FLORA MALESIANA SERIES I — SPERMATOPHYTA Flowering Plants Volume 12, part | Meliaceae Index to revised families in Series I (Spermatophyta) Aceraceae 4; 3, Actinidiaceae s.s. 4: Aizoaceae 4: Alismataceae 5: 317; 6: Alliaceae 11: Alseuosmiaceae 10: Amaranthaceae 4: 69, 593; 6: 915; 8: Amaryllidaceae 11: Anacardiaceae 8: Ancistrocladaceae 4: Aponogetonaceae 42013 7: Araliaceae—I Waite Araucariaceae 10: Aristolochiaceae 10: Balanophoraceae TS TOP Basellaceae 5: Bat(id)aceae Sp Betulaceae 5: 207; 6: Bignoniaceae 8: 114; 9: Bixaceae s.s. 4: Burmanniaceae ASOD St 552098 Burseraceae 5: 209, 6: 917; 7: 820; 9: Butomaceae BR Byblidaceae 1B Callitrichaceae 4: Campanulaceae 6: 107, 928; 9: Cannab(in)aceae 4: Cappar(id)aceae 6: 61; 7: Caprifoliaceae 4: 175; 6: 928; 9: Cardiopteridaceae a Celastraceae 6: 227, 389, Centrolepidaceae 5: Ceratophyllaceae 4: Chenopodiaceae 4: 99,594; 6: 932; 9: Chloranthaceae 10: Chrysobalanaceae 10: Clethraceae We Cochlospermaceae 4: Combretaceae 4: 533; 5: 564; 6: Coniferales 10: 592 37 267 915 375 335 Connaraceae 594955) 62/9333 19« Convolvulaceae 4: 388, 5: 558; 6: 7282359: Coriariaceae 11: Cornaceae 8: Corynocarpaceae 4: 262; 5: 4: 197; 9: Cruciferae 10: Crypteroniaceae 8: Ctenolophonaceae 10: Cupressaceae 10: Cyperaceae 7: 435; 9: Datiscaceae 4: Dichapetalaceae 5: 305; 6: Dilleniaceae 4: 141; 7: Dioscoreaceae 4: Dipsacaceae 4: Dipterocarpaceae 9: Droseraceae AS WTIS SA S112 Se Elaeagnaceae 10: Elatinaceae 4: Epacridaceae 6: Ericaceae 6: 469, 8: 549; 9: 562; 10: Erythroxylaceae 5: 543; 8: Fagaceae T2699: Flacourtiaceae Se jl 6: 943; 7: 827; 9: Flagellariaceae 4: 245; 9: Geraniaceae 6: 445; 9: Gnetaceae 4: 336; 6: Gonystylaceae 4: Goodeniaceae S5355 6: 949; 7: 827; 9: Haemodoraceae Ss Leo: Haloragaceae TBS, Hamamelidaceae Si Hippocrateaceae 6: Hydrocharitaceae SH 6: 952; 7: 9: 566; 10: Hydrophyllaceae 4: Crassulaceae LIBRARY be if a 936 558 385 85 557 558 541 187 629 442 107 382 941 824 293 290 237 562 151 203 422 943 716 549 563 565 563 564 565 944 349 567 566 717 828 363 389 381 828 PLT 207 Hypericaceae 8:35.10; Icacinaceae Tina lae Iridaceae 8: 77; 10: Ixonanthaceae 10: Juglandaceae 6: Juncaceae 4: 210; 9: Juncaginaceae 4: Labiatae 8: 301; 3: Legum.-Mimosoideae 11: Leeaceae Te Lemnaceae ie Lentibulariaceae 8: Liliaceae s.s. 9: Linaceae 10: Loganiaceae 6: 293, 9595 nee Lophopyxidaceae ai Magnoliaceae 10: Malpighiaceae Bi Martyniaceae 4: Meliaceae 12: Menispermaceae 10: Mimosaceae ie Monimiaceae 10: Moringaceae 4: Myoporaceae 4: Myricaceae 4: Najadaceae 6: Nyctaginaceae 6: Nyssaceae 4: Ochnaceae as Olacaceae 102-0 Onagraceae 8: Opiliaceae 10: Oxalidaceae FP sessile Papaveraceae 5: Passifloraceae We Pedaliaceae 4: 216; 7: Pentaphragmataceae 4: Pentaphylacaceae Sh Pentastemonaceae Tits Philydraceae 4: Phytolaccaceae 4: Pinaceae 10: Pittosporaceae 5: 345; 6: Plumbaginaceae 4: Podocarpaceae 10: all (continued on inside back cover) THE NEW YORK BOTANICAL GARDEN BRONX, NEW YORK 10458 FLORA MALESIANA SERIES I — SPERMATOPHYTA Volume 12 — part 1 — 1995 Meliaceae by D.J. Mabberley, C.M. Pannell, A.M. Sing This work was supported by the Netherlands Organization for Pure Scientific Research (ZWO), Greshoff’s Rumphiusfonds, and the Foundation Flora Malesiana in the Netherlands, and by the National Environment Research Council (NERC), the Science and Engineering Research Council (SERC), and the Leverhulme Trust in the United Kingdom. CIP-GEGEVENS KONINKLIJKE BIBLIOTHEEK, DEN HAAG Flora Flora Malesiana. Series I, Spermatophyta : Flowering plants. - Leiden : Rijksherbarium / Hortus Botanicus, Leiden University. Vol. 12, pt. 1: Meliaceae / by D.J. Mabberley, C. M. Pannell, A.M. Sing. - Ill. Comp. and publ. under the auspices of Foundation Flora Malesiana. - Met index, lit. opg. ISBN 90-71236-26-9 Trefw.: flora ; Zuidoost-Azié. All rights \reserved © 1995 Foundation Flora Malesiana No part of the material protected by this copyright notice may be reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, or by any information storage and retrieval system, without written permission from the copyright owner. FLORA MALESIANA SERIES I — SPERMATOPHYTA Volume 12 1995 — 1996 Published for Foundation Flora Malesiana by Rijksherbarium/ Hortus Botanicus * Leiden University * The Netherlands Contents Part 1 (1995) I SANS CTA fl 7 ARR a RA EP oh. 8. a 1-388 by D.J. Mabberley, C.M. Pannell & A.M. Sing index tO:SClENTIIC NAMES. aes De eee 8 2 ls pa 389-407 Part 2 (1996) Caesalpiniaceae (Leguminosae—Caesalpinioideae) ...... 409-730 by Ding Hou, K. Larsen & S.S. Larsen SCION OPICSIOCEGE id acne e SSP ee DOE 731-736 by J.E. Laferriére LCT HCNCLAGCE ACS ee A es hee I Ee 137-761 by B.E.E. Duyfjes POW ICCG eo aN oe ing RI cues EI 763-774 by K. Larsen TRG eT TOMSGICNINIC NOINES. shen 5s cA: 25 ee eee ae 775-784 if Ta 1) : hi Fa \oe ‘ wy “ni ww wis | fe / s0ee/ iti ef Am} ' AE | id: Ve ivy iv rs 5 7 Tra TTA S ‘ all) Vee Flora Malesiana ser. I, Vol. 12 (1) (1995) 1-407 MELIACEAE (D.J. Mabberley, C.M. Pannell (Aglaia) & A.M. Sing, Oxford University Herbaria, United Kingdom)! Meliaceae Juss., Gen. Pl. (1789) 263 (‘Meliae’); T.D. Penn. & Styles, Blumea 22 (1975) 419-540: Mabb. in FI. Nouv.-Caléd. et Dép. 15 (1988) 17—89; Mabb. & Pannell in Tree Fl. Malaya 4 (1989) 199-260. Cedrelaceae R.Br. in Flinders, Voy. Terra Austr. 2, App. (1814) 64 (‘Cedreleae’). Azedarachaceae Schultes in Roem. & Schultes, Syst. Veg. 5 (1819) xxxviii (‘familia Azedaracharum’). Aitoniaceae (Harv.) Harv., Fl. Cap. 1 (1860) 243 (‘Aitonieae’). Trees, treelets, often pachycaul or, more rarely, shrubs or suckering shrublets, mono- podial or sympodial, rarely with Terminalia-branching (Vavaea), dioecious, poly ga- mous, monoecious or with all flowers hermaphrodite. /ndumentum of simple, bifid or stellate hairs or stellate or peltate scales or sometimes mixtures of these, sometimes with small glands. Buds naked or with scale-leaves (in Malesia only in subfam. Swietenioi- deae). Leaves exstipulate (occasionally pseudostipules present), spirally arranged, rare- ly decussate, pinnate (sometimes with a terminal ‘bud’, i.e. pseudogemmula), trifolio- late, with a single blade (simple or unifoliolate) or rarely bipinnate (Melia); rachis very rarely winged; leaflets usually entire, rarely lobed or serrate (or spinous, not in Malesia), sometimes with minute black glandular dots. /nflorescences thyrsoid, racemose or spi- cate, sometimes reduced to fascicles or solitary flowers, axillary, supra-axillary, rami- florous, cauliflorous to groundlevel or rarely epiphyllous (Chisocheton). Flowers? her- maphrodite and/or more usually, unisexual, with well developed rudiments of