lythraceae); Whitmore, Tantra & Sutisna l.c. 331. 

Medium-sized to big trees; crown monopodial, rather open, ultimate branches drooping; 

bole with insignificant concave buttresses at base. Bark smooth, pale ochreous-cream or 

grey; inner bark pale yellow, soft, fibrous. Sapwood white, soft. Twigs 4-angled or winged, 

often with an interpetiolar ridge. Leaves oblong-ovate, base heart-shaped, glaucous 

beneath, coriaceous; lateral veins prominent beneath, looped at the margin forming intramarginal 

veins; petioles short. Flowers 5-many, in terminal corymbs, 4-8-merous; calyx 

fleshy, green, cup-shaped with triangular segments; petals shortly clawed, white or 

yellowish, broad, wrinkled; stamens 12 or many, I-seriate; anther recurved over one end of 

the connective; ovary 4-8-celled; stigma stout, lobed. Fruits 4-8-valved, loculicidally 

dehiscent. Seeds tailed at both ends by the extended testa. 

Distribution. 2 species, from eastern Himalaya to New Guinea. Only 1 species (D. 

moluccana) is found in Sabah and Sarawak and is common. 

445


Ecology. Mainly in mixed evergreen and deciduous forests at 15-1200 m. Usually 

occurring as scattered individuals on brownish yellow sandy clay loam soils or yellow 

podsols. The seeds are readily dispersed by wind. 

Taxonomy. Based on the revision of Sonneratiaceae by Backer & van Steenis (l.c. 288), 

Duabanga consists of two species, D. grandijlora and D. moluccana. In 1967, Jayaweera 

(l.c. 89), added one new species, D. taylorii Jayaweera. However, detailed study of D. 

taylorii by Geesink (Blumea 2, 18 (1970) 454) reveals that this species is a primary hybrid 

between D. grandiflora and D. moluccana which is believed to have originated in the 

Bogor Botanic Garden in the 1850s. 

Uses. In Sabah and Sarawak, its non-durable and soft wood has no commercial value. 

However, in other regions, the wood is used for house and boat-building and also for 

firewood. 

Duabanga moluccana Blume 

(of the Moluccas islands) 

Fig. 1. 

Mus. Bot. Lugd. Bat. 1 (1849) 109; Merrill, PEB (1929) 212; Holthuis & Lam l.c. 216; Backer & 

van Steenis I.c. 288; Jayaweera I.c. 91; Cockburn I.c. 223; Anderson I.c. 323; Ashton I.c. 375; Corner 

l.c. 476; Whitmore, Tantra & Sutisna l.c. 331. Type: Forstein, s.n., Moluccas, Ambon (holotype L). 

Synonym: D. borneensis R. Knuth, Fedde Rep. 38 (1935) 121. 

Tree to 45 m tall and 100 cm diameter, with or without buttresses; trunk columnar. Bark 

smooth becoming fissured or scaly with age, lenticellate, grey or brownish; inner bark 

yellowish or pale brown, soft and fibrous. Sapwood cream, darkening purplish on 

exposure. Twigs 3-8 mm diameter, slender, 4-angled, becoming terete with age, hairy 

when young, glabrescent. Leaves ovate, oblong or oblanceolate, 10-26 x 4-13 cm, dark 

green above, paler beneath, leathery; base shallowly heart-shaped, apex acuminate; midrib 

prominent beneath; lateral veins 11-26 pairs, joined near the margin to form an intramarginal 

vein; petioles 5-10 mm long. Inflorescences few to many-flowered, dense or 

rather lax, pubescent at first, becoming glabrous. Flower-buds 2 x 1.5 cm, ovoid, ribbed, 

acute. Flowers 4-merous, with musty odour; calyx-tube c. 2 cm across at anthesis, lobes 4, 

c. 1-2 cm deep; petals 4, c. 2 x 1.2 cm, yelloWish or cream, falling off early; stamens 12, 

anthers yellow turning to pale brown; style c. 40 mm long, dark red, stigma dark green. 

Fruits ovoid or oblong, 4-valved, 1.5-3 x 1-2 cm. Seeds c. 0.7 mm long. 

Vernacular names. Sabah--magas (Kadazan). Sarawak-benung kasung (Bidayuh), sawih 

(!ban). 

Distribution. Java, Borneo (Sabah, Sarawak, Brunei, Kalimantan), Celebes, Moluccas, and 

New Guinea. 

Ecology. In Sabah and Sarawak, widely distributed at 15-750 m, on damp clay-rich fertile 

soils, especially in high light intensity areas such as river-banks, forest-edges, logged-over 

forests, road-sides, abandoned cultivation sites and also on limestone hills. 

Uses. The timber is little used at present. Recorded uses include temporary light 

construction, floaters for extracting timber logs, fish-net floats and dug-out boats. It grows 

446


fast alld may have potential to be planted for pulping materials or for core-stock in plywood 

manufacturing. 

2. SONNERATIA L.j, nom. cons. 

(Pierre Sonnerat, 18th century French explorer-naturalist) 

SuppL PI. Syst.'Veg. (1781) 38; Merri1l.e. (1921) 416; Rid1ey I.e. 825; Watson I.e. 121; Burkilll.e. 

2051; Backer & van Steenis I.e. 282; Wyatt-Smith, MF 16 (1953); Kochummen & Wyatt-Smith I.e. 

315; \Vhitmore I.e. 444; Cockbum I.e. 224; Anderson I.e. 323; Duke & Jackes, Blumea 32 (1987) 

277; Ashton I.e. 375; Corner I.e. 476 (under Lythraceae); Whitmore, Tantra & Sutisna I.e. 33l. 

Synonyms: Blatti Adall,s., Fam. 2 (1763) 88; Pagapate Sonnerat, Voy. Nouv. Guin. (1776) 16. 

Small to medium-sized glabrous trees, with sympodial crown and low branching; bole 

fluted at base, surrounded by spindle-shaped breathing roots (pneumatophores), buttresses 

absen t. Bark smooth, becoming scaly or finely fissued with age, lenticellatc, pale greygreen 

to dark brownish grey; inner bark pinkish or brownish, rather watery. Young twigs 

(branchlets) drooping or erect, distinctly jointed above the nodes, 4-angled, becoming 

terete with age. Flowers in terminal clusters of 1-3, 4-8-merous; calyx cup-shaped, 4-8- 

lobed, segments ovate or triangular, often reddish within; petals absent or 4-8, narrow, 

and falling off early; stamens many, falling off early; ovary 10- to many-celled; style 

slender, sinuous in bud. Fruits indehiscent berries, depressed-globose, tipped by the 

persistent style and seated on the persistent star-like calyx-cup, pericarp leathery. Seeds 

many, embedded in foul-smelling pulp, not tailed at both ends; embryo straight. 

Distribution. 5 species distributed along sea-shores of tropical East Mrica, Asia, Malesia, 

Australia and Pacific Islands. 2 species occur in Sabah and three in Sarawak. 

Ecology. Mangrove trees, found on sandy and muddy tidal flats, estuaries, brackish watercourses, 

river-banks and creeks and coral-terraces at tide level. The flowers are ephemeral 

and pollinated by nectar-feeding bats and birds. The fruits are water-dispersed. Hybrids 

have been recorded between S. alba x S. caseolaris and S. alba x S. ovata along the Brunei 

coasts, and their status has been substantiated by palynological (Muller, Ancient Pacific 

Floras (1964) 33), morphological and cytological (Muller & Hou-Liu, Blumea 14 (1966) 

337) evidence. 

Uses. The timber is classified as moderately hard to hard and moderately heavy to heavy. 

However, it is only locally used for firewood and occasionally for house-posts. Burgess 

(TBS (1966) 461) claimed that the wood of S. caseolaris could be pulped by the sulphate 

process to give a pulp with similar strength properties to commercial eucalypt pulp. The 

fruits of some species are edible. In Sarawak, fruits of S. caseolaris and S. ovata are edible. 

Key to Sonneratia species 

1. Leaves elliptic to elliptic-ovate. Young twigs drooping. Petals linear-lanceolate, dark 

red; filaments bright pink at base, upper part white ................................ 2. S. caseolaris 

Leaves obovate, broadly ovate to suborbicular. Young twigs erect. Petals (if present) 

linear, white; filaments white ...................................................................................... 2 

447


f 'om 

~'mm[r 

~ ~~ ;1 

I 

"~,, il 

B J ~ 

E 

Fig. 2. Sonneratia ova ta, A, flowering leafy twig; B, pistil with other floral parts removed; C, 

stamen; D, longitudinal section of flower; E, fruit (All from living specimens,) 

448


2. Leaves obovate, base cuneate. Petals linear, white ........................................... 1. S. alba 

Leaves broadly ovate to suborbicular, base rounded or sub cordate. Petals absent.. .......... 

..... 3. S. ovata 

1. Sonneratia alba J. Smith 

(Latin, albus = white; the flowers) 

in Rees, Cyc!. 33 (1819) 2; Beccari, Nelle For. di Borneo (1902) 579; Merrill, Interp. Rumph. Herb. 

Amb. (1917) 383, I.e. (1921) 418; Backer & van Steenis I.e. 285; Wyatt-Smith I.e. 214; Browne I.e. 

323; Duke & Iackes I.e. 286; Ashton I.e. 379; Corner I.e. 476; Whitmore, Tantra & Sutisna I.e. 33l. 

Type: Rumph. Herb. Amb. 3 (1754) 111, t. 73, Ambon. Synonyms: Rhizophora easeolaris 1. in 

Stickman, Herb. Amb. (1754) 13, in part, Blatti alba (J. Smith) Kuntze, Rev. Gen. 1 (1891) 238. 

Tree to 40 m tall and 70 cm diameter; stems usually crooked; crown rather open, more or 

less broadly oblong, low-branching. Young twigs erect. Bark slightly cracked to fissured, 

grey to brownish above normal tide mark, lenticels usually present; inner bark not 

laminated, mottled cream and pinkish brown. Leaves fleshy, broadly obovate to rarely 

ovate, 3.5-10 x 2-7.5 cm; base cuneate, apex broadly rounded; lateral veins 7-9 pairs, 

thin, inconspicuous; petioles 7-13 mm, stout. Flower-buds oblong, 2-2.5 x 1.5 cm. 

Flowers solitary or in terminal clusters of 2-3, 6-7(-8)-merous; calyx 1.5-2.5 cm long, 

cup ob conical, ribbed or angular, angles as many as segments and alternating with them, 

segments 6-7, ovate-oblong, 1.3-2 cm long, inner surface red, especially at base; petals 6- 

7, linear, inconspicous and almost resemble the filaments, white, pink at base; filaments 

white; ovary 14-18-celled. Fruits 4-5 cm long, 3-4 cm across, dark green, subtended by 

the ribbed calyx-cup. 

Vernacular names. Sabah-pedada (Malay, Kadazan). Sarawak-perepat (Malay, lban). 

Distribution. Tropical East Africa, N Madagascar, Seychelles, Sri Lanka, Andamans, 

continental SE Asia, throughout Malesia to Australia and the W Pacific Islands. In Sabah, 

a common component of mangroves in Kudat, Sandakan, Lahat Datu and Semporna. In 

Sarawak, very common in Buntal Bay from Santubong to Bako National Park and 

elsewhere such as the Lawas mangroves. Also in Brunei and Kalimantan. 

Ecology. A pioneer mangrove species found on muddy and shallow parts of calm seashores 

and along the mouths of tidal creeks, forming dense pure stands on intertidal sites. 

Rarely found far from river mouths and sandy beaches. 

Uses. In Sabah and Sarawak, the wood is used as firewood. Elsewhere, such as in Minahasa 

(NE Celebes), the wood is valued for ship and house-building. 

2. Sonneratia caseolaris (L.) Engl. 

(Latin, caseus = cheese; the fruit which resembles a small, depressed rounded piece of 

cheese) 

in Eng1er & Prantl, Nachtr. (1897) 261; Merrill I.e. (1917) 383, I.e. (1921) 418; Backer & van 

Steenis I.e. 283; Wyatt-Smith I.e. 213; Browne I.e. 248; Whitmore I.e. 445; Cockburn I.e. 225; 

449


Anderson I.e. 323; Duke & Jackes I.e. 289; Ashton I.e. 379; Corner I.e. 476; Whitmore, Tantra & 

Sutisna I.e. 331. Basionym: Rhizophora easeolaris 1., p.p. in Stickman, Herb. Amb. (1754) 13. 

Type: Rumph. Herb. Amb. 3 (1754) 112, t. 74-75, Ambon. Synonyms: S. acida 1., Suppl. 252; 

Aubletia easeolaris (1.) Gaertn., Fruct. (1788) 379; Blatti easeolaris (1.) Kuntze, I.e. 238; Blatti 

aeide (1.) Lam., Enc. 1 (1789) 429; S. pagatpat Blanco, Fl. Filip. (1837) 424; Blatti pagatpat 

(Blanco) Niedenzu in Eng1er & Prantl, Pfl. Fam. 3,7 (1891) 21; S. ovalis Korth., Ned. Kruidk. Arch. 

1 (1846) 198; S. evenia Blume, Mus. Bot. Ind. Bat. 1 (1851) 337; S. laneeolata Blume I.e. (1851) 

337; S. obovata B1ume I.e. (1851) 337; S. negleeta B1ume I.e. (1851) 338. 

Tree to 20 m tall and 30 cm diameter; stems frequently leaning and crooked; crown rather 

open. Bark smooth, greenish grey and lenticellate when young, becoming irregularly 

fissured and pale brown with age; inner bark cream or pinkish brown, faintly laminated; 

exudate watery. Young twigs drooping, yellowish brown turning to greenish brown with 

age. Leaves elliptic to elliptic-ovate, 3.8-9.5 x 2-8 cm; base cuneate, apex blunt or 

subacuminate; lateral veins 8-12 pairs, thin and inconspicuous; petiole 7-11 mm long, 

reddish. Flower buds broadly ovoid or oblong to subglobose, 2-2.5 x 1-1.5 cm. Flowers 6- 

8-merous; calyx at anthesis shallowly cup-shaped, smooth, segments 6-8, usually longer 

than the tube, cream or pale yellow inside, base crimson, forming a crimson ring around 

the fruit; petals linear-lanceolate, 6-8, 20-25 x 1.5-2 mm, dark red; filaments white, 

bright pink at base; ovary 16-21-celled. Fruits c. 4 cm long, 7.5 cm across, dark green, 

crowned with persistent conically thickened style base, subtended by the flattened, green 

calyx-tube with horizontally spreading lobes. 

Vernacular names. Sabab--perepat (Malay, Kadazan). Sarawak-pedada (Iban, Malay). 

Distribution. Sri Lanka, tropical SE Asia to New Guinea, Northern Australia, the Solomon 

Island, and New Hebrides. In Sabah and Kalimantan, very common in mangrove front. In 

Sarawak and Brunei, common along tidal river-banks and mangroves, especially on deep 

muddy river-banks, and from the limit of brackish water down to the lower end of the 

Nypa-zone near the river-mouths. 

Ecology. Individual trees of S. caseolaris are usually found scattered among other 

mangrove species (e.g., Avicennia species) rather than forming a pure stand. 

Uses. The young, sour-tasting berries are edible and are also used in traditional medicine, 

chiefly externally but also internally (Burkilll.c. 2088). Burkill & Haniff (Gard. Bull. S.S. 

6 (1930) 204) reported that leaves pounded with rice were applied for small-pox. Pectin is 

locally extracted from the fruits. The wood is used as fuel, and the pneumatophores, after 

boiling in water, furnish an inferior substitute for cork. 

3. Sonneratia ovata Backer 

(Latin, ovatus = oval; the leaf shape) 

Fig. 2. 

Bull. Jard. Bot. Btzg. 3, 2 (1920) 329; van Steenis, Bull. Jard. Bot. Btzg. 3, 12 (1931) 162; Backer & 

van Steenis I.e. 162; Wyatt-Smith I.e. 214; Whitmore I.e. 445; Anderson I.e. 323; Duke & Jackes I.e. 

299; Ashton I.e. 380; Corner I.e. 476; Whitmore, Tantra & Sutisna I.e. 331. Type: Backer 21422, 

Java (lectotype L). 

450


Tree to 20 m tall and 20 cm diameter; stems short and usually twisted. Bark grey, smooth 

to slightly fissured, lenticellate; inner bark pale brown to reddish, faintly laminated. Young 

twigs erect, greyish brown. Leaves usually ovate to orbicular or broadly ovate, 2.5-7.5 x 

3-9 cm; base rounded or sub cordate, apex broadly rounded; adaxial surface slightly 

corrugated by the fine but conspicuous 9-10 pairs of lateral veins; petioles 4-11 mm long. 

Flowers usually in terminal clusters of 2-3(-4), occasionally solitary; buds broadly ovoid, 

apex rounded or obtuse, finely verruculose, 1.5-2 x 1-1.5 cm; calyx cup-shaped, tapering 

abruptly into a stalk-like base, ribbed, segments usually 6, ovate-triangular, 13-15 mm 

long, inner surface tinged reddish at base; petals absent; filaments white; ovary 13-15- 

celled. Fruits c. 4 cm long, 7 cm across, dark green when young, turning to yellowish 

green when ripe. 

Vernacular name. Sarawak-rogam (Malay). 

Distribution. China, Thailand and throughout Malesia. In Sarawak, Brunei and Kalimantan 

very local. In Sabah, has not been recorded with certainty. 

Ecology. Occasionally found on banks of tidal rivers and muddy soils inundated only by 

spring tides. 

Uses. In Sarawak, the mature fruit is edible and eaten as ulam (a salad) by the Malays. 

451


STAPHYLEACEAE 

J. T. Pereira 




Merrill, EB (1921) 355; Rid1ey, FMP 1 (1922) 511; van Steenis, Nova Guinea 10,2 (1959) 211; van 

der Linden, FM 1,6 (1960) 49; Airy-Shaw, Kew Bull. 18,2 (1965) 252; Backer & BakhuizenJ, FJ 2 

(1965) 145; Sponberg, Journ. Am. Arb. 52 (1971) 196; Whitmore, TFM 1 (1972) 446; Anderson, 

CLTS (1980) 324; Dickison, Journ. Am. Arb. (1984) 149; Whitmore, Tantra & Sutisna, CLK 2, 1 

(1990) 332; Webster, Ann. Missouri Bot. Gard. 81 (1994) 54. 

Trees or shrubs, non-Iaticiferous. Leaves simple to compound (imparipinnate to paripinnate), 

oppposite or alternate, leaflet margin toothed; stipules usually falling off early .. 

Inflorescences terminal, subterminal or axillary panicles or racemes. Flowers bisexual, 

sometimes unisexual, actinomorphic (with regular symmetry); sepals 5, imbricate; petals 5, 

imbricate; stamens 5, alternating with the petals outside the disc; anthers 2-celled, 

longitudinally dehiscent, dorsifixed; disc present or absent; ovary superior, 2-4-celled, 

styles 2-4, free or fused together, stigmas capitate or coherent, forming a peItate structure; 

ovules one or more per cell, placentation axile or basal. Fruit an indehiscent berry, follicle 

or inflated capsule. Seeds one to several, arillate or exarillate; embryo straight; cotyledons 

flat; endosperm fleshy or horny. 

Distribution. 5 genera with about 60 species. Staphylea (11 species), Tapiscia (1 species) 

and Euscaphis (1 species) are found in the north temperate region. Huertea (4 species) 

occurs from West Indies to Peru. Turpinia has 30-40 species distributed from the Indomalesian 

region to Japan and tropical America. In Sabah and Sarawak, Turpinia is 

represented by 6 species all of which occur in Sabah, although only 2 species are found in 


Uses. Some species of the temperate Staphylea and Euscaphis are cultivated as ornamental 

plants. Timbers of a few species of Turpinia are occasionally utilised locally. The fruit of 

Euscaphis japonica, a common tree or shrub in Japan and Central China, is used for 

preparing a drug. 

Taxonomy. In early publications, the family was considered closely affiliated to the 

Rhamnaceae. Later ~uthors (c! van Steenis I.c. 211, van der Linden I.c. 49, Spongberg I.c. 

196) placed it near to the Aceraceae, Celastraceae, Cunoniaceae, and Sapindaceae. Its 

affinity to the Cunoniaceae was based on the evidence derived from morphology, anatomy 

and embryology (Hallier, 1908; Whitmore l.c. 446; Thorne, 1976; Dickison I.c. 149). 

Anatomically, there are major differences between the Staphyleaceae and the Sapindaceae 

(Metcalfe & Chalk, Anat. Dicot. 1 (1957) 443) although the Staphyleaceae have some 

453


similarities to the woody Saxifragaceae in terms of nodal and leaf anatomy (Dickinson, Bot. 

Gaz. 148,4 (1987) 475). Whitmore (l.c. 446) included Bischojia under the Staphyleaceae, 

which had been initially placed in the Euphorbiaceae by Decaisne (1842) and later isolated 

as a separate family, the Bischofiaceae, by Airy-Shaw (1965). However, evidence from 

embryology and anatomy appears to support its inclusion in the Euphorbiaceae (Webster, 

1994). The Indochinese Tricaphis was also included in the Staphyleaceae by Gagnepain 

(Not. Syst. 13 (1948) 190) but van Steenis (1960) excluded it, and suggested that it 

belonged to either the Anacardiaceae or Sapindaceae. Hence, the taxonomic status of this 

family, particularly the position of certain genera, remains controversial. 

TURPINIA Vent., nom. cons. 

(P.J.Fr. Turpin, 1775-1840, an eminent French botanical artist) 

Choix (1803) 31, t. 31; Ridley I.e. (1922) 511; Merrill & Perry, Joum. Am. Arb. 22 (1941) 543; 

Metcalfe & Chalk I.e. 443; van Steenis I.e. 211; van der Linden I.e. 49; Backer & BakhuizenJ I.e. 

145; Burkill, EPMP 2 (1966) 2234; Grey-Wilson, Kew Bull. 26,1 (1972) 141; Whitmore I.e. 446; 

Anderson I. e. 324; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 332; Pereira, Sandakania 5 (1994) 

15. Synonyms: Trieeros (non Griff) Lour., FI. Coch. 1 (1790) 184; Dalrympelea Roxb., PI. Corom. 

3 (1820) 76, t. 279; Oehranthe Lindl., Bot. Reg. 8 (1836) t. 1819; Hasskarlia Meisn., PI. Vasc. Gen. 

2 (1843) 348; Kaembaehia (non O. Kuntz.) Schltr., Bot. Jahrb. 52 (1914) 15l. 

