An Illustrated Guide to Diversity - Trees of Tropical Asia LaFrankie

TREES OF TROPICAL ASIA 

An Illustrated Guide 

to Diversity 

James V. LaFrankie, Jr. 

Comprising an Annotated Checklist 

of 157 Families and 880 Genera of 

Trees Native to the Region 

With Notes on their Ecology and Field Recognition 

Arranged in Accordance with the New Phylogeny 

With Illustrations by the Author 

Supplemented with Photographs by 

Other Regional Biologists 

BLACK TREE PUBLICATIONS, INC. 

P H I L I P P I N E S

10 

Preface ....................................................................................................................................................4 

Acknowledgments .........................................................................................................................................9 

THEORY 

Introduction To The Essay ...........................................................................................................................16 

Synopsis of The Essay ..................................................................................................................................17 

The Names of Trees .....................................................................................................................................17 

The Common Language of Names .............................................................................................17 

The Linnaean Revolution ...........................................................................................................18 

From Linnaeus To Cronquist .....................................................................................................19 

The New Phylogeny: A Revolution in Methods and Goals .........................................................21 

Addendum I: On Common Names in Tropical Asia ....................................................................................23 

Addendum II: The International Code of Botanical Nomenclature .............................................................25 

Addendum III: On Species ..........................................................................................................................29 

Addendum IV: Definitions ..........................................................................................................................31 

Addendum V: Cladistic Analysis .................................................................................................................32 

Addendum VI: Polyphyletic & Paraphyletic ...............................................................................................33 

Addendum VII: Parallel Nomenclatures of the 21st Century .......................................................................34 

PRACTICE 

CONTENTS 

The Diversity of Trees: Where Are We? ........................................................................................................39 

Getting Started: Recognition Versus Analysis...............................................................................................40 

Getting Started: The Problem ......................................................................................................................41 

Leaf Stalk, Node and Field Characters of the Main Families ........................................................................42 

Exudate ..................................................................................................................................................43 

Odor ..................................................................................................................................................44 

On Notebooks and Photographs .................................................................................................................45 

SURVEY 

Terms of Regional Geography .....................................................................................................................52 

Sources ..................................................................................................................................................57 

PTERIDOPHYTES 

CYATHEACEAE ........................................................................................................................................59 

POLYPODIACEAE ....................................................................................................................................60 

GYMNOSPERMS 

CYCADACEAE ..........................................................................................................................................62 

TAXACEAE ................................................................................................................................................63 

CUPRESSACEAE ......................................................................................................................................64 

ARAUCARIACEAE ...................................................................................................................................65 

PODOCARPACEAE ..................................................................................................................................67 

GNETACEAE ............................................................................................................................................71 

PINACEAE.................................................................................................................................................72

MAGNOLIIDS 

AUSTROBAILEYALES 

ILLICIACEAE ..........................................................................................................................75 

CHLORANTHALES 

CHLORANTHACEAE ............................................................................................................76 

MAGNOLIALES 

MYRISTICACEAE ...................................................................................................................79 

MAGNOLIACEAE ..................................................................................................................88 

HIMANTANDRACEAE ..........................................................................................................90 

ANNONACEAE ......................................................................................................................91 

LAURALES 

MONIMIACEAE ...................................................................................................................113 

HERNANDIACEAE ..............................................................................................................115 

LAURACEAE .........................................................................................................................116 

CANELLALES 

WINTERACEAE ...................................................................................................................129 

PIPERALES 

ARISTOLOCHIACEAE ........................................................................................................130 

PIPERACEAE .........................................................................................................................131 

MONOCOTS 

PANDANALES 

PANDANACEAE ...................................................................................................................132 

ASPARAGALES 

RUSCACEAE .........................................................................................................................134 

ARECALES 

ARECACEAE .........................................................................................................................136 

BASAL TRICOLPATES 

RANUNCULALES 

MENISPERMACEAE ............................................................................................................147 

BERBERIDACEAE ................................................................................................................148 

SABIALES 

SABIACEAE ...........................................................................................................................149 

PROTEALES 

PLATANACEAE ....................................................................................................................150 

PROTEACEAE ......................................................................................................................151 

BUXALES 

BUXACEAE ...........................................................................................................................153 

DILLENIALES 

DILLENIACEAE ...................................................................................................................154 

OTHER ROSIDS 

VITALES 

LEEACEAE ............................................................................................................................158 

CARYOPHYLLALES 

PLUMBAGINACEAE ............................................................................................................161 

POLYGONACEAE ................................................................................................................161 

NYCTAGINACEAE ...............................................................................................................162 

SANTALALES 

OLACACEAE ........................................................................................................................164 

SCHOEPFIACEAE ................................................................................................................169 

OPILIACEAE .........................................................................................................................169 

SANTALACEAE ....................................................................................................................171 

SAXIFRAGALES 

HAMAMELIDACEAE ...........................................................................................................172 

11

12 

ROSIDS 

MYRTALES 

COMBRETACEAE ................................................................................................................178 

LYTHRACEAE ......................................................................................................................182 

MYRTACEAE ........................................................................................................................186 

MEMECYLACEAE ................................................................................................................196 

MELASTOMATACEAE.........................................................................................................199 

CRYPTERONIACEAE ..........................................................................................................206 

CROSSOSOMATALES 

STAPHYLEACEAE ................................................................................................................207 

UNPLACED 

BISCHOFIACEAE .................................................................................................................208 

ROSIDS (FABIDS) 

CELASTRALES 

CELASTRACEAE ..................................................................................................................209 

MALPIGHIALES 

ACHARIACEAE ....................................................................................................................219 

SAMYDACEAE ......................................................................................................................224 

SCYPHOSTEGIACEAE ........................................................................................................226 

SALICACEAE ........................................................................................................................227 

VIOLACEAE ..........................................................................................................................232 

PASSIFLORACEAE ...............................................................................................................233 

POTTINGERIACEAE ...........................................................................................................234 

PUTRANJIVACEAE ..............................................................................................................235 

PANDACEAE ........................................................................................................................239 

PICRODENDRACEAE .........................................................................................................240 

PHYLLANTHACEAE............................................................................................................241 

PERACEAE ............................................................................................................................256 

EUPHORBIACEAE ...............................................................................................................258 

OCHNACEAE .......................................................................................................................291 

IRVINGIACEAE ....................................................................................................................294 

IXONANTHACEAE..............................................................................................................295 

CENTROPLACACEAE .........................................................................................................297 

DICHAPETALACEAE ...........................................................................................................299 

TRIGONIACEAE ..................................................................................................................299 

CHRYSOBALANACEAE.......................................................................................................300 

CTENOLOPHONACEAE ....................................................................................................306 

ERYTHROXYLACEAE ..........................................................................................................307 

RHIZOPHORACEAE ...........................................................................................................308 

CLUSIACEAE ........................................................................................................................315 

BONNETIACEAE .................................................................................................................319 

CALOPHYLLACEAE ............................................................................................................320 

HYPERICACEAE ..................................................................................................................326 

OXALIDALES 

CONNARACEAE ..................................................................................................................328 

OXALIDACEAE ....................................................................................................................330 

CUNONIACEAE ...................................................................................................................332 

ELAEOCARPACEAE .............................................................................................................333 

FABALES 

POLYGALACEAE ..................................................................................................................337 

FABACEAE ............................................................................................................................341 

ROSALES 

ROSACEAE ............................................................................................................................376 

ELAEAGNACEAE .................................................................................................................379 

RHAMNACEAE ....................................................................................................................380 

ULMACEAE ..........................................................................................................................384 

CANNABACEAE ...................................................................................................................385 

MORACEAE ..........................................................................................................................389 

URTICACEAE ......................................................................................................................403

CUCURBITALES 

ANISOPHYLLEACEAE .........................................................................................................406 

TETRAMELACEAE ..............................................................................................................409 

FAGALES 

FAGACEAE ............................................................................................................................412 

MYRICACEAE ......................................................................................................................418 

JUGLANDACEAE .................................................................................................................419 

BETULACEAE.......................................................................................................................420 

CASUARINACEAE ...............................................................................................................421 

ROSIDS (MALVIDS) 

HUERTALES 

DIPENTODONTACEAE ......................................................................................................423 

TAPISCIACEAE .....................................................................................................................423 

BRASSICALES 

MORINGACEAE ..................................................................................................................424 

SALVADORACEAE ...............................................................................................................425 

CAPPARACEAE .....................................................................................................................426 

MALVALES 

THYMELAEACEAE ..............................................................................................................428 

BIXACEAE .............................................................................................................................434 

DIPTEROCARPACEAE ........................................................................................................435 

MUNTINGIACEAE ..............................................................................................................462 

MALVACEAE .........................................................................................................................462 

SAPINDALES 

BURSERACEAE ....................................................................................................................492 

ANACARDIACEAE ...............................................................................................................501 

SAPINDACEAE .....................................................................................................................520 

SIMAROUBACEAE ...............................................................................................................537 

MELIACEAE ..........................................................................................................................540 

RUTACEAE ...........................................................................................................................553 

OTHER ASTERIDS 

CORNALES 

ALANGIACEAE ....................................................................................................................562 

NYSSACEAE ..........................................................................................................................565 

HYDRANGEACEAE .............................................................................................................567 

ERICALES 

TETRAMERISTACEAE ........................................................................................................571 

BARRINGTONIACEAE .......................................................................................................572 

SAPOTACEAE .......................................................................................................................577 

EBENACEAE .........................................................................................................................587 

MAESACEAE .........................................................................................................................591 

MYRSINACEAE ....................................................................................................................592 

PENTAPHYLACACEAE........................................................................................................596 

THEACEAE ...........................................................................................................................599 

SYMPLOCACEAE .................................................................................................................602 

CLETHRACEAE ...................................................................................................................604 

STYRACACEAE ....................................................................................................................605 

ERICACEAE ..........................................................................................................................607 

ACTINIDIACEAE .................................................................................................................611 