opposite sex. Calyx usually + lobed, sometimes with discrete sepals, these occasionally spirally arranged and transitional to bracteoles (Dysoxylum), sometimes truncate or closed in bud and circumscissile at base at anthesis. Petals 3—7(—14), in 1 whorl, rarely (some Chisocheton species) in a spiral to give up to 2 (apparent) whorls, green, white, cream, pink to claret and violet or yellow (Aglaia). Stamens usually partially or completely united by a tube with or without lobes; anthers 3—10(—30) in 1 or, rarely, 2 or more whorls, sometimes locellate, at tips of filaments or at the margin of the tube or within its throat. Disk (at least sometimes nectariferous) around ovary, cushion-like, tubular or absent. Ovary (1—)2—6(—20)-locular, each locule with 1—many ovules; stylehead dis- coid to capitate. Fruit a capsule, berry or drupe. Seed with fleshy aril or sarcotesta or a combination of these or winged and then attached to a woody columella, or with corky outer layers, or very rarely without any of them, endosperm usually absent; cotyledons 1) With contributions by P. Baas, Leiden (vegetative anatomy), T.P. Clark, Oxford (Pseudoclausena, Walsura), J.M. Edmonds, Oxford (Toona), R.W.J.M. van der Ham, Leiden (palynology), R. Heg- nauer, Leiden (phytochemistry). The drawings are by Ms. Rosemary Wise at Oxford and Ms. Ruth van Crevel at Leiden. The photographs are from various sources, as indicated. 2) As it seems that male flowers may fall before females or hermaphrodites, it is often difficult in the absence of field studies to ascertain the true sexual arrangements of any particular species. Sizes of floral parts throughout the text are those of dried material, which sizes are often less than half those of fresh material. NO Flora Malesiana ser. I, Vol. 12 (1) (1995) collateral, superposed or, rarely, oblique, emergent or not at germination, when scale leaves are sometimes produced before first foliage leaves, which may be opposite or spirally arranged, simple or pinnate with later ones simple to bipinnate. 2n = 16—c. 360. INTRODUCTORY NOTE The bulk of this text was prepared by D.J.M. in 1982/83 at Leiden and, in conse- quence, largely follows the original style of Flora Malesiana. As there are no modern monographs of several of the genera, the text therefore provides the only available guide to those concerned with Meliaceae in these genera beyond the Flora Malesiana area. For that reason non-Malesian synonyms and references have been included. DISTRIBUTION Throughout the Tropics and Subtropics, with weak penetration into temperate zones, the family comprises 50—52 genera with about 550 species. Since Pennington & Styles’s generic monograph (1975), Megaphyllaea Hemsl. is considered to belong in Chisocheton Blume and Pseudocarapa Hemsl. in Dysoxylum Blume, while Heynea Roxb. ex Sims is considered distinct from Trichilia L. and Pseu- doclausena T.P. Clark has been segregated from Walsura Roxb.; Turraea breviflora may represent an undescribed genus and Naregamia Wight & Arn. may not be generically distinct from Turraea L., nor may Pseudobersama Verdc. from Trichilia L. The family is best represented in the Malesian region for, although Africa is almost as diversified in terms of the number of genera, the Malay Peninsula alone has more spe- cies (91 in 17 genera) than the whole of Africa (84 species) and, furthermore, begins to approach the specific richness of the whole neotropical region (122), where only eight genera are found, so that the Malesian region is over twice as rich in genera and almost as much so in species as the Neotropics. The largest genus, Aglaia, is centred on Ma- lesia, the great bulk of the rest of the species being in the closely interrelated genera Dysoxylum and Chisocheton. FOSSILS Fossils with features which, if they were found in a living plant, would place them in the concept of the modern Meliaceae are known from the Upper Cretaceous, some of the earliest ones being referred to the genus Guarea. Those referred to Cedrela are recorded from the Palaeocene (Raven & Axelrod 1974) and Pliocene (Graham 1991), the closely allied Indomalesian Toona is from the Eocene (London Clay) of England and those re- ferred to the African Entandrophragma C.DC. are recorded from the Miocene of Kenya (Chesters 1957). The volcanic basalt deposits of the Cretaceous/Tertiary boundary, the Deccan Intertrappean beds of Mandla District, have yielded fossils referred to Aglai- oxylon and Heyneoxylon, allied to Aglaia and Heynea respectively (Trivedi & Kiran Srivastava 1982), while ‘Carapa spp.’ (i.e. Xylocarpus spp.?) are known from the so- called peat bed near Calcutta, 22° 08'N (Ghosh & Negi 1958). See also Miller (1990). Mabberley et al. — Meliaceae 3 References: Chesters, K.I.M., Paleontographica B 101 (1957) 30-71. — Ghosh, S.S. & B.S. Negi, Curr. Sci. 27 (1958) 359-360. — Graham, A., Ann. Missouri Bot. Gard. 78 (1991) 201. — Miller, N.G., J. Arnold Arbor. 71 (1990) 453-486. — Raven, P.H. & D.I. Axelrod, Ann. Missouri Bot. Gard. 61 (1974) 587. — Trivedi, B.S. & Kiran Srivastava in Nautiyal (ed.), Phyton. Studies on living and fossil plants. Pant Commem. Vol. (1982) 255. HABITAT Meliaceae are very common trees of the canopy and understorey of lowland primary forest throughout Malesia, making up between 2.3% (Van Steenis 1984) and 17% of all trees over 10 cm bole diameter (Whitten et al. 1984) in the forests of Sumatra for exam- ple. They are absent from only the driest zones, though in Australia, for example, they are found even there, as species of Owenia F. Muell. They are represented by species of Xylocarpus on rocky shores and in mangrove swamps, into the upper reaches of which penetrates Aglaia cuculata. They are poorly represented at higher altitudes although some Dysoxylum species and Toona sinensis are sometimes conspicuous in lower mon- tane forest. The family is represented in freshwater swamp forest by Sandoricum bor- neense and Chisocheton amabilis and includes some species restricted to limestone, like Chisocheton ruber of Sarawak, while Walsura monophylla is restricted to ultramafics. Along rivers in west Malesia are a number of rheophytic species of Aglaia and Dysoxy- lum angustifolium while Aglaia cucullata is believed to be found along tidal riverbanks. Sandoricum borneense seems always to grow along riverbanks too. A few species are tolerant of more open conditions and will colonize large gaps in forest or are frequently encountered in secondary forest e.g. Aglaia argentea, Chisocheton tomentosus, Toona spp. and Chukrasia tabularis, which even colonizes bare ground along road cuttings in the