Trees or shrubs. Twigs smooth, terete, pithy, glabrous or hairy. Stipules interpetiolar, entire 

or bifid at apex, falling off early (except in T. stipulacea), leaving prominent scars. Leaves 

opposite-decussate, unifoliolate, trifoliolate to pinnate (imparipinnate or paripinnate), 

petioles wrinkled at base; glands present at base of leaflets and at the insertion of 

petiolules to the rachis; leaflets chartaceous to coriaceous, margin serrate to crenate, midrib 

prominent. Inflorescences 

• 

axillary, terminal to subterminal panicles or racemes. Flowers 

bisexual; outer sepals broader than the inner ones, ovate, margin ciliate; petals of equal 

size, longer than sepals, margin ciliate; stamens of equal size, broad at the base, glabrous, 

anthers oval, apex sometimes apiculate; disc lobed, fleshy; ovary superior, (2-)3(-4)-celled; 

styles closely adnate together forming a peltate stigma; ovules one to numerous in each 

cell, in axile plancentation in 2 rows. Fruit a globose to sub globose or trilobed, indehiscent, 

fleshy berry, typically crowned by three radial lines or horn-like structures (remains of the 

styles) on top; fruit wall thin to thick, fleshy. Seeds mostly angular, one to a few in each 

cell, pale to dark brown; cotyledon flat, roundish. 

Distribution. About 30-40 species from Sri Lanka to southern Japan, southward and 

eastward to Malesia, west Pacific and the central and south American tropics (West Indies, 

Columbia, Ecuador, Peru). In Sabah and Sarawak, the genus is represented by 6 species, 2 

of which are endemic to Sabah, viz., Turpinia nitida and Turpinia stipulacea. 

Ecology. Evergreen understorey trees or shrubs of lowland tropical to subtropical rain 

forests to montane zones reaching c. 2400 m in New Guinea and Mt. Kinabalu and to 3000 

m in the Himalayas and Yunnan. Fruits are probably dispersed by birds and other animals 

and the hard-coated seeds can probably withstand acidity and digestion in intestines during 

dispersal by animals. 

454

STAPHYLEACEAE (PERElRA) 

Uses. Turpinia species produce timber of a low durability, hence usage in construction is 

exceptional. The wood is used for packaging cases in Indonesia (Burkill I.c. 2234). 

Turpinia nepalensis, common in western China, produces a useful tough wood. Most 

Turpinia species are grown as pioneer trees for reafforestation due to their rapid growth on 

eroded mountain slopes in central Java. 

Key to Turpinia species 

(based mainly on leaf characters) 

1. Stipules 14-25 mm long, conspicuous and persistent on apices ofleafy twigs ............... . 

.... ...... ....... ................... ....... .... ...... ....... ............. ... ................................ . 6. T. stipulacea 

Stipules not exceeding 10 mm in length, not very conspicuous, falling off early from 

leafy twigs ................................................................................................................... 2 

2. Lower surface of leaflets sparsely to densely short-hairy on the midrib, lateral veins 

and blade .................................................................................................. 3. T. grandis 

Lower surface of leaflets glabrous ................................................................................ 3 

3. Intercostal veins pseudoscalariform (with many almost straight cross-veins between the 

lateral veins) ..................................................................... 5. T. sphaerocarpa (in part) 

Intercostal veins reticulate (with wide areoles, or obscurely or distinctly tessellate, with 

no or very few straight cross-veins between the lateral veins) ...................................... .4 

4. Leaflets thickly coriaceous, margin strongly recurved when dry ................... 4. T. nitida 

Leaflets chartaceous or coriaceous, margins not strongly recurved when dry ................ 5 

5. Leaflets narrowly acuminate to caudate, acumen usually 1-2 cm ......... 1. T. borneensis 

Leaflets shortly acuminate to cuspidate, acumen to c. 1 cm long, often shorter.. ........... 6 

6. Leaves typically 20-46 cm long (rachis 10-18 cm long) .... 5. T. sphaerocarpa (in part) 

Leaves often less than 20 cm long (rachis less than 10 cm long) .................................. 7 

7. Stipules 2.5-3.5 x 2-2.5 mm. Leaflets coriaceous and slightly shiny on lower or both 

surfaces. Fruit globose to subglobose, without short apical horns, but with radial lines 

at the apex. (Plants restricted to limestone habitats.) .............................. 2. T. calciphila 

Stipules 5-7 x 3-4 mm. Leaflets chartaceous and not shiny. Fruit slightly trilobed at 

the apex, with short apical horns. (Plant of non-limestone habitats.) ............. . 

.... ...... ...... ..... ...... ...... ..... ........ ...... ...................... ... ... .......... 5 T. sphaerocarpa (in part) 

Key to Turpinia species 

(based mainly on fruits and flowers) 

1. Fruits slightly to distinctly trilobed at the apex, typically with short apical horns ......... 2 

Fruits globose to subglobose, without short apical horns .............................................. 3 

455


2. Petals 2-4 x 1.5-2 mm; anthers 0.8-1 mm long; pistils 2.5-3.5 mm long. Seeds 5-8 

mm across. Leaflets oblong, elliptic-oblong to lanceolate or ovate, typically shiny 

above, coriaceous, margin strongly recurved ................................................ 4. T. nitida 

Petals 1.8-2 x 1-1.1 mm; anthers 0.5 mm long; pistils 2-2.2 mm long. Seeds 2-4 mm 

across. Leaflets ovate-elliptic to elliptic, not shiny above, chartaceous, margin not 

recurved .................................................................................. 5. T sphaerocarpa (in part) 

3. Fruit-wall brittle, 0.1-0.3 mm thick (Plants restricted to limestone habitats) ................. . 

...... . 2 .. T. calciphila 

Fruit-wall not brittle, thicker than 0.5 mm (Plants not restricted to limestone 

habitats) ...................................................................................................................... .4 

4. Ovules (2-)3-4 per cell .................. ........ 5 

Ovules (4-)6-8(-9) per cell .......... ... 6 

5. Petals 2.5-4.5 mm long; stamens 2-3 mm long; pistils 2-3 mm long, ovary 1-1.5 mm 

across. Stipules 14-25 mm long, conspicuous and persistent on apices of leafy 

twigs ..................................................................................................... 6. T. stipulacea 

Petals 1.2-2 mm long; stamens 0.8-1.8 mm long; pistils 1-2 mm long, ovary 0.6-1 

mm across. Stipules 3-8 mm long, not very conspicuous, falling off early from leafy 

twigs .................................................................................................... 1. T. borneensis 

6. Seeds 5-14 per fruit. Lower surface of leaflets sparsely to densely short-hairy on the 

midrib, lateral veins and blade. Leaf rachises 3-8 cm long ........................................... . 

................ 3. T. grandis 

Seeds 1-7 per fruit. Lower surface of leaflets completely glabrous. Leaf rachises 10-18 

cm long ............................................................................. 5. T. sphaerocarpa (in part) 

1. Turpinia borneensis (Merr. & Perry) B.L. Linden 


I.e. 49; Anderson I.e. 324; Whitmore, Tantra & Sutisna I.e. 332. Basionym: Turpinia montana var. 

bomeensis Merr. & Perry I.e. 549. Type: 1. & M. S. Clemens 29391 bis, British North Borneo, 

Kinabalu, Tenompok (holotype DC; isotypes BO, K). 

Tree to 20 m tall and 20 cm diameter. Bark shallowly fissured, pale brown; inner bark 

greyish. Sapwood brownish. Stipules 3-8 x 2-4 mm, not very conspicuous, falling off early 

from leafy twigs, glabrous, margin sparsely hairy to entire. Leaves 3-5( -7)-foliolate, (13-) 

16-35(-40) cm long; petioles 3-11 cm long, 1-2 mm thick, glabrous; leaf-rachis 4.5-13 

cm long, glabrous; petiolules of lateral leaflets 2-15 mm long, glabrous; leaflets drying 

pale green above, chartaceous, glabrous, not shiny, ovate-elliptic to lanceolate, 5-18 x 2-7 

cm; base cuneate, sometimes rounded or oblique, margin finely serrate, not recurved, apex 

narrowly acuminate to caudate, acumen (0.5-)1-2 cm long; lateral veins 7-11 pairs, 

distinctly looping at the margin, prominent on lower surface; intercostal veins reticulate or 

distinctly tessellate and prominent, with no or very few straight cross-veins between the 

lateral veins on both surfaces. Inflorescences axillary panicles, to 30 cm long, rachis 

subglabrous. Flowers cream to pale yellow, sepals ovate, thin, 1.2-2 x 1-1.8 mm; petals 

obovate, 1.2-2 x 0.8-1.5 mm; stamens 0.8-1.8 mm long, glabrous, filaments 0.5-1.5 mm 

456

STAPHYLEACEAE (PEREIRA) 

long, glabrous, anthers 0.4-0.5 mm long, apiculate or not; pistils 1-2 mm long, styles 0.6- 

1.8 mm long; ovary 0.6-1 mm across, 3 (-4)-celled, each cell with 3-4 ovules. Fruits 

globose, 8-18 mm diameter, without short apical horns but with three radial lines at the 

apex, wrinkled; wall thin, not brittle, 0.5-1 mm thick. Seeds pale to dark brown, 4-10 x 6- 

10 mm, 1-4 in each fruit. 

Distribution. Borneo (Kalimantan and Sabah) and the Philippines; not yet recorded in 


Ecology. Primary and secondary mixed dipterocarp forest to 1800 m. Flowering has been 

recorded from November to March whereas fruiting has been noted from May to October. 

2. Turpinia calciphila 1. T. Pereira 

(Latin, calx = lime; Greek, philos = loving; referring to the habitat) 

Sandakania 5 (1994) 18. Type: Anderson S. 4710, Sarawak, Baram, Gunung Api (holotype SAR; 

isotypes BO, K, L, SING). 

Tree to 25 m tall and 50 cm diameter. Bark creamy yellow, slightly flaky. Stipules 2.5-3.5 

x 2-2.5 mm, not very conspicuous, falling off early from leafY twigs, glabrous, margin 

hairy. Leaves 1-3-foliolate, 9-16(-25) cm long, petioles 1.5-4(-7.5) cm long, 1-2 mm 

thick, glabrous; leaf-rachis 1.5-4 cm long, glabrous; petiolules of the lateral leaflets 5-14 

mm long, glabrous; leaflets drying pale green to brown above, coriaceous, glabrous, 

slightly shiny on lower or both surfaces, ovate to ovate-elliptic, 6.5-15 x 3.5-6.5 cm; base 

rounded to slightly cuneate, sometimes oblique, margin shallowly serrate, sometimes 

slightly undulate, not recurved, apex shortly acuminate, acumen 0.4-1 cm long; midrib 

and lateral veins raised and distinct on lower surface; lateral veins 6-8 pairs, ascending, 

parallel, open or looping towards the margin; intercostal veins reticulate or distinctly 

tessellate beneath, with no or very few straight cross-veins between the lateral veins. 

Inflorescences axillary panicles to 25 cm long, rachis densely hairy at the distal end. 

Flowers not known in mature state. Fruits globose to subglobose, 5-10 mm diameter, 

without short apical horns but with three radial lines at the apex, smooth; wall 0.1-0.3 mm 

thick, brittle. Seeds dark brown to reddish brown, 4-8 x 5-8 mm, 1-6 in each fruit. 

Vernacular name. Sarawak-Iaba (lban). 

Distribution. Endemic to Sarawak (4th Division, Baram, Gunung Api and Gunung Buda; 

1st Division, near Bau, Bt. Buan and Bt. Gebung). 

Ecology. So far only known from the lower, mesic slopes of limestone screes, to 900 m. 

3. Turpinia grandis B.L. Linden 

(Latin, grandis = large, big; the leaves) 

l.c. 49; Whitmore, Tantra & Sutisna l.c. 332. Type: Endert 4669, E Borneo, W Koetai, Kiau River, 

700 malt. (holotype L; isotypes BO, K). 

Tree to 20 m tall and 30 cm diameter. Bark smooth, greyish; inner bark yellowish. 

Sapwood pale white. Stipules 4 x 3-5 mm, not very conspicuous, falling off early from 

457


B \\ 

I 

3 cm """1 

,>;~ 

n 

A 

Fig. 1. Turpinia sphaerocarpa var. microcerotis. A, leafy twig; B, part of inflorescence; C, flower in 

longitudinal section; D, fruit. CA & D from J. & M.S. Clemens 28707, B & C from Kostermans 

10409.) 

458

STAPHYLEACEAE (PEREIRA) 

leafY twigs, densely hairy, margin hairy. Leaves 1-3-foliolate, 20-39 cm long, petioles 2.5 

-9 cm long, 2-4 mm thick, minutely hairy; leaf-rachis 3-8 cm long, short-hairy; petiolules 

of lateral leaflets 5-15 mm long, minutely hairy; leaflets drying dark brown above, ovate to 

ovate-oblong, 14-25 x 7-13 cm, thickly chartaceous, sparsely to densely short-hairy 

beneath; base narrowly rounded to cuneate, sometimes asymmetric, margin sharply serrate, 

not recurved, apex shortly acuminate, acumen 0.5-1 cm; midrib sparsely to densely shorthairy; 

lateral veins 7-12 pairs, ascending, parallel, open or sometimes looping towards the 

margin, distinct beneath, sparsely to densely short-hairy; intercostal veins pseudoscalariform 

(with many almost straight cross-veins between the lateral veins). 

Inflorescences axillary or terminal panicles to 35 cm long, rachis densely to sparsely hairy. 

Flowers pale yellowish; sepals ovate, thin, 1.8-2.5 x 1-2 mm; petals linear obovate, 2-3 x 

1-1.5 mm; stamens 2.5-3 mm long, glabrous, filaments 2-3 mm long, anthers 0.5-0.6 mm 

long, not apiculate; pistils 2-3 mm long, styles 1.5-2 mm long, ovary 1-1.1 mm across, 3- 

celled, each cell with (5-)6-8(-9) ovules. Fruits globose, 10-20 mm diameter, without 

short apical horns but with three radial lines at the apex, wrinkled, wall not brittle, 1.5-2 

mm thick. Seeds pale brown, 5-8 x 5-8 mm, 5-14 in each fruit. 

Distribution. Endemic to Borneo (Kalimantan, Sabah, Sarawak). In Sabah, collected from 

Tambunan (SAN 113560 and SAN 113536) and Gunung Lotung, Lamag (SAN 83167) and 

in Sarawak from the 4th Division, Baram, Kelabit Highlands (s. 35520). 

Ecology. Scattered on ridges, forested river-banks and deep ravines. Flowers recorded 

around May and fruits from October to November. 

4. Turpinia nitida Merr. & Perry 

(Latin, nitidus = shiny or polished; the leaf surface) 

l.c. 543; B.L. Linden l.c. 49; Whitmore, Tantra & Sutisna l.c. 33. Type: J. & M. S. Clemens 30840, 

British North Borneo, Mt. Kinabalu, Penibukan (holotype UC; isotypes BO, K). 

Tree to 15 m tall and 40 cm diameter. Bark fissured or lenticellate, greyish brown to black; 

inner bark reddish brown. Sapwood pale yellow. Stipules 3.5-10 x 2-5 mm, not very 

conspicuous, falling off early from leafY twigs, glabrous, margin entire to sparsely hairy. 

Leaves 1-3-5-7-foliolate, (15-)18-35(-39) cm long; petioles 3-13 cm long, 2-4(-6) mm 

thick, glabrous; leaf-rachis 2-18 cm long, glabrous; petiolules of lateral leaflets 1-25 mm 

long, glabrous; leaflets, drying pale green, sometimes pale brown, oblong, elliptic-oblong 

to lanceolate or ovate, 6-21 x 3-11 cm, thickly coriaceous, glabrous, typically shiny on 

upper surface; base rounded to acute, sometimes asymmetric, margin sharply to shallowly 

serrate, strongly recurved, apex acuminate to cuspidate, acumen 0.5-1.5(-2) cm long; 

lateral veins 6-11 pairs, ascending arcuately half way, parallel, open or sometimes looping 

towards the margin, obscure on upper surface; intercostal veins reticulate or distinctly 

tessellate, with no or very few straight cross-veins between the lateral veins on lower 

surface. Inflorescences axillary or terminal panicles, to 45 cm long, rachis glabrous or 

sparsely hairy. Flowers yellowish to pink-tinged, sepals oblong to ovate, 1.5-3.5 x 1.5-2.5 

mm, thin; petals spathulate, 2-4 x 1.5-2 mm; stamens 1.8-4 mm long, filaments 1.5-8 

459


mm long, anthers 0.8-1 mm long, slightly to distinctly apiculate; pistils 2.5-3.5 mm long, 

styles 2-3 mm long; ovary 1-2 mm across, (2-)3(-4)-celled, each cell with (3-)4-7(-8) 

ovules. Fruits 3-lobed, typically with three short apical horns at the apex, smooth, woody, 

8-20 mm diameter, wall not brittle, l.5-5 mm thick. Seeds pale brown, 5-8 mm across, 1- 

7 in each fruit. 

Distribution. Endemic to Sabah (Ranau, Sandakan, Kudat and Lahad Datu), not yet known 

in Sarawak. 

Ecology. Montane forest to 2400 m and also in primary and secondary forest on hills near 

streams. Frequently found on ultramafic soils. Flowers documented around November to 

May, fruits mostly during July to September. 

5. Turpinia sphaerocarpa Hassk. Fig. 1. 

(Greek, sphaero = globose, spherical, karpos = fruit; the fruit shape) 

Flora 25, 2 (1842) Beibl. 1, 42; Rid1ey, J. Str. Br. R. As. Soc. 82 (1920) 179, J. Ma!. Br. R. As. Soc. 

1 (1923) 58; Merrill & Perry I.e. 543; B.L. Linden I.e. 49; Whitmore i.e. 448; Anderson I.e. 324; 

Whitrnore, Tantra & Sutisna I.e. 332; Pereira I.e. 21. Type: s. colI., s.n., Herbarium Reinwardtianum 

in Aead. Lugduno-Batavo (ho1otype L). Synonyms: Dalrymplea javaniea Hassk., Pl. Jav. Rar. 

(1848) 439; Turpinia latifolia Wall. ex Ridl. I.e. (1920) 178; T. laxiflora Rid!. I.e. (1920) 179. 

Tree to 35 m tall and 90 cm in diameter; buttresses to 6 m high. Bark slightly fissured, 

flaky to smooth, greyish brown, yellowish brown to pale brown; inner bark yellowish to 

brown. Sapwood yellow to white. Stipules 4-7 x 3-5 mm, not very conspicuous, falling off 

early from leafY twigs, glabrous to minutely hairy, margin entire to sparsely hairy. Leaves 

3-5-7-9-foliolate, 9-46 cm long; petioles (2-)4-11 cm long, 1-5 mm thick, glabrous; 

leaf-rachis 2-18 cm long, glabrous; petiolules of the lateral leaflets 1-15 mm long, 

glabrous; leaflets drying greenish brown to dark purplish brown, ovate-elliptic, elliptic, 

ovate, sometimes lanceolate to oblanceo1ate, 5 19 x 22.5 11 cm, chartaceous to coriaceous, 

glabrous, not shiny to shiny above; base rounded to cuneate sometimes asymmetric, margin 

shallowly to deeply serrate, not re curved, apex shortly acuminate to cuspidate, acumen 0.4- 

1 cm long; midrib raised and distinct on lower surface; lateral veins 5-10 pairs, ascending, 

parallel, open or looping towards the margin; intercostal veins pseudoscalariform (with 

many almost straight cross-veins between the lateral veins) to reticulate with wide areoles, 

obscurely or distinctly tessellate (with no or very few straight cross-veins between the 

lateral veins). Inflorescences axillary to terminal panicles, 17-40 cm long, rachis sparsely 

to densely hairy at the distal end. Flowers with sepals ovate, thin, 1-2 x 1-2 mm; petals 

spathulate to linear obovate, l.7-3.5(-4) x 0.8-l.5 mm, thin; stamens l.5-3.5 mm long, 

filaments 1-3 mm long, anthers 0.4-0.6 mm long, apiculate or not; pistils l.5-3.5 mm 

long, styles 1 3 mm long, ovary 0.8 1.2( 1.5) mm across, (2 ) 3-celled, each cell with (4-)5- 

7(-9) ovules. Fruits 5-18 mm diameter, slightly trilobed at the apex and with three short 

apical horns, or globose and without short apical horns but sometimes with three radial 

lines at the apex, smooth to wrinkled upon drying; wall not bri~tle, 0.5-3 mm thick (rarely 

0.1-0.3 mm). Seeds pale brown io dark brown, 2-9 x 2-9 mm, 1-7 in each fruit. 

460

STAPHYLEACEAE (PERElRA) 


Leaf-rachises 10-18 cm long. Leaves typically 20-46 cm long. Leaflets (7-)10-19 x (3-)5 

-11 cm. Fruits globose, without short apical horns .............................................................. . 

var. sphaerocarpa 

Synonym: Dalrymplea javaniea Hassk., I.e. 439; Turpinia latifolia Wall. ex Ridl. I.e. (1920) 

178; Turpinia laxiflora Ridl. I.e. (1920) 179. 

Sumatra, Peninsular Malaysia, Singapore, Borneo, Java, Philippines, Lesser Sunda 

Islands & Moluccas. Common in Sabah and Sarawak and scattered in primary and 

secondary lowland to submontane forests to 1500 m, near river banks, also occurring 

in limestone forest. Found on various types of high nutrient soils. Vernacular names: 

Sarawak-maba (Than), entrang (Sg. Sinjan, Kuching), tait berak (Kelabit). 

Leaf-rachises 2-8 cm long. Leaves 9-18(-20) cm long. Leaflets 5-10 x 2.5-5 cm. Fruits 

slightly trilobed at the apex, with three short horns .............................................................. . 

var. microcerotis 1. T. Pereira 

Sandakania 5 (1994) 2l. Type: J. & M. S. Clemens 28707, British North Borneo, Ranau, 

Kinabalu, Tenompok (holotype SING; isotypes BO, K). 

Borneo (Sabah and Kalimantan). In Sabah collected from Tenompok and Mt. Kinabalu, 

Ranau. Not yet recorded for Sarawak. Lowlands, on sandy loam soils (Kalimantan) to 

montane forest to c. 1500 m (Sabah). Vernacular name: Sabah--tapong-tapong 

(Dusun). 

6. Turpinia stipulacea B.L. Linden 

(Latin, stipula = stipule; the conspicuous stipules) 

I.c. 49; Whitmore, Tantra & Sutisna l.c. 332. Type: Carr SFN 27516, British North Borneo, Mt. 

Kinabalu, near Tibabah River (holotype SING). 

Tree to 10 m tall and 10 cm diameter. Bark thin, greyish, slightly fissured; inner bark dull 

pale brownish. Sapwood pale yellow. Stipules 14 25 x 4 15 mm, conspicuous and persistent 

on apices of leafy twigs, woody, glabrous, margin entire. Leaves 7-9-11-foliolate, 20-43 

cm long; petioles 5-8 cm long, stout, glabrous; leaf-rachis 20-30 cm long; petiolules of 

lateral leaflets 3-11 cm, stout, glabrous; leaflets drying reddish brown to greenish brown 

above, ovate-elliptic, 7-14 x 2-7 cm, chartaceous to thinly coriaceous, glabrous; base 

cuneate to rounded, margin shallowly undulate to deeply serrate, apex shortly acuminate, 

acumen 0.5-0.8 cm long; lateral veins 8-11 pairs, ascending, parallel, or open towards the 

margin, distinct on lower surface; intercostal veins pseudoscalariform (with many almost 

straight cross-veins between the lateral veins). Inflorescences axillary panicles, to 35 cm 

long, rachis subglabrous. Flowers with sepals oblong to ovate, 2-3.5 x 1.5-2.5 mm, thick; 

petals linear to elliptic, 2.5-4.5 x 1-1.5 mm, thin; stamens 2-3 mm long, glabrous, 

filaments 1.8-2.8 mm, anthers 0.4-0.7 mm long, distinctly apiculate; pistils 2-3 mm long, 

styles c. 2 mm long; ovary 1-1.5 mm across, 3-celled, each cell with (2-)3-4 ovules. 