UNPLACED 

POLYOSMACEAE .................................................................................................................613 

13

14 

ASTERIDS (LAMIIDS) 

GARRYALES 

ICACINACEAE .....................................................................................................................614 

GENTIANALES 

PTELEOCARPACEAE ..........................................................................................................617 

RUBIACEAE ..........................................................................................................................618 

GENTIANACEAE .................................................................................................................645 

LOGANIACEAE ....................................................................................................................647 

APOCYNACEAE ...................................................................................................................649 

LAMIALES 

OLEACEAE ...........................................................................................................................660 

CARLEMANNIACEAE .........................................................................................................663 

ACANTHACEAE ..................................................................................................................663 

LAMIACEAE .........................................................................................................................666 

VERBENACEAE ....................................................................................................................677 

BIGNONIACEAE ..................................................................................................................678 

GESNERIACEAE ..................................................................................................................686 

SCROPHULARIACEAE ........................................................................................................687 

PAULOWNIACEAE ..............................................................................................................688 

UNPLACED 

BORAGINACEAE .................................................................................................................688 

SOLANALES 

CONVOLVULACEAE ..........................................................................................................691 

SOLANACEAE ......................................................................................................................692 

ASTERIDS (CAMPANULIDS) 

AQUIFOLIALES 

CARDIOPTERIDACEAE .....................................................................................................694 

STEMONURACEAE .............................................................................................................695 

AQUIFOLIACEAE ................................................................................................................699 

APIALES 

TORRICELLIACEAE ............................................................................................................701 

PITTOSPORACEAE .............................................................................................................702 

ARALIACEAE ........................................................................................................................703 

MYODOCARPACEAE ..........................................................................................................710 

MACKINLAYACEAE ............................................................................................................710 

ASTERALES 

GOODENIACEAE ................................................................................................................712 

ASTERACEAE .......................................................................................................................713 

DIPSACALES 

ADOXACEAE ........................................................................................................................714 

REFERENCES .........................................................................................................................................717 

INDEX TO MALAY NAMES ..................................................................................................................731 

INDEX TO FILIPINO NAMES ..............................................................................................................734 

INDEX TO VIETNAMESE NAMES ......................................................................................................736 

INDEX TO THAI NAMES .....................................................................................................................740 

INDEX TO GENUS HEADINGS...........................................................................................................744 

INDEX TO FAMILY HEADINGS ..........................................................................................................748

Such contrasting features among these basal families 

of the angiosperms raise numerous issues that impinge 

upon our understanding of how flowering plants 

evolved. How did the ‘stipule’ of Magnolias arise? Does 

it relate in any way to stipules in other angiosperms? 

By what changes could the small and strange unisexual 

? 

MYRISTICACEAE 

NAME: From the genus Myristica, as below. In English 

these trees constitute the nutmeg family, from the seed 

of commerce, the name of which is likely a corruption 

of nois-muscade or musk-nut. A few individual species 

are distinguished with local names, especially in those 

peripheral locations with only a few indigenous species. 

However, in general, vernacular names apply to the family 

as a whole and are mostly based on some derivation 

of darah, blood, in reference to the characteristic red 

exudates: chendarahan, penarahan and darah-darah (Malay, 

generally), kumpang (Iban), duguan widely in Philippines, 

but some Knema as tambalau. 

OVERVIEW: The wild nutmegs form a cohesive and 

easily recognized family of roughly 20 genera and 500 

species with five genera and maybe 300 species in tropi- 

1 Sauquet, H, et al. 2003. Botanical Journal of the Linnean Society. 

142: 125–186. 

2 Doyle, J. et al. 2004. International Journal of Plant Science. 

165: 55–67. 

Family General Description Trees of Tropical Asia 

Myristicaceae 20/500, pantropical, different genera 

in different continents, trees, 

red exudate, leaf exstipulate with 

short simple leaf stalk, unisexual 

flowers, dioecious, single-seeded 

arillate fruit. 

MAGNOLIIDS: MAGNOLIALES 

Phylogeny of Magnoliales 

Magnoliaceae 2/222, Asian-American, no exudate, 

conical encircling stipule, leaf 

stalk basally swollen or tapered 

toward the blade, often joined to 

well-developed encircling stipule, 

bisexual flower. 

Degeneriaceae 1/2, Fiji. 0. 

Himantandraceae Monogeneric, Galbulimima, tropical 

Australia, Asian. 

Eupomatiaceae 1/3, small trees and herbs, New 

Guinea to E Australia. 

Annonaceae 129/2200, different genera in different 

continents (except Anaxagorea, 

Xylopia, Popowia) no exudate, leaf 

exstipulate with simple leaf stalk, 

bisexual flowers, multiple carpels. 

# 

5/300, species-rich and abundant especially 

wet equatorial lowlands. 

79 

flowers of Myristicaceae and the large bisexual flowers of 

Magnolias evolve from a common ancestor? If Myristicaceae 

are sister to the other families, then why are fossil 

nutmegs so recent (no earlier than Miocene, some 10-20 

million years ago) and why is the molecular divergence 

among the nutmegs so modest? 

1/20, diversity of form increases toward 

China, consistent in equatorial lowlands 

but at low abundance, locally abundant 

- especially seasonal and lower montane. 

2, from Sulawesi, New Guinea, Australia. 

0. 

32/860, species-rich and exceedingly 

abundant trees and lianas in most forests 

below 800 m; not mangrove, poor in 

secondary forests. 

cal Asia. The family as a whole is strongly monophyletic 

1,2 . In distribution, the nutmegs are pantropical but 

with different genera in the different continents. The individual 

Asian genera appear to be clearly monophyletic, 

but the Asian taxa as a whole may or may not form a 

clade. There is modest evidence that independent clades 

diverged in America, Africa and Asia at an early age. 

However, the low molecular divergence among extant 

lineages and fossil evidence indicating a Miocene origin 

contradicts that scenario. 

Within Asia, the center of diversity lies in Borneo, 

and yet the genera differ greatly in details of distribution. 

Myristica is most species-rich in New Guinea and diminishes 

to the west, whereas Knema is richest in the Malay 

Peninsula and the wet parts of Borneo and has only a 

single species in New Guinea. 

The nutmegs are medium-sized evergreen dioecious 

trees (Endocomia is monoecious) with tiered branching, 

alternate simple entire leaves lacking stipules and bearing 

a bitter red sap and a distinctive single-seeded red 

arillate fruit. 

In stature, the Myristicaceae are exclusively woody, 

perennial upright trees. The range of stature is great: 

Myristica gigantea may reach 45 m in height while some

7 

Knema 

12 

1 2 3 

4 

11 

Knema. 1-3, variation in mature bark; 1, K. hookeriana; 2, K. glauca; 3, K. conferta; 4-8, leaf blade of Knema is glaucous below, the 

nerves in a fish bone arrangement, the fruit globose, orange with crusty red granules; 4, K. latifolia, Lambir, Sarawak; 5 & 8, K. glomerata, 

southern Vietnam; 6, K. pulchra, Lambir, Sarawak; 7, the twig apex of K. laurina, red hairy and typical of the family; 9-10, K. 

laurina, XS fruit 15 mm wide, natural and iodine stained, showing starch (of the genus) and ruminate endosperm (of the family); 11, 

K. intermedia, Pasoh, Malaya, red granular triangular buds, flower 7 mm wide, petals white within, spreading, staminal column is red; 

12, K. hookeriana, Pasoh, Malaya, flower 12 mm wide, red within and weakly opening. 

5 

10 

6 

8 

9 

87

94 

Annonaceae: Vegetative Form 

1 

2 

est. The American genus, Annona, contributes many 

commercial fruits as described below. The flowers of the 

native Cananga odorata are the source of the fragrant 

ylang-ylang oil. As ornamentals, the family has been 

much neglected, which is surprising because the flowers 

are fragrant while the trees demand little care. The 

Indian tree, Polyalthia longifolia is planted everywhere in 

Asia specifically because it is tough, drought tolerant and 

insect resistant. 

ANAXAGOREA. [Commemorates Anaxagoras, 

d. 428 BC, one of the first Athenian philosophers.] 

About 20 species, more than half in Central and South 

America, and then five to nine closely related species in 

lowland forest and lower wet mountains of Asia from Sri 

Lanka and Myanmar to Indochina, and throughout the 

Sunda Shelf to the Philippines. 

These are all small trees, not much over 2-5 cm DBH, 

sometimes forming an undergrowth no more than one 

m tall at maturity. The blade is broad and the venation 

is a loose open reticulation. The distribution is patchy, at 

local scales as well as regionally. Where they occur, Anaxagorea 

can be the most abundant of small trees, a local 

abundance presumably driven by the ballistic dispersal. 

And yet we find many seemingly appropriate forests 

where they are absent altogether. 

The flower bears three valvate sepals and three or six 

valvate petals in two whorls (some variants with parts in 

fours), the inner slightly smaller that outer, somewhat 

spoon-shaped, the many stamens with extrorse anthers 

some sterile or not, with connectives slightly bent outwards 

with an extended tongue-shaped apex; the carpels 

4 

Annonaceae: Vegetative form. 1 & 2, the upright, orthotropic axis (gray) with leaves (green) in spiral arrangement; in the axil of each 

leaf is a cluster of buds, the most eccentric of which develops as a lateral shoot (red) overtopping the growth of the apical meristem. 3 

& 4, in lateral branches the leaves are two-ranked, with axillary buds singly or in clusters; 5 & 6, the thick bark of the twigs peels easily 

to reveal a fishnet pattern of fibers intervening among the wide rays. 

3 

have a U-shaped opening at the stigmatic part with a 

groove on anterior side. The mature carpel is stalked and 

club-shaped; it splits as it dries and pushes out the shiny 

slippery black seed to a distance of 2-3 m. 