Malay Peninsula. In Africa, the commercially significant swietenioid Meliaceae are emergent trees, some of the biggest in the continent and, in Java, 58-metre tall specimens of the melioid Dysoxylum mollissimum were some of the island’s tallest trees, but these have long been removed, like the commercial mahoganies of South America, now severely depleted, and the red cedar, Toona ciliata, in northeastern Australia. References: Steenis, C.G.G.J. van, Blumea 29 (1984) 405. — Whitten, A.J., et al., The ecology of Sumatra (1984) 262. POLLINATION Most species appear to be insect-pollinated, the agents possibly being bees, stingless sweatbees or syrphids in those cases examined (Aglaia spp., Xylocarpus spp.), while some species are strongly scented particularly in the evening, which, with their white flowers, suggests moth pollination in e.g. Chisocheton and Dysoxylum spp. as re- corded in neotropical Cedrela and Guarea spp. Some species of these two Old World genera have long flagelliform inflorescences suggesting bat pollination, though the Bornean species are known to be visited by spiderhunters; whether these birds are effi- cacious pollinators is not known. 4 Flora Malesiana ser. I, Vol. 12 (1) (1995) DISPERSAL The fruits of species of Swietenioideae are dry dehiscent capsules, the winged seeds of Toona and Chukrasia being wind-dispersed, the irregular corky-coated ones of Xylo- carpus being dispersed by saltwater in which they float just below the surface. The indehiscent fleshy fruits of Azadirachta indica are known to be dispersed by bats and baboons in Africa, those of Melia azedarach by birds. Those species of Aglaia, Chisocheton, Dysoxylum (see Whittaker & Turner 1994) and Aphanamixis with dehiscent capsules and arillate or otherwise fleshy seeds are known to be taken by birds, apparently attracted by the contrasting colours of seeds and/or peri- carp, while those species with indehiscent fruits are, at least in Aglaia and Reinwardtio- dendron, eaten by primates which pass the seed divested of its fleshy layers (Pannell & Koziot 1987). The bird and primate ‘syndromes’ are associated in a sample of Aglaia spp. at least, with characteristic chemistry as might be expected, those taken by birds being high in lipid, those by primates higher in free carbohydrates, but it is clear that few if any tree species is restricted to any one species, or indeed group, of vertebrate dispersers. Moreover, as there are no native primates in some parts of the ranges of certain species, Aglaia elaeagnoidea in New Caledonia for example, it must be con- cluded that bats or birds are active dispersal-agents even for those species which are thought to be dispersed by primates or other terrestrial mammals elsewhere. The seeds of many species are taken by squirrels but whether these animals do more than harm is unknown. For further details on Aglaia, see below. A parallel to the situation within Aglaia appears to be bird-dispersed Heynea and the closely allied but mammal-dispersed Walsura (Clark & Mabberley 1991). In New Guinea, birds of paradise are the only known dispersal agents for Chisocheton lasiocarpus (‘C. weinlandii’ ) seeds (Beehler 1983). Seeds of Dysoxylum angustifolium, and possibly Sandoricum borneense, are believ- ed to be dispersed by fish like those of the neotropical Guarea guidonia (L.) Sleumer (Pennington 1981). This ‘syndrome’ has yet to be analysed chemically. References: Beehler, B., Auk 100 (1983) 1. — Clark, T.P. & D.J. Mabberley, Nat. Monsp., h.s. (1991) 582. — Pannell, C.M. & M.J. Koziol, Phil. Trans. Roy. Soc. Lond. B 316 (1987) 303-333. — Pennington, T.D. in Fl. Neotrop., Monogr. 28 (1981) 11 [Swietenioideae by B.T. Styles]. — Whittaker, R.J. & B.D. Turner, J. Trop. Ecol. 10 (1994) 167-181. MORPHOLOGY AND ANATOMY Growth form — In terms of ‘architecture’, most Meliaceae correspond to the Models of Corner (unbranched, inflorescences lateral) and related structures with this pattern variously reiterated (Model of Champagnat) in the scheme of Hallé & Oldeman (1970), from pachycaul to leptocaul, while the genus Vavaea has Terminalia-branching (Model of Aubréville) and the Xylocarpus architecture corresponds to the Model of Rauh (Tom- linson 1986). References: Hallé, F. & R.A. A. Oldeman, Essai sur l’architecture ... (1970). — Tomlinson, P.B., The botany of mangroves (1986). Mabberley et al. — Meliaceae 5 Vegetative anatomy (P. Baas) — Leaf anatomy: Metcalfe & Chalk (1950) provide a sum- mary of the leaf anatomical range of the family based on the older literature. There are only very few more recent studies on the leaf anatomy of genera of Meliaceae which are also represented in Malesia (e.g., Farooqui 1981; Inamdar et al. 1986; Lersten & Pohl 1984). The indumentum includes simple unicellular hairs, two-armed hairs, stellate hairs, peltate scales, and variously shaped glandular hairs. Mucilage cells occur in the epidermis of Chukrasia and some other genera. Stomata are confined to the lower leaf surface and are usually anomocytic. Adaxial hypodermis is recorded in Aglaia and San- doricum. Mesophyll is dorsiventral. Vascular system of petiole and midrib is diverse, present as a single arc or closed cylinder, with or without medullary bundles. Secretory cells of various shapes and sizes occur in the mesophyll of many taxa. Crystals are com- mon, both solitary prismatic and clustered. A more comprehensive survey of the leaf anatomy of the Meliaceae is needed to assess its diagnostic and systematic significance. Wood anatomy: The wood anatomy of the Meliaceae is well documented. Key publi- cations are Moll & Janssonius (1908), Kribs (1930), Desch (1951), Metcalfe & Chalk (1950), Pennington & Styles (1975), and Ilic (1991). For a full annotated bibliography of the wood anatomy of the Meliaceae see Gregory (1994). The following concise de- scription of the woods of the Malesian genera is largely derived from these key publi- cations. Growth rings ranging from absent to distinct, and, if present, usually marked by mar- ginal parenchyma bands. Vessels mostly diffuse, but wood semi-ring-porous or ring- porous in Melia. Vessels of medium diameter to wide (maximum tangential diameter ranging from 80—320 um) and occurring in low frequency (2—20/sq.mm). Intervessel pits nonvestured, alternate, small (2—4 um) to medium-sized (4—8 um), vessel-ray and vessel-parenchyma pits usually similar but half-bordered. Helical vessel wall thickenings recorded in narrow vessels of Melia and Azadirachta. Tyloses absent. Dark-coloured vessel contents (gum) often present. Ground tissue of either non-septate or septate fibres with minutely