Fruits globose, (6-)10-16 mm diameter, withOut short apical horns but with three radial 

lines at the apex, wrinkled, wall not brittle, 0.5-1.5 mm thick. Seeds dark brown, 6-10 x 6 

-10 mm, 1-4 in each fruit. 

461


Distribution. Endemic to Sabah (Ranau only). 

Ecology. Occurs along streams or ridges on montane forest to 2100 m.This species 

superficially resembles Turpinia sphaerocarpa in some vegetative characters. It can be 

distinguished by the larger and more persistent stipules and smaller number of ovules. 

462


STYRACACEAE 

R. Kiew 

Universiti Pertanian Malaysia, 

Serdang, Malaysia 

Merrill, EB (1921) 485; Backer & BakhuizenJ, FJ 2 (1965) 203; van Steenis, FM 1,4 (1949) 49, 

FM 1, 6 (1972) 976; Cockburn, TS 2 (1980) 103; Anderslln, CLTS (1980) 327; Whitmore, Tantra & 

Sutisna, CLK 2, 1 (1990) 340. 

Trees or shrubs. Leaves simple, spirally arranged or pseudo-alternate, margin entire or 

toothed, mostly with stellate or lepidote indumentum, without stipules. Inflorescences 

racemes or panicles, axillary or terminal. Flowers regular; calyx tubular more or less 

adnate to the ovary, lobes if present valvate; corolla mostly joined into a basal tube (rarely 

of free petals), lobes 4-7, valvate or imbricate; stamens equal and alternate or double the 

number of petals, mostly adnate to the tube; ovary superior (rarely semi-inferior), 3-5- 

celled, style 1, stigma punctiform to 3-5-lobed. Fruits capsules (rarely berries), dehiscent 

or not, pericarp often thick and woody or corky, calyx persistent. Seeds 1 to many per 

locule, with endosperm. 

Distribution. 12 genera and c. 190 species, mostly in the northern hemisphere, especially 

E Asia (absent from Australia and the Central Pacific). In Malesia, represented by 2 

genera, Bruinsmia and Styrax, with the greatest diversity in Sumatra and no species as yet 

being found in the Philippines, C and E Java and the Lesser Sunda Islands. Two species are 

known from Borneo, one, Styrax agreste, occupies the southern half of the island and has 

not yet been collected from Sabah and Sarawak, although it may be expected; the other, 

Bruinsmia styracoides, is less collected from Sarawak than from Sabah. 

Ecology. Lowland and submontane forests to 1600 m. 

Uses. Styrax benzoin is the source of benzoin resin (kemenyan), which is used for 

medicinal and a variety of other purposes. 

Taxonomy. Symplocos is treated as a separate family as it does not have the stellate hairs, 

nor fasciculate stamens, linear anthers and the half or wholly superior ovary characteristic 

of the Styracaceae. 

BRUINSMIA Boerl. & Koord. 

(A.E.J. Bruinsma, 1852-1943, Dutch East India Forest Service) 

Nat. Tijd. Ned. Ind. 53(1893) 68; Perkins, Pfl. R. 30 (1907) 14, 88; van Steenis, Bull. Jard. Bot. 

Btzg. 3, 12 (1932) 215, I.e. (1949) 49; van Steenis & BakhuizenJ, Bot. Jahrb. 86 (1967) 393; 

Cockburn, TS 2 (1980) 103. 

463


Fig. 1. Bruinsmia styracoides. A, flowering leafy twig; B, flower; C, fruit. (All from van Balgooy 

5182.) 

464

STYRACACEAE (KIEW) 

Glabrate trees with flattened-angular twigs. Leaf margin entire to coarsely crenate. 

Inflorescences compound, dichotomous, terminal or axillary panicles; pedicels articulated 

at apex. Flowers dull green or white; corolla-lobes free without a staminal tube; ovary 

glabrous, 3-5(-6)-loculed. Fruits fleshy, indehiscent, style base persistent. Seeds minute, 

i-many per locule. 

Distribution. 2 species, one in Borneo; the other, B. polysperma (Clarke) Steenis, is known 

from India (Assam), Burma and Thailand (Chiangmai). 

Taxonomy. Bruinsmia is distinguished from Styrax by the leaves, which are glabrescent or 

sparsely stellate hairy, by the compound dichotomous inflorescences (as opposed to 

racemose-paniculate), the articulate flower-stalks, the 3- to 5-celled ovary and fruit, and by 

the many seeds in the fruit. 

Bruinsmia styracoides Boerl. & Koord. Fig. 1. 

(resembling the classic Greek tree, Styrax ofJicinalis, whiCh yields the resin storax) 

I.e. 68; Perkins I.e. 14,88; van Steenis I.e. (1932) 215, I.e. (1949) 49; Cockbum I.e. 103; Anderson 

I.e. 327; Whitmore, Tantra & K. Sutisna I.e. 340. Type: Koorders 8529fJ and 8530fJ (1893), Java 

(syntypes BO, K, L). 

Tree to 45 m tall and 30-200 cm diameter; clear bole 23 m; crown large and spreading, 

flowering at 5 m; trees over 25 m with buttresses c. 65-100 cm high, 5-7 cm thick and 65- 

80 cm out. Bark smooth becoming slightly fissured with age, grey to grey-brown; inner 

bark soft and stringy, pale brown becoming reddish brown on exposure with a jelly-like 

exudate that reddens on exposure. Sapwood white. Twigs flattened-angular at nodes. 

Indumentum of young twigs, lower leaf surface and panicle dense, comprising stellate 

hairs, persisting only on inflorescences. Leaves ovate to oblong, 7.5-19 x 3-10 cm, 

chartaceous; base usually rounded, margin serrate, apex acute to acuminate; lateral veins 

6-10 pairs; stalk grooved, 0.5-1.5 cm long. Panicles terminal and 4.5-25 cm long or 

axillary and 2.5-7 cm long, with dense yellow-brown pubescence; pedicels c. 1 mm, 

lengthening to 2-3 mm in fruit, articulated; bracteoles narrow, 1-2.5 mm long. Flowers: 

calyx broadly cup-shaped, entire or shortly 5-toothed, 1.2-3 mm long and 5 mm wide; 

corolla lobes 5(-6), free, ovate-oblong, 9-10 x 4-4.5 mm, imbricate, pale yellow, silvery 

hairy on both sides; stamens yellow-orange, 10(-12), 5 longer (6-6.5 mm) alternate, 5 

shorter (5-5.5 mm) epipetalous, or 10 subequal (3.5-4 mm), joined to the base of corolla; 

ovary 2-3 x 2.5-3.5 mm, imperfectly 5(-6)-locular; style 3-7 mm long, stigma capitate; 

ovules many. Fruits dark green, on dish-like calyx 7 mm wide, globular to pear-shaped, 

6-iO x 6-9 mm, attenuated to persistent style base 0.7-6 mm long; fruit-wall indehiscent, 

thick and pulpy. Seeds many per locule, c. 1.5 mm long, black, 4-angular and very finely 

corrugated. 

Vernacular names. Sabah-tingo-tingo (Dusun Kundasan).· Sarawak-empudu (? Than). 

Distribution. Sumatra, W Java, Borneo, Philippines (Mindanao), Celebes, Moluccas 

(Bacan Is.) and New Guinea. In Sabah and Sarawak, widely distributed but more common 

in Sabah. 

465


Ecology. On steep hill sides or ridges in primary, occasionally in disturbed forest, from the 

lowland at 450 m to montane forest at 2000 m. Most flowering and fruiting occurs between 

May and November, buds and mature fruits develop simultaneously. 

466


TRIGONIACEAE 

K.M. Wong 




Merrill, PEB (1929) 132; Masamune; EPB (1942) 378; van Steenis, FM 1, 4 (1949) 59; 

Kochummen, TFM 1 (1972) 449; Anderson, CLTS (1980) 340; Ashton, MNDTS (1988) 469; 

Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 361. 

Trees or shrubs. Leaves simple, entire; stipules small, caducous. Inflorescences racemose 

or paniculate, axillary and terminal. Flowers bisexual, bilaterally symmetrical; sepals 5, 

free or joined at the base, imbricate, unequal; petals 3-5, free, imbricate, unequal; stamens 

3-12, grouped to one side opposite the anterior petals, filaments fused at the base, anthers 

2-celled and dehiscing lengthwise; ovary superior, 3-locular, woolly; style simple; ovules 

I-several per locule, placentation axile. Fruit a 3-lobed structure splitting into 3 flat, 

winged fruit portions (samaras). Seeds compressed, hairy, endosperm none. 

Distribution. 3 genera and 44 species, in tropical South America and Malesia. In Malesia, 

only the monotypic genus Trigoniastrum is found. 

Taxonomy. Trigoniaceae is generally accepted as most closely related to the Polygalaceae. 

TRIGONIASTRUM Miq. 

(Latin, trigonus = 3-angled, aster = star; the fruit form) 

Fl. Ind. Bat. Suppl. (1860) 394; Merrill I.e. 132; Masamune I.e. 378; van Steenis I.e. 59; 

Kochummen I.e. 449; Anderson I.e. 340; Ashton I.e. 469; Whitmore, Tantra & Sutisna I.e. 361. 

Trees. Leaves alternate and distichous on branches. Flowers with 5 petals, the posterior 

one largest and saccate and with a concave gland at the mouth of the sac, the lateral ones 

spathulate and spreading, the anterior two closer together and obliquely held; stamens 6; 

ovules solitary in each locule. 

Distribution. Monotypic, confined to Sumatra, Peninsular Malaysia and Borneo. 

Trigoniastrum hypoleucum Miq. Fig. 1. 

(Greek, hypo = lower side, leuco = white or glaucous; the leaf surface) 

I.e. 394; Merrilll.e. 132; Masamune I.e. 378; van Steenis I.e. 59; Kochummen I.e. 449; Anderson I.e. 

340; Ashton I.e. 469; Whitmore, Tantra & Sutisna I.e. 361. Type: Teijsmann, s.n., Sumatra, 

467


3mm 

D-~~~,>,." 