Anaxagorea 

Anaxagorea javanica, individual stalked carpels, dehiscent with 

ballistically dispersed seeds, 2 cm DBH trunk to the right; inset, 

A. luzonensis, flower. 

5 

6

Cananga 

2 

3 

Anaxagorea is of special phylogenetic interest in that it 

appears to be basal to the rest of the family, a result a little 

surprising in that the dehiscent fruit and ballistic dispersal 

would normally be presumed to be highly derived. 

The disjunct occurrence of species between America and 

Asia is itself rare in the Annonaceae, shared only with 

the pantropical Xylopia - but, unlike the heterogeneous 

Xylopia, the species of Anaxagorea are astonishingly similar 

to one another. Such a lack of divergence is surprising 

in the purportedly basal genus. 

The seeds smell of camphor and according to 

BURKILL loc. cit. were once used to preserve clothes. 

2 - Annonaceae Basal Grade - 2 

CANANGA. [From the Malay name.] A genus of 

two species, native to seasonal places in tropical Asia, 

6 

7 

8 

Cananga odorata. 1, long drooping lateral branch with two-ranked leaves and numerous axillary flowers in various stages of development; 

2 & 3, flower with nine perianth parts, usually described as three short triangular sepals and six equal petals; 4, numerous 

stamens spirally arranged about the numerous free carpels; 5, in a later stage the stamens have dehisced, the stigmatic surface now 

becomes sticky; 6, the mature carpels; 7, a single carpel partially dissected to show the several flat seeds embedded in the flesh; 8, the 

pimpled seed with a small white aril at the base. 

4 

1 

5 

95 

but cultivated with mixed success elsewhere. The flowers 

bear three valvate sepals, six valvate petals in two 

roughly equal whorls, connectives with lance-shaped 

apex, the mature carpels oblong, stalked with several flat 

pimpled seeds that bear a small white aril-like structure 

at the base. The complex history of the name Cananga 

has been recently reviewed. 

Cananga odorata is a small tree of poor form, a weak 

main trunk and numerous long swooping branches, almost 

liana-like and can be trained as such. It is known 

widely as ylang-ylang or kenanga. The flowers are sold on 

the street, especially in the Philippines, and also grown 

commercially for two perfumery products. Ylang-ylang 

oil is now grown mostly in Madagascar and Comoros, 

while kenanga oil is mostly grown and used locally in 

Turner, I. et al. 2009. Gardens’ Bulletin Singapore. 61: 189-204.

156 

Dillenia 

4 

1 

2 

6 

Dillenia - vegetative form. 1, D. grandifolia, wet ground in a forest gap in Malaysia, with great sweeping prop roots, saplings and 

seedlings are rare anywhere nearby; 2, the paired stipules extend from the leaf stalk and clasp one another over the twig apex, here 

broad and round in D megalantha, Philippines; 3, the stipules are scarcely developed in D. indica, although the leaf stalk is u-shaped 

with a thin dry flange; 4, the leaf of D. indica is characteristic of the genus with many strong straight regular nerves; 5, the nerves 

continue directly through the margin and extend beyond as fine teeth; 6, the bark of Dillenia is varied but most always thin, an here 

in D. indica is bright red and papery; 7, juvenile leaves are sometimes exceedingly long as in the elephant Dillenia, D. reticulata on a 

roadside in Malaya. 

3 

5

Leea 

1 

3 

9 

10 

2 

Leea. All photographs from the Philippines. 1-4, L. guineensis; 1-2, a small ill-formed tree of open roadsides, the red-flowered inflorescence 

is visible; 3, the flowers with spreading petals and thick staminal column of contrasting white color, in some the anthers 

immature, in others the anthers dehisced and reflexed; 4, a weevil on flowers of L. guineensis; 5-6, L. aculeata, spiny stem, once-cut leaf 

(with red fruit) and inset of white flowers; 7, 4-petaled flower of L. acuminatissima; 8-10, L. congesta, a single-stemmed tree, once-cut 

leaf, broad stipule and large, heavy golden fruit. (Photograph 4, © Jeanmaire Molina; 5-6, 8-10, © Leonardo L. Co; 7, © Ulysses 

Ferreras.) 

8 

7 

4 

5 

6 

159

174 

Liquidambar 

1 

3 

2 

Liquidambar excelsa (= Altingia excelsa), Java; 1, bark; 2, flower; 3, spirally arranged leaves, pointed buds, infructescence; 4-5, leaves 

of juveniles. (All photographs © Professor Eizi Suzuki.) 

Exbucklandia 

2 

4 

3 

Exbucklandia populnea, here from the Central Highlands of Vietnam; 1, the long-stalked palmately nerved leaf from vigorous stump 

sprouts; 2, note the leaf shape and nervation, the strong thumbnail-like stipule protecting the twig apex; 3, the paired stipules, with 

fused margins; 4, the stipular scar along old twigs; 5, the fruit is a woody head of several flowers, about 3 cm across. 

5 

4 

5 

1

Sonneratia 

1 

5 

4 

Sonneratia alba. 1-5, Hundred Islands National Park, Philippines; 1, thick rounded blades from short thick leaf stalks, opposite decussate; 

2, the flower without petals, the stamens numerous, opening after dark in a powder-puff display, bat pollinated; 3, the twig apex 

hidden and protected by the clasping base of the leaf stalk, a pair of stipule-like glands associated with each leaf; 4, the exerted twig 

with a scar; 5, dense pointed pneumataphores surround the patches; 6, in fruit, Palawan, Philippines. (Photograph 5, © Leonardo L. 

Co.) 

of the two, perhaps originated in 

the Bogor Botanic Garden in the 

1850s. The Malay name for the 

genus is berambang-bukit, the Iban 

sawih. 

In habit, these trees are recognized 

by the long drooping branches 

with new leaves that flush red. 

They might be mistaken from afar 

for Neolamarckia (Rubiaceae) on 

account of the habit, habitat, and 

the broad opposite leaves, but the 

twig here is four-angled, and there 

is a ridge between the petioles, not 

a well-developed stipule. Duabanga 

are fast growing light-wooded trees 

and indicators of rich damp soils, 

especially prevalent along sunlit 

river banks and exposed alluvium. 

They were long thought to be the 

dry-land relative of the mangrove 

species of Sonneratia - the two often 

linked within the family Sonneratiaceae 

- but the molecular 

studies cited above demonstrate 

the falsity of the claim and that the 

3 

Pemphis 

2 

185 

Pemphis acidus, Hundred Islands National Park, Philippines; growing on the raised coral 

beds overlooking the ocean. 

6

Memecylon 

8 

3 

4 

5 

6 

Memecylon. 1, species of larger stature have a characteristic brown bark with narrow and sharp furrows; 2, most are small singlestemmed 

trees (inset), the leaves are opposite, short-stalked, elliptic or ovate, mostly with widely spaced pinnate nerves either visible 

or obscure, the fruit typically axillary, in this species orange; 3, the leaves along the twig are all the same size, shiny, glabrous, with 

entire margins; 4, the node has a characteristic scar between the leaves, the twig bark is typically red, striate and flaky; 5-7, the flowers 

are small (usually less than 5 mm) compared to melastomes, with short fleshy corolla parts, white or violet, the stamens blue or violet, 

usually obvious in aggregate, from axillary clusters; 8, the fruit is from an inferior ovary, calyx remnants persistent, sometimes blueblack 

as in melastomes. 

2 

7 

1 

197

220 

Hydnocarpus 

6 

2 

3 

5 

4 1 

7 

Hydnocarpus. 1, bark and fallen leaf of H. pinguis, Lambir, Sarawak; 2-6, H. alcalae, endemic to Luzon, Philippines; 2-3, the staminate 

flower and bud, note the calyx lobes, free and overlapping in bud, reflexed in bloom; 4-5, the alternating leaves of lateral shoots, and 

the branches and scale leaves of upright shoots; 6, the lower trunk with flowers from the lower trunk, often lying on the ground; 7-8, 

H. borneensis, Lambir, Sarawak; 7, herbarium specimen showing the typical twice-swollen leaf stalk and asymmetric leaf base; 8, apex 

of upright shoot with reduced foliage leaves and characteristic terminal bud kicked to one side; 9, H. kunstleri, Lambir, Sarawak, flowering 

specimen; 10, fruit, possibly H. sumatranus, Palawan, Philippines. 

8 

9 

10

Drypetes 

1 

7 

3 

9 

8 

2 

12 

Drypetes, diversity of vegetative form; 1-3, D. myrmecophila, Lambir, Sarawak; 1, a large cauliflorous species; 2, the leaves are very large 

with a strongly unequal base; 3, the twigs are hollow with small ants within; 4, a dry leaf of D. pendula, from Malaya, a species similar 

to D. myrmecophila, but bears fruit among the branches and lacks ants in the twig; 5-6, leaf and bark of D. longifolia, a widespread 

species typical of the genus; 7-9, a distinctive species, D. polyneura, with deeply fluted bole, here of 42 cm DBH; 8, the leaf dries black, 

the margin with strong well-spaced teeth along the margin; 9, the new leaves flush white and develop chlorophyll only after they reach 

full size; 10-11, D. laevis, a species with untoothed elliptic leaves; 12, the node of D. falcata with almost no hint of stipular scar; 13, 

the node of D. longifolia, with stipules at the twig apex, then quickly falling; 14, the node of D. eriocarpa, with palmate and persistent 

stipules. (Photograph 13, © Leonardo L. Co.) 

4 

13 

5 

11 

14 

237 

6 

10

242 

known genera will take years to implement and so I 

leave them intact for now. Otherwise the arrangement of 

genera follows these recent phylogenetic studies except 

for the exclusion of Bischofia javanica which I place in its 

own family, next to the Staphyleaceae. 

The individual genera of the family differ in several 

vegetative features including their basic architecture 

which still needs a better survey of detail. The Phyllanthus 

F-clade is characterized by orthotropic axes with 

only spirally arranged scale leaves, while the branches 

that grow from the axil of the scale leaves are plagiotropic 

and bear numerous photosynthetic leaves in distichous 

array, the entire lateral branch sometimes abscises 

as if it were a compound leaf. The growth form of most 

Aporusa and Baccaurea in the T-clade are of more ordinary 

construction. 