bordered to simple pits mainly confined to the radial walls (libriform fibres), thin- to thick-walled (thick-walled in e.g. the heavy timbers of Dysoxylum, and Reinwardtiodendron). Parenchyma mainly paratracheal: vasicentric, aliform, confluent and/or banded, in various combinations. Apotracheal parenchyma absent or present as marginal (zonate) bands and/or diffuse strands. Rays narrow (1—2[—3]-seriate) in many genera of the Melioideae, but broader in Aglaia p.p., Azadirachta, Chisocheton p.p., Melia, Trichilia p.p., Sandoricum, and Walsura; rays in the Swietenioideae 3—6- (1—7-) seriate. Ray composition often varying from weakly heterocellular (Kribs Heterogeneous II) to homocellular within individual genera. Crystals mainly in axial (often chambered) parenchyma in the Melioideae; restricted to marginal ray cells in most Swietenioideae. Silica bodies recorded in Aphanamixis p.p., Azadirachta, Chisocheton, Dysoxylum p.p., and Trichilia p.p. Traumatic gum ducts forming tangential bands occasionally pres- ent, e.g. in Azadirachta, Dysoxylum, Melia, and Sandoricum. Rays and axial parenchy- ma storied in a few genera, e.g. Chukrasia and Xylocarpus. Pennington & Styles (1975) critically discussed the taxonomic value of the wood anatomical diversity in the Meliaceae and concluded that the secondary xylem provides 6 Flora Malesiana ser. I, Vol. 12 (1) (1995) good characters for subfamily delimitation, and that the association of non-septate fibres with terminal parenchyma bands helps to characterize the tribes Turraeeae, Melieae, Tri- chilieae (except Cipadessa) and Sandoriceae, whilst septate fibres in combination with absence or sporadic occurrence of marginal parenchyma characterize the remaining tribes. Many individual genera and species overlap in their wood anatomical range, so that only few genera can be distinguished by their wood anatomy alone. References: Desch, H.E., Manual of Malayan timbers Vol. 1, Malayan For. Rec. 15 (1951). — Farooqui, P., Indian For. 107 (1981) 237-242 (epidermis of Azadirachta). — Gregory, M., Biblio- graphy of systematic wood anatomy of Dicotyledons, [AWA J. Suppl. 1 (1994), [AWA/Rijksher- barium/Hortus Botanicus, Leiden. — Ilic, J., CSIRO Atlas of hardwoods, Crawford House & CSIRO, Melbourne (1991). — Inamdar, J.A., R.B. Subramaniam & J.S.S. Mohan, Ann. Bot. 58 (1986) 425 —429 (resin glands in Azadirachta). — Kribs, D. A., Amer. J. Bot. 17 (1930) 724 —738 (wood anat- omy). — Lersten, N.R. & R.W. Pohl, Ann. Bot. 56 (1984) 363 — 366 (extrafloral nectaries in Cipades- sa). — Metcalfe, C.R. & L. Chalk, Anatomy of the Dicotyledons 1 (1950). — Moll, J.W. & H.J. Janssonius, Mikrographie des Holzes der auf Java vorkommenden Baumarten, vol. 2 (1908) 110-215. — Pennington, T.D. & B.T. Styles, Blumea 22 (1975) 419-540 (wood anatomy and classification). Palynology (R.W.J.M. van der Ham) — Comparative accounts of the pollen morphol- ogy of the Meliaceae include those of Erdtman (1952; 18 genera), Chang & Wang (1956; 14 genera from China), Pennington & Styles (1975; 50 genera), Oltmann (1975; 12 gen- era from Madagascar) and Durdana Yunus & Nair (1989; 14 genera from India). That of Erdtman is very brief and hardly illustrated. Pennington & Styles offer short de- scriptions of nearly all genera, but they do no give any illustration or documentation of the extensive material studied. Durdana Yunus & Nair (1.c.), Straka & Friedrich (1984; 5 genera from Madagascar), El Ghazali (1993; Khaya, Pseudocedrela, Trichilia), Tissot et al. (1994; Aphanamixis, Trichilia, Walsura), and Bonnefille & Riollet (1980; Trichilia) provide the only scanning electron micrographs available to date. The Meliaceae are a stenopalynous family. The pollen grains are nearly always iso- polar, radially symmetric, suboblate to subprolate monads. Rarely the grains are oblate or prolate, and very rarely the pollen is shed in rhomboidal tetrads (Dysoxylum champt- onii Hook. f. & Thoms. ex Thw., Sri Lanka, and allies). The equatorial outline is + cir- cular to obtusely angular. The polar axis (P) is 13—62 um (mostly 25—40 um), the equa- torial diameter (E) is 1O—77 um (mostly 20—40 ppm). Pennington & Styles (1.c.) recorded much higher values for the average overall grain size (10-130 tm; in many genera > 40 uum). Relatively small pollen grains (P usually < 30 «m) occur in the tribe Aglaieae (e.g. Aglaia, Aphanamixis, Lansium, Reinwardtiodendron). The tribes Turraeeae (e.g. Munronia, Turraea), Melieae (e.g. Azadirachta, Melia) and Vavaeeae (Vavaea) show relatively large pollen (P > 40 um). The aperture system is tri-, tetra-, or pentacolporate, the tetra-aperturate condition being most common. Pollen of the tribes Turraeeae (except Munronia) and Vavaeeae is usually tricolporate. The ectoapertures are short to long meridional colpi; in Ag/aia the colpi are very short and indistinct. Tetracolporate pollen is rarely loxocolpate. The colpus ends are sometimes bifurcate. The endoapertures are circular to lalongate pori. Pollen of most genera has distinctly thickened ecto- and endoaperture margins (costae). Fastigiate apertures are known in several species of Munronia and Turraea. Mabberley et al. — Meliaceae 7) Exine thickness is 1—3 um. Little is known of the exine structure. The sexine and nexine can mostly be distinguished with light microscopy, but the infratectum is rather thin, and probably usually columellate. The ornamentation is mostly psilate to scabrate, with or without small perforations. Sometimes the exine surface is irregularly undulate to verrucate-rugulate or + reticulate (Carapa, Munronia, Trichilia, Xylocarpus). Finely striate ornamentation was observed in the pollen of Lansium (Durdana Yunus & Nair, l.c.) and striate ornamentation in that of Ekebergia (Straka & Friedrich, |.c.). As far as Meliaceae pollen is known, there are no significant differences between the subfamilies. Pennington & Styles (I.c.) considered prolate-spheroidal to subprolate, tetracolporate, psilate pollen constant in the Swietenioideae, but a literature survey has shown that oblate-spheroidal pollen, pentacolporate pollen and scabrate pollen also oc- cur in this subfamily (e.g. Erdtman, |.c.; Chang & Wang, I.c.; Sowunmi 1974; Dur- dana Yunus & Nair, I.c.). In a few cases pollen morphology proved to be valuable in transferring or segregating genera (see Pennington & Styles, |.c.). Turraea is a com- paratively eurypalynous genus, showing, besides meliaceous types, pollen unique in the family. Pollen morphology confirms that Nymania and Pterorhachis belong to the Meliaceae. Pollen like that of Cedrelopsis and Ptaeroxylon, which is tricolporate