2"~~ 

tTI,¥c 

~~~,J 

2cm 

3mm 

l0~ \CL-- 

5mm 

F 

Fig. 1. Trigoniastrum hypoleucum. A, flowering twig with larger leaf form; B, twig with smaller leaf 

form; C, detail of lower leaf surface; D, gland on leaf margin; E, fruiting twig; F, detail of 

inflorescence bract; G, flower with petals removed; H, posterior petal with pouch-like base. CA, C, D, 

F, G, H from SAN 89752; B from SAN 84757; E from SAN 83338.) 

468

TRIGONIACEAE (WONG) 

Lampong, Mengala (holotype L; isotype K). Synonym: Isopterys penangiana Wall. ex A.W. Benn. in 

HookerJ, Fl. Br. Ind. 1 (1872) 208. 

Small to medium-sized tree, to 30 m tall and 50 cm diameter, sometimes with buttresses up 

to 1 m high. Bark pale greenish grey to brown, smooth to lenticellate; inner bark pale 

greenish yellow to yellowish brown, exuding a yellowish sticky sap which turns reddish on 

exposure. Sapwood yellowish ochre, hard. Leaves elliptic, 5-20 x l.5-8 cm, chartaceous to 

thickly coriaceous, glaucous beneath, glabrous or yellow-hairy on the veins below and 

midrib on upper surface; base cuneate to rounded, apex acute to cuspidate; midrib sunken 

to flat on upper side, prominent on lower surface, drying pale brown to dark purplish; 

lateral veins 5-8 pairs, distinct, looping coarsely towards the margin; intercostal veins 

reticulate, prominent on the lower leaf-surface and sometimes the upper surface; stalks 5-9 

mm long. Inflorescences 7-15 cm long 1 

paniculate, the axes scantily to densely yellow 

short-hairy all over. Flowers fragrant; sepals ovate-acute, c. 2.5 x l.5 mm, pale green, 

yellow short-hairy; posterior petal white with a yellOWish basal sac-like part c. 2 mm 

across and a reflexed distal part c. 2.5 mm long and 2 mm wide, the concave gland at the 

mouth of the sac to 0.8 mm across; lateral petals spathulate, 3.5-4 x l.5 mm, creamy 

yellow; anterior petals oblong, oblique, 4-5 mm long, white; staminal tube c. 1 mm long, 

anthers c. 0.5 mm long; ovary pale yellow woolly, to 0.5 mm across, style c. l.5 mm long 

and subglabrous. Fruit a 3-lobed structure; each lobe a samara, ovate, flattened and winglike, 

the inner basal portion straight and fused with the other 2 lobes, the whole lobe to 6 

cm long and 2.5 cm wide, yellowish green ripening pale brown. 

Vernacular names. Sabah-none generally, but once (SAN 32418) documented as miapa 

(Dusun Bundu Tuhan). Sarawak-atap (Kenyah), lia (!ban), ngilis (!ban), nyalin (!ban: in 

common with Xanthophyllum which it resembles), nyalin bintek (!ban), tulang kwe 

(Kenyah). Brunei-mengilas babi (!ban). 

Distribution. As for genus. In Sabah and Sarawak in all districts. Also in Brunei and 

Kalimantan. 

Ecology. Although widespread in Borneo, the species is nowhere common. It has a wide 

ecological amplitUde and has been collected from peat swamp, freshwater swamp, heath 

(kerangas), alluvial and mixed dipterocarp forests in the lowlands and hills, to submontane 

forest at about 1200 m. Ashton (I.c.) points out there may be two ecotypes, one a small tree 

with thick leaves found in peat swamp, heath and mixed dipterocarp forests on poor sandy 

soils, and the other a larger tree with thinner, narrower leaves in mixed dipterocarp forest 

on richer sites. Van Steenis (I.c.) records flowering and fruiting specimens collected in 

every month of the year; it is possible, though, that there are local variations in the 

phenology of the species. The winged fruits are wind-dispersed. 

Uses. None recorded, but the wood has been tested and is listed as being hard or very hard 

and heavy or very heavy, with a moderately fine texture (Wong (1982) DMT). 

The leafy twigs alone are easily confused with those of Xanthophyllum (Polygalaceae), 

although the flowers and fruits readily distinguish it. In many Xanthophyllum species, the 

leaf blades have distinct glands and the ultimate leafy twigs are green, and so are different. 

469

Anderson, J.A.R 1980. A Checklist of the Trees of Sarawak. Forestry Dept., Sarawak. 

CLTS 

Ashton, P.S. 1988. Manual of the Non-Dipterocarp Trees of Sarawak. Forestry Dept., 


MNDTS 

Backer, C.A & RC. Bakhuizen v.d. Brink, Jr. 1965. Flora of Java. Noordhoff, Groningen, 

the Netherlands. 

FJ 

471 

Masamune, G. 1942. Enumeratio Phanerogamarum Bornearum. Taihoku. 

EPB 

Mabberley, D.J. 1987. The Plant Book. Cambridge University Press, Cambridge. 

PB 

Keng, H. 1969. Orders and Families of Malayan Seed Plants. University of Malaya Press, 

Kuala Lumpur. 

OFMSP 

Keith, H.G. 1937. The Timbers of North Borneo. Govt. Col. N. Borneo, Sandakan. 

TNB 

Flora Malesiana 

FM 

Corner, E.J.H. 1988. Wayside Trees of Malaya. 2 Vols. Malayan Nature Society, Kuala 

Lumpur. 

WSTM 

Cockburn, P.F. 1976 & 1980. Trees of Sabah, Vols. 1 & 2. Forestry Dept., Sabah. 

TS 

Burkill, I.H. 1966. A Dictionary of the Economic Products of the Malay Peninsula. Kuala 

Lumpur. 

EPMP 

Burgess, P.F. 1966. Timbers of Sabah. Forestry Dept., Sabah. 

TBS 

Browne, F.G. 1955. Forest Trees of Sarawak and Brunei. Govt. Press, Sarawak. 

FTSB 

ABBREVIA TIONS OF 

FREQUENTL Y CITED REFERENCES

FMK 

Whitmore, T.C. (ed.). 1972 & 1973. Tree Flora of Malaya, Vols. 1 & 2. Longman, 

Malaysia. 

TFM 

Whitmore, T.e. & I.G.M. Tantra. 1986. Tree Flora of Indonesia. Check List for 

Sumatra. Forest Research & Development Centre, Bogor. 

CLS 

Whitmore, T.C., I.G.M.Tantra & U. Sutisna. 1990. Tree Flora of Indonesia. Check List 

for Kalimantan, Part 1 & Parts 2.1-2.2. Forest Research & Development Centre, Bogor. 

CLK 

Willis, J.e. 1973. A Dictionary of the Flowering Plants & Ferns, revised by H.K. Airy 

Shaw. Cambridge University Press, Cambridge. 

DFPF 

472 

Stapf, O. 1894. On the Flora of Mount Kinabalu, in North Borneo. Transactions of the 

Linnean Society, London, Ser. 2,4, pp. 69-263. 

(PRO SEA, to be listed without editor or author names, but to include volume and page 

numbers). 

Ng, F.S.P. 1991 & 1992. Manual of Forest Fruits, Seeds and Seedlings. Malaysian Forest 

Records, No. 34. Vols. 1 & 2. FRlM, Malaysia. 

MFFSS 

Merrill, E.D. 1929. Plantae Elmerianae Borneenses. University of California Publications 

Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. J. Str. Br. Roy. As. 

Ridley, H.N. 1923-1925. The Flora of the Malay Peninsula, Vols. 1-5. Reeves, London. 

FMP 

Ng, F.S.P. (ed.). 1978 & 1989. Tree Flora of Malaya, Vols. 3 & 4. Longman, Malaysia. 

TFM 

Smythies, B.E. 1965. Common Sarawak Trees. Borneo Literature Bureau, Sarawak. 

CST 

Plant Resources of South East Asia. Wageningen.(Various authors and volumes). 

Perry, L.M. 1980. Medicinal Plants of East and Southeast Asia. Cambridge. 

in Botany, Vol. 15. University of California Press, Berkeley. 

PEB 

Soc., Sp. No. 

EB 

MPESA

Wong, T.M. 1982. A Dictionary of Malaysian Timbers. Malayan Forest Records No. 30. 

Forest Research Institute, Kepong, Kuala Lumpur. 

DMT 

COMMONLY USED 

ABBREVIATIONS FOR LOCALITIES 

English 

Malay 

Word 

Central 

Division 

East 

Ab b reviation Word Abbreviation 

C Bukit Bt. 

Div. Gunung G. 

E Kampung Kg. 

I 

Forest Reserve FR Sungai Sg. 

Island Is. Tanjung Tg. 

Mount 

National Park 

North-East 

North-West 

Mt. 

NP 

NE 

NW 

River R. 

South 

South-East 

South-West 

West 

S 

SE 

SW 

W 

473

GLOSSARY 

abaxial 

accrescent 

achene 

acrid 

acrodromous 

acroscopic 

actinomorphic 

acumen 

acuminate 

acute 

adaxial 

adherent 

adnate 

alate 

anastomosing 

anatropous 

androdioecious 

androecium 

androgynophore 

anisophylly 

annular 

annulus 

anterior 

anther 

anthesis 

antidysenteric 

antiemetic 

anti helminthic 

apetalous 

apiculate 

arachnoid 

arboreous 

arcuate 

areolation 

areole 

aril 

the side of an organ facing away from the distal portion of the 

axis which bears it 

increasing in size with age, e.g., the calyx of some plants after 

flowering 

a small, hard, dry, non-splitting fruit 

sharp, irritating to the taste 

with veins converging and uniting at the apex of the leaf 

towards the apex or distal part of an organ; e.g., flowers in an 

inflorescence 

(flowers) radially symmetrical 

a tapering point or tip 

drawn out into a long point; having a gradually tapering point; 

pointed; e.g., leaf apex 

ending in a sharp tapering point; e.g., leaf apex 

the side of an organ facing the distal portion of the axis which 

bears it 

in contact but not fused together; e.g., floral parts 

attached to some other organ 

winged; as a stem or petiole 

joining of veins to form a network 

(ovules) with hilum and micropyle close together and chalaza 

at the other end 

plants with male or hermaphrodite flowers 

male reproductive organs of a plant 

common structure or stalk that supports androecium and 

gynoecium 

the occurrence of leaves with distinct form and size 

ring-like 

ring 

frontal 

the part of a stamen which contains pollen 

(flowers) that period between the opening of the bud and the 

withering of the stigma and/or stamen 

against dysentery 

preventing vomiting 

working against worms 

without petals 

with an abrupt small tip, as a leaf 

like a cobweb 

with the habit of a tree; tree-like 

curved or shaped like a bow; arc-shaped 

of fine veins anastomosing and enclosing tiny spaces 

a tiny space marked on a surface; e.g., on leaves 

a fleshy expansion of the funicle, arising from the placenta, 

and enveloping a seed 

474

arillate 

arillode 

arilloid 

articulated 

attenuate 

auricle 

auriculate 

axile 

axillar/axillary 

ballistic 

basal 

basifixed 

berry 

bifid 

bifoliolate 

bilocular 

biseriate 

bisexual 

bistipulate 

bitegmic 

bract 

bracteate 

bracteolate 

bractcole 

bud 

bullate 

caducous 

calcareous 

calyptra 

calyx 

campanulate 

camptotropous 

canaliculate 

capitate 

capsular 

capsule 

carinate 

carminative 

carpel 

carpellate 

having / bearing an aril 

a false aril 

resembling an aril 

jointed 

gradually narrowing to a tip or base; e.g., leaf-blade 

ear-like lobe or appendage 

with auricles; e.g., leaf'with expanded bases surrounding stcm 

(ovule placentation) with ovules attached to central axis within 

the ovary 

(inflorescence; bud) borne in the axil, i.e. the junction bctween 

leaf-stalk and stem 

(seed dispersal) referring to the fruit which discharges its 

seeds elastically 

at or near the base of an organ 

(stamens) having filament attached to anther base 

non-spliting pulpy fruit with many seeds 

forked; divided nearly to middle line 

with two leaflets 

having two cavities or chambers; eg., anther or ovary 

in two series 

having both male and female reproductive organs; hermaphrodite 

with two stipules 

(ovules) with two layers of integuments 

a modified leaf in whose axil a flower or branch axis arises 

having bracts 

with bracteoles 

a secondary bract as one on a pedicel of a flower, usually 

smaller than bract 

a rudimentary state of a stem or branch; an unexpanded flower 

blistered like a savoy-cabbage leaf 

fall off very early; deciduous; e.g., stipules 

growing on soil derived from decomposition of calcareous 

rocks; limy 

a cap-shaped structure 

the outer whorl of floral envelope; collective name for sepals 

bell-shaped 

(ovule) curved or bent like a horse-shoe 

longitudinally channelled or grooved 

enlarged or swollen at tip; arranged into a mass at apex, as 

some inflorescences 

possessing fruit of capsule-type 

a dry, dehiscent fruit 

having a ridge or a keel 

relieving flatulence 

a simple pistil formed by a single leaf-like, ovule-bearing 

structure; a basic ovule-bearing chamber of the ovary 

with carpels 

475

catkin 

concolorous 

concrescent 

cone 

conical! coniform 

connate 

connective 

connivent 

contorted 

convex 

convolute 

cordate 

coriaceous 

corolla 

corymb 

corymbiform/corymbose 

crassinucellate 

crenate 

firm and tough but elastic like cartilage 

a wart or protuberance near the hilum of a seed 

the early leaf-forms of a plant or shoot, as cotyledons, budscales, 

rhiz~me scales, etc. 

a deciduous spike, consisting of unisexual apetalous flowers 

with a slender tail-like appendage 

having flowers on the main stem 

that part of an ovule or a seed where the nucellus joins the 

integuments 

papery 

fringed with hairs 

ciliate but hairs minute 

dehising in transverse circular line, the top separating like the 

lid of a pill box; e.g., in fruits 

the incorporation of one part with another, as the petals to 

form a tubular corolla 

a severe spasmodic abdominal pain 

positioned side by side 

mucilaginous hairs on the buds of many flowering plants 

which secrete gum 

a persistent central axis around which the carpels of some 

fruits are arranged 

having a form of a column; pillar-like 

the face by which one carpel joins another as in the 

Umbelliferae (cabbage family) 

having an outline or surface curved like the interior of a circle 

or sphere 

uniform in tint 

growing together 

a dry multiple fruit- or seed-bearing structure; e.g., in pine tree 

cone-shaped 

united to similar structures 

the portion of a stamen distinct from the filament which 

connects the two halves of an anther 

coming in contact; converging 

twisted 

having a more or less rounded surface 

rolled around 

heart-shaped 

leathery in texture 

the inner series of floral envelope, composed of petals; 

collective name for petals 

a flat-topped or convex and open flower-cluster of the 

indeterminate or centripetal order 

arranged in corymbs 

(ovule) having a thick nucellus 

toothed with the teeth rounded at apex 

476 

chartaceous 

ciliate 

ciliolate 

circumscissile 

commissure 

cartilaginous 

caulitlorous 

columnar 

columella 

colic 

collateral 

colleter 

cataphyll 

coherent 

concave 

caruncle 

caudate 

chalaza

crenulate 

crescent 

cristate 

crustaceous 

crypts 

cryptocotylar 

cucullate 

cuneate 

cupular 

cupuliform 

cuspidate 

cyme 

cymose 

cymules 

deciduous 

decurrent 

decussate 

dehisce 

dehiscent 

deltate/deltoid 

dentate 

denticulate 

dichotomous 

didynamous 

dimorphic 

dioecious 

discoid 

dispersal 

distichous 

distylous 

diuretic 

divaricate 

divergent 

domatia 

dorsifixed 

drupaceous 

drupe 

crenate with small teeth 

a structure having a shape like the moon as seen in the first or 

the last quarter of the month 

crested; having a tesseJled margin 

brittle in texture 

stomatal pits; also applied to sunken glands, receptacles for 

secretions in a plant with dotted leaves 

(seed) with hidden cotyledons during germination 

hooded or hood-shaped 

wedge-shaped 

furnished with, or subtended by a cup-like structure 

cup-shaped 

tipped with a sharp, rigid point 

a flower-cluster of determinate or centrifugal type, often 

convex or flat -topped 

bearing cymes; cyme-like 

a small cyme or portion of one 

falling off, as petals fall after flowering or leaves in autumn or 

dry season 

running down, as when leaves are prolonged beyond their 

insertion, and thus run down the stem 

in pairs that alternate at right angle, thus in four rows 

gape or burst open spontaneously when ripe, as seed capsules 

or anthers 

opening by definite pores or slits at maturity to release the 

content; as in fruits or anthers 

shaped like a Greek letter Delta; equilaterally triangular 

toothed 

minutely toothed 

two-forked, the branches equal or nearly so 

(flower) having four stamens arranged in pairs, two long and 

two short 

in two forms 

(plant) having unisexual flowers, with the staminate and 

pistillate flowers borne on separate individuals 

resembling a disk 

the various ways by which seeds or fruits are scattered away 

from the mother-trees 

disposed in two vertical ranks, as in leaves 

having two styles 

promoting the secretion of urine 

extremely divergent 

separated or turning in different directions 

dome-like, usually hairy projections sheltering parasites 

(stamen) having filament attached to the back of an anther 

resembling a drupe 

one-seeded, indehiscent fruit with the pericarp fleshy or 

leathery and the seed-coat hard and stony 

477

druplet 

dyspeptic 

eglandular 

ejaculatory apparatus 

ellipsoid 

elliptic 

emarginate 

emetic 

endocarp 

endocarpid 

endosperm 

entire 

eophyll 

ephemeral 

epicarp 

epigeal 

epipetalous 

epiphyllous 

epiphytic 

episepalous 

epitropous 

estipulate 

exarillate 

excentric 

excrescence 

exfoliate 

exocarp 

exserted 

exstipulate 

extra 

extrastaminal 

extrorse 

exudate 

falcate 

fascicle 

fasciculate 

febrifuge 

ferruginous 

filament 

filamentous 

filiform 

a diminutive drupe 

stomach discomfort; e.g., indigestion 

without glands 

a structure that forcibly throws out endogenously formed 

content; e.g., in fruit 

a tridimensional structure elliptic in outline; e.g., seeds 

shaped like an ellipse, oblong with regularly rounded ends; 

e.g., leaf-blade 

having a notch cut out, usually at the extremity 

that causes vomiting 

the inner layer of a pericarp 

indehiscent kernel or pyrene embedded in certain fruits 

albumen deposited within the embryo-sac of a seed 

having an even margin, without lobes or teeth; e.g., leaf-blade 

early form of leaf 

lasting for a day or less 

the external layer of a pericarp 

(seed germination) with the cotyledons lifted above the ground 

surface by an elongating axis 

(stamens) borne upon the petals 

borne upon the leaves 

relating to epiphytes 

borne upon the sepals 

denotes an anatropous ovule with its raphe averse when 

ascending, adverse when suspended 

without stipules 

without aril 

out of the centre; one-sided 

growing in an unnatural way, as a wart or other outgrowth 

to come away in scales or flakes 

the outer layer of a pericarp 

projecting beyond the surrounding organ, as stamens beyond 

the tube of the corolla 

without stipules 

outside or beyond 

arising or situated outside the stamens 

directed outwards, as the dehiscence of an anther 

discharge from incision or pore; e.g., gum, latex, moisture, 

resin, etc. 

sickle-shaped 

a close bundle or cluster of flowers, leaves, etc. 

connected or drawn into a fascicle 

a medicine that reduces fever 

rust-coloured 

the stalk of an anther 

having a form of a filament 

thread -shaped 

478

fimbriate 

fissured 

flavedo 

foliaceous 

follicle 

follicular 

fulvous 

funicle/funiculus 

furfuraceous 

fusiform 

gametophyte 

germicidal 

germicide 

gibbous 

glabrate/ glabrous 

glabrescent 

glandulous 

glaucous 

globose/globular 

granular 

gynobasic 

gynoecium 

gynophore 

herbaceous 

hermaphrodite 

hesperidium 

heterostylous 

hilum 

hirsute 

hirsutulous 

homostylous 

hyaline 

hypanthium 

hypocotyl 

hypogeaIlhypogeous 

hypogynous 

with the margin bordered by long slender processes 

deeply grooved or furrowed 

yellowness 

having the texture or sp.ape of a leaf 

a fruit of one carpel, opening by a ventral suture to which the 

seeds are attached 

shaped like a follicle 

dull yellow; tawny 

the cord or thread which connects the ovule or seed to the 

placenta 

scurfy; having soft scales 

spindle-shaped 

the generation of a plant that bears the sexual organs, 

producing gametes, in turn giving rise to the sporophyte 

ability to destroy germs 

a substance that destroys germs 

enlarged, humped or swollen on one side 

smooth; without pubescence 

becoming glabrous, or slightly so 

possessing glands 

bluish green 

rounded or spherical in shape 

like grains or granules 

denotes a style that adheres by its base to a prolongation 

upwards of the torus between carpels 

an organ in which female cells are formed 

a stalk supporting the ovary/pistil/carpel 

having the texture, colour and properties of a herb 

(flower) with both stamens and pistils; having the 

characteristics of both sexes 

many-celled, few-seeded indehiscent fruit, having the epicarp 

and mesocarp joined together, and the endocarp projecting 

into the interior as membranous partitions which divide the 

pulp into chambers; such as the orange 

(flowers) with different types (length) of styles 

the scar left on a seed where formerly attached to the funicle 

or placenta 

coarse-hairy 

minutely hirsute 

with uniform styles 

colourless or translucent 

an enlargement or elongation of the torus or floral axis below 

the calyx 

the portion of the axis below cotyledons in an embryo 

(seed germination) with cotyledons not lifted above the ground 

surface by an elongating axis 

free from but inserted below the gynoecium or pistil; e.g., 

petals and sepals 

479

idioblast 

imbricate 

imparipinnate 

incipient bract 

indehiscent 

indumentum 

induplicate 

indusium 

inferior 

inflexed 

inflorescence 

infructescence 

insecticidal 

intercostal . 

interpetiolar 

intervenium 

intra 

intramarginal 

intrapetiolar 

intrastaminal 

introrse 

invaginating 

juvenile 

lacerate 

laciniate 

lamellate 

lamina 

laminate 

lanate 

lanceolate 

laticiferous 

latrorse 

laxative 

lenticel 

lenticellate 

lenticular 

lepidote 

ligulate 

ligule 

a special cell in a tissue that markedly differs from the rest in 

form, size, or contents 

overlapping 

pinnate with an odd terminal leaflet 

bract in an initial stage of development 

not opening by valves or along regular lines 

hairy covering 

with the margins bent inwards, and the external face of these 

edges, applied to each other, without twisting 

an epidermal outgrowth covering and protecting another 

structure; a ring of collecting hairs below the stigma 

(ovary) arising or growing at a level below the insertion of 

other floral parts 

turned abruptly or bent inwards 

the deposition of the flowers on the floral axis; flower cluster 

the inflorescence in a fruiting stage; collective fruits 

ability to destroy insects 

between veins of a leaf 

between the petioles 

a portion of parenchyma between the veins of a leaf 

within 

placed within the margin near the edge 

within the petiole or between it and the stem 

within the stamens, as the disc of Anacardiaceae (the mango 

family) 

turned inwards, towards the axis 

involuting or drawn into a sheath 

young stage of growth 

torn or irregularly cleft 

incised or slashed into narrow lobes 

made up of thin plates 

the blade of a leaf 

consisting of plates or layers 

clothed with woolly and intergrown hairs 

lance-shaped; narrow and tapering to each end 

latex -bearing 

turned or directed towards the side of an organ; e.g., anthers of 

a flower 

tending to stimulate or facilitate evacuation of the bowel 

lenticular corky spots on young bark, corresponding to 

epidermal stomata 

having lenticels 

like a doubly convex lens 

beset with small scurfy scales 

furnished with a ligule 

a strap-shaped body/structure, such as the limb of the ray 

florets in Compositae (sun-flower family) 

480

Iysigenous 

megasporophyII 

membranous 

merism 

mesocarp 

micropyle 

obtuse 

ochraceous 

olivaceous 

orbicular 

orthotropic 

orthotropous 

oblong 

obovate 

obpyriform 

obturator 

obcordate 

oblanceolate 

monotypic 

mucilaginous 

mucro 

mucronate 

mucronulate 

multiflorous 

nectariferous 

nectarivorous 

nocturnal 

nucellus 

monoembryonic 

monogeneric 

monopodial 

microsporangium 

microsporophyII 

monoecious 

linear 

lobulate 

locular/loculate 

locule/ loculus 

loculicidal 

481 

of cells 

seed- or ovule-bearing leaf-like structure 

thin and semi-transparent 

repetition of parts to form a symmetry or pattern 

the middle layer of a pericarp 

aperture at the apex of an ovule through which pollen-tube 

enters the embryo-sac 

a microspore- or pollen-bearing structure or organ 

a leaf-like structure bearing microsporangium 

(plant) having stamens and pistils in separate flowers, but 

borne on the same plant 

with only one embryo 

a family having a single genus 

(tree) with the branches or appendages arising from a simple 

axis 

a genus with a single species 

slimy, composed of mucilage 

a sharp point abruptly terminating an organ 

abruptly terminated by a short and straight point 

tipped by a diminutive mucro 

many-flowered 

nectar -bearing 

nectar-feeding 

(flowers) night-blooming 

the body of the ovule containing the embryo-sac; the kernel of 

an ovule 

inversely heart -shaped 

inversely lanceolate; tapering towards the base more than 

towards the apex 

much longer than broad, with nearly parallel sides 

inversely egg-shaped, with narrow end attached to stalk 

inversely pear-shaped 

a wart-like protuberance of the placenta covering the 

micropyle 

with blunt end 

yellow with a tinge of red 

olive-coloured 

of a flat body with a circular outline 

assuming a vertical position 

(ovule) having a straight axis, the chalaza being at the 

insertion and the micropyle at the opposite end 

narrow, several times longer than wide 

having small lobes 

divided into locules or cavities 

the cell or cavity of an,ovary or an anther 

dehiscence of a fruit down the center of a compartment or a 

locule 

a state when a cavity is formed by a degeneration or dissolving

ovary 

ovate 

ovoid 

ovule 

palmate 

palmatifid 

p alynological 

panicle 

paniculate 

papilla 

papillate/papillose 

paraphyll 

parenchyma 

parenchymatous 

parietal 

paripinnate 

patelliform 

pedicel 

peduncle 

pedunculate 

pellucid 

peltate 

pendulous 

percurrent 

perennial 

perianth 

pericarp 

perigynous 

petal 

petiole 

petiolule 

phanerocotylar 

pilose 

pinnate 

pistil 

pistillode 

that part of the pistil which contains the ovules 

egg-shaped (two dimensional), the broad end attached to the 

stalk 

somewhat egg-shaped (three dimensional) 

the female gamete- or egg-cell-bearing tissue of seed-bearing 

plants; the organ which after fertilization develops into a seed 

(leaf-blade) divided into lobes which arise from a common 

centre 

(leaf-blade) cut in a palmate fashion nearly to the petiole 

pertaining to pollen and spores 

a loose flower-cluster, as a branched raceme or corymb 

having flowers clustered in a panicle 

soft superficial glands or protuberances 

having papillae 

leaf-like expansion produced near the leaves 

soft, succulent tissue in plant composed of more or less 

isodiametric, thin-walled cells 

consisting of parenchyma 

(ovules) borne on or belonging to the inner wall of an ovary 

pinnate, with an equal number of leaflets, that is without a 

terminal one 

shaped like a small flat dish 

the ultimate stalk of a single flower in an inflorescence 

the stalk of a solitary flower or an inflorescence 

borne upon a peduncle 

wholly or partially transparent 

shaped like a small shield; fastened to stalk at a point within 

margin 

hanging or pendent 

extending throughout the entire length 

persisting throughout the year or for a number of years 

the floral envelopes of which calyx and corolla cannot be 

distinguished 

fruit-wall 

(flowers) having sepals, petals and stamens adnate with the 

lower part of the pistil 

one of the leafy expansions in the floral whorl constituting the 

corolla 

leaf-stalk 

stalk of a leaflet in a compound leaf 

(germination) with exposed cotyledons 

having soft distinct hairs 

compound leaf with the leaflets arranged on each side of the 

midrib or rachis 

the female organ of a flower, consisting of ovary, style and 

stigma 

rudi.mentary pistil 

482

placenta 

placentation 

plagiotropic 

pleurisy 

plicate 

plumose 

pneumatophore 

polyadelphous 

polyembryonic 