Pollination is thought to be of a generalized insect 

sort, effected without specialization, but only a few species 

have been studied. Such presumptions about pollination 

proved surprisingly false among he Phyllanthus 

group (see notes under that genus). Dispersal is varied 

with many seeds brightly arillate, a few with a fleshy outer 

wall, at least some species of Cleistanthus with ballistic 

dispersal, while many species bear seeds with a small 

fatty body that aids secondary dispersal by ants. 

Phylogeny of Phyllanthaceae 

The phylogeny below is based on the papers cited in the family introduction 1-5 , and is restricted to the genera represented 

by trees within our region. Note that Bischofia is excluded and that the paraphyly of Phyllanthus is accepted 

but not implemented. The two main clades in the family are indicated: the T-clade is characterized by tanniniferous 

epidermal cells and often elongate racemose inflorescences; the F-clade is characterized by clustered (fasciculate) 

inflorescences. 

Tanniniferous Clade 

Fasciculate Clade 

GENUS DIVERSITY AND DISTRIBUTION 

Antidesma 100, Paleotropical, chiefly Asian, in our region about 70 as small trees 

and shrubs. 

Hymenocardia 9, chiefly Africa, 1 in our region, a small shrub. 

Aporusa 90, tropical Asia, trees of small to medium stature. 

Baccaurea 80, tropical Asia, trees of small to medium stature. 

Ashtonia 2, Sundaic Region, trees of medium stature. 

Dicoelia 1, Sundaic, a small tree. 

Margaritaria 14, pantropical, 2 in our region, medium trees. 

Flueggea 15, Paleotropical, about 4 in our region, small trees and shrubs, dry 

seasonal lands. 

Phyllanthus 

Phyllanthus 600 as a narrow but highly paraphyletic genus, herbs to small trees; the 

other three genera are nested within. 

Breynia 50, including Sauropus, herbs and shrubs. 

Glochidion 200, chiefly in our region, shrubs to medium trees, the core Asian group is 

possibly monophyletic but nested within Phyllanthus. 

Cleistanthus 140, especially tropical Asia. 

Bridelia 60, paleotropical, especially Asia with about 15 species, small trees, a few 

lianas, possibly nested within Cleistanthus. 

Actephila 20, paleotropical, about 6 in our region, small shrubs. 

Chorisandrachne 10, warm Asia, also Central America. 

Leptopus 20, Asia, shrubs. 

These small trees are an ecologically critical part of the 

forest understory, joining the Annonaceae and Rubiaceae 

as a source of food for wildlife. As such, they should 

be exploited in early stages of reforestation. Antidesma 

in particular has the virtues of a ready supply of seeds, 

light tolerance, and fruiting that is early, frequent and 

abundant. Otherwise, the Phyllanthaceae are of minor 

economic impact, providing little more than a few minor 

fruits. 

The family has benefitted from extensive collectionbased 

research at Leiden and Kew; the resulting monographs 

(cited under the genera) provide a good framework 

for regional study. What the family now needs is 

a new crop of students in Asia prepared to conduct field 

studies on these abundant and ecologically important 

trees. This is a reasonable ambition in that these genera 

are abundant and species-rich in most of tropical Asia, 

they are small in stature, flower often, are easy to observe 

and collect, and they make good specimens. 

2 - Antidesma Group - 2 

The singular genus Antidesma, which is readily 

distinguished by flower, fruit and leaf was linked 

in the cited molecular studies with the odd pair of

Antidesma 

1 

4 

2 

10 

7 

3 

9 8 

Antidesma. 1, bark of A. coriaceum, typical of the genus, thin, tan-white and brown, 25 cm DBH; 2-3, A. bunius; 2, habit; 3, leaf and 

fruiting stalk, maturation from green to red and purple; 4, A. sootepensis, in flower, Thailand, the bark, with narrow shaggy strips; 5, 

axillary floral stalks of A. cumingii, Philippines; 6, large leafy stipules of A. pendulum; 7, odd heart-shaped stipules of A. neurocarpum, 

Borneo; 9-10, leaf, double-swollen stalk and spreading whiskered stipules of A. coriaceum, all unusual in the genus; 11, diagram of 

pistillate flower, A. bunius. 

6 

5 

243

248 

Baccaurea 

2 

1 

10 

9 

Baccaurea. 1, B. sarawakensis, Borneo, Baccaurea are mostly small trees with thin bark that is variously cracked, peeling, smooth or 

thinly fissured; 2, B. tetrandra, Philippines, the leaves are in upturned spirally arranged clusters, the leaf stalk swollen above and below, 

the stipule conspicuous but small, the flowers and fruit typically from below the leaf clusters or on the twigs or trunk; 3, B. racemsosa, 

Malay Peninsula, the fruit, similar to Aporusa, is often colorful, with a more or less dehiscent outer wall, while the seed wears an arillate 

jacket, in this species a bright blue; 4, the densely cauliflorous tree of B. ramiflora; 5, the fruit of B. angulata, of Borneo, sold in markets, 

is divergent in the elongate apex and sharp ridges; 6, large fruit from species such as B. macrocarpa are sold in markets as tampoi, 

for the edible aril, here in Borneo; 7-10, the widespread and abundant B. parvifolia is distinctive in the secondary sex characters; the 

staminate tree (7) bears flowers all along the trunk, the staminate flower (8) is typical of the genus in the petals being so much larger 

than those of Aporusa; the pistillate tree (9) bears flowers only at the base, the fruit are usually single-seeded indehiscent berries that lie 

in a great mass around the base of the tree (10). Photograph 2, © Leonardo L. Co.) 

5 

3 

7 

6 

8 

4

260 

In tropical Asia, we find many species of Euphorbiaceae 

and Malvaceae that share a similar appearance: large 

palmately nerved leaf blades with long leaf stalks and 

stipulate nodes. This is particularly true in open sunny 

habitats in strongly dry seasonal places. Below I compare 

two small trees often found growing side-by side. 

Euphorbiaceae 

The upper leaf 

stalk is weakly 

swollen, without a 

hard central core, 

with a trace of 

whitish exudate. 

The twig bark 

is weakly fibrous, 

the fibers 

straight, without 

mucilage. 

Euphorbiaceae versus Malvaceae 

On the left is Melanolepis multiglandulosa (Euphorbiaceae); 

on the right is Kleinhovia hospita (Malvaceae). 

One other difference not illustrated is the tendency of 

Euphorbiaceae to bear leaf blades and leaf stalks of different 

sizes at maturity, whereas in Malvaceae the mature 

leaves are of uniform size. 

Malvaceae 

The upper leaf 

stalk is swollen, 

with a hard central 

core, without 

exudate, often 

with mucilage. 

The twig bark is 

tough, fibrous, 

the fibers in a 

fish-net pattern, 

mucilage is common.

270 

Macaranga 

5 

12 

10 

11 

8 

Macaranga. 1-6, M. tanarius, Philippines, a species of full-sun, of poor soils and without ants; 1, a tree 6-years old, tiered branches at 

discrete levels form a flat-topped crown; 2, the leaf clusters seen from below; 3, the fast-growing trunk with prominent scars from the 

leaf and stipule; 4, the stipules of a young twig, the stem with a white waxy covering; 5, the peltate leaf; 6, leaf margin with small glandular 

teeth; 7-9, M. grandifolia, Philippines; 7, superficially similar in the large leaves to the unrelated M. gigantifolia of the Sundaic 

Region, but here peculiar for the genus in the inflorescence which is dense and sessile; 8-9, the pistilae flowers; 10-11, M. recurvata in 

Malaya, a species characteristic of gaps in swampy forest; 10, with a white bark and prop roots; 11, the dehiscent fruit with red-arillate 

seeds. 

6 

3 

9 

2 

1 

7 

4

322 

Calophyllum 

1 2 

9 

7 

Calophyllum. 1-3: variation in bark form, 1, C. macrocarpum, the most characteristic bark type with flat plates divided by diamond 

shaped fissures; 2, C. pulcherimum, a red fibrous bark; 3, the gray shaggy bark of C. inophyllum; 4, typical pendent clusters of globose 

fruit, C. inophyllum; 5, the apical bud typically naked, pointed, exerted, crusted with red excretions; 6, leaf surface with closely spaced 

corrugated nerves; 7-8, C. macrocarpum; 7, flowers bisexual with white petals, numerous stamens; 8, fruit usually green with scant 

drops of yellow exudate, with a single large seed; 9, the fruit typically in a sparse infructescence, in some species the fruit is yellow at 

maturity, sometimes pointed, here a possibly new species from Lambir. 

6 

8 

3 

4 

5

Calophyllum 

6 

1 

5 

2 

7 

Diversity of leaf form in Calophyllum in Lambir, Sarawak The blade is simple and generally elliptic, but varies in relative width and 

length, and includes lanceolate (5) and oblanceolate (1); the nerves vary in density, conspicuity and the angle they make with the midrib; 

the leaf stalk varies in length; the twig varies from terete to strongly 4-winged; the apex varies in species-specific details of form and 

pubescence; and note in (5) and (6) the contrast between leaves of juvenile and adult trees. (Species identity: 1, C. inophylloide; 2, C. 

confertum; 3, C. ferrugineum; 4, C. aff. woodii; 5-6, C. castaneum; 7, C. pulcherimum; 8, C. macrocarpum; 9, C. alboramulum. However, 

note that you cannot be certain of identity by matching leaves to photographs alone.) 

These are small to medium trees, The plants appear 

to be androdioecious but are evidently cryptically dioecious 

1 . The leaves are opposite, short stalked, entire, 

leathery; fine venation densely and evenly boxed-reticulate 

and with translucent gland dots. The leaf pairs 

when young appear to bear the equivalent of Rubiaceaelike 

stipules, with alternately long and short internodes, 

creating a somewhat whorled appearance to the leaf arrangement. 