and distinctly reticulate with finely reticulate colpus margins (Leroy et al. 1990), does not occur in the Meliaceae, but is common in the Rutaceae and Simaroubaceae (Zanthoxylum type). This was one of the reasons for removing these genera from the Meliaceae (see Pennington & Styles, l.c.). Pollen with 4 or 5 colporate, costate apertures, like that of many melia- ceous genera, though usually with a + reticulate ornamentation, also occur in a number of genera of the subfamily Aurantioideae of the Rutaceae. Fossil pollen attributed to modern Meliaceae is known from a few Tertiary localities (Muller 1981). The 3—5-colporate pollen genus Meliapollis is not necessarily related to modern Meliaceae pollen (Thanikaimoni et al. 1984). References: Bonnefille, R. & G. Riollet, Pollens des savannes d’ Afrique orientale (1980). — Chang, K.T. & F.H. Wang, Acta Bot. Sinica 5 (1956) 253-266. — Durdana Yunus & P.K.K. Nair, Pollen morphology of Indian Geraniales (1989). — El Ghazali, G.E.B., Review Palaeobot. Palynol. 76 (1993) 99-345. — Erdtman, G., Pollen morphology and plant taxonomy (1952). — Leroy, J.-F., D. Lobreau- Callen & M. Lescot, Bull. Mus. Nat. Hist. Nat. Paris, IV, 12 (1990) 43—57. — Muller, J., Bot. Re- view 47 (1981) 1-142. — Oltmann, O., Pollen et Spores 17 (1975) 15-42. — Pennington, T.D. & B.T. Styles, Blumea 22 (1975) 419-540. — Sowunmi, M.A., Grana 13 (1974) 145-186. — Straka, H. & B. Friedrich, Trop. Subtrop. Pflanzenwelt 51 (1984) 27-43. — Thanikaimoni, G., C. Caratini, B.S. Venkatachala, C.G.K. Ramanujam & R.K. Kar, Trav. Sect. Sci. Tech. Inst. Fr. Pondichéry 19 (1984) 1-92. — Tissot, C., H. Chikhi & T.S. Nayar, Publ. Dép. Ecol. Inst. Fr. Pondichéry 35 (1994) 1-133. Seeds — The seeds of Meliaceae are some of the most diverse and intricate in structure so far investigated. The following is largely taken from Cheek’s thesis (1989). The seeds are usually pendulous and epitropous [but apotropous in the Australian Synoum (Guareeae)| in relation to the placenta. They are usually anatropous (but hemi-anatro- pous in most Turraeeae and Cipadessa), occasionally orthotropous as in all Chisocheton and some Guarea spp., campylotropous in e.g. Nymania (Turraeaeae), usually with a 8 Flora Malesiana ser. I, Vol. 12 (1) (1995) distinct funicle and raphe. Integument vascularization occurs in a few bitegmic genera, e.g. Chisocheton and Heynea (vascularized tegmen), Dysoxylum and Trichilia (vascu- larized testa) and is usual in pachychalazal seedcoats. In the Guareeae, arillate seeds characterize genera with bitegmic seeds, sarcotestal ones the unitegmic genera, while both conditions are found in Chisocheton and Dysoxylum, the genus with the greatest variation in seed anatomy investigated by Cheek. The single most taxonomically useful layer of the seedcoat in the majority of genera is the exotegmen: dimensions of the fibres and their wall characters but also the number of cell layers. However, in Dysoxylum, for example, the type of seed appendage and its organization are more helpful in recog- nizing species groupings as the exotegmen is reduced and poorly developed there. From his survey of the family, Cheek concludes that the ancestral meliaceous seed was com- paratively large and bitegmic, with a small chalaza and a funicular-raphe-aril. From such can be derived the unitegmic sarcotestal state and the unitegmic ‘pseudoarillate’ state seen in Malesian species. Reference: Cheek, M.R., The systematic seed anatomy of the Meliaceae. Unpubl. D.Phil. thesis Oxford Univ. (1989). CYTOTAXONOMY There is a wide range of somatic chromosome numbers from 2n = 16 (Sandoricum koe- tjape) to c. 360 in some African Trichilia spp., with polyploid series in some genera, e.g. Aphanamixis and Aglaia and within species, e.g. Chisocheton cumingianus. Much is summarized by Khosla & Styles (1975) and Styles & Khosla (1976) with further dis- cussion by Datta & Samanta (1977) and counts for Aglaia provided by Pannell (1992) and Chisocheton by Mabberley (1979). References: Datta, P.C. & P. Samanta, Cytologia 42 (1977) 197. — Khosla, P.K. & B.T. Styles, Silvae Genet. 24 (1975) 73-83. — Mabberley, D.J., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 301— 386. — Pannell, C.M., Kew Bull., Add. Series 16 (1992). — Styles, B.T. & P.K. Khosla, Cytol. & reprod. biol. of Meliaceae, in: J. Burley & B.T. Styles (eds.), Tropical trees: variation, breeding and conservation (1976) 61—67. PHYTOCHEMISTRY AND CHEMOTAXONOMY (R. Hegnauer) Meliaceae belong to Rutales s.str. comprising Rutaceae, Meliaceae, Simaroubaceae, Cneoraceae and Burseraceae (Waterman & Grundon 1983). A chemical key character of this order (if Burseraceae are excluded) is the synthesis and accumulation of bitter and biologically active nortriterpenoids called limonoids, meliacins and quassinoids depend- ing on structural features and occurrence. These oligonortriterpenoids derive from tetra- cyclic triterpenes which are known as protolimonoids. Moreover, each family of this order has its own main patterns of secondary metabolites. Recently, Meliaceae have in- terested many phytochemists and biologists, because they yield many compounds with insect-antifeeding, insect-repellent or/and insecticidal properties, and because many members of this family are highly esteemed in traditional medicine of most continents. Mabberley et al. — Meliaceae 9 A number of phytochemical review treatments of Meliaceae are available in recent literature, e.g. Hegnauer 1969, 1983, 1990; Waterman & Grundon 1983. Furthermore, many reviews are devoted to the chemically complex limonoids, which in the case of Meliaceae are sometimes called meliacins; see e.g. Arnason et al. 1989, 1993: Cham- pagne et al. 1992; Ley et al. 1993; Rembold 1983, 1989; Rembold & Puhlman 1993; Saxena 1989; Taylor 1983; Triterpenes 1984-1994. References alluded to in the above reviews are only exceptionally repeated in the fol- lowing phytochemical survey in which metabolites of Meliaceae are arranged following Hegnauer (1983). A. Essential oils — Idioblasts containing oleo-resins seem to be ubiquitous. Usually these excretory cells contain mono- and sesquiterpenoids as their main steam-volatile constituents. Some of the sesquiterpenoids, like the aphanamols of Aphanamixis poly- stachya (A. grandifolia), a 3-oxo-a-bourbonene derivative from Lansium domesticum and 8-hydroxycalamenene from seeds of Dysoxylum spp. have piscicidal properties. Dysoxylum spectabile (A. Juss.) Hook. f. from New Zealand produces a fruit oil con- taining an ant-repellent sesquiterpenoid which can be interpreted as a hydrated B-santa- lene (Russell et al. 1994). Some