polygamodioecious 

polygamous 

posterior 

prominulous 

protandrous 

proximal 

pseudopuberulous 

pubescence 

pubescent 

pulp-vesicle 

pulpy 

pulvinate 

pulvinus 

punctate 

punctiform 

punctulate 

purgative 

pustular 

pustule 

pyrene 

pyriform 

raceme 

racemose 

rachis 

radicle 

ramiflorous 

raphe 

receptacle 

reniform 

the organ which bears ovules in an ovary 

the manner by which ovules are attached to the ovary 

having oblique or horizontal direction of growth 

inflammation of the pl~ura 

folded into plaits, usually lengthwise 

feathery; feather-like 

an aerating root; e.g., in Rhizophoraceae 

having stamens grouped into several bundles 

having more than one embryo in a seed 

(plant) having bisexual flowers and unisexual flowers on 

different individuals of the same species 

(plant) having bisexual flowers and unisexual flowers on the 

same, or on different individuals of the same species 

on the side next or close to the axis 

slightly raised, standing out a little from the surface 

(flowers) having anthers mature before the pistils in the same 

flower 

the part nearest to the axis 

false or resembling 

slightly hairy 

hairiness 

clothed with soft hair 

air cavity or small bladder in the pulp of a fruit; e.g., Citrus 

fruit 

with juicy or fleshy tissue 

cushion-shaped 

the swollen base of a leaf-stalk 

marked with dots, depressions or translucent glands 

in the form of a point or a dot 

with minute dots 

strongly laxative 

having slight elevation like blisters 

a blister-like prominence; pimple 

a stone or kernel of a fruit; nutlet 

pear-shaped 

an indeterminate or centripetal inflorescence with lengthened 

axis and equally stalk;ed flowers 

raceme-like or having racemes 

the central elongated axis of an inflorescence or a compound 

leaf 

the rudimentary root of a plant embryo 

flowering on the branches 

a ridge of fibrous vascular tissue connecting the base of 

nucellus with the placenta 

that part of flower stalk (usually fleshy) which bears other 

floral organs 

kidney-shaped 

483

esinous 

reticulate 

retrorse 

retuse 

revolute 

rheumatism 

rhomboid 

rosette 

rugose 

rugulose 

ruminate 

saccate 

sagittate 

samara 

sarcotesta 

sarcotheca 

scabies 

scabrous 

scalariform 

scale 

scandent 

schizocarp 

scurfy 

semi-inferior 

semi-superior 

sepal 

septicidal 

seriate 

sericeous 

serrate 

serrulate 

sessile 

setaceous 

sinuate/sinuous 

spathaceous 

spathe 

spathulate 

spicate 

spike 

stamen 

producing resin or covered with resin 

netted like net-work, as venation of a leaf 

(anther) turned or directed backwards 

with a shallow notch at a rounded apex 

rolled back from the margin or apex 

any disease marked by inflammation and pain in the joints , 

muscles or fibrous tissue 

approaching a rhombic outline; quadrangular with the lateral 

angles obtuse 

arranged in a condensed circular fashion 

covered with wrinkles 

somewhat wrinkled 

(endosperm) having a mottled appearance through infolding of 

inner seed-coat 

sac-shaped or pouch-shaped 

shaped like the barbed head of an arrow 

a winged, non-spliting, flat, one-seeded fruit 

the fleshy outer portion of seed-coat 

fleshy pollen case or theca 

a contagious skin disease causing severe itching 

rough to the touch due to the presence of stiff hairs, scales, or 

points 

having markings suggestive of a ladder 

any thin scarious body, usually a degenerate leaf, sometimes of 

epidermal origin 

climbing 

a dry fruit which splits into two or more one-seeded portions 

covered with bran-like scales 

partially inferior 

partially superior 

leaf-like divisions of a calyx 

dehiscing through the dissepiments or lines of junction 

disposed in series of rows, either transverse or longitudinal 

clothed with close-pressed soft and straight pubescence 

beset with sharp teeth directed upwards on the margin 

serrate but the teeth minute 

without stalk 

bristle-like 

with a deep wavy margin 

resembling or bearing a spathe 

a large bract sheathing or enclosing an inflorescence or a 

flower cluster 

oblong, with the basal end attenuate like a druggist's spatula 

disposed in or resembling a spike 

an indeterminate inflorescence, with sessile flowers seated 

along a common axis 

the male organ of a flower, consisting of a stalk or a filament 

and-an anther containing pollen 

484

staminate 

staminode 

stellate 

stigma 

stigmatic 

stigmatose 

stimulant 

stipe 

stipellae 

stipitate 

stipular 

stipulary 

stipule 

stomatal crypts 

strand 

striate 

strigose 

style 

subulate 

sulcate 

superior 

suture 

syncarpous 

tanniferous 

tendril 

tenuinucellate 

tepal 

terete 

tessellate 

testa 

tetrad 

theca 

thyrse 

thyrsoid 

tomentellous 

tomentose 

tomentum 

having stamens 

rudimentary sterile stamen 

star-like or star-shaped with slender segments radiating out 

from a common centre, 

that portion of a pistil or style which receives the pollen 

relating to or having the function of a stigma 

provided with stigmas or having conspicuous stigmas 

a stimulus-producing agent 

stalk 

a minute stipule on a partial petiole of a compound leaf 

having a stipe or a special stalk 

having stipules or relating to stipules 

occupying the place of stipules 

an appendage of a leaf on each side of the leaf-insertion 

simple glandular pits or cavities on stomata 

a bundle of vascular tissue, resembling a cord 

marked with fine longitudinal parallel lines, as grooves or 

ridges 

beset with sharp-pointed appressed straight and stiff hairs or 

bristles 

the slender upper part of an ovary supporting a stigma 

awl-shaped; narrowing and tapering from the base to a fine 

point 

grooved or furrowed 

(ovary) with all the floral envelopes inserted below it, on the 

torus 

a line or a junction of two parts immovably connected; line of 

dehiscence 

composed of two or more united carpels 

producing tannin 

a slender stem or leaf-outgrowth which functions as a 

climbing apparatus 

(ovule) with a thin nucellus 

free segment of a perianth not differentiated into sepals and 

petals 

cylindrical and usually tapering, rounded in cross-section 

checkered 

the outer coat of a seed, usually hard and brittle 

a group of four; a body formed of four cells, as in the 

formation of pollen in the pollen-mother-cells 

a pollen-case 

a mixed inflorescence with the main axis indeterminate, and 

the secondary and ultimate axes cymose 

resembling a thyrse 

sparingly or minutely tomentose 

densely pubescent with matted wool, or short hairs 

pubescence of matted interwoven hairs 

485

torulose 

torus 

translucent 

trapezoid 

triad 

trichome 

trichotomous 

trifid 

trifoliolate 

tristylous 

truncate 

tubercle 

tuberculate 

turbinate 

umbel 

undulate 

ungulate 

uniseriate 

unisexual 

unitegmic 

urceolate 

valvate 

vein 

velutinous 

velvety 

venation 

vermifuge 

verrucose 

verruculose 

versatile 

vessel 

vestigial 

villous 

viviparous 

zygomorphic 

cylindrical with swollen portions at intervals; beaded 

same as receptacle 

transmitting rays of light without being transparent 

having a shape of a trapezium 

group of three 

hair-like outgrowth of the epidermis 

three-forked, branching into three divisions 

three-cleft 

with three leaflets 

with three styles 

squared at the tip or base as if cut off with a straight blade 

a small rounded protuberance 

having tubercles 

top-shaped; inversely conical 

an indeterminate inflorescence in which a cluster of pedicels 

spring from the same point, like the ribs of an umbrelia 

wavy 

hoofed; clawed 

in one horizontal row or series 

(flower) with either stamens or pistils only, not both 

(ovule) having one layer of integument 

pitcher-shaped, hollow and contracted at the mouth like an 

urn or a pitcher 

opening by valves, as in most dehiscent fruits or anthers; when 

parts of a flower-bud meet exactly without overlapping 

a thread or strand of vascular tissue in a flat organ, as a leaf 

velvety due to a coating of fine soft hairs 

densely covered with fine short soft erect hairs 

the disposition of veins; the mode of veining 

that expels intestinal worms 

warty 

very warty; much covered with warts 

swinging freely on its support, as many anthers on their 

filaments 

any tube or canal with properly defined walls in which fluids 

circulate 

small and imperfectly developed; the remaining trace of an 

organ which fully developed in some ancestral form 

covered with long fine hairs 

germinating or sprouting from seed or bud while still attached 

to the parent plant 

not radially symmetrical 

486

INDEX TO SCIENTIFIC NAMES 

(compiled by Rusea Go) 

Names in italics here refer to synonyms; family names given in upper case are those of families 

revised in this volume. Pages with illustrations are listed in italics. 

Acer 

Acer caesiaefolium 

Acer caesium 

Acer curranii 3 

Acer javanicum 3 

Acer laurinum 2, 3 

A~~w~ 3 

Acer philippinum 3 

ACERACEAE V, 1,453 

Acioa heteropetala 163 

Acronychia 352, 353, 356, 358, 381 

Acronychia apiculata 358 

Acronychia arborea 358 

Acronychia laurifolia 358 

Acronychia pedunculata 358,359 

Acronychia porteri 381,382 

Adenanthera triphysa 424 

Adolphia 306 

Aegiphila viburnifolia 113 

Agathidanthes 253 

Agathidanthes javanica 255 

Agathis XLVII, 27, 28 

Agathis alba 29 

Agathis beccarii 29 

Agathis borneensis XLV, 28, 29, 31 

Agathis dammara 29 

Agathis endertii 28, 29 

Agathis kinabaluensis 29,30 

Agathis latifolia 29 

Agathis lenticula 28, 30 

Agathis loranthiifolia 29 

Agathis macrostachys 29 

Agathis orbicula 28, 32 

Agathis rhomboidalis 29 

Agathisanthes 253 

Agathisanthes javanica 255 

Agelaea 188 

Agelaea borneensis 188 

Agelaea insignis 188 

Agelaea macrophylla 187, 188 

Agelaea trinervis 188 

1 

3 

3 

Ailanthus 421, 422, 423 

Ailanthus blancoi 423 

Ailanthus integrifolia 423 

Ailanthus peekelii 423 

Ailanthus philippinensis 424 

Ailanthus triphysa 423, 424, 425 

ALANGIACEAE V, 5,199 

Alangium 5,6, 199 

Alangium sect. Alangium 6 

Alangium sect. Angolam 6 

Alangium sect. Canostigma 6 

Alangium sect. Marlea 6 

Alangium sect. Rhytidandra 6 

Alangium begoniaefolium 10, 13 

Alangium bogoriense 10 

Alangium borneense 10 

Alangium chinensis var. tomentosum 10 

Alangium circulare XLIX, 7 

Alangium ebenaceum 10 

Alangium ebenaceum "var. E" 10 

Alangium ebenaceum "var. C" 10 

Alangium ebenaceum "var. D" 10 

Alangium ebenaceum "var. E" 10 

Alangium ebenaceum "var. G" 10 

Alangium ebenaceum var. tutela 10 

Alangium griffithii 7, 8 

Alangium havilandii 7, 8 

Alangium hirsutum 12 

Alangium javanicum 7, 9 

Alangiumjavanicum "form E" 10 

Alangiumjavanicum "form C" 10 

Alangiumjavanicum "form D" 10 

Alangiumjavanicum var. ebenaceum 10,11 

Alangium javanicum var. javanicum 10, 11 

Alangiumjavanicum var. meyeri 10,11 

Alangium kinabaluense 14 

Alangium kurzii 7, 10 

Alangium lamarckii 12 

Alangium longiflorum 6, 12 

Alangium meyeri 10 

Alangium mezianum 10 

487

A1angium nobile 7, 13 

Alangium ridleyi 10 

Alangium rotundatum 13 

A1angium rotundifo1ium 7, 13 

Alangium salvifolium subsp. hexapetalum 12 

A1angium scandens 6 

Alangium sessiliflorum 10 

Alangium tutela 10 

A1angium sp. 1 6 

A1eurocanthus wog1umi 407 

Allanthospermum 422,421,427 

Allantospermum borneense XLVII, 427 

Allantospermum borneense 

subsp. borneense 

Allantospermum borneense 

subsp. rostratmm 

Allantospermum multicau1e 

A1phitonia 

A1phitonia exce1sa 

Alphitonia incana 

Alphitonia philippinensis 

Ampacus accedens 

Ampacus alba 

Ampacus aromatica 

Ampacus glabra 

Ampacus macrophylla 

Ampacus robusta 

Ampacus roxburghiana 

Ampacus triphylla 

Amyris graveolens 

Anacardiaceae 

426,429 

429 

427 

306, 307 

308,309 

308 

308 

386 

391 

392 

390 

386 

390 

392 

394 

370 

XIII, XIV, XVIII, 

46,352,354,480 

Anaco1osa 271,274 

Anacolosa arborea 274 

Anaco1osa fmtescens 272, 274 

Anacolosa heptandra 274 

Anacolosa luzoniensis 274 

Anacolosa maingayi 284 

Anacolosa sp. 274 

Angelesia 165 

Angelesia splendens 165 

Anisophyllea 15,16,17,25,322 

Anisophyllea beccariana 17, 18 

Anisophyllea chartacea 17, 18 

Anisophyllea corneri 17, 19 

Anisophyllea disticha 16, 17, 19, 23 

Anisophyllea fermginea XLVII, 17, 21 

Anisophyllea glandulifolia 25 

Anisophyllea globosa 17, 20, 22 

Anisophyllea impressinervia 17, 22 

Anisophyllea nitida 17, 23 

Anisophyllea rhomboidea XLIX, 17, 19, 23 

ANISOPHYLLEACEAE V, 15, 16,322 

Anisophylleaceae tribe Anisophylleae 15 

Anisophyllum 16 

Anisophyllum trapezoidale 19 

Anisoptera grossivenia XL V, XL VI, XL VII 

Araliaceae 199 

Ara1idium 199 

Araucaria 27 

ARAUCARIACEAE V, 27 

Ardisia ochracea 127 

Arrabidaea magnifica 33 

Ata1antia 411 

Atalantia disticha 412 

Atalantia disticha var. paniculata 412 

Atalantia maritima 412 

, Atalantia nitida 411 

488 

Atalantia paniculata 

Atuna 

Atuna cordata 

Atuna elata 

Atuna excelsa 

Atuna nannodes 

Atuna racemosa 

Atuna racemosa subsp. exce1sa 

Atuna racemosa subsp. racemosa 

Atuna villamilii 

Aubletia caseolaris 

Aucuba 

Aucuba japonica 

Averrhoa 

Averrhoa bilimbi 

Averrhoa carambo1a 

A verrhoaceae 

Avicennia 

Barringtonia 

Begoniaceae 

Berchemia 

Bergera koenigii 

Bhesa 

Bhesa panicu1ata 

Bhesa robusta 

Bignonia indica 

Bignonia pentandra 

Bignonia spathacea 

412 

156, 157, 158 

L, 158, 159 

161 

161 

158 

158, 160 

161 

156, 161 

161 

450 

199 

199 

287,289 

287,289 

289 

288 

324,450 

421,442 

209 

306 

406 

107, 109, III 

Jl 0, 111.112 

111,112 

38 

38 

37

Bignonia tripinnata 38 

BIGNONIACEAE V, 33 

Biophytum 287,288 

Biophytum sensitivum 288 

Bischofia 454 

Bischofiaceae 454 

Blatti 447 

Blatti acide 450 

Blatti alba 449 

Blatti caseolaris 450 

Blatti pagatpat 450 

Bombacaceae 

XIII, XVIII 

Borneodendron aenigmaticum XLVIII 

Boswellia 46 

Boswellia sacra 46 

Brackenridgea 257,258,259 

Brackenridgea sect. Brackenridgea 259 

Brackenridgea sect. Notochnella 259 

Brackenridgea denticulata 259 

Brackenridgea hookeri 259,261 

Brackenridgea palustris 259,260 

Brackenridgea serrulata 260 

Brassicaceae 101, 102 

Brongniartia 246 

Brongniartia coriacea 247 

Brucea 421,422,429 

Brucea amarissima 431 

Bruceajavanica 428, 429 

Brucea sumatrana 431 

Brucea sumatrensis 

Bruguiera 

Bruguiera 10-angulata 

Bruguiera angularis 

Bruguiera australis 

Bruguiera capensi 

Bruguiera caryophylloides 

Bruguiera conjugata 

Bruguiera cylindrica 

Bruguiera decandra 

Bruguiera eriopetala 

Bruguiera gyqmorrhiza 

Bruguiera malabarica 

Bruguiera oxyphylla 

Bruguiera parietosa 

Bruguiera parviflora 

Bruguiera rheedii 

Bruguiera ritchiei 

Bruguiera rumphii 

431 

321,322,323 

326 

326 

326 

324 

324 

325 

324, 325, 326 

335 

326 

323, 324 

324, 326 

326 

326 

324,325,326,327 

324 

325 

324 

Bruguiera sexangula 323, 326 

Bruguiera wighti 324 

Bruinsmia 463,465 

Brutnsmia polysperma 465 

Bruinsmia styracoides 463, 464, 465 

Bulbophyllum 

xl 

Burkillanthus 261,352,353,357 

Burkillanthus malaccensis 261,360 

BURSERACEAE 

V, XIV, XIX, 

Burseraceae tribe Bursereae 

Burseraceae tribe Canarieae 

Calophyllum 

Calophyllum nodosum 

Calosanthes indica 

Campanulaceae 

Campylocerum 

Camunium 

45,46,352,421 

46 

46 

XIV, XIX 

XLVIII 

38 

213 

265 

406 

Canariopsis aspera 51 

Canarium 45, 46, 47, 48, 65, 84, 174 

Canarium sect. Canarium 48 

Canarium sect. Pimela 48 

Canarium sect. Tenuipyrena 65 

Canarium subg. Mricanarium 48 

Canarium subg. Canariellum 48 

Canarium subg. Canarium 48 

Canarium acutum 58 

Canarium album 46 

Canarium angulatum 68 

Canarium apertum 46, 50 

Canarium asperum L, 49, 51 

Canarium asperum subsp. asperum 51 

Canarium asperum subsp. asperum 

var. asperum 51 

Canarium asperum subsp. asperum 

var. clementis 51 

Canarium asperum subsp. papuanum 51 

Canarium beccarii 59 

Canarium brunneum 99 

Canarium caudatum 49,51 

Canarium caudatum forma auricuIiferum 52 

Canarium caudatum forma caudatum 52 

Canarium connarifolium 

Canarium costatum 

Canarium crassifolium 

Canarium cuspidatum 

Canarium decumanum 

Canarium denticulatum 

83 

66 

73 

73 

L,49,52,62,63 

49, 53 

489

Canarium denticulatum subsp. denticulatum 

53 

Canarium denticulatum subsp. kostermansii 

53 

Canarium dichotomum 

Canarium divergens 

Canarium endertii 

Canarium expansum 

Canarium fissistipulum 

Canarium jlavum 

Canarium fragile 

Canarium fusco-calycinum 

Canarium giganteum 

Canarium glaucum 

49, 53 

XLVII, 49, 54 

53 

67 

53 

58 

69 

50, 54 

58 

58 

Canarium grandifolium XLVII, 50, 55, 62 

Canarium hirsutum 50,55 

Canarium hirsutum subsp. hirsutum 

var. hirsutum forma scabrum 

Canarium hirtellum 

Canarium hirtipetalum 

Canarium his pi dum 

Canarium incurvatum 

Canarium indicum 

Canarium kedondon 

56 

62 

92 

55 

68 

46,48 

73 

Canarium kinabaluensis 49,56 

Canarium korthalsii 97 

Canarium kostermansii 49, 56 

Canarium kunstleri 53 

Canarium laciniatum 53 

Canarium laevigatum 90 

Canarium latistipulatum 50, 57,64 

Canarium laxum 69 

Canarium littorale 49, 50, 57 

Canarium littorale forma littorale 58 

Canarium littorale forma pruinosum 58 

Canarium littorale forma purpurascens 58 

Canarium littorale forma rufum 58 

Canarium littorale forma tomentosum 58 

Canarium luzonicum 46, 48 

Canarium megalanthum 48, 50, 58 

Canarium merrillii 50, 59 

Canarium micrantherum 97 

Canarium minahassae 73 

Canarium molle 51 

Canarium montanum 73 

Canarium motleyanum 62 

Canarium multifidum 59 

Canarium nitens 68 

Canarium nitidum 61 

Canarium odontophyllum 48, 50, 59, 60 

Canarium ovatum 46,48 

Canarium palawanense 59 

Canarium parvifolium 61 

Canarium patentinervium 49, 61, 63 

Canarium pauciflorum 52, 86 

Canarium pilosum 49,50,61 

Canarium pilosum subsp. borneensis 62 

Canarium pilosum subsp. pilosum 62 

Canarium pilosum var. hirtellum 62 

Canarium pimela 48,63,64 

Canarium planchonii 95 

Canarium pruinosum 58 

Canarium pseudodecumanum 48,49,62 

Canarium pseudopatentinervium 49,63 

Canarium pseudopimela 49,63 

Canarium reticulatum 73 

Canarium rubiginosum 

Canarium rufum 

Canarium rugosum 

Canarium rugosum var. sumatranum 

Canarium sarawakanum 

Canarium serricuspe 

Canarium serrulatum 

Canarium subcordatum 

Canarium villosum 

73, 94 

58 

74 

74 

50, 64 

58 

50, 58 

55 

51 

Canarium virgatum 74 

Canarium vulgare 46,48 

Cantanola 188 

CAPPARACEAE V, 101, 102 

Capparaceae subfam. Cleomoideae 102 

Capparis 

102 

Capparis magna 103 

Capparis spinosa 101 

Capusia 154 

Carallia 321,322,328 

Carallia arguta 330 

Carallia borneensis 332,329,331 

Carallia brachiata 329, 330 

Carallia calycina 330 

Carallia celebica 330 

Carallia cerisopsifolia 330 

Carallia con finis 330 

Carallia coriifolia 329, 332 

Carallia cuprea 330 

Carallia cuspidata 330 

490

Carallia floribunda 

Carallia lucida 

Carallia mindanaensis 

Carallia multijlora 

Carallia scortechinii 

Carallia spinulosa 

Carallia timorensis 

Carallia viridifolia 

Carallia sp. 1 





Cardiocarpus amarus 

Cardiophora hindsii 

Carya 

Caryospermum 

Caryospermum philippinensis 

Cassine 

Cassine vibumifolia 

Casuarina equisetifolia 

Ceanothus 

Ceanothus asiaticus 

CELASTRACEAE 

330 

330 

329 

330 

330 

330 

330 

330 

329, 332 

329, 333 

328,333 

329, 333 

328,334 

442 

442 

233 

150 

150 

113 

108, 113, 115 

XLIV 

305 

311 

V, XIV, XVIII, 

107,322,453 

Celastrales 322 

Celastrus 108, 154 

Celastrus alpestris 150 

Celastrus robustus 112 

Celtis angllstifolia 318 

Celtis grewioides 318 

Cenarium 47 

Ceratostachys 253 

Ceriops 321, 322, 334, 335 

Ceriops boviniana 335 

Ceriops candolleana 335 

Ceriops decandra 335 

Ceriops fosteniana 335 

Ceriops lucida 336 

Ceriops paucijlora 335 

Ceriops roxbllrghiana 335 

Ceriops tagal 335, 337 

Ceriops timoriensis 336 

Ceriops zippeliana 335 

Chalcas koenigii 406 

Chalcas paniculata 406 

Chamaepericlymenum suecicum 199 

CHRYSOBALANACEAE V, 155, 156 

Cinnamomum 433 

Citrofortllnella microcarpa 364 

Citrus 351, 352, 353, 355 

356,357,362,363,483 

Citrus acida 

Citrus angulata 

Citrus aurantifolia 

365 

395 

365 

Citrus aurantium subsp. saponacea 366 

Citrus aurantium var. grandis 364 

Citrus aurantium var. medica 363 

Citrus aurantium var. sinensis 365 

Citrus decumana 

Citrus deliciosa 

Citrus grandis 

Citrus halimii 

Citrus hystrix 

Citrus limon 

Citrus limonum 

Citrus macroptera 

Citrus macroptera var. macroptera 

Citrus madurensis 

Citrus malaccensis 

Citrus maxima 

Citrus medica 

Citrus medica var. limon 

Citrus microcarpa 

Citrus mitis 

Citrus nobilis 

Citrus papeda 

Citrus papuana 

Citrus reticulata 

Citrus reticlllata var. austera 

Citrus sinensis 

364 

364 

363,364 

362, 364 

365, 366 

364 

364 

362, 366 

366,367 

364 

261 

364 

363 

364 

364 

364 

364 

365 

366 

363, 364 

364 

365 

Citrus torosa 365 

Clausena 352,353,354,356,369,371 

Clausena calciphila 369,370 

Clausena excavata 368, 369, 370 

Clausena excavata var. excavata 370 

Clausena javanensis 

Clausena lansium 

Clausena wampi 

Cleome 

Clethra 

Clethra canescens 

Clethra canescens var. canescens 

Clethra canescens var. clementis 

Clethra canescens var. ledermannii 

Clethra canescens var. luzonica 

370 

369 

369 

102 

181 

181, 183 

183 

183 

183 

183 

491

Clethra canescens var. novoguinensis 

Clethra ciementis 

Clethra elongata 

Clethra longispicata 

Clethra pachyphylla 

CLETHRACEAE 

Clusiaceae 

Cnestis 

Colubrina 

Colubrina anomala 

Colubrina arborescens 

Colubrina asiatica 

Colubrina beccariana 

Colubrina elliptica 

Colubrina excelsa 

Colubrina ferruginosa 

Combretocarpus 

Combretocarpus motleyi 

Combretocarpus rotundatus 

Combretum 

183 

183 

184 

183, 184 

182, 183,184 

V, 181 

XIV, XLVII 

188 

305,306,311 

312 

311 

311 

310,311,312 

311 

308 

311 

15, 16, 25, 26, 322 

25 

XLVIII, 24, 25 

25 

Commiphora myrra 46 

Commiphora opobalsamum 46 

Compositae 480 

Coniferae 28 

CONNARACEAE V, 187,288 

Connaropsis 289 

Connaropsis acuminata 290 

Connaropsis diversifolia 290 

Connaropsis glauca 291 

Connaropsis grandiflora 290 

Connaropsis grifJithii 290 

Connarus 83, 187, 188, 189, 190 

Connarus agamae L, 189, 190, 193, 195 

Connarus borneensis 192 

Connarus cullionensis 192 

Connarus densiflorus 191 

Connarus ellipticus 190 

Connarus euphlebius 191 

Connarus falcatus 191 

Connarus grandis 190 

Connarus hebephyllus 192 

Connarus impressinervis 190 

Connarus jackiana 192 

Connarus lucens 191 

Connarus monocarpus 191 

Connarus monocarpus subsp. malayensis 191 

Connarus mutabillis 192 

Connarus odoratus 192 

Connarus pachyphyllus 191 

Connarus plumoso-stellatus 192 

Corinarus semidecandrus 192 

Connarus stellatus 192 

Connarus urdanetensis 197 

Connarus villosus 192 

Connarus winkleri 191 

Connarus sp. A. 189, 190 

Connarus sp. B. 189, 190 

Cookia chlorosperma 372 

Cookia cyanocarpa 374 

Cookia wampi 369 

CORNACEAE V, 5,199,200,253 

Cornales 5, 322 

Cornus mas 199 

Cotylelobium lanceolatum 

XL V 

Craterianthus 341 

Crateva 101, 102, 103 

Crateva brownii 104 

Crateva lophosperma 103 

Crateva macrocarpa 104 

Crateva magna 103,105 

Crateva membranifolia 104 

Crateva nurvala 103 

Crateva religiosa 103, 104,105, 106 

Cratoxylum 219,220,221 

CratoJo.,"ylum sect. Cratoxylum 221 

Cratoxylum sect. Isopterygium 221 

Cratoxylum sect. Tridesmos 221 

Cratoxylum arborescens 219,221,223 

Cratoxylum celebicum 226 

Cratoxylum cochinchinense 

219,221,222,223 

Cratoxylum cochinchinense var. calcareum 

225 

Cratoxylum formosum 219,221,223,224 

Cratoxylum glaucum 219,221,225 

Cratoxylum hypericum 226 

Cratoxylum ligustrinum 223 

Cratoxylum maingayi 219,221,223,225 

Cratoxylum myrtifolium 223 

Cratoxylum polyanthum 223 

Cratoxylum procerum 225 

Cratoxylum sumatranum 221, 226 

Cratoxylum sumatranum subsp. blancoi 226 

Cratoxylum sumatranum 

subsp. sumatranum 226 

Crossostylis 322 

Cucurbitaceae 209 

492

Cunoniaceae 

Curtisia 

Curtisia faginea 

322,453 

199 

199 

Curtisiana 65 

Curtisiana penangensis 69 

Cyclandrophora 157 

Cyclandrophora asperula 161 

Cyclandrophora e/ata 161 

Cyclandrophora excelsa 161 

Cyclandrophora glaberrima 161 

Cyclandrophora nannodes 159 

Cyclandrophora villamilii 161 

Dacrydium comosum 

XLIX 

Dacryodes 46, 47, 65, 76, 84, 87 

Dacryodes angulata 68 

Dacryodes costata 65,66,67 

Dacryodes elmeri L, 65, 67 

Dacryodes expansa 66, 67 

Dacryodes incurvata 66, 68 

Dacryodes kostermansii 71 

Dacryodes laxa 65, 68 

Dacryodes longifolia 66, 69 

Dacryodes longifolia var. penangensis 69 

Dacryodes macrocarpa 66, 69 

Dacryodes macrocarpa var. genuina 71 

Dacryodes macrocarpa var. kostermansii 71 

Dacryodes macrocarpa var. macrocarpa 71 

Dacryodes macrocarpa var. patentinervia 71 

Dacryodes nervosa 66, 72 

Dacryodes rostrata 65, 66, 70, 72 

Dacryodes rostrata f. cuspidata 65, 73 

Dacryodes rostrata f genuina 73 

Dacryodes rostrata f. rostrata 73 

Dacryodes rubiginosa XLIX, 66, 73 

Dacryodes rugosa 65,66,67, 74 

Dacryodes rugosa var. genuina 74 

Dacryodes rugosa var. rugosa 74 

Dacryodes rugosa var. virgata 74 

Dacryodes scan dens 62 

Dactylocladus stenostachys XL VIII 

Dalrymplea 454 

Dalrympleajavanica 460,461 

Dammara 28 

Dammara loranthiifolia 29 

Dapania 288 

Dapania grandifolia 288 

Daphniphyllopsis 253 

Daphniphyllopsis capitata 255 

Datisca 

DATISCACEAE 

Deplanchea 

Deplanchea bancana 

Deplanchea coriacea 

Derris 

Diacaecarpium rotundifolium 

Diatoma brachiata 

Dichapetalum tetramerum 

Diemenia 

Dilleniales 

209 

V,209 

35,36 

33, 34, 36 

36 

187 

13 

330 

196 

169 

187 

Diplanthera bancana 36 

Dipterocarpaceae XIII, XVIII, 201 

Dipterocarpus beccarii 

XL V 

Dipterocarpus caudiferus 

XL VI 

Dipterocarpus crinitus XL V, XL VI, XL VII 

Dipterocarpus globosus XL VI, XL VII 

Dipterocarpus lowii 

XL VII 

Dipterocarpus mandus 

L 

Dipterocarpus pachyphyllus 

L 

Dipterocarpus rigidus 

XL VII 

Dipterocarpus sarawakensis XL VII 

Dipterocarpus validus 

L 

Dipteronia 1 

Discaria 306 

Dolichandrone 35, 37 

Dolichandrone longissima 37 

Dolichandrone rheedii 37 

Dolichandrone spathacea 33, 35, 37,39 

Dracontomelon cuspidatum 73 

Driessenia microthrix 25 

Drimys piperita 

XLIX 

Dryobalanops aromatica 

XLVI, XL VII, XLIX 

Dryobalanops beccarii XL VII, XL VIII 

Dryobalanops fusca 

XLIX 

Dryobalanops keithii 

L 

Dryobalanops lanceolata 

XL V, XL VI, XL VIII 

Dryobalanops rappa 

225 

Dryptopetalum coriaceum 

339 

Duabanga 

443,445,446 

Duabanga borneensis 

446 

Duabanga grandiflora 

446 

Duabanga moluccana 444,445,446 

Duabanga taylorii 

446 

Durio 

XIII, XVIII 

Durio bukitrayaensis 

XVIII 

493

Dysoxylum dasyphyllum 

Ebenaceae 

423 

XLVII 

Elaeocarpaceae 322 

Elaeodendron 113 

Elaeodendron subrotundum 113 

Elepanthus longifolius 197 

Ellipanthus 187, 188, 195, 196 

Ellipanthus beccarii 196 

Ellipanthus beccarii var. beccarii XLIX, 196 

Ellipanthus beccarii var. peltatus 196 

Ellipanthus burebidensis 197 

Ellipanthus luzoniensis 197 

Ellipanthus mindanaensis 196 

Ellipanthus sarawakensis 197 

Ellipanthus tomentosus 194, 196 

Ellipanthus tomentosus subsp. tomentosus 

var. luzoniensis 197 

Ellipanthus urdanetensis 197 

Ellipanthus vidalii 197 

Elodea Jormosa 224 