But if you have seen the dry leaf once, you 

will not likely mistake it for anything else. The ovary is 

two-loculed, each locule with two ovules, the style very 

short in contrast to Calophyllum, The fruit is an indehiscent 

berry, variably thick and fleshy wall, with one to 

four large seeds. 

1 Dunthorn, M. 2004. Plant Systematics and Evolution. 249: 

191-196. 

bintanggor 

3 

8 

9 

4 

323 

Wood sample of bintanggor from the Sarawak Timber Council, 

probably Calophyllum soulatri, which is a common timber from 

peat swamps.

348 

Sindora 

8 

6 

ed, but see the images of type specimen at the Singapore 

Botanical Gardens web site.) 

SINDORA. [From a native name.] Sindora is a genus 

chiefly of tropical Asia, with perhaps 20 species, although 

a single species is recorded for Africa. Only three 

are noted for Thailand, five to seven in the Sundaic Re- 

3 

7 

Sindaora. 1-5, S. coriacea; 1,5, Sarawak, 2-4, Malaya; 2, canopy high and wide, deciduous; 3, canopy with nearly mature fruit sticking 

up exposed to the wind; 5-7, S. beccariana, Sarawak; 5, fallen leaf and flower; 6, mature tree, 95 cm DBH; 7, fallen fruit; 8, S. velutina, 

Malaya, line drawing from voucher specimen, showing leaflet shape and prominent stipules. A variant form of Sindora siamensis, perhaps 

better called S. maritima, fruiting at a height of less tha 4 m, seaside in central Vietnam. 

5 

9 

2 

4 

gion with two species found as far north as Luzon, Philippines, 

chiefly along the east coast. The Malay name 

seperti applies to both the tree and the timber, while in 

Borneo the Iban name is tampar. The name kayu-galu is 

common in southern Philippines. 

The leaflet of Sindora is characteristic with the strictly 

paired asymmetric shape; the venation of the leaves is 

1

350 

merbau 

Merbau timber, Intsia palembanica, from a wood sample of the 

Sarawak Timber Council. 

AFZELIA. [Commemorates Adam Afzelius, d. 1837, 

student of Linnaeus, botanist in Sierra Leone.] A genus 

of 13-20 species, in Africa and Asia, with two species 

especially notable in our region: Afzelia rhomboidea in 

Java, Borneo and the Philippines, and A. xylocarpa of 

Mainland SE Asia. The genus differs from Intsia in bearing 

seven rather than three or fewer stamens, and in the 

Afzelia 

thicker seed pod with arillate seeds. In Malay languages 

it shares with Intsia the name merbau, while the Tagalog 

for Afzelia rhomboidea is tindalo, one of the most characteristic 

timber trees of the Philippines. These are slow 

growing trees of seasonal climates. The wood is comparable 

to that of merbau, and is equally regarded as one of 

the choice timbers for fine furniture. Heavy, hard, beautifully 

figured in shades of red and brown. The trunk of 

Afzelia xylocarpa is readily recognized in lowland forest 

of the Mainland because it seems to invariably develop 

two or three great branches no more than four or five 

meters above the ground. The flowers are remarkable 

in the brightly colored flag petal and the extraordinary 

length of the stamens. I find no record of the likely pollinator 

although large bees are obvious visitors. The arilcovered 

seeds are greedily taken by a variety of wildlife 

and especially by gibbons. 

ENDERTIA. [Commemorates F. Endert, d. 1953, 

senior Dutch forester between 1915 and 1950.] Another 

monotypic genus, represented by Endertia spectabilis, 

which was only formally described in 1947. This is a 

large trees, the leaflet blade is pale glaucous below, and 

as the species epithet suggests, it flowers in profusion. 

While evidently restricted to eastern Borneo, and some- 

Afxelia. Left, A. xylocarpa, the typical trunk of a forest grown tree in Vietnam is very large with large branches that fork close to the 

ground, old fruit pods litter the ground; from Thailand, the arillate seeds are a favorite food of gibbons; right, illustration of leaves and 

fruit of A. rhomboidea in the Philippines, the flower bears exceptionally long stamens and style, the pollinator unknown. (Illustration 

from from BLANCO loc. cit.; photograph of flower, © Ulysses Ferreras.)

ROSIDS (MALVIDS): MALVALES 

DIPTEROCARPACEAE 

NAME: From the genus Dipterocarpus, as below. The 

family as a whole has no common name, although in 

English dipterocarps in lower case implies the family 

broadly. A great many individual genera and species have 

names as treated below. 

OVERVIEW: The family is usually regarded as comprising 

17 genera and 500 species. Molecular data, while 

encumbered by sparse sampling of the larger genera, 

nonetheless demonstrates that the family as a whole 

is strongly monophyletic. Two small subfamilies are 

found respectively in Africa and tropical America while 

the bulk of the family consists of the exclusively Asian 

subfamily Dipterocarpoideae, with 13 genera and about 

470 species. The subfamily shows good genus-level representation 

throughout tropical Asia west of New Guinea, 

while adding a significant pocket of endemism in Sri 

Lanka. The genera are individually uniform, strongly 

monophyletic and the molecular data generally conform 

to the traditional view of generic circumscription, with 

important exceptions in the species-rich and abundant 

assemblage under the tribe Shoreeae which is monophyletic 

as a whole but finds that Hopea is nested within 

Shorea. The tribe requires a new structure at the genus 

level. (See the following separate sections on dipterocarp 

phylogeny and the later section on phylogeny of the 

tribe Shoreeae.) 

The best introduction to the family remains Colin 

Symington’s classic account of Dipterocarpaceae in the 

Malay Peninsula 1 . The volume is a readable yet exact account 

of the many species; but more than that, it continues 

to inspire each new generation as a demonstration 

of how forest botany in the tropics should begin with 

the living tree. Two recent reviews of the family provide 

a modern account 2,3 . At the species level, Symington’s 

treatment for Malaya is complemented by other good accounts 

for the several regions within tropical Asia; these 

are listed in a separate section below. No book treats the 

family in its entirety, nor even for tropical Asia broadly. 

In stature, dipterocarps include so many species of 

very large size, and these are so ubiquitous and abundant, 

that we might wrongly conclude that the family 

is exclusively of the upper canopy and emergent forest 

layer. In Vatica, Hopea and a few types of Shorea we find 

some relatively small understory trees. Dipterocarps 

can be said to be without exudate but they are strongly 

resinous; the resin exudes slowly from cuts, often forming 

distinctive patches on the side of the mature trunk, 

sometimes strongly smelling of turpentine, while the dry 

resin, dammar, burns with a rich odor of incense. The 

1 Symington, C. 1943. Foresters’ Manual of Dipterocarps. Malay 

Forest Records 16. 

2 Appanah, S. et al. [eds.]. 1998. A Review of Dipterocarps: Taxonomy, 

Ecology and Silviculture. Center for International 

Forestry Research, Bogor, Indonesia. 

3 Ashton, P. S. 2002. The Families and Genera of Vascular Plants. 

5: 182-197. 

435 

Dipterocarpus alatus planted in Ho Chi Minh City, Vietnam in 

the late 1890s, now with a diameter of 1.8 meters. The two trees 

illustrate the typical form of mature trees of Dipterocarpus. They 

make an excellent tree for urban landscape, but are rarely so used 

outside of Vietnam and Chiang Mai, Thailand. 

presence and form of buttresses varies with genera and 

sections. Leaves are borne in a spiral phyllotaxy, but at 

maturity usually array themselves in a plane along the 

horizontal branches. Many dipterocarps bear glands in 

the leaves, although these seem to function only at the 

youngest stages of development. Stipules are likewise 

characteristic of the family and protect the growing apex 

in a mechanical way. The leaf stalk is conspicuous, often 

long and decidedly pulvinate at the blade junction. 

The blade is entire (Dipterocarpus leaves are sometimes 

wavy) and pinnately nerved, often with strong conspicuously 

scalariform venation, and varied domatia as pits 

and tufts. While only a few species can be said to have 

a densely hairy leaf surface, most dipterocarps bear some 

sort of hair, sometimes simple, often stellate. The typical 

dipterocarp flower is small in size (except in many 

Dipterocarpus), bisexual, regular, aggregated in terminal 

or axillary bracteate panicles, scented, with parts in fives, 

including a five-part calyx, often with basal tube that is 

persistent and distinguished by two to five of the calyx 

lobes that enlarge as wings in ways that are diagnostic 

among genera. The corolla is of five parts, more or less 

spiral and partially fused at the base; stamens often number 

15, but vary among genera, dehiscing by longitudinal 

slits; the gynoecium with two to five fused carpels, 

styles one to three with a three-lobed stigma; ovules two 

to four per locule. The fruit is characteristic, a dry inde- 

. . . continued on page 452

442 

Dipterocarpus 

1 

7 

6 

2 

5 

Dipterocarpus. 1, flower of D. grandiflorus, Philippines, showing the five overlapping corolla lobes, clustered stamens and central 

exerted style; 2, flower of D. crinitus; 3, the fruit of Dipterocarpus is distinctive with the two calyx lobes expanded as wings, often red 

before drying, species vary in the length of the wings, size of the seed and details of the calyx with regard to ridges and wings; 4, D. 

confertus, 86 cm DBH, large columnar trees with low plank buttresses and a dense bed of persistent dry leaves around the base; 5, the 

dry leaves are distinctive in the strong regular nerves that nearly touch the wavy margin and the plicate leaf surface, 6, the fresh leaves 

can appear more pliable; 7, the inflorescence is often sparsely flowered in arrays of axillary racemes; 8, indumentum when present is 

often in star-shaped clusters. (Photograph 1, © Ulysses Ferreras.) 