taxa have a distinct onion- or garlic-like smell and are assumed to produce volatile sulphur compounds. In fact, cyclic tri- and tetrasulfides have been isolated from fresh leaves of Azadirachta indica and leaves of Dysoxylum mollissimum subsp. molle (D. richii) have yielded dysoxysulfone, C5H204Ss. B. Arylpropanoids and arylpropanoid-dimers (lignanoids) — The ubiquitous cin- namic acids are not lacking in Meliaceae, but usually they are not accumulated in large amounts, neither free nor as esters. p-Coumaric and caffeic acids have been detected in several species, and Azadirachta indica produces methyl grevillate; grevillic acid is 2,5- dihydroxycinnamic acid. Cinnamic acid itself occurs sometimes as an amide (see below, sub G—H) or esterified with limonoids. Contrary to what occurs in some Rutaceae, Meliaceae obviously do not accumulate appreciable amounts of steam-volatile phenyl- propanoids in their essential oils. The methylether of lariciresinol, a lignan, has been isolated from leaves of Turraea nilotica Kotschy & Peyr. (Africa), and small amounts of a benzofuranoid neolignan having insect-antifeedant properties and called melianoninol has recently been isolated from fruits of Melia azedarach (Xu et al. 1992). C. Acetogenins (= Polyketides) — Accumulation of pure acetogenins is exceptional in the family. 5,7-Dihydroxy-2-methylchromone (= noreugenin, a pentaketide) occurs free and combined in 8-position with 3-hydroxy-N-methylpiperidine to the alkaloid rohi- tukine in leaves of Aphanamixis polystachya (Amoora rohituka). 4-Methoxy-5-methyl- coumarins are assumed to be produced along the polyketide pathway. Ekersenin from Ekebergia capensis Sparrm. (E. senegalensis) and siderin from Toona ciliata (Cedrela toona) belong to this group and are probably also pentaketides. D. Coumarins — True coumarins are lactones of cis-2-hydroxycinnamic acids; they represent a characteristic group of arylpropanoids. Simple coumarins like umbelliferone, aesculetin, scopoletin, scoparone and isofraxidin have been isolated from several species. Rutaceous-type derivatives of isoprenylated coumarins, however, are of rare occurrence in Meliaceae. Such coumarins have been isolated recently from seeds of a variety of 10 Flora Malesiana ser. I, Vol. 12 (1) (1995) Toona ciliata (bergaptene, a furanocoumarin) and from bark of Ekebergia capensis (xan- thoxyletin, a 2,2-dimethylchromenoid coumarin). E. Flavonoids — Flavonoids are of mixed biogenetic origin (p-coumaric acid + 3 ace- tates). Their patterns depend on plant parts and taxa. The patterns known from Rutales (including Meliaceae) were discussed by Harborne (1983). The flavonol quercetin is of common occurrence in Meliaceae; it is sometimes accompanied by kaempferol and/or myricetin. Rutin, the 3-rutinoside of quercetin, has been isolated from several taxa. O-Polymethylation and C-prenylation of flavonoids, which are characteristic of many rutaceous plants, seem to be rather exceptional in Meliaceae. Examples are syringetin and 2,3-dihydrosyringetin from wood of Soymida febrifuga (Roxb.) A. Juss. (India), tabularin (= 5,7-dihydroxy-6,2’,4’,5’-tetramethoxyflavone) from leaves of Chukrasia tabularis and the 8,3’-diprenylated 5,7-dihydroxy-4’-methoxyflavanone from leaves of Azadirachta indica. A C-methylated flavonoid, the 3-xyloside of 8-methylquercetin, has been isolated from root bark of Aphanamixis polystachya. A new type of flavonoids has recently been reported from the bark of Aglaia tomentosa (A. ferruginea) by Dean et al. (1993); ferrugin, Cog6H2606, can be interpreted as a condensation product of 5,7- dimethoxyisoflavone with a Cg-Co- or a decarboxylated C¢-C3-unit; if this biogenetic assumption is correct, it represents an isoflavano-norlignan. Hitherto flavones have sel- dom been observed in the family. The sporadic occurrence of flavones is illustrated by the aforementioned tabularin and by the apigenin glycosides of Melia azedarach (Tang & Eisenbrand 1992). Flavanones and chalcones are also rare Meliaceae constituents. F. Tannins and tannin precursors — Many Meliaceae are tanniferous plants. Tannins accumulate mainly in barks and roots and belong to the proanthocyanidin group (= con- densed tannins). Some species produce barks with up to 20% tannins and are used lo- cally for tanning purposes. Catechins and gallocatechins have been isolated from barks of Dysoxylum spectabile, Melia azedarach and Azadirachta indica, and wood of Toona ciliata (Cedrela toona) has yielded a procyanidin. Bark of Azadirachta indica also con- tains gallic acid. This suggests that condensed tannins may be accompanied by gallo- tannins in some taxa. Ellagic acid and ellagitannins have not yet been detected in the family. G-H. Protoalkaloids and alkaloids — \n contrast to Rutaceae no family-specific types of alkaloids are known from Meliaceae. In this summary of present knowledge seven types of alkaloid-like substances will be discussed below. True alkaloids are re- presented by four distinct types. 1) Ekeberginine from Ekebergia capensis (E. senegal- ensis) is a carbazole alkaloid; it is a position isomer of indizoline of Clausena indica Oliv. (Rutaceae). Ekeberginine and xanthoxyletin mentioned sub coumarins strengthen the biochemical link between Rutaceae and Meliaceae indicated by limonoids. 2) The antifeedant 2-chinolone alkaloid N-methylflindersine from roots of Xylocarpus grana- tum forms another biochemical link between Rutaceae and Meliaceae, just as does 3) acetonyldihydrochelerythrine, a phenanthridine alkaloid isolated from the same plant material. 4) Alkaloids of Dysoxylum lenticellare Gillespie (Fiji), e.g. dysoxyline, 3-epi- schelhammericine, dysazecine and others all belong to the phenylethylisoquinoline group of alkaloids occurring erratically in Spermatophytes. A recent addition to these alkaloids Mabberley et al. — Meliaceae 1] of D. lenticellare are the homoerythrinane 2a-methoxycomosivine and the secohomo- erythrinanes or lenticillaranes lenticillarine, 2-hydroxy- and 2-methoxylenticillarine (Aladesanmi et al. 1994). The remaining three types of nitrogen-containing secondary metabolites are rather alkaloid-like compounds than true alkaloids. 5) Pyridine deriva- tives such as 3-hydroxypyridine isolated in trace amounts from the bark of Entandro- phragma cylindricum (Sprague) Sprague (Africa) and two diesters of the limonoid phragmalin with one molecule of nicotinic acid (giving basic properties to these esters) and one molecule of isobutyric acid isolated from the bark of E. caudatum Sprague. 