Elodea sumatrana 226 

Embelia urophylla 171 

Engelhardia XLVIII, 233, 235 

Engelhardia aceriflora 244 

Engelhardia apoensis 37,235 

Engelhardia chrysolepis 242 

Engelhardia danumensis 235,236,237,238 

Engelhardia Jenzelii 242 

Engelhardia Jormosana 242 

Engelhardia kinabaluensis 235,237,238 

Engelhardia lepidota 241 

Engelhardia mendalomensis 235,237,239, 

Engelhardia mersingensis 235, 237, 240, 242 

Engelhardia nudiflora 243 

Engelhardia palembanica 243 

Engelhardia parvifolia 243 

Engelhardia permicrophylla 243 

Engelhardia philippinensis 244 

Engelhardia pterococca 242 

Engelhardia pterococca var. aceriflora 244 

Engelhardia rigida 235,237,238, 

239,240,241 

Engelhardia rigida var. rigida 241 

Engelhardia rigida var. subsimplicifolia 240 

Engelhardia roxburghiana 234, 235, 

237,240,241 

Engelhardia serrata 235, 237, 242 

Engelhardia serrata var. cambodica 243 

Engelhardia serrata var. nudiflara 243 

Engelhardia serrata var. parvifolia 243 

Engelhardia serrata var. serrata 243 

Engelhardia spicata 235,237,244 

Engelhardia spicata var. aceriflora 244 

Engelhardia spicata var. colebrookeana 244 

Engelhardia spicata var. Jormosana 242 

Engelhardia spicata var. spicata 244 

Engelhardia subsimplicifolia 240 

Engelhardia wallichiana 242 

Engelhardia zambalensis 241 

Engelhardtia 233,234 

Ericaceae 

XLVII 

Ericales 181 

Erythropalum 273 

Erythropalum scandens 273 

Eugenia XIV, 202 

Eugenia bankense 

XL VIII 

Euodia 384 

Euodia accedens 386 

Euodia alba 391 

Euodia arborea 393 

Euodia aromatica 392 

Euodia bonwickii 388 

Euodia concinna 393 

Euodia conJusa 389 

Euodia glabra 389, 390 

Euodia kingii 390 

Euodia krukovii 390 

Euodia latiJolia 392 

Euodia lunu-ankenda 392 

Euodia macrophylla 386 

Euodia malayana 393 

Euodia obtusifolia 393 

Euodia punctata 393 

Euodia ridleyi 389, 390 

Euodia robusta 390 

Euodia roxburghiana 392 

Euodia schullei var. ridleyi 389 

Euodia speciosa 388 

Euodia suaveolens var. ridleyi 389 

Euodia subunifoliolata 394 

Euodia tenuistyla 389 

Euodia triphylla 394 

Euodia triphylla var. pubescens 393 

Euodia vilamilii 388 

Euonymus 107, Ill, 114, 117, 121 

Euonymus acuminifolius 117 

Euonymus acuminifolius var. borneensis 117 

494

Euonymus alatus 119 

Euonymus castaneifolius 116, 117, 118 

Euonymus cochinchinensis 117, 118 

Euonymus coriaceus 119 

Euonymus elmeri 119 

E uonymus glandu10sus 117, 119 

Euonymus javanicus 114, 117, 119 

Euonymus micropetalus 119 

Euonymus moultonii 118 

Euonymus pahangensis 119 

Euonymus philippinensis 118 

Euonymus viburnifolius 113 

Euphorbiaceae 

Eurycoma 

Eurycoma longifolia 

Eurycoma merguensis 

Euscaphis 

Euscaphis japonica 

Euthemis 

Euthemis ciliata 

Euthemis hackenbergii 

Euthemis 1eucocarpa 

Euthemis minor 

Euthemis obtusifolia 

Euthemis robusta 

Evodia accedens 

Evodia nervosa 

Exitelia corymbosa 

Fagaceae 

Fagara 

Fagara avicennae 

Fagura glabra 

Fagara lunu-ankenda 

Fagara myriacantha 

/

Griselinia 

Griselinia littoralis 

199 

199 

Guaiacum abilo 75 

Guttiferae XIV, XIX, XLVII, 219, 220 

Gynotroches 321, 322, 323, 336 

Gynotroches axillaris 

Gynotroches dryptopetalum 

Gynotroches lanceolata 

Gynotroches micrantha 

Gynotroches parvifolia 

Gynotroches puberula 

Gynotroches reticulata 

Haloragis disticha 

Haplolobus 

Haplolobus beccarii 

Haplolobus bintuluensis 

Haplolobus borneensis 

Haplolobus inaequifolius 

Haplolobus kapitensis 

Haplolobus leenhoutsii 

Haplolobus sarawakanus 

Harmandia 

Harmandia kunstleri 

Harmandia mekongensis 

Harrisonia 

Harrisonia perforata 

Hasskarlia 

Hebonga obliqua 

Hemisantiria 

Hemisantiria nitida 

Hemisantiria rostrata 

Hemisantiria rugosa 

Hemisantria ? n. sp. 

Hippocratea maingayi 

Hippoxylum indicum 

Hopea badiifolia 

Hopea beccariana 

Hopea bullatifolia 

Hopea centipeda 

Hopea dasyrrachis 

Hopea fluvialis 

Hopea megacarpa 

Hopea tenuinervula 

Hopea treubii 

Huertea 

HYPERICACEAE 

Hypericaceae tribe Cratoxyleae 

Hypericaceae tribe Hypericeae 

338, 339 

339 

339 

339 

339 

339 

339 

19 

47, 75, 76 

XLIX, 76 

76, 79 

86 

XLIX, 76, 79 

76,78,80 

76,78,80 

76, 81 

273,275 

277 

276,277 

421,422 

422 

454 

424 

65 

68 

72 

74 

67 

128 

38 

L 

XLVII 

L 

L 

L 

L 

L 

XLVII 

XLVII 

453 

V, 219, 220 

220 

220 

Hypericaceae tribe Vismieae 

Hypericum 

Hypericum arborescens 

Hypericum cochinchinense 

Hypericum japonicum 

Hypericum petiolulatum 

Icacinaceae 

Jcicaster 

Jcicaster planchonii 

Illex daphniphylloides 

ILLlCIACEAE 

Illiciales 

Illicium 

Illicium sect. Badiami 

Illicium sect. Cymbostemon 

Illicium sect. Illicium 

Illicium anisatum 

Illicium cauliflorum 

Illicium kinabaluense 

Illicium religiosum 

Illicium stapfii 

Illicium verum 

Illicium sp. 

Indovethia 

Indovethia calophylla 

Irvingel/a harmandiana 

Irvingella malayana 

Irvingella oliveri 

Irvingia 

Irvingia harmandiana 

Irvingia longipedicellata 

Irvingia malayana 

Irvingia oliveri 

Irvingiaceae 

Isopterys penangiana 

Itea javanica 

Ixonanthaceae 

Jacaranda filicifolia 

Jacaranda rhombifolia 

Jambolifera pedunculata 

JambolzJera porteri 

JUGLANDACEAE 

Juglans 

Juglans pterococca 

Kaernbachia 

Kandelia 

Kandelia candel 

Kanilia caryophylloides 

220 

219,220 

223 

223 

219, 220 

219,220 

271 

83 

95 

255 

V, 227 

227 

227, 229 

229 

229 

229 

229 

229,230,231 

229, 230 

229 

228, 229,230, 231 

229 

230 

258 

258 

434 

434 

434 

421,422,433 

434 

434 

432, 434 

434 

421 

469 

299 

422 

33 

33 

358 

382 

V, 233 

233 

242 

454 

321, 322, 339 

340, 341 

324 

496

Kanilia parviflora 325 

Kibara 245,246 

Kibara angustifolia 247 

Kibara blumei 247 

Kibara chartacea 247 

Kibara coriacea 246 

Kibara cuspidata 247 

Kibara depauperata 247 

Kibara ellipsoidea 247 

Kibara grandifolia 247 

Kibara macrophylla 247 

Kibara mollis 247 

Kibara motleyi 247 

Kibara obtusa 

L,246,247,249 

Kibara serrulata 247 

Kibara streimannii 246 

Kibara tomentosa 247 

Kibara trichantha 247 

Kokoona 108, Ill, 122, 123, 124, l30, 153 

Kokoona coriacea 123, 124 

Kokoona lanceolata 127 

Kokoona leucoclada 123, 124, 125 

Kokoona littoralis 123, 124, 125 

Kokoona littoralis var. bakoensis 127 

Kokoona littoralis var. littoralis 127 

Kokoona littoral is var. longifolia 127 

Kokoona ochracea 123, 124, 127 

Kokoona ovatolanceolata 

123, 124, 126, 128, 129 

Kokoona reflexa 

Kokoona sabahana 

Kokoona scortechinii 

Koompassia 

Koompassia excelsa 

Koompassia malaccensis 

Kostermanthus 

Kostermanthus heteropetalus 

Kostermanthus malayanus 

Krugiodendron 

Kurrimia 

Kurrimia luzonica 

Kurrimia maingayi 

Kurrimia minor 

Kurrimia paniculata 

Kurrimia pulcherrima 

Labiatae 

Lauraceae 

Laurus caesia 

123, 124, 128 

L, 124, 128 

127, 128 

XLVI 

212 

187 

156, 157, 161 

162, 163 

161 

305 

III 

112 

112 

112 

112 

112 

35 

201 

3 

Lavellea 

Leguminosae 

Lepidobotryaceae 

Lepidobotrys 

Lepidocarpa costata 

Leptospermum flavescens 

Licania 

Licania splendens 

Limonellus angulosus 

Limonia 

Limonia aurantifolia 

Limcnia corymbosa 

Limonia disticha 

Limonia lucida 

Limonia minuta 

Limonia parviflora 

Litsea palustris 

Lobelia 

Lobelia frutescens 

Lobelia p!umieri 

Loeseneriella 

Lophopetalum 

283 

187,212 

287 

287 

l75 

XLIX 

156, 157, 165 

164, 165 

395 

351, 352, 353 

365 

411 

411 

406 

400 

372 

XLVIII 

213 

217 

217 

109 

108, Ill, 123, 129, 130, 131, 153 

Lophopetalum beccarianum 

130, 131, 132, 133 

Lophopetalum coriacea 124 

Lophopetalum curtisii 135 

Lophopetalum dubium 127 

Lophopetalum fimbriatum 138 

Lophopetalum fuscescens 134 

Lophopetalum glabrum 130, 131, l32, 133 

Lophopetalum havilandii 132 

Lophopetalum intermedium 134 

Lophopetalumjavanicum 131,132,134 

Lophopetalum littoralis 127 

Lophopetalum maingayi 127 

Lophopetalum multinervium 131, l32, 134 

Lophopetalum oblongifolium 134 

Lophopetalum oblongum 134 

Lophopetalum pachyphyllum 

XLIX, l31, 135 

Lophopetalum pallidum 

Lophopetalum paucinervium 

Lophopetalum reflexum 

Lophopetalum rigidum 

Lophopetalum scortechinii 

Lophopetalum sessilifolium 

Lophopetalum subovatum 

108, 131, 135, 

134 

128 

131, 136 

132 

130, 131, 137 

130, l31, 137 

497

Lophopetalum subsessile 

l36 

Lophopetalum wightianum l31, l3 8 

Lophopetalum winkleri 

l38 

Lunasia 352, 353, 354, 355, 379 

Lunasia amara 378, 379 

Lunasia amara var. amara 380 

Lunasia amara var. babuyanica 380 

Lunasia gigantifolia 379 

Lunasia reticulata 379 

Lusa radja 431 

Luvunga 

352,353,357 

445,447 

212 

Lythraceae 

Macaranga 

Mac1urodendron 

352,353,354,356,380,381 

Mac1urodendron parviflorum XLIX, 381 

Mac1urodendron porteri 381,382,383 

Mac1urodendron pubescens L, 381, 382 

Macrosolen rotundatus 25 

Magnoliaceae 227 

Magnoliales 227 

Malpighiales 288 

Mangifera 

XIII, XVIII 

Mangium 345 

Manotes asiatica 433 

Mansoa hymensala 33 

Manungala pendula 439 

Maranthes 156, 157, 167 

Maranthes corymbosa 166, 168 

Marlea begonifolia 

l3 

Marlea densiflora 8 

Marlea ebenacea 10 

Marlea grifjithii 8 

Marlea nobilis 13 

Marlea rotundifolia 

l3 

Mastixia 199,200,201 

Mastixia subg. Manglesia 200 

Mastixia subg. Mastixia 200 

Mastixia subg. Mastixia series Alternae 200 

Mastixia subg. Mastixia series Oppositae 

200 

Mastixia subg. Pentamastixia 

Mastixia subg. Tetramastixia 

Mastixia acuminatissima 

Mastixia bracteata 

Mastixia caesia 

Mastixia caudatifolia 

200 

200 

205 

201 

205 

205 

Mastixia clarkeana 208 

Mastixia clarkeana var. macrophylla 208 

Mastixia cuspidata 201, 202 

Mastixia cuspidata var. margarethae 

Mastixia eugenioides 

Mastixia glauca 

A1astixia kimanila 

205,201 

201, 202 

XLIX, 201, 203 

205 

Mastixia korthalsiana 206 

Mastixia korthalsiana var. macrophylla 208 

Mastixia laxa 205 

Mastixia laxa var. angustifolia 205 

Mastixia macrocarpa 201,203 

Mastixia maingayi 206 

Mastixia maingayi var. subtomentosa 206 

Mastixia margarathae 205 

Mastixia megacarpa 204 

Mastixia parvifolia 204 

Mastixia pentandra 201, 204 

Mastixia pentandra subsp. cambodiana 204 

Mastixia pentandra subsp. chinensis 204 

. Mastixia pentandra subsp. mo1uccana 204 

Mastixia pentandra subsp. pentandra 204 

Mastixia pentandra subsp. philippinensis 204 

Mastixia pentandra subsp. scortechinii 204 

Mastixia pen tan dra var. cuspidata 201 

Mastixia premnoides 208 

Mastixia propinqua 206 

Mastixia rostrata 201,204,205 

Mastixia rostrata subsp. caudatifolia 

202, 205, 207 

Mastixia rostrata subsp. rostrata 205 

Mastixia scortechinii 204 

Mastixia trichotoma 201,205 

Mastixia trichotoma var. benculuana 208 

Mastixia trichotoma var. c1arkeana 208 

Mastixia trichotoma var. korthalsiana 206 

Mastixia trichotoma var. maingayi 206 

Mastixia trichotoma var. rhynchocarpa 208 

Mastixia trichotoma var. simalurana 208 

Mastixia trichotoma var. tenuis 208 

Matthaea 245, 246, 248 

Matthaea calophylla 248 

Matthaea ellipsoidea 248 

Matthaea latifolia 248 

Matthaea sancta 248,250 

Maytenus 108 

Maytenus emarginatus 108 

498

Meesia 265 

Meesia serrata 265 

Melastomataceae 25 

Meliaceae XXXII, 46,352,421 

Melicope 104,352,353,355,384,385 

Melicope accendens 387, 386 

Melicope bonwickii L, 385, 388 

Melicope clemensiae 385, 388 

Melicope conferta 358 

Melicope confusa 385, 389 

Melicope denhamii L, 385, 389 

Melicope glabra 385, 390 

Melicope hookeri 386,390 

Melicopeincana 386,391 

Melicopejugosa L, 385, 391,393 

Melicope latifolia 385, 392 

Melicope lunu-ankenda 386, 392 

Melicope sororia L,393 

Melicope subunifoliolata L, 385, 394 

Melicope tetrandra 414 

Melicope triphylla 385, 394 

Melicope unifoliolata 382 

Menispermaceae 

XXVIII 

Merope 352,353, 355, 357, 395 

Merope angulata 395, 397, 405 

Merrillia 

352,353,354,356,396 

Merrillia caloxylon 398,399 

Micromelum 352, 353, 354, 356, 389 

Micromelum glabrescens 400 

Micromelum minutum 400,402 

Micromelum minutum var. minutum 

400,401 

Micromelum pubescens 400 

Microtropia 138 

Microtropis 107, 109, 138, 139, 140 

Microtropis argentea 139, 140, 141 

Microtropis bicolor 149 

Microtropis borneensis 139, 140, 141, 142 

Microtropis fascicularis 139, 140, 143; 144 

Microtropis g~andifolia 139, 140, 144 

Microtropis grandifolia var. grandifolia 144 

Microtropis grandifolia var. longipetiolatus 

144 

Microtropis keningauensis 139, 140, 144 

Microtropis kinabaluensis 139, 141, 144, 146 

Microtropis kinabaluensis var. acuminata 

144 

Microtropis ovata 139, 140, 145 

Microtropis pauciflora 149 

Microtropis platyphylla 139, 141, 142, 145 

MicIotropis rigida 139, 140, 146 

Microtropis sabahensis L, 139, 140, 146,147 

Microtropis sarawakensis 139, 140, 141, 147 

Microtropis sterrophylla 144 

Microtropis suborbiculata 149 

Microtropis sumatrana 13 9, 141, 147 

Microtropis valida 139, 140, 149 

Microtropis wallichiana 139, 140, 149 

Millingtonia hortensis 33 

Millingtonia pinnata 42 

Monanthocitrus 352, 353, 355, 357, 402 

Monanthocitrus bispinosa 402 

Monanthocitrus oblanceolata L, 402, 403 

MONIMIACEAE V, 245 

Monimiaceae subfam. Mollinedieae 245 

Moraceae 

)CL 

Morinda 434 

Murraya 352, 353, 354, 356, 

371,396,398,405,406 

Murraya sect. Bergera 406 

Murraya sect. Murraya 396, 398, 406 

Murraya burmanni 370 

Murraya calaxylon 398 

Murraya exotica 405,406.407 

Murraya foetidissima 406 

Murraya koenigii 405,406 

Murraya odorata 406 

Murraya paniculata 404, 405,406,407 

Murraya paniculata var. exotica 406 

Murraya paniculata var. paniculata 406, 407 

Murraya sumatrana 406 

Myricaceae 233 

Myrtaceae 

XIV, )CL VII 

Myrtales 156, 322 

Neckia 258 

Neckia serrata 

258 

Neolamarckia cadamba 

212 

Nepenthes 

XXXVIII 

Nyctocalos 

35 

Nypa 

450 

Nypa fruticans 

396 

Nyssa 

199,253 

Nyssa arborea 

255 

Nyssa bifida 

255 

Nyssa javanica 

253,254,255 

499

Nyssa sessiliflora 

NYSSACEAE 

Ochanostachys 

Ochanostachys amentacea 

Oehanostaehys baneana 

OCHNACEAE 

255 

V, 199,253 

271,274,277 

278,279 

278 

V, 257 

257 

Ochnaceae subfam. Ochnoideae 

Ochnaceae subfam. Sauvagesioideae 257 

Oehranthe 454 

Octomeles 209 

Octomeles sumatrana XLVl, 210, 2Jl, 212 

OLACACEAE V,271 

01acales 271 

01acineae 

271 

01ax 

273 

01ax imbricata 

273 

Opiliaceae 

271 

Orchidaceae 

XIII, XXXVl 

Oreomunnea 

234 

Oreorhamnus 

312 

Oroxylum 

35,38 

Oroxylum indicum 33,35,38,40 

Ouratia angustifolia 

265 

Ouratia beeeariana 

265 

Ouratia borneensis 

265 

Ouratia megaearpa 

265 

Ouratia neriifolia 

265 

OXALIDACEAE 

V, 287, 288 

Oxalis 

287,288 

Oxalis corniculata 

288 

Oxalis corymbosa 

288 

Paehylobus 

65 

Pagapate 

447 

Paliurus 

305, 306 

Paliurus spina-christin 

305 

Paraeelastrus 

138 

Paramignya 

352,353,357 

Paramignya angulata 

395 

Paramignya longispina 

395 

Parashorea malaanonan 

L 

Parashorea tomentella 

L 

Parastemon 155,156, 157,169 

Parastemon grandifructus 157, 169, l71 

Parastemon spieatum 171 

Parastemon urophyllus 169, 170, 171 

Parinari 156, 157, 161, 167, 172, l73 

Parinari sect. Sareostegia 167 

Parinari subg. Cyclandrophora 157 

Parinari subg. Exitelia 

Parinari subg. Sareostegia 

Parinari argenteo-sericea 

Parinari ashtonii 

Parinari bieolor 

Parinari canarioides 

Parinari costata 

Parinari costata subsp. costata 

Parinari costata subsp. rubiginosa 

Parinari elmeri 

Parinari gigantea 

Parinari heteropetaia 

Parinari heteropetaium 

Parinari metallica 

Parinari myriandra 

Parinari nannodes 

Parinari oblongifolia 

Parinari rigida 

Parinari rubiginosum 

Parinarium 

Parinarium subg. Petroearya 

Parinarium asperulum 

Parinarium borneensis 

167 

167 

L,I73 

179 

175 

173, 174 

173, l75 

175 

175 

l73, l77 

178 

161 

163 

XL VlI, 173, l78 

161 

158 

173, 176, l78 

173, 179 

l75 

172 

172 

161 

l78 

Parinarium eorymbosum 168 

Parinarium eostatum var. rubiginosum 175 

Parinarium elatum 161 

Parinarium giaberrimum 161 

Parinarium grifJithianum 168 

Parinarium myriandrum 163 

Parinarium scab rum 161 

Parinarium villamilii 161 

Pellacalyx 321,322,323,341,343 

Pellacalyx axillaris 342, 343 

Pellacalyx cristatus 343,344 

Pellacalyx lobbii 343, 344 

Pellacalyx symphiodiscus 343,345 

Perrottetia 107, 109, 150 

Perrottetia alpestris 150 

Perrottetia alpestris subsp philippinensis 

148, 150 

Perrottetia philippinensis subsp. alpestris 

150 

Perrottetia philippinensis subsp. moluccana 

150 

Petalinia 

Petalinia bancana 

Petroearya 

Petroearya exeelsa 

277 

278 

172 

161 

500

PhylIoc1adaceae 

PhylIoc1adus hypophyllus 

Picrasma 

Picrasma andamanica 

Picrasma denhamii 

Picrasma javanica 

Picrasma nepalensis 

Picrasma philippinensis 

Picroxylon siamense 

Pilocarpus amara 

28 

XLIX 

421,422,435 

435 

389 

435,437 

435 

435 

433 

379 

Pimela dichotoma 53 

Pinus dammara 29 

PITTOSPORACEAE V, 297 

PHtosporum 297, 298, 299 

Pittosporum accuminatissimum 303 

Pitfosporum clementis 302 

Pitfosporum comptum 302 

Pitfosporum epiphyticum 303 

Pittosporum ferrugineum 298,299 

Pitfosporum javanicum 299 

Pittosporum linearifolium 

L, 298,299, 300,301 

Pittosporum longisepa1um 298,299,300 

Piftosporum nativitatis 299 

Pittosporum ramiflorum 298, 299, 302 

Pitfosporum ramiflorum var. parviflorum 

Pittosporum resiniferum 

Pitfosporum rufescens 

Pittosporum si1amense 

Pitfosporum versteeghii 

Plaesiantha 

Plaesiantha lobbii 

P1eiospermium 

302 

L, 298, 299, 303 

299 

298,299,303 

299 

341 

344 

352,353,354,355,356,357,407,409 

P1eiospermium 1atia1atum 409 

P1eiospermium longisepalum 408, 409,410 

Podocarpaceae XLVII, 28 

Po1ygalaceae 154,467,469 

Po1yosma hookeri 14 

Protium 47, 81 

Protium connarifo1ium L, 82, 83 

Protium philippinensis 83 

Pseudellipanthus 195 

Pseudellipanthus beccarii 196 

Pseudellipanthus peltatus 196 

Pseuditea javanica 299 

Pterocymbium tubu1atum 

XL VI 

Pterospermum javanicum 

XLVI 

Pterilema 234 

Pterilema aceriflorum 244 

Quassia 422, 436 

Quassia borneensis XL VII, 436, 439 

Quassia indica 421,438,439 

Quinaria lansium 369 

Radermachera 33,35,41 

Radermachera corymbosa 42 

Radermachera lobbii 42 

Radermachera lobbii subsp. acuminata 42 

Radermachera pinnata 41,42 

Radermachera pinnata subsp. acuminata 

42,43 

Radermachera pinnata subsp. pinnata 42 

Radermachera ramiflora 41,42,44 

Radermachera whitfordii 42 

Radermachera sp. A. 41,44 

Rafflesia 

XIV, XIX 

Rafflesiaceae 

XIV, XIX 

Reissantia 108 

Rex amaroris 

Reynosia 

RHM1NACEAE 

Rhamnaceae tribe ColIetieae 

Rhamnaceae tribe Gouanieae 

442 

305 

V, 305,453 

306 

306 

Rhamnaceae tribe Rhamneae 306 

Rhamnaceae tribe Venti1agineae 306 

Rhamnaceae tribe Zizipheae 306 

Rhamnus 306,312 

Rhamnus borneensis 313, 315 

Rhamnus borneensis var. borneensis 3 l3 

Rhamnus cathartica 313 

Rhamnus incanus 308 

Rhamnus lancifolia 313 

Rhizophora 

321,322,323,325,326,345,347 

Rhizophora apiculata 346, 347 

Rhizophora australis 326 

Rhizophora candel 341 

Rhizophora candelaria 347 

Rhizophora caryophylloides 324 

Rhizophora caseolaris 449,450 

Rhizophora ceratophylloides 324 

Rhizophora conjugata 325, 347 

Rhizophora cylindrica 324, 325, 326 

Rhizophora decandra 335 

Rhizophora eriopetala 326 

501

Rhizophora glomerulata 

Rhizophora gymnorrhiza 

Rhizophora lingissima 

Rhizophora macrorrhiza 

Rhizophora mangle 

Rhizophora mucronata 

Rhizophora mucronata var. stylosa 

Rhizophora palun 

Rhizophora parvijlora 

Rhizophora plicata 

Rhizophora polyandra 

Rhizophora sexangula 

Rhizophora stylosa 

Rhizophora tagal 

Rhizophora timorensis 

Rhizophora tinctoria 

RHIZOPHORACEAE 

335 

324 

348 

348 

348 

347,348 

348 

324 

325 

326 

326 

326 

347, 348 

335 

335 

324 

V, XIV, XVIII, 6, 15, 321, 322, 483 

Rhizophoraceae tribe Gynotrocheae 322 

Rhizophoraceae tribe Macarisieae 322 

Rhizophoraceae tribe. Rhizophoreae 322 

Rhizophorales 322 

Rhododendron quadrasianum XLIX 

Rhus javanica 429 

Rosaceae 155, 156 

Rosales 156, 187 

Roucheria 289 

Roucheria macrophylla 292 

Rourea 189,288 

Rourea diversifolia 290 

Rourea mimosoides 187 

Roureopsis 188 

Roureopsis acutipetala 188 

Roureopsis emarginata 188 

Rubiaceae 5,212,322,434 

RUTACEAE V, 351, 352, 421 

Rutaceae subfam. Aurantioideae 352,353 

Rutaceae subfam. Flindersioideae 353 

Rutaceae subfam. Rhabdodendroideae 353 

Rutaceae subfam. Rutoideae 352,353 

Rutaceae subfam. Toddalioideae 352, 353 

Rutales 288 

Sageretia 

Sagittipetalum 

Sagitlipetalum mindanaensis 

Sagittipetalum palawanense 

Salacia 

Salacia bartletli 

306 

328 

329 

329 

109, 153 

274 

Samadera 

Samadera brevipetala 

Samadera indica 

Santalales 

436 

439 

439 

271 

Santalodes 189 

Santalodes diversifolium 290 

Santaloides 189 

Santiria 45,46,47, 76, 83, 84, 93 

Santiria sect. Icicopsis 84 

Santiria sect. Santiria 84 

Santiria apiculata 84, 85, 87, 89 

Santiria apiculata var. apiculata 86 

Santiria apiculata var. pilosa 86 

Santiria beccarii 86 

Santiria borneensis 

Santiria brachystachys 

Santiria caesia 

Santiria conferta 

Santiria conferta var. wrayi 

Santiria costata 

Santiria dacryodifolia 

Santiria fasciculata 

Santiria glabra 

Santiria glabrifolia 

Santiria grandiflora 

Santiria griffithii 

Santiria havilandii 

88 

86 

93 

85, 86 

86 

66 

85,87 

74 

86 

90 

84, 87 

83, 84, 85, 88, 94 

95 

Santiria impressinervis 85, 89 

Santiria kalkmaniana 85,89,91 

Santiria laevigata 84, 85, 90 

Santiria laevigata forma glabrifolia 90 

Santiria laevigata forma laevigata 90 

Santiria laevigata forma typica 90 

Santiria lat!folia 93 

Santiria laxa 69 

Santiria long!folia 69 

Santiria macrocarpa 71 

Santiria megaphylla XL VII, 85, 92 

Santiria minimijlora 94 

Santiria minutijlora 86 

Santiria mollis 84, 92 

Santiria mollissima 97 

Santiria montana 

Santiria multijlora 

Santiria nervosa 

Santiria nigricans 

Santiria nitida 

Santiria oblongifolia 

73 

97 

72 

85,93 

68 

84,85,93 

502

Santiria pedicel/ata 94 

Santiria pilosa 86 

Santiria planchonii 95 

Santiria rostrata 72 

Santiria rubiginosa 73, 83, 84, 85, 94 

Santiria rubiginosa var. IatipetioIata 94 

Santiria rubiginosa var. pedicellata 94 

Santiria rubiginosa var. rubiginosa 95 

Santiria rugosa 74 

Santiria samarensis 73 

Santiria sarawakana 84, 95 

Santiria serrulata 50 

Santiria tomentosa 84, 88, 95 

Santiria vio/acea 90 

Santiria virgata 47 

Santiria wrayi 86 

Sapindaceae 1,187,352,453,454 

SapindaIes 352 

Sapindus 376 

Sarawakodendron 107, 109, 153 

Sarawakodendron filamentosum 

XLVII, 151,153 

Sarcodiscus 246 

Sarcodiscus chloranthiformis 247 

Sarcotheca 287, 288, 289, 290 

Sarcotheca,.acuminata 290 

Sarcotheca diversifolia 289,290 

Sarcotheca gIauca 290, 291,293 

Sarcotheca macrophylla XLIX, 290, 292 

Sarcotheca oblongifolia 294 

Sarcotheca ochracea 290,294 

Sarcotheca rubrinervis L, 290, 294 

Sarcotheca subtriplinervis 290 

Saxifragaceae 454 

ScaevoIa 213,214 

ScaevoIa chanii 214, 215 

Scaevola frutescens 217 

Scaevola koenigii 217 

Scaevola leschenaultii 217 

Scaevola macrocalyx 

Scaevola merrillii 

Scaevola merrillii var. mollis 

. ScaevoIa micrantha 

ScaevoIa muIuensis 

Scaevola pedunculata 

Scaevola piliplena 

ScaevoIa pIumieri 

ScaevoIa sericea 

217 

216 

216 

L, 214, 216 

214,217 

216 

217 

214 

214,217 

Scaevola taccada 

ScaevoIa verticillata 

Schisandraceae 

SchismatogIottidinae 

Schuurmansia 

Schuurmansia angustifolia 

Schuurmansia borneensis 

Schuurmansia eIegans 

Schuurmansia parvijlora 

Schuurmansiella 

Schuurmansiella angustifolia 

Sciadicarpus 

Sciadicarpus brongniartii 

Sclerostylis lanceolata 

Sclerostylis spinosa 

Scorodocarpus 

Scorodocarpusborneensis 

ScrophuIariaceae 

Scutinanthe 

Scutinanthe brunnea 

Severinia 

217 

214,218 

227 

XXXVIII 

257, 258, 276, 

268 

268 

266,268 

268 

258,268 

XLIX, 268, 269 

246 

247 

374 

395 

274,278 

271, 280,281 

35 

45,46,47,97 

96,97,99 

352,353,354,356,357,410,411,412 

Severinia disticha 410, 411, 412, 413 

Severinia panicuIata 410,411,412 

Shorea acuta 

XL VII 

Shorea agamii subsp. diminuta L 

Shorea aIbida XLVI, XLVIII, 26, 225 

Shorea aImon 

L 

Shorea aIutacea 

XLIX 

Shorea asahi 

L 

Shorea bakoensis 

XLIX 

Shorea chanii 

L 

Shorea collaris 

L 

Shorea crassa 

XLVII 

Shorea curtisii XL VI, XL VII, XLIX 

Shorea cuspidata 

XLVII, XLIX 

Shorea dasyphylla 

XLIX 

Shorea deaIbata 

XLIX 

Shorea elliptica 

XLVII, XLIX 

Shorea falcifera XL VI, XL VII, XLIX 

Shorea falciferoides 

L 

Shorea flaviflora 

XLVIII 

Shorea flemmichii 

XLVII 

Shorea genicuIata 

XLVII 

Shorea iIiasii 

L 

Shorea induplicata 

XLIX 

Shorea kunstIeri 

XLVII 

503

Sonneratia evenia 450 

Sonneratia lanceolata 450 

Sonneratia neglecta 

450 

450 

450 

Sonneratia ovata 443,447,448,449,450 

Sonneratia obovata 

Sonneratia ovalis 

Sonneratia pagatpat 450 

SONNERA TlACEAE V, 443, 445, 446, 

504 

Tapiscia 

Tecoma stans 

Temminckia 

Temminckia micrantha 

Terminthodia 

Terminthodia viridiflora 

Tetracrypta 

Tetractomia 

Tetractomia beccarii 

Tetractomia holttumi 

Tetractomia latifolia 

Tetractomia montana 

Tetractomia obovata 

Tetractomia parviflora 

Tetractomia tetrandrum 

Tetradium 

Tetramelaceae 

284 

284 

284 

284 

284 

V,463 

414 

414 

414 

414 

414 

414 

384 

209 

414,415 

463,465 

463 

463 

9 

465 

328 

330 

463 

XIV, XIX 

XLVIII 

33 

188 

453 

33 

213 

216 

412 

414 

16 

283,284 

282,283 

33 

330 

453 

V, 453, 454 

274 

212 

101, 102 

102 

271,273,274,283 

421,440,442 

352,353,355,412 

421,422,441 

Strombosia latifolia 

Strombosia lucida 

Strombosia maingayi 

Strombosia muItiflora 

Strombosia rotundifolia 

Soulamea 

Soulamea amara 

Spathodea campanulata 

Stemonurus frutescens 

Stalagmites lamponga 

Sterculiaceae 

Stixis 

Stixis ovata 

Strombosia 

Strombosia ceylanica 

Strombosia javanica 

STAPHYLEACEAE 

Symplocos 

Syzygium 

Syzygium bankense 

Tabebuia rosea 

Symmetria 

Symmetria obovata 

STYRACACEAE 

Styrax javanicum 

Styrax officinalis 

Taeniochlaena 

Styrax 

Styrax agreste 

Styrax benzoin 

Staphylea 

Smythea 306, 307 

Siphonodon 107, 108, 109, 154 

Siphonodon celastrineus 152, 154 

Siphonodon pyriformis 154 

SIMAROUBACEAE V, 46, 352, 421, 422 

XLV, XLVI, XLVII, XLVIII, L 

Shorea resinosa XLIX 

Shorea richetia XLIX 

Shorea rubella XL VII 

Shorea slootenii XL VII 

Shorea splendida XLIX 

Shorea stenoptera XLIX 

Shorea subcylindrica XLIX 

Shorea superba XL VI 

Shorea symingtonii L 

Shorea waltonii L 

Solenospermum 130 

Solenospermum apiculatum 137 

Solenospermum aquatile 134 

Solenospermum javanicum 134 

Solenospermum littorale 127 

Solenospermum oblongifolium 134 

Solenospermum paIlidum 135 

Solenospermum paucinervium 134 

Solenostigma angustifolium 318 

XLVII 

XLV, XLVIII 

XLVII 

XLIX 

XLVI, XLVII 

XLVIII 

XLVII 

XLIX 

Sonneratia 443,445,447 

Sonneratia acida 450 

Sonneratia alba 449 

Sonneratia alba x Sonneratia caseolaris 447 

Sonneratia alba x Sonneratia ovata 447 

Sonneratia caseolaris 443, 447, 449, 450, 

Shorea ladiana 

Shorea laevis 

. Shorea laxa 

Shorea lunduensis 

Shorea macroptera 

Shorea multiflora 

Shorea ovata 

Shorea pallidifolia 

Shorea parvifolia

Tetrameles 

Tetramerista 

Tetrameristaceae 

Theaceae 

Toddalia 

Tricaphis 

209 

257 

257 

XLVII 

352,353,355 

454 