8 

3 

4

1 

Dipterocarpus 

7 

9 

3 

2 

8 

Dipterocarpus, diversity of form. 1-3, D. acutangulus; 1, low buttressed trunk with shaggy bark, 80 cm DBH; 2, small leaves, finger for 

scale, with sharply ascending nerves; 3, juvenile twig with bristle-bush apex; 4, D. caudatus, trail side tree at Bukit Timah, Singapore, 

with low plank buttresses, smooth elliptic leaf; 5-6, D. geniculatus; 5, columnar trunk, still immature at 35 cm DBH, with distinctive 

flaking bark; 6, the dry leaf with long geniculate leaf stalk (see inflorescence of the same species on previous plate); 7-8, D. cornutus; 

7, pale columnar trunk with low buttresses, 89 cm DBH, the inset showing the large strongly nerved leaf and red-winged fruit; 8, 

the stipule of the juvenile is diagnostically long, pendent and bears clusters of golden stellate hairs; 9, the twig apex of D. kunstleri 

juvenile, long smooth and differs from the adult which bears large paired stipules like banana peels; 10, D. grandifolius, with typical 

columnar bole and pale tan flaky bark (arm for scale), inset of fallen leaf and fruit; 11, D. crinitis, 131 cm DBH, distinctive in the 

broad base, rounded nearly fluted buttresses. 

10 

4 

5 

6 

11 

443

464 

The modern view of the Malvaceae echoes the 19th 

century practice of aligning these many genera in a single 

family, an arrangement in contrast with the 20th century 

practice of limiting the Malvaceae to the immediate 

associates of Malva, while recognizing three other major 

families: Sterculiaceae, Tiliaceae and Bombacaceae, all 

four in a narrowly conceived Malvales. The change is not 

merely the raising or lowering of taxa along the hierarchy 

- which is trivial - but represents a real change for the 

better by the realignment of so many genera. Durians finally 

break away from the utterly different Bombax trees, 

Grewia is segregated from Pentace and allies while Tilia 

itself is properly isolated, and the very distant elaeocarps 

are altogether dismissed from the Malvaceae. 

Phylogeny of Malvaceae 

GENUS NOTES 

GREWIA GROUP - Grewioideae - 25/770, Pantropical. 

Grewia 90, Paleotropics, especially Africa. 

Microcos 60 India to Australia. 

Colona SE. Asia, India to the Solomons. 

Trichospermum 30, Pantropical. 

LEPTONYCHIA GROUP - Byttnerioideae - 20/650, Pantropical, chiefly American. 

Leptonychia 28, Africa, Africa, 3 in Asia. 

Byttneria 132, Pantropical, herbs and lianas, maybe 1 tree in Asia. 

Commersonia 9 Australia, 1 widespread, weedy in Tropical Asia. 

Abroma 5, Asia to Australia. 

Kleinhovia 1, Indian Ocean to Polynesia. 

STERCULIA GROUP - Sterculioideae - 12/430. Pantropical. 

Sterculia Pantropical; 19 in Borneo. 

Heritiera 30, Africa and Asia. 

Scaphium 12, Mainland SE Asia to Borneo. . 

Pterocymbium 15, Tropical Asia. 

Pterygota 12, Pantropical. 

Firmiana 12, Africa and Asia. 

PENTACE GROUP - Brownlowioideae - 8/70, mainly Palaeotropics. 

Pentace 25, tropical Asia. 

Brownlowia 20, India to the Solomon Islands. 

Diplodiscus 7, Philippines, Sabah, Malaya, Sri Lanka 

Jarandersonia 3-5, Borneo. 

Berrya 3, India, Sri Lanka to Pacific. 

Pityranthe 2, 1 Sri Lanka, 1 in S China and Mainland SE Asia. 

PTEROSPERMUM GROUP - Dombeyoideae - 8/70, especailly Africa to Iddia. 

Pterospermum 20, India to China and the Pacific. 

Burretiodendron 4, tropical China and northern Mainland SE Asia 

Eriolaena 17, India to China. 

Excentrodendron 2, China and Vietnam. 

Paradombeya 1 China. 1 Myanmar and Thailand. 

Schoutenia 9, N Thailand to Java and N. Australia; not Philippines. 

TILIA GROUP - Tilioideae - 3/50, N Temperate, Central America. 

Craigia 2, China, Vietnam. 

HELICTERES GROUP - Helicteroideae, in part - 6/150, Pantropical. 

Helicteres 60, tropical Asia and America, 3-5 in dry seasonal tropical Asia 

Reevesia 18, S China and E Himalayas, 1 S to Vietnam, cultivated. 

DURIO GROUP - Helicteroideae, in part - 4/50, tropical Asia. 

Durio 30, Sundaic Region. 

Neesia 7, tropical Asia 

Coelostegia 5, tropical Asia 

Kostermansia 1 tropical Asia 

Ten clades equivalent to subfamilies are usually recognized. 

All ten are represented with at least one species 

of tree in tropical Asia, and only Tilia is not treated in 

this volume, but even one species of that genus reaches 

the northern extremity of Vietnam. Below, I provide an 

arrangement of the subfamilies, listing only those genera 

represented by trees in our region. The new arrangement 

is based largely on molecular studies cited in the family 

introduction. Note that although the individual groupings 

are relatively strong, their relationship to one another remains 

unclear. I provide informal group names following 

our more conspicuous genera of trees, and provide the 

formal subfamily beside each group name. 

HIBISCUS GROUP - Malvoideae -, 78/1800, cosmopolitan. 

Camptostemon 2, Borneo, Philippines. 

Hibiscus 250, pantropical, maybe 12 trees in tropical Asia. 

Thespesia Pantropical, several in New Guinea, 2 in tropical Asia. 

Kydia 1, India to Mainland SE Asia. 

BOMBAX GROUP - Bombacoideae - 12/120, tropical, especially Africa and America. 

Bombax 20, Pantropical 

Ceiba American, 1 in Africa, cultivated in tropical Asia.

468 

Colona 

Colona. 1-2, C auriculata, Vietnam coastal lowlands, flower and fruit with negligible wings; 2, the habit of single arching stem with 

large pendent leaves and numerous flowers; 3, C. evrardii, Vietnam, winged stipules, asymmetric 3-nerved leaf base; 4, C. serratifolia, 

Philippines, showing habit, flower and winged fruit. (Illustration from BLANCO loc. cit.) 

2 - Leptonychia Group - 2 

Subfamily Byttnerioideae 

This subfamily has been recently reorganized based 

on molecular work 11 . It now includes 26 genera 

and 650 species, pantropical, but especially rich 

in South America. The flowers are bisexual with 

calyx and corolla, five or more stamens. The complex 

hooded petals seen in Theobroma are evidently 

pleisiomorphic and the smaller flatter or scale-like 

petals are derived. Most of these genera were formerly 

in the Sterculiaceae. Included here is the 

Neotropical Theobroma cacao, the source of chocolate, 

sometimes in home gardens, especially in the 

Philippines, a few plantations in Malaysia, but the 

quality of the seed is low. 

LEPTONYCHIA. [Greek, thin nail-shaped, in reference 

to the petal.] A genus chiefly of Africa with nearly 

40 species; about three in tropical Asia. These are common 

and relatively abundant species that include small 

understory trees and medium-sized lower canopy trees. 

The leaves are entire with a three-nerved base, borne on 

a long stalk that is sharply pulvinate above. The flowers 

are borne in small axillary cymes; they bear five se- 

11 Whitlock, B. et al. 2003. Systematic Botany. 26: 420-437. 

2 

1 

3 

pals and five petals, neither showy, the filament of the 

stamens are united in the basal half. The stamens are in 

three series with fertile stamens in the middle ring and 

sterile stamens outside and in. The fruit is one-threecelled 

capsule, thin walled and breaking open when dry 

to reveal brightly arillate seeds. 

Leptonychia caudata is a small tree with thin papery 

leaves and shiny black-violet twigs. It is abundant and 

widespread in lowland equatorial forests from Malaya, 

Sumatra and Borneo; the same name is applied broadly 

even though the small trees vary in detail from place 

to place. Leptonychia banahaensis is an abundant lower 

canopy tree in the Philippines. 

ABROMA. [Greek, inedible, uncertain reference.] A 

genus of only two species. The first is Abroma fastuosa of 

Australia and New Guinea. The second comprises small 

weedy shrubs of variant forms that are usually lumped 

under the name Abroma angusta. It is found in most of 

tropical Asia from the Mainland to Australia. The Malay 

is rami sengat. It never much exceeds three to four m in 

height; the branches arch outward to display the broad 

leaves. The leaves and stems are covered with bristly irritating 

hairs. It is sometimes cultivated under the name 

devil’s-cotton. The flower and the fruit are both odd. The 

large maroon corolla forms something of a chamber and 

4

474 

Scaphium 

1 

2 

3 

Scaphium. 1-3, S. linearicarpum, Pasoh Forest, Malaya; 1, the mature tree, 52 cm DBH above the narrow buttresses, and bark dippled 

red and brown; 2, the seed germinating in a mass of water-retaining mucilage; 3, staminate flowers, about 4 mm across; 4-6, S. macropodum, 

Pasoh Forest, Malaya; 4, the characteristic deeply furrowed bark of the mature tree, 31 cm DBH; 5, the array of deeply digitate 

leaves of the sapling; 6, pistillate flower with arched stigma. 

Scaphium 

1 2 

4 

5 

Scaphium. Changes in leaf form with matuation. 1-3, S, borneensis, Lambir Forest, Sarawak; 1, sapling, the first leaf is entire; 2, at 

2 cm DBH the leaf is 3-lobed; 3, the leaf of the mature tree is a small entire blade with a 3-nerved base; 4-7, S. macropodum; 4, the 

seedling leaves change from entire to trilobed; 5, at 2 cm DBH, the leaf is deeply digitate with 5 or 7 lobes; 6-7, at maturity the 

leaf is entire with a 3-nerved base; the base is either rounded (6, Pasoh Forest Malaya) or square to nearly invaginated (7, Lambir 

Forest). 