6) Complex cinnamic acid amides of Aglaia species, e.g. pyrimidatin, Co9H22N20o, a bisamide of putrescine, from leaves of A. silvestris (A. pyramidata); related bisamides are piriferine from A. edulis (A. pirifera) and odorine, 5’-epiodorine and odorinol from A. odorata and A. elaeagnoideae (A. roxburghiana) (Saifah & Puripattanavong 1993). Rocaglamide, a more complex amide, was originally isolated from A. rimosa (A. ellip- tifolia) and later together with the similar compounds desmethylrocaglamide, rocaglaol and methyl rocaglate from A. odorata (Janprasert et al. 1993; Ishibashi et al. 1993). Roots of A. odorata yielded a still more complex rocaglamide-type compound, C3;H2gN2O¢, which contains a pyrimidinone ring, probably derived from the putrescine amide of rocaglaic acid (Kokpol et al. 1994). Rocaglamide-type compounds have insecticidal properties. The genus Aglaia is believed to be very promising for the development of new insecticides of plant origin (Arnason et al. 1993; Satosook et al. 1994). 7) The last type of alkaloid-like compounds of Meliaceae was already mentioned sub C, i.e. rohi- tukine. Alkaloid and alkaloid-like constituents seem to occur rather frequently in the family; most of them have not yet been investigated. Present knowledge suggests that these compounds characterize individual species and genera rather than larger taxonomic entities. I. Bitter tetranortriterpenoids (limonoids or meliacins) and their tetracyclic triterpen- oid precursors — Probably every meliaceous plant accumulates one or several types of limonoids. Agostinho et al. (1994) recently discussed the infra-familiar taxonomic mean- ing of meliacins (compare also Silva et al. 1984 and 1988) and their possible role in plant defence. References to this much studied class of compounds have already been given in the introductory notes. Some additional information is included in the follow- ing remarks. Precursors of meliacins or limonoids are euphane(tirucallane)-type tetra- cyclic triterpenoids like odoratone from Cedrela odorata or melianone and melianol from Melia azedarach; they have a polyoxygenated side-chain at C-17. Limonoid biosynthesis starts with the so-called apo-rearrangement, i.e. the migration of a double bond and one methyl group. Examples of such apo-euphanoid or apo-tirucallanoid protolimonoids are grandifoliolenon from wood of Khaya grandifolia C. DC. (Africa), 21-O-acetyltoosen- dantriol from Melia azedarach (M. toosendan) (Tang & Eisenbrand 1992) and the dyso- rones from leaves of the New Caledonian Dysoxylum roseum (Baillon) C.DC. (Ade- sanya et al. 1991). The next biogenetic step leading to limonoids is an oxidative loss of four C-atoms of the side-chain (C-24 to C-27). Much chemical, chemo-ecological and pharmacological work has been performed with limonoids of the neem tree (= margosa tree) (Van der Nat et al. 1991; Stone 1992; Govindachari 1992). Recently one of its 12 Flora Malesiana ser. I, Vol. 12 (1) (1995) main anti-insect principles, azadirachtin-A was got into crystalline form. This made it possible to confirm its highly complex structure by X-ray diffraction studies (Govinda- chari et al. 1994; Kabaleeswaran et al. 1994). Neem seed oil also contains limonoids with antifeedant activity against white ants (Reticulitermes speratus: Ishida et al. 1992). Six Chisocheton species cultivated in the Botanic Gardens Bogor, Java, have been investigated for their insect-deterrent properties. Five of them furnished leaf and twig extracts with antifeedant activity; two meliacins with a y-lactone ring were isolated from C. pentandrus (C. microcarpus) (Gunning et al. 1994). Tang & Eisenbrand (1992) re- viewed the chemical constituents and their biological properties of Melia azedarach (incl. M. toosendan). New limonoid antifeedants have been isolated from root bark of Chinese plants and from stem bark of plants of Melia azedarach growing in Okinawa (Nakatani et al. 1994). Limonoids of Trichilia species have been shown to inhibit growth of lepidopteran larvae when incorporated in their diet and after topical application; one of the active principles of 7. hirta L. (America) was identified with hirtin (Satasook et al. 1994). Xylocarpus granatum and X. rumphii (*X. moluccensis’) growing in Fiji yield- ed the new limonoids xyloccensin-I and -J (Alvi et al. 1991). J. Diterpenoids and triterpenoids — Isoprenoid metabolism of Meliaceae yields li- monoids and protolimonoids in the first instance, but diterpenes and ‘normal’ triter- penes are by no means lacking. The acyclic diterpene geranylgeraniol has been isolated from the wood of Toona ciliata (Cedrela toona). Diterpenoids have also been isolated from barks of Azadirachta indica (sugiol, nimbiol, a bisnorditerpene, and nimbidiol, a trisnorditerpene), from leaves of Aphanamixis polystachya (aphanamixol) and from leaves of Dysoxylum lenticellare (phyllocladene, 8-hydroxysandaracopimarene, ferru- ginol and ferrubietolide, a dimeric abietane-type diterpene). The non-protolimonoid triterpenes are represented in the family by several types. The sapelenins of Entandro- phragma cylindricum are tetra- or hexahydroxy derivatives of partially hydrogenated squalene; sapelenins-A and -C are monoacetates (Ngnokam et al. 1993). The tetracyclic triterpenes are represented in the family by three types. Heyneic acid from Heynea tri- juga, kulinone and kulactone from Melia azedarach and the rather widespread cycloarte- nol and 24-methylenecycloartanol belong to the euphane-tirucallane group. The dam- marane group is represented by aglaiol from leaves of Aglaia odorata and richenone, richenol, richenoic acid and five other compounds isolated from leaves of Dysoxylum mollissimum subsp. molle (D. richii); richenoic acid, eichlerianic acid and shoreic acid are A-ring seco-dammaranes (Aalbersberg & Singh 1990). Ocotillone, cabraleone and eichlerianic acid and other dammarane-type triterpenes had previously been isolated from the wood of Cabralea canjerana (Vell.) Mart. (C. eichleriana) and subsp. polytri- cha (A. Juss.) T.D. Penn. (C. polytricha). Recently, stem bark of Aglaia leucophylla (Malaysia) yielded ocotillone, cabraleone, ocotillol, cabralealactone, eichlerianic acid and a new 3,4-seco-tirucallane-type C3° diacid and its monomethyl! ester (Benosman et al. 1994). Onocerane-type tetracyclic triterpenes occur erratically in tracheophytes. In Meliaceae they are represented by onoceradienone and the seco-derivatives lansic and lansiolic acids (Lansium domesticum). The more or less ubiquitous pentacyclic triter- penes of the lupeol- and a- and B-amyrin-series have been hitherto isolated from a rela- Mabberley et al. — Meliaceae 13 tively small number of