~~~ 4~ 

Trichocarya 169 

TRIGONIACEAE V, 467 

Trigoniastrum 467 

Trigoniastrum hypoleucum 467,468 

Trigonochlamys 83 

Trigonochlamys grandifolia 55 

Trigonochlamys grijjithii 88 

Triomma 45, 46, 99 

Triomma malaccensis 98, 100 

Triphasia 352, 351, 353 

Triplochiton scleroxylon 212 

Turpinia 453, 454, 455 

Turpinia borneensis L, 455, 456 

Turpinia calciphila 455, 456, 457 

Turpinia grandis 455, 456, 457 

Turpinia latifolia 460,461 

Turpinia laxiflora 460,461 

Turpinia montana var. borneensis 456 

Turpinia nepalensis 455 

Turpinia nitida L, 454, 455, 456, 459 

Turpinia sphaerocarpa 455, 456, 460, 462 

Turpinia sphaerocarpa var. microcerotis 

458,461 

Turpinia sphaerocarpa var. sphaerocarpa 

461 

Turpinia stipulacea 

U mbelliferae 

Upuna borneensis 

Vatica 

Vatica endertii 

Vatica compressa 

Vatica dulitensis 

Vatica granulata 

454,455,456,461 

476 

XLIX 

201 

L 

XLIX 

XLVIII 

L 

Vatica pedicellata 

XLIX 

Vatica umbonata 

XLVIII 

Ventilago 

306, 307 

Verbenaceae 

35 

Viburnum 

113 

Vitex premnoides 

208 

Weinmannia blumei 

XLIX 

Wenzelia 

402 

Winteraceae 227 

Xanthomyrtus spp. 

XLIX 

Xanthophyllum 154, 171,469 

Xanthophyllum subglobosum 154 

Ximenia 273 

Ximenia americana 271,273 

Ximenia borneensis 281 

Zanthoxylon aromaticum 392 

Zanthoxylum 352, 353, 354, 355, 416 

Zanthoxylum avicennae 

Zanthoxylum diversifolium 

Zanthoxylum iwahigense 

Zanthoxylum macrophyllum 

Zanthoxylum myriacanthum 

Zanthoxylum nitidum 

Zanthoxylum roxburghianum 

Zanthoxylum scandens 

Zanthoxylum triphyllum 

Ziziphus 

Ziziphus angustifolius 

Ziziphus borneensis 

Ziziphus calophylla 

Ziziphus crebrivenosa 

Ziziphus cumingiana 

Ziziphus forbesii 

Ziziphus grewioides 

Ziziphus havilandii 

Ziziphus horsfieldii 

Ziziphus inermis 

Ziziphus jujuba 

Ziziphus mauritiana 

Ziziphus suluensis 

416,417 

417 

417 

386 

416,418,419 

417 

392 

417 

394 

306,314, 

316,317,318 

317 

318 

318 

317 

318 

318 

317 

317 

318 

314 

314 

318 

505

INDEX TO VERNACULAR NAMES 

(compiled by Rusea Go) 

Scientific names in brackets are the correct names corresponding to vernacular names listed here. 

aam (Gomphia serrata (Gaertn.) Kanis) 

adas china (Illicium verum Hook. f) 

akar malam (Agelaea macrophylla (Zo11.) Leenh.) 

ala (Illicium stapjii Merr.) 

alan (Shorea albida Symington) 

ambibiliw (Kibara coriacea (Blume) Tulasne) 

ambun ambun (Anisophyllea disticha (Jack) Baill.) 

antimagas gimbaan (Gomphia serrata (Gaertn.) Kanis) 

ara bukit (Pittosporum ferrugineum Aiton) 

aremajuh (Sarcotheca glauca (Hook. f) Hallier f) 

asam-asam (Dacryodes macrocarpa (King) H.J. Lam) 

asam daham (Sarcotheca glauca (Hook. f) Ha11ier f) 

asam piai (Sarcotheca glauca (Hook. f) Hallier f) 

asem-asem (Sarcotheca rubrinervis Hallier f) 

atap (Trigoniastrum hypoleucum Miq.) 

bajan (Kokoona spp.) 

bajan paya (Kokoona ovatolanceolata Rid!.) 

bakau (Rhizophora spp.) 

bakau aleh aleh (Kandelia candel (L.) Dmce) 

bakau kurap (Rhizophora mucronata Lam.) 

bakau lali (Ceriops decandra (Griff.) Ding Hou) 

bakau minyak (Rhizophora apiculata Blume) 

balak bekatan (Combretocarpus rotundatus (Miq.) Danser) 

balik angin (Alphitonia exelsa (Fenzl) Reiss ex End!,) 

balimbing (Picrasmajavanica Blume) 

bangkawang (Maranthes corymbosa Blume) 

bangkita (Rhizophora apiculata Blume) 

bantis (Mastixia trichotoma Blume) 

barat-barat (Cassine viburnifolia (Juss.) Ding Hou) 

bawang hutan (Scorodocarpus borneensis (Baill.) Becc.) 

bedara (Eurycoma longifolia Jack) 

bedung (Pittosporum resiniferum Hems!.) 

belian landak (Anisophyllea ferruginea Ding Hou) 

belimbing (Averrhoa spp.) 

belimbing bulat (Sarcotheca diversifolia (Miq.) Hallier f) 

belimbing manik (Sarcotheca macrophylla Blume) 

benuang (Octomeles sumatrana Miq.) 

benung kasung (Duabanga moluccana Blume) 

bems bems (Kandelia candel (L.) Dmce) 

bems kurong (Bruguiera gymnorrhiza (L.) Lam.) 

bems lenggadai (Bruguiera parviflora (Roxb.) Wight & Arn. ex Griff.) 

bems merah (Bruguiera gymnorrhiza (L.) Lam.) 