5 

6 

3 

6 

7 

4

528 

Nephelium 

1 

7 

8 

9 

11 

Nephelium. 1-8 N. lappaceum; 1, fruit with fleshy aril attached to fibrous endocarp; 2, leaf; 3, lower surface of leaflet, glaucous, typical 

venation, axillary domatia; 4, twig with ridges and leaf base; 5, rambutan in Thailand; 6, yellow form of rambutan in Sarawak; 7, 

inflorescence; 8, pistillate flower, 2-lobed ovary, long spreading stigma, small or absent staminodes; 9, N. maingayi bark, reddish, warty 

lenticellate; 10, fruit of N. ramboutan-ake, the Malaysian pulasan, Sarawak; 11, transever view of the seed and aril. 

4 

5 

3 

10 

2 

6

530 

Dimocarpus 

1 

Dimocarpus. 1-4, D. longan; 1, cultivated tree, Philippines; 2, lower leaf surface; 3, wild tree with distinctive bark, among the most 

abundant species at Huai Kha Khaeng, Thailand, with inset showing the leaf; 4, mature fruit, from a market in Sarawak. (Photographs 

1-2, © Ulysses Ferreras.) 

Xerospermum 

1 

3 

Xerospermum. 1-4, X. noronhianum; 1-2, Pasoh; 1, characteristic dark red and black bark with small round flakes that are colored 

orange-red; 2, the fruits remain green for nearly a month, then mature to yellow, Malaya; 3-4, southern Vietnam; 3, the fruits dehisce 

at maturity exposing the orange aril; 4, fruit scattered below a tree. 

3 

2 

4 

4 

2

572 

OTHER ASTERIDS: ERICALES 

BARRINGTONIACEAE 

NAME: From the genus Barringtonia, as below. No 

common names apply to the family as a whole, but we 

might call it the putat or cutnut family. Note that if this 

family is treated under a broad Lecythidaceae, then the 

name at the subfamily rank is Planchonioideae. 

OVERVIEW: The trees of this small family represent a 

monophyletic group of five genera and maybe 50 species. 

They fall within the broadly circumscribed Lecythidaceae, 

a DNA-based monophyly without a morphological 

synapomorphy, consisting of five clades that are perhaps 

best treated as five families 1 . The Barringtoniaceae are 

strictly Paleotropical in distribution and predominantly 

tropical Asian. They are distinguished by the important 

if obscure synapomorphy of pollen that is syntricolpate 

with a thickened colpus ridge. All species in the family 

are trees although a few are understory plants of the 

schopfbaum-habit, that is, with a single thick stem and a 

dense rosette of leaves. The flowers are radially symmetrical 

while the seeds are variably reduced to as few as one. 

Ecologically, the Barringtoniaceae are of two types. First, 

are those of the lowland equatorial forest, especially Barringtonia, 

for which a few species are usually abundant 

in most lowland forests, together with a few Planchonia 

species. Second, are the genera and species of the dryseasonal 

forests, which include Careya, Petersianthus, 

Chydenanthus, and a few species of Barringtonia. The 

family is rare in equatorial mountains; for example, only 

two species of Barringtonia are claimed for Kinabalu. 

All are trees with moderately thick twigs, spirally arranged 

leaves, the blade of which is entire with a strongly 

serrate margin, or at least a few distant and obscure teeth 

near the apex. The node is clean. (The paired ‘stipules’ 

if they are such, are little more than dark patches.) The 

twig apex is typical of Ericales with a long thin pointed 

tip. The flowers are mostly large powder-puffs, bisexual, 

with a large thick persistent calyx. The radially symmetric 

flower, nectary disc, long slender style, and few 

seeded indehiscent fruit collectively distinguish the family 

from the Neotropical Lecythidaceae. 

The genera - all species-poor other than Barringtonia 

- differ most notably from one another in details of the 

gynoecium, but it would be good to know if Barringtonia 

(with four ovules per locule pendent from the apex, 

and one mature seed) is monophyletic or if these small 

genera lie nested within. Petersianthus has a winged fruit, 

while Chydenanthus bears an articulated floral stalk, the 

ovary with one to two ovules from the base. The other 

three genera all bear numerous ovules arrayed along the 

axis of the ovary, differing among one another as follows: 

Careya (abundant dry-seasonal) ovules from the axis 

1 Mori, S. et al. 2007. American Journal of Botany. 94: 289-301. 

2 El-Sheriff, I. et al. 2006. Folia Malayana 7: 41-54. 

3 Payens, J. 1967. Blumea. 15: 157–263. 

midpoint, carpels three, seed arillate; Planchonia (common, 

lowlands of Sundaic Region) likewise, but the mature 

seed unique in the family in that the embryo bears 

well-developed plicate cotyledons. 

I include in Barringtonia the Whitmore genus Abdulmajidia, 

the only distinction being the presence of 

more than one seed. His genus included two new species, 

neither known in the field, and each represented by 

not much more than the type specimens from Maxwell 

Hill in Perak (A. maxwelliana) and Bukit Chini and Gunung 

Pulai in Johore (A. chanii). In 2006, three more 

species were added 2 . Combinations under Barringtonia 

have not been published. 

There is a strong but superficial similarity to those 

Myrtaceae with spiral leaves while the powder-puff 

flowers do look like a Syzygium. However, in Barringtoniaceae 

the many stamens of the flower mature from the 

center of the flower outwards, while in Myrtaceae they 

mature from the outer edge towards the center. 

The economic potential of these trees has been badly 

neglected. They are sturdy and insect-resistant trees with 

deep green foliage and attractive night-blooming flowers, 

and should be far more common in cultivation, both in 

cities and towns. They could also figure in reforestation. 

Several have useful timber, and a few bear edible nuts. 

BARRINGTONIA. [Commemorates Daines 

Barrington, d. 1800, English naturalist.] The genus is 

known from Africa to Madagascar, India to Australia 

and the Pacific. Over 150 basionyms have been published, 

these represent perhaps 51 species following the 

last monograph 1 . However, as CORNER loc. cit. noted 

long ago, the variant and localized forms in Asia are not 

very well resolved. We count about 17 species in Malaya, 

perhaps 10 species in Indochina, 13 in Brunei, and 

one species reaches Yunnan in the form of the endemic 

B. fusicarpa. Local names apply to the genus as a whole, 

although sometimes with additional adjectives for the 

FIELD RECOGNITION 

Clean glabrous trees with no exudate. 

Leaves of medium to large size in spiral clusters from relatively 

thick twigs, a candelabra appearance . 

Stipules are present, but are a difficult character in that they are 

greatly reduced or abortive in many species and represented 

by no more than a tiny bit of black tissue on each side of the 

leaf base, and only in the youngest leaves. 

The leaf blade is usually glabrous, oblanceolate, with small teeth 

in upper portion (but not in the coastal B. asiatica). 

The leaf stalk is broadly thickened at the base, typically with two 

ridges 

The twigs are hollow with cortical vascular bundles, which in 

some genera we find the xylem is to the outside and the 

phloem to the inside, reversing the normal order. 

FIELD CONFUSION 

The general habit of leaf and twig is fairly widespread in 

Anacardiaceae (e.g., Campnosperma, but these they have 

solid twigs, leaf stalk with terete and swollen base, no hint 

of stipules), a few stalkless pachycaul Sterculia (these with 

3-nerved leaf), and Sapotaceae (these with white exudate).

Barringtonia 

6 

1 

2 

7 

8 

Barringtonia. 1-2, B. curranii, Borneo, 32 cm DBH; 3-5, B. macrostachya, Pasoh, Malaya; 3, fallen stamens, 7:30 AM; 4; leaf and 

flower buds; 5, leaf and bark of mature tree, 16 cm DBH; 6-9, B. racemosa; 6, apex with characteristic leaf base; 7, toothed leaf margin; 

8, single seeded fruit, thin papery testa, small undifferentiated embryo; 9, from Mt. Hamaguitan, Philippines, typical habit, new leaves 

red below; 10, B. procera, one of the cutnuts, here from Solomon Islands; (Photograph 9, © Leonardo L. Co; 10, © Roger Leakey.) 

9 

5 

3 

10 

573 

4

594 

Ardisia 

8 

6 

4 

7 

10 

Ardisia. 1, A. pyramidalis, in the Philippines, a medium multi-branched shrub with spirally arranged leaves and terminal panicles; 2-3, 

two other types of floral architecture, 2, a common type with terminal pendent inflorescence of flowers borne in cycles from a persistent 

main axis; 3, an open lax and pendent inflorescence; 4, the characteristic base of lateral branches, flattened and swollen apically 

and basally; 5, the twig apex, mottled green and white with finely red hairy slender point; 6-7, the leaf margin is often glandular and 

sometimes invaginated; 8-9, the flowers typically pink, with a single exerted style and a cone of stamens, are readily recognized; 10, the 

mature fruit vary from dull green and red to brilliant red. (Photographs 8, 10 © Leonardo L. Co.) 

5 

9 

2 

3 

1

740 

INDEX TO THAI NAMES 

These are the names in Thai for some of the more common trees found in Thailand. The list represents a standard 

list of names developed by Dr. Sarayudh Bunyavechewin of the Royal Thai Forest Department. A more exhaustive 

list of the many local and dialect names of plants in Thailand is found in Tem Smitinand’s published lexicon of Thai 

plant names 1 . Note that here the Thai name refers to a particular species, whereas the page reference is to the genus 

heading, and the full species name is given even if this particular species is not mentioned in the text. 

1 Smitinand, T. 2001. Thai Plant Names. (Revised Edition.) The Forest Herbarium, Royal Forest Department, Thailand. 810 pp. 

กรวยป่า Casearia grewiifolia 225 

กรวยป่าไม่มีขน Casearia graveolens 225 

กระโดนแดง Tristaniopsis burmanica 188 

กระท้อน Sandoricum koetjepe 545 

กระท่อมหมู Mitragyna rotundifolia 632 

กระบกกรัง Hopea helferi 457 

กระบาก Anisoptera costata 444 

กระพี้ 

Sapindus rarak 533 

กฤษณา Aquilaria crassna 430 

กล้วยขี้ 

Polyalthia cerasoides 103 

กล้วยไม้ลูกเล็ก Mitrephora vandaeflora 110 

กล้วยไม้ลูกใหญ่ Alphonsea ventricosa 108 

กล้วยฤาษี Diospyros glandulosa 587 

ก่อข้าว Castanopsis inermis 416 

ก่อแซะ Anacolosa ilicoides 168 

ก่อดาน Castanopsis purpurea 416 

ก่อดำ Lithocarpus truncatus 416 

ก่อเดือย Castanopsis acuminatissima 416 

ก่อตาหมู Lithocarpus thomsonii 416 

ก่อนก Lithocarpus polystachyus 416 

ก่อน้ำ Lithocarpus annamensis 416 

ก่อใบเลื่อม 

Castanopsis tribuloides 416 

ก่อแป้น Castanopsis diversifolia 416 

กอมขม Picrasma javanica 539 

ก่อลิ้ม 

Castanopsis indica 416 

ก่อสีเสียด Quercus brandisiana 416 

ก่อหม่น Lithocarpus elegans 416 

ก่อหยุม Castanopsis argyrophylla 416 

ก่อหรั่ง 

Castanopsis armata 416 

ก่อหัวหมู Lithocarpus sootepensis 416 

ก่อเหล็ก Lithocarpus vestitus 416 

ก่อแอบ Quercus vestita 416 

กะทังใบใหญ่ Litsea grandis 123 

กะอวม Acronychia pedunculata 555 

กะออก Artocarpus elasticus 392 

กางขี้มอด 

Albizia odoratissima 372 

กางหลวง Albizia chinensis 372 

ก้านเหลือง Gonocaryum lobbianum 694 

ก้ามกุ้ง 

Beilschmiedia velutinosa 120 

ก่ายสามใบ Melicope roxburghiana 557 

การบูรป่า Cinnamomum caudablm 122 

กาสามปีก Vitex peduncularis 668 

กำจัดต้น Zanthoxylum limonella 554 

กำยาน Styrax benzoides 605 

กุ๊ก 

Lannea coromandelica 519 

เก็ดดำ Dalbergia cultrata 354 

เกล็ดแรด Sterculia villosa 471 

ไก๋แดง Ternstroemia gymnanthera 597 

ขนุนปาน Artocarpus rigidus 392 

ขมิ้น 

Adina (Metadina trichotoma) 630 

ขางขาว Xanthophyllum flavescens 338 

ขี้หนอน 

Zollingeria dongnaiensis 533 

ขี้หนอนคาย 

Celtis tetrandra 387 

เข็มป่า Pavetta indica 640 

แข้งกวาง Wendlandia tinctoria 633 

แข้งกวางดง Wendlandia paniculata 633 

คอไก่ Tarennoidea wallichii 642 

คอแลน Nephelium hypoleucum 527 

ค่างหมอบ Psydrax glabum 636 

ค่าหด Engelhardtia spicata 419 

คำมอกหลวง Gardenia sootepensis 640 

คูน Cassia fistula 364 

เคียนทราย Shorea gratissima 448 

เคียนทราย Shorea gratissima 448 

เคี่ยมคนอง 

Shorea henryana 448 

แคยอดดำ Stereospermum colais 682

744 

ABROMA 468 

ABUTILON 487 

ACACIA 372 

ACALYPHA 275 

ACANTHUS 664 

ACER 522 

ACERATIUM 337 

ACROCARPUS 365 

ACRONYCHIA 555 

ACTEPHILA 256 

ACTINODAPHNE 123 

ADENANTHERA 367 

ADINA 630 

ADINANDRA 597 

ADONIDIA 143 

AEGIALITIS 161 

AEGICERAS 595 

AEGLE 561 

AESCULUS 524 

AFZELIA 350 

AGATHIS 65 

AGLAIA 552 

AGROSTISTACHYS 263 

AHERNIA 228 

AIDIA 641 

AILANTHUS 538 

ALANGIUM 562 

ALBIZIA 372 

ALCHORNEA 263 

ALECTRYON 533 

ALEISANTHIA 637 

ALEURITES 282 

ALLANTOSPERMUM 297 

ALLOPHYLUS 536 

ALNIPHYLLUM 606 

ALNUS 420 

ALPHITONIA 381 

ALPHONSEA 108 

ALSEODAPHNE 128 

ALSTONIA 650 

AMARACARPUS 625 

AMESIODENDRON 526 

AMHERSTIA 352 

AMMORA 552 

ANACARDIUM 506 

ANACOLOSA 168 

ANAXAGOREA 94 

ANDROTIUM 515 

ANERINCLEISTUS 202 

ANISOPHYLLEA 406 

ANISOPTERA 444 

ANNESLEA 596 

ANNONA 101 

ANOGEISSUS 182 

ANTHEROPORUM 359 

ANTHEROSTELE 626 

ANTIARIS 397 

ANTIDESMA 244 

ANTIRHEA 627 

APHANAMIXIS 552 

APHANANTHE 385 

APODYTES 615 

APORUSA 245 

AQUILARIA 430 

ARALIA 705 

ARALIDIUM 701 

ARAUCARIA 66 

ARCHIDENDRON 370 

ARDISIA 593 

ARECA 143 

ARENGA 141 

ARFEULLIA 525 

ARTHROPHYLLUM 706 

ARTOCARPUS 392 

ARYTERA 533 

ASCARINA 78 

ASHTONIA 249 

INDEX TO GENUS HEADINGS 

ASTROCALYX 200 

ASTRONIA 200 

ASTROTHALAMUS 403 

ATALANTIA 561 

ATALAYA 533 

ATUNA 302 

AULANDRA 583 

AUSTROBUXUS 240 

AVERRHOA 330 

AVICENNIA 664 

AXIANDRA 190 

AXINANDRA 207 

AZADIRACHTA 544 

AZIMA 425 

BACCAUREA 245 

BAECKEA 188 

BALAKATA 288 

BALIOSPERMUM 281 

BARRINGTONIA 572 

BAUHINIA 345 

BECCARIANTHUS 200 

BEILSCHMIEDIA 120 

BENNETTIODENDRON 228 

BERBERIS 148 

BERRYA 477 

BETULA 420 

BHESA 297 

BISCHOFIA 208 

BIXA 434 

BLACHIA 281 

BLASTUS 201 

BLEEKRODEA 390 

BLUMEODENDRON 264 

BOMBAX 488 

BONIODENDRON 526 

BORASSODENDRON 141 

BORASSUS 141 

BORNEODENDRON 283 

BOTRYOPHORA 266 

BOUEA 506 

BOUGAINVILLEA 162 

BRACHYTOME 643 

BRACKENRIDGEA 292 

BRANDISIA 688 

BRASSAIOPSIS 709 

BREYNIA 253 

BRIDELIA 255 

BROOKEA 687 

BROUSSONETIA 390 

BROWNLOWIA 478 

BRUCEA 539 

BRUGMANSIA 693 

BRUGUIERA 310 

BRUINSMIA 606 

BUCHANANIA 515 

BUCKLEYA 172 

BUDDLEJA 687 

BURKILLANTHUS 559 

BURRETIODENDRON 481 

BUTEA 359 

BUXUS 154 

CAESALPINIA 365 

CALLERYA 360 

CALLICARPA 675 

CALOCEDRUS 64 

CALOCHLAENA 60 

CALOPHYLLUM 320 

CALOTROPIS 657 

CAMELLIA 600 

CAMPNOSPERMA 511 

CAMPTOSTEMON 487 

CAMPYLOSPERMUM 292 

CANANGA 95 

CANARIUM 494 

CANTLEYA 697 

CAPPARIS 426 

CARALLIA 308 

CAREYA 576 

CARISSA 654 

CARLEMANNIA 663 

CARPINUS 420 

CARYODAPHNOPSIS 118 

CARYOTA 142 

CASEARIA 225 

CASSIA 364 

CASTANEA 416 

CASTANOPSIS 416 

CASUARINA 421 

CATUNAREGAM 640 

CECROPIA 405 

CEIBA 489 

CELASTRUS 215 

CELTIS 387 

CEPHALOMAPPA 275 

CERBERA 656 

CERIOPS 310 

CERISCOIDES 642 

CHAETOCARPUS 258 

CHAMPEREIA 170 

CHASSALIA 623 

CHEILOSA 261 

CHIONANTHUS 662 

CHISOCHETON 547 

CHLORANTHUS 78 

CHOEROSPONDIAS 519 

CHONDROSTYLIS 264 

CHORISANDRACHNE 256 

CHRYSOPHYLLUM 586 

CHUKRASIA 543 

CHYDENANTHUS 576 

CIBOTIUM 60 

CINNADENIA 128 

CINNAMOMUM 122 

CIPADESSA 547 

CITRONELLA 694 

CITRUS 558 

CLADOGYNOS 277 

CLAOXYLON 276 

CLAUSENA 558 

CLEIDIOCARPON 276 

CLEIDION 264 

CLEISTANTHUS 255 

CLERODENDRUM 672 

CLETHRA 604 

CLEYERA 599 

COCCULUS 147 

COCHLOSPERMUM 434 

COCOS 144 

CODIAEUM 279 

CODIOCARPUS 699 

COELOSTEGIA 486 

COFFEA 643 

COLONA 465 

COLUBRINA 381 

COMBRETOCARPUS 406 

COMBRETUM 182 

COMMERSONIA 469 

CONNARUS 329 

COPTOSAPELTA 644 

CORDIA 689 

CORDYLINE 134 

CORDYLOBLASTE 602 

CORYPHA 138 

COTYLELOBIUM 447 

CRAIBIODENDRON 610 

CRATEVA 426 

CRATOXYLUM 326 

CROTON 278 

CRUDIA 352 

CRYPTERONIA 206 

CRYPTOCARYA 119 

CTENOLOPHON 306 

CUBILIA 531 

CUPRESSUS 64

An Illustrated Guide to Diversity - Trees of Tropical Asia LaFrankie

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