267 

229 

187 

230 

XLVI, 26, 265 

247 

21 

267 

300 

292 

71 

292 

292 

295 

469 

122 

128 

345 

341 

348 

335 

348 

26 

308 

436 

168 

348 

208 

114 

281 

433 

303 

21, 275 

289 

291 

292 

210 

446 

341 

325 

326 

325 

506

ems ngayong (Bruguiera cylindrica (L.) Blume) 

bems puteh (Bruguiera cylindrica (L.) Blume) 

bems putut (Bruguiera cylindrica (L.) Blume) 

beus (Bruguiera spp; Kandelia candeI (L.) Dmce) 

biansu gunong (Mastixia cuspidata Blume) 

biku-biku (Bhesa paniculata Am.) 

bindang (Agathis borneensis Warb.) 

binkuli (Oroxylum indicum (L.) Vent.) 

binong (Octomeles sumatrana Miq.) 

binuang (Octomeles sumatrana Miq.) 

binutan (Radermachera pinnata (Blanco) Merr.) 

bogua (Harrisonia perforata (Blanco) Seemen) 

bowlong (Illicium stapjii Men.) 

buah piang (Sarcotheca diversifolia (Miq.) Hallierf) 

bulok (Agathis endertii Meijer Drees & Agathis orbicula de Laub.) 

bulu (Agathis borneensis Warb.) 

bulu bulu (Gynotroches axilIaris Blume) 

bunga lawang (Illicium verum Hook. f) 

butun laut (Scaevola sericea Vahl) 

chinaga-lampong (Gomphia serrata (Gaertn.) Kanis) 

dadam (Alangium havilandii Bloemb.) 

dajak (AnisophyIIea ferruginea Ding Hou) 

danguh (PeIIacalyx axilIaris Korth.) 

dat (Cratoxylum arborescens (Vahl) Blume) 

dat tetong (Cratoxylumformosanum (Jack) Dyer) 

daun kari (Murraya koenigii (L.) Spreng.) 

derum (Cratoxylum spp.) 

di'it (Cratoxylum arborescens (Vahl) Blume) 

empudu (Bruinsmia styracoides Boerl. & Koord.) 

engkop engkop (AnisophyIIea nitida Madani) 

entalun (Engelhardia serrata Blume) 

entemu (Cratoxylumformosanum (Jack) Dyer) 

entrang (Turpinia sphaerocarpa Hassk.) 

gadong hutan (Alangium griffithii (Clarke) Harms) 

geroking (Cratoxylum arborescens (Vahl) Blume) 

geronggan lompong (Cratoxylum glaucum Korth.) 

geronggang (Cratoxylum spp.) 

geronggang gajah (Cratoxylum arborescens (Vahl) Blume) 

geronggang puteh (Cratoxylum gIaucum Korth.) 

geronggang timau (Cratoxylum gIaucum Korth.) 

gerunggang (Cratoxylum maingayi Dyer) 

gilas (Parastemon urophyIIus (Wall. ex A.DC.) A.DC.) 

gimurai (Oroxylum indicum (L.) Vent.) 

gitan gizu (Sarcotheca glauca (Hook. f) Hallier f) 

ibajantan (Sarcotheca diversifolia (Miq.) Hallierf) 

iba talon (Sarcotheca rubrinervis Hallier f) 

icerawas bumng (Dacryodes macrocarpa (King) H.J. Lam var. macrocarpa) 

idat (Cratoxylum arborescens (Vahl) Blume) 

507 

324 

324 

326 

323,324,326,341 

202 

112 

29 

41 

210 

210,212 

42 

422 

230 

291 

30, 32 

29 

339 

229 

218 

267 

9 

21 

344 

223 

224 

406 

219,220 

223 

465 

23 

243 

219,224 

461 

8 

223 

225 

219,220,223,224,225 

223 

225 

225 

226 

172 

41 

292 

291 

295 

71 

223

ikor mata (Sarcotheca ochracea Hallier f) 

imah (Ochanostachys amantacea Mast.) 

ionadiandau (Eurycoma longifolia Jack) 

ira prumpuan (Sarcotheca rubrinervis Hallier f) 

itan beruang (Mastixia trichotoma Blume) 

jadam (Alangium javanicum (Blume) Wangerin) 

jadam paya (Alangium havilandii Bloemb.) 

jaloot (Bruceajavanica (L.) Merr.) 

jambul merah (Jacaranda rhombifolia G.F.W. Meijer) 

jampang (Tetractomia tetrandrum (Roxb.) Merr.) 

jampang rusa (Tetractomia tetrandrum (Roxb.) Merr.) 

jenangan (Alangium havilandii Bloemb.) 

jerit (Anacolosa frutescens (Blume) Blume) 

jiwang (Sarcotheca diversifolia (Miq.) Hallierf) 

kacang-kacang (Quassia indica (Gaertn.) Noot.) 

kait-kait (Harrisonia perforata (Blanco) Merr.) 

kajo jelan (Cratoxylum formosanum (Jack) Dyer) 

kambayau (Dacryodes rostrata (Blume) H.J. Lam f. rostrata) 

kandis batu (Gynotroches axil/aris Blume) 

kandis daham (Sarcotheca glauca (Hook. f) Hallier f) 

kata mudung (Cratoxylum arborescens (Vahl) Blume) 

kayu ajung (Parastemon grandifructus Prance) 

kayujadi (Agathis borneensis Warb.) 

kayu labu (Ziziphus angustifolius (Miq.) Hatusima ex Steenis) 

kayu pahit (Picrasmajavanica Blume) 

kayu runap (Anisophyllea disticha (Jack) Baill.) 

kayu tom (Combretocarpus rotundatus (Miq.) Danser) 

kayu tulang (Allanthospermum borneensis Forman) 

kayu wulu (Mastixia pentandra Blume) 

kebuan (Crateva magna (Lour.) DC.) 

kedondong (Canarium spp.; Dacryodes spp.) 

kedondong asam (Triomma malaccensis Hook. f) 

keladang (Gomphia serrata (Gaertn.) Kanis) 

kelaju (Dolichandrone spathacea (L.f) K. Schum.) 

kelalud (Gynotroches axil/aris Blume) 

kelana (Ellipanthus beccarii Pierre var. peltatus (Schellenb.) Leenh.) 

kelapahit (Picrasma javanica Blume; Quassia indica (Gaertn.) Noot.) 

kelin (Ellipanthus tomentosus Kurz) 

kelutak (Gomphia serrata (Gaertn.) Kanis) 

kembayau (Dacryodes spp.; Dacryodes rostrata (Blume) H.J. Lam) 

kemenyan (Stryrax benzoin Dryand.) 

kemuning (Murraya paniculata (L.) Jack) 

kemuning hutan (Carallia borneensis Oliver) 

kenari (Canarium vulgare Leenh.) 

kepajang

keramoh (Daryodes rostrata f. cuspidata (Blume) H.J. Lam) 

kerantai (Santiria spp.) 

kerapa-kerapa (Sarcotheca diversifolia (Miq.) Hallierf) 

keruas (Dacryodes macrocarpa (King) H.J. Lam var. macrocarpa) 

keruntum (Combretocarpus rotundifolius (Miq.) Danser) 

kimayau (Dacryodes spp.) 

kiraV (Cratoxylum glaucum Korth.) 

kolarnbang (Gomphia serrata (Gaertn.) Kanis) 

kolintuhan (Clethra spp.) 

kondolon (Alangium spp.) 

kuinin (Bruceajavanica (L.) Merr.) 

kujuk langit (Radermachera pinnata (Blanco) Seemen) 

kukualang (Harrisonia perforata (Blanco) Merr.) 

kupi kupi (Gynotroches axillaris Blume) 

kurong (Bruguiera gymnorrhiza (L.) Lam.) 

laba (Turpinia calciphila J.T. Pereira) 

labak (Kibara coriacea (Blume) Tulasne) 

labakan (Cratoxylum arborescens (Vahl) Blume) 

ladin (Gomphia serrata (Gaertn.) Kanis) 

ladit (Anacolosa frutescens (Blume) Blume) 

lampyos (Sarcotheca rubrinervis Hallier f) 

langarai (Bruguiera parviflora (Roxb.) Wight & Am. ex Griff.) 

lanjing lanjing (Bruguiera parviflora (Roxb.) Wight & Am. ex Griff.) 

lemeng (Octomeles sumatrana Miq.) 

lenggadai (Bruguiera spp.) 

lia (Trigoniastrum hypoleucum Miq.) 

limau besar (Citrus grandis (L.) Osbeck) 

limau buaya (Merope angulata (Willd.) Swingle) 

limau hantu (Burkillanthus malaccensis (Rid!.) Swingle; 

Citrus macroptera Montr.) 

limau hutan (Citrus halimii Stone; Citrus macroptera Montr.) 

limau kesturi (Citrus microcarpa Bunge) 

limau laut (Merope angulata (Willd.) Swingle) 

limau limau (Pleiospermium longisepalum Swingle) 

limau manis (Citrus reticulata Blanco; Citrus sinensis (L.) Osbeck) 

limau mata kerbau (Citrus limon (L.) Burm.f) 

limau nipis (Citrus aurantifolia (Christm. & Panz.) Swingle) 

limau purut (Citrus hystrix DC.) 

limau· susu (Citrus medica L.) 

limo antu (Pleiospermium latialatum Swingle) 

limo bali (Pleipspermium latialatum Swingle) 

limo to' (Pleiospermium latialatum Swingle) 

linggayong (Kandelia candel (L) Druce) 

linggayong laut (Kandelia candel (L) Druce) 

longugan (Illicium kinabaluensis A.C.Sm.) 

maba (Turpinia sphaerocarpa Hassk.) 

machit laling (Cratoxylum cochinchinense (Lour.) Blume) 

madak (Cratoxylum arborescens (Vahl) Blume) 

65, 73 

83 

291 

71 

26 

65 

225 

267 

181 

5 

431 

42 

422 

339 

325 

457 

247 

223 

267 

275 

295 

326 

326 

210 

323, 326 

469 

364 

396 

362, 366 

366,410 

364 

396 

410 

364, 365 

364 

365 

365 

363 

409 

409 

409 

341 

341 

230 

461 

224 

223 

509

magas (Duabanga moluccana Blume) 

majang-majang (Gomphia serrata (Gaertn.) Kanis) 

mamungal (Quassia borneensis Noot.) 

manat (Cratoxylum arborescens (Vahl) Blume) 

mandailas (Parastemon grandifructus Prance; 

Parastemon urophyllus (Wall. ex ADC.) A DC.) 

manggilan (Agathis borneensis Warb.) 

manuggal (Quassia borneensis Noot.; Quassia indica (Gaertn.) Noot.) 

mara (Bruceajavanica (L.) Merr.) 

marapangi (Alangium kurzii Craib; Alangium rotundifolium (Hassk.) Bloemb.) 

mata buaya (Bruguiera sexangula (Lour.) Poir) 

mata ulat (}:.okoona ~pp.) 

mayam kampong (Alangium grifJithii (Clarke) Harms) 

medang (Lauraceae species; Sarcotheca glauca (Hook. f) Hallier f) 

medang kanigara (Mastixia trichotoma Blume) 

medang pahit (Quassia borneensis Noot.) 

medang rawang (Tetratomia tetrandrum (Roxb.) Men.) 

medang surungan (Mastixia pentandra Blume) 

melan (Cratoxylumformosanum (Jack) Dyer) 

mempulak (Crateva magna (Lour.) DC.) 

mendailas (Parastemon grandifructus Prance) 

menengang (Anisophyllea spp.) 

mengilas (Parastemon urophyllus (Wall. ex ADC.) ADC.) 

mengilas babi (Trigoniastrum hypoleucum Miq.) 

mengkudu (lrvingia malayana Oliver ex AW. Benn.; Morinda spp.) 

meransi (Carallia sp. 1) 

merawai (Carallia sp. 1) 

merbatu (Atuna spp.; Kostermanthus heteropetalus (Scort. ex King) Prance; 

446 

246 

439 

223 

171 

29 

441 

431 

12, 14 

326 

122 

8 

201,292 

208 

439 

414 

204 

224 

104 

172 

16 

172 

469 

434 

332 

332 

Maranthes corymbosa Blume; Parinari spp.) 157, 165, 168, 172 

merbuloh (Pellacalyx spp.) 341 

merinang (Ellipanthus beccarii Pierre var. peltatus (Scheenb.) Leenh.) 196 

mertama (Anisophyllea corneri Ding Hou; Anisophyl/ea disticha (Jack) Baill.) 16, 19,21 

mertilan (Cratoxylum arborescens (Vahl) Blume; 

Cratoxylum cochinchinense (Lour.) Blume) 

miapa (Trigoniastrum hypoleucum Miq.) 

midong (Alangium spp.) 

mirinos (Cratoxylumformosanum (Jack) Dyer) 

momon (Engelhardia serrata Blume) 

mopu (Anisophyllea spp.) 

murai (Oroxylum indicum (L.) Vent.) 

mutun (Combretocarpus rotundatus (Miq.) Danser) 

ngali-nut (Canarium indicum L.) 

ngayong (Bruguiera cylindrica (L.) Blume) 

ngilas (Parastemon grandifructus Prance) 

ngilis (Trigoniastrum hypoleucum Miq.) 

nonok (Pittosporum ferrugineum Aiton) 

nuad-mandau (Eurycoma longifolia Jack) 

nyalin (Allantospermum borneense Forman; Trigoniastrum hypoleucum Miq.) 

510 

223,224 

469 

5 

224 

243 

16 

41 

26 

46 

324 

171 

469 

300 

433 

429,469

nyalin bahe (Cratoxylumformosanum (Jack) Dyer) 

nyal in bintek (Trigoniastrum hypoleucum Miq.) 

obar-obar (Colubrina beccariana Warb.) 

odok-odok (Colubrina beccariana Warb.) 

owl (Engelhardia serrata Blume) 

pahau pahau (Gynotroches axillaris Blume) 

pait-pait (Bruceajavanica (L.) Merr.; Quassia borneensis Noot.) 

pam.atudon (Canarium decumanum Gaertn.) 

pangos (Crateva magna (Lour.) DC.) 

panguban (Picrasmajavanica Blume) 

para bileh (Pleiospermium latialatum Swingle) 

parang nyabor (Oroxylum indicum (L.) Vent.) 

paraIlgpamol (Oroxylum indicum (L.) Vent.) 

paserujan gunugo (Pellacalyx lobbii (Hook f) Schimp.) 

pasil-pasil (Ziziphus angustifolius (Miq.) Hatusima ex Steenis) 

pasob-pasoh (Dacryodes macrocarpa (King) HJ. Lam) 

pati yata (Alphitonia exelsa (Fenzl) Reiss ex End!,) 

patok entilit (Mastixia rostrata Blume) 

patok tilan (Cratoxylumformosanum (Jack) Dyer; Cratoxylum maingayi Dyer) 

patoli entelit (Jrvingia malayana Oliver ex. A.W. Benn.) 

pauh kijang (lrvingia malayana Oliver ex. A.W. Benn.) 

paya.5 (Bruceajavanica (L.) Blume) 

pechi mata (Sarcotheca ochracea Hallier f) 

pedarla (Sonneratia alba 1. Smith; Sonneratia caseolaris (L.) Engl.) 

pei (Anisophyllea beccariana Baill.) 

perapan macas (Sarcotheca diversifolia (Miq.) Hallierf) 

. perdu (Aeer laurinum Hassk) 

perepat (Sonneratia alba 1. Smith; Sonneratia caseolaris (L.) Engl.) 

perepat hutan (Combretocarpus rotundatus (Miq.) Danser) 

perepat paya (Combretocarpus rotundatus (Miq.) Danser) 

perepat perepat (Combretocarpus rotundatus (Miq.) Danser) 

perupok (Lophopetalum spp.) 

perupok kuning (Kokoona ochracea (Elmer) Men.) 

petaling (Ochanostachys amentacea Mast.) 

piang (Sarcotheca spp.; Sarcotheca glauca (Hook. f) Hallierf) 

pidang (Cratoxylum glaucum Korth.) 

pinggoh (Sarcotheca rubrinervis Hallierf) 

pinguh (Sarcotheca rubrinervis Hallier f) 

pokudata (Alphitonia excelsa (Fenzl) Reiss ex Endl.) 

priabu (Mastixia trichotoma Blume) 

pusing-pusing (Engelhardia serrata Blume) 

putut (Bruguiera gymnorrhiza (L.) Lam.; Bruguiera sexangula (Lour.) Poir) 

quintalai (Gomphia serrata (Gaertn.) Kanis) 

rabong (Caralfia brachiata (Lour.) Men.) 

radipah (Carallia brachiata (Lour.) Men.) 

raja tugag (Cratoxylumformosanum (Jack) Dyer) 

rangkas-rangkas (Sarcotheca glauca (Hook f) Hallier f) 

rasak batu (Kostermanthus heteropetalus (Scot. ex King) Prance) 

224 

469 

312 

312 

243 

339 

431 

53 

104 

436 

409 

41 

41 

345 

319 

71 

308 

434 

224,226 

205 

433,434 

431 

294 

449,450 

18 

291 

3 

449,450 

26 

26 

26 

129 

128 

278 

289,292 

225 

295 

295 

308 

208 

243 

325, 326 

267 

330,333 

330 . 

224 

292 

165 

511

asu (Pellacalyx lobbii (Hook. f) Schimp.) 

rawang (Maclurodendron porteri (Hook. f) T.G. Hartley) 

rawang mata (Tetractomia tetrandrum (Roxb.) Merr.) 

rawang paya (Tetractomia tetrandrum (Roxb.) Merr.) 

rogam (Sonneratia ovata Backer) 

sabal (Dacryodes macrocarpa (King) H.J. Lam var. patentinervia Leenh.) 

sabutun (Combretocarpus rotundatus (Miq.) Danser) 

sagad-berauh (Ochanostachys amentacea Mast.) 

sagan-berauh (Scorodocarpus borneensis (Baill.) Becc.) 

saipang (Pi ttosporum ferrugineum Aiton) 

salang (Agathis borneensis Warb.) 

salangugapit (Anacolosafrutescens (Blume) Blume) 

salong banggi (Dacyodes rostrata var. cuspidata (Blume)H.J. Lam) 

sambar bubu (Gynotroches axil/aris Blume; Pellacalyx spp.) 

sampaluan (Licania 'splendens (Korth.) Prance) 

sangkuak (Anisophyllea ferruginea Ding Hou) 

sansanglang (Engelhardia spp.) 

santikal (Ochanostachys amentacea Mast.) 

sasgah (Crateva magna (Lour.) DC.) 

satu inchi (Alangiumjavanicum (Blume) Wangerin) 

sawar bubu (Pellacalyx spp.) 

sawih (Duabanga moluccana Blume) 

sebelu (Grateva magna (Lour.) DC.) 

segera (Glycosmis chlorosperma (Blume) Spreng.) 

seladah (Dacryodes spp.; Dacryodes macrocarpa (King) H.J. Lam 

var. patentinervia Leenh.; Santiria spp.) 

selangan biabas (Cratoxylum cochinchinense (Lour.) Blume) 

selangan tandok (Irvingia malayana Oliver ex A.W. Benn.) 

sempetan (Mastixia eugenioides Matthew) 

sengkanyat (Eurycoma longifolia Jack) 

sengkayap (Eurycoma longifolia Jack) 

serang (Crateva magna (Lour.) DC.; Melicope spp.) 

seraya (Shorea curtisii Dyer ex King) 

serunai (Bhesa paniculata Arn.) 

serungan (Cratoxylum arborescens (Vahl) Blume; 

Cratoxylum cochinchinense (Lour.) Blume; 

Cratoxylum sumatranum (Jack) Blume) 

serungan batu (Cratoxylum spp.) 

serungan labakan (Cratoxylum arborescens (Vahl) Blume) 

serungan mampat (Cratoxylum sumatranum (Jack) Blume) 

sibut (Dacryodes macrocarpa (King) HJ. Lam var. patentinervia Leenh.) 

sidodot (Cratoxylum formosanum (Jack) Dyer) 

simun (Bhesa paniculata Am.) 

sindok (Scorodocarpus borneensis (Baill.) Becc.) 

sindu (Scorodocarpus borneensis (Baill.) Becc.) 

sireh sireh (Anisophyllea corneri Ding Hou) 

si sit (Alangium havilandii Bloemb.) 

soipang (Pittosporum ferrugineum Aiton) 

512 

345 

382 

414 

414 

451 

71 

26 

278 

281 

300 

29 

275 

73 

341 

167 

21 

233 

278 

104 

9 

339, 341 

446 

104 

373 

65, 71, 73, 83 

224 

434 

202 

433 

433 

104,384 

XLVI 

112 

219,223,224,226 

219 

223 

226 

71 

224 

112 

281 

281 

16, 19 

9 

300

t-izo:s-izo~ (Sarcotheca glauca (Gaertn.) Kanis) 

tabarus (S.arcotheca spp.; Sarcotheca rubrinervis Hallier f) 

tabaus (Sa.rcotheca diversifolia (Miq.) Hallier f) 

tagal (Ceriops tagal (Pers.) C.B. Rob.) 

taikakang (Cratoxylum cochinchinense (Lour.) Blume) 

tait berak (Turpinia sphaerocarpa Hassk.) 

tampaluan (Licania splendens (Korth.) Prance) 

tangal (Ceriops tagal (Pers.) C.B. Rob.) 

tanggal (Ochanostachys amentacea Mast.) 

tangilan (Agathis lenticula de Laub.) 

tansanglang (Engelhardia spp.) 

tanug (Ceriops tagal (Pers.) C.B. Rob.) 

tapang (Koompassia malaccensis Maingay ex Benth.) 

tapong-tapong (Turpinia sphaerocarpa Hassk.) 

tat (Cratoxylum arborescens (Vahl) Blume) 

tebarus (Sarcotheca diversifolia (Miq.) Hallierf) 

tengar (Ceriops decandra (Griff.) Ding Hou; Ceriops tagal (Pers.) c.B. Rob.) 

tengar samak (Ceriops tagal (Pers.) c.B. Rob.) 

tengar tikus (Ceriops decandra (Griff.) Ding Hou) 

tengnilan (Jrvingia malayana Oliver ex A.W. Benn.) 

tengoda (A nisaphyllea ferruginea Ding Hou) 

tenug (Ceriops tagal (Pers.) C.B. Rob.) 

tepanga (Engelhardia serrata Blume) 

teran~ bulu (Nyssajavanica (Blume) Wangerin) 

tigarun (Crateva magna (Lour.) DC.) 

tikolod (Carallia sp. 1) 

timau. (Cratoxylum glaucum Korth.) 

tingo-tingo (Bruinsmia styracoides Boer!. & Koord.) 

tombuid (Eurycoma longifolia Jack) 

tongkat ali (Bruceajavanica (L.) Merr.;Eurycoma longifolia Jack) 

tongkat iangit (Eurycoma longifolia Jack) 

trodub (Scorodocarpus borneensis (BaiII.) Becc.) 

tubu

11frE 111 ° E 112° E I 

5° N] I 

4°N - 

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,0 

00 

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) South China Sea .~ I 

SABAH ~'\:jtz,~ , 

(MALAYSIA) A -(ja 

PENINSULAR oP ~ , 

MALAYSIA • ':> l-t----+---I----+---l--I ' 

SARAWAK 

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\\ BORNEO 

-- Java Sea U ~ 

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1----+-----1-- SOUTH C:~IINA SEA 

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Mt. S;aoh (.,. t ,,'" KALlMAN' 

1° N I - I "'- ...... -+-+++ 

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110° E 

~ !l Mt. Apeng 

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I 

112° E 

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114° E 115° E ~" ........ j HRUNEif 

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) . Jr _ r 

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V x , 

L·- I L.. " 

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;:: ~~~~~==~-----t---~'tJ--~ 1- 

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NDONESIA) 

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113° E 114° E 115° E 

I 10 N

Tree Flora of Sabah and Sarawak, Volume I - ITTO

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