Meliaceae. Examples are katonic acid from wood of Lansium domesticum, walsurenol from leaves of Walsura tubulata Hiern (India), a- and B-amy- rin, hederagenin and lupeol benzoate from wood of Swietenia mahagoni and betulin and betulinic acid from roots of Amoora canarana (Turcz.) Hiern (= Aglaia lawii). K. Saponins — The usual saponins are aphrogenic and hemolytically active glyco- sides with one or two sugar chains attached to an alcoholic hydroxyl of an aglycone (= sapogenin). Sapogenins are triterpenoids or steroids (usually Co7). Sapogenins com- bined with a single hexose or pentose are saponin-like compounds rather than true sa- ponins. Common phytosterols, e.g. sitosterol and stigmasterol, and polyhydroxy- pregnanes also occur frequently as monosides or biosides and may have saponin-like properties. Present knowledge suggests that true saponins are of rare occurrence in Meliaceae, but that saponin-like constituents are rather common. Examples of such saponin-like glycosides are 3-mono- and biosides of stigmastadienol, poriferasterol and aglaiol (Aphanamixis polystachya), the lansiosides of fruits of Lansium domesticum and a norcycloswietenol-3-glucoside from wood of Swietenia mahagoni. New additions to saponin-like glycosides of Meliaceae are the cumingianosides of Dysoxylum cumingianum; they are acetylated monoglucosides of tetracyclic triterpenes with a 14,18-cyclo-apoeuphane-type skeleton and are accompanied by a similar trisnor-com- pound (cumindysoside-A; C25—Co7 lost) and tetranor-compound (cumindysoside-B; Co4—Co7 lost) (Kashiwada et al. 1992). Toosendanoside, the 2-glucoside of a tetra- hydroxypregnane from Melia azedarach (M. toosendan) (Nakanishi et al. 1988) seems to belong to the same group. Recently a number of glucosides or glucuronides of limonoids have been detected in the family (e.g. Tang & Eisenbrand 1992). Glycosidic limonoids (or meliacins) may also possess saponin-like properties. L. Unclassified secondary metabolites — A number of erratically occurring com- pounds should be mentioned here. p-Hydroxyacetophenone was isolated from stems of Dysoxylum lenticellare. Bark of Khaya senegalensis A. Juss. (Africa) contains the aller- genic yellow quinone 2,6-dimethoxy-p-benzoquinone which occurs in many members of Simaroubaceae. Glycosylated anthraquinones of uncertain biogenetic origin have been isolated from Aphanamixis polystachya and Melia azedarach. M. Mucilages — Gummosis is not uncommon in Meliaceae. Gums exuded by Aza- dirachta indica, Carapa procera DC. (tropical America & Africa), Cedrela odorata and several Khaya species have been studied chemically. They are acidic heteropolysaccha- rides which are partially acetylated and release easily acetic acid during purification. The gum exudate of the neem tree has been thoroughly investigated and shown to be a proteo- glycane (heteropolysaccharides firmly combined with proteins) (Van der Nat et al. 1991). N. Storage products of seeds — Many Meliaceae produce endospermless or endo- sperm-poor seeds which have their reserves stored in the cotyledons which are also rich in large excretory cells. There is no family-characteristic pattern of seed storage prod- ucts. Many species store mainly oil and proteins. Others have predominantly oil or pre- dominantly proteins. Starch is absent in ripe seeds of many taxa, but was reported to be present in large amounts in ‘Amoora’ (Netolitzki 1926). The chemistry of seed oils varies within the family. Some taxa, notably Entandrophragma (three species investi- 14 Flora Malesiana ser. I, Vol. 12 (1) (1995) gated) have vaccenic acid as a main component. Seed oils of Melia (3 species) contain more than 60% of linolenic acid, and a number of taxa have seed oils with over 50% of saturated fatty acids (12:0—18:0). Obviously fatty acid patterns of seed oils are rather characteristic at infrageneric and generic levels than at suprageneric levels. O. Inorganic compounds (see also the paragraph on Vegetative anatomy, p. 5) — Some Meliaceae accumulate small to moderate amounts of silica. Silica deposition in leaves has been reported for six species of Cedrela and for one species of each of the genera Soymida, Swietenia, Carapa and Xylocarpus; 25 species had leaves without sili- cated cell walls (Edman 1936). In woods of some species silica bodies occur in ray and parenchyma cells. Such woods contain between 0.05 and 3.4% of silica and belong to the genera Aphanamixis, Chisocheton, Dysoxylum (exceptionally), Entandrophragma (p.p.), Guarea (p.p.) and Trichilia (most species) (Amos 1952; Savard et al. 1954; Ter Welle 1976). The character may have some taxonomic importance up to generic level. Chemotaxonomy — By their highly versatile limonoid metabolism Meliaceae are clearly related to Rutaceae and other families of Rutales. Each family of Rutales, how- ever, has distinct features of secondary metabolism. Meliaceae, for example, lack totally or nearly totally classes of compounds present in many Rutaceae (prenylated coumarins; anthranilic acid derived alkaloids) and Simaroubaceae (canthinone-type alkaloids). For infrafamiliar taxonomy the limonoids (or meliacins) offer possibilities. Taxonomists should be very cautious and careful, however, in using them, because our knowledge is still extremely fragmentary. Far-reaching phylogenetic conclusions (Silva et al. 1984; Agostinho et al. 1994) are premature. Everyone who intends to make taxonomic use of secondary metabolites in Rutales should start with reading Waterman (1990). References: Aalbersberg, W. & Y. Singh, Phytochemistry 30 (1991) 921. — Adesanya, S.A., et al., J. Nat. Prod. 54 (1991) 1588. — Agostinho, S.M.M., et al., Limonoids from Toona ciliata and speculations on their chemosystematic and ecological significance, Biochem. Syst. Ecol. 22 (1994) 323-328. — Aladesanmi, A.J., et al., Phytochemistry 35 (1994) 1361. — Alvi, K.A., et al., Tetra- hedron 47 (1991) 8943. Amos, G.L., Silica in timbers, Bull. No. 267 CSIRO, Australia, Mel- bourne (1952) 55 pp. — Arnason, J.T., et al. (eds.), Insecticides of plant origin, ACS Symposium Series 387, Amer. Chem. Soc. Washington (1989). — Arnason, J.T., et al., Insecticides in tropical plants with non-neurotoxic modes of action, pp. 107-131 (Meliaceae pp. 112—124) in K.R. Downum et al. (1993). — Benosman, A., et al., Phytochemistry 37 (1994) 1143. — Champagne, D.E., et al., Insecticidal activity of phytochemicals and extracts of the Meliaceae, pp. 95-109 in J.T. Arnason et al. (1989). — Champagne, D.E., et al., Biological activity of limonoids from the Rutales. Review article nr. 65, Phytochemistry 31 (1992) 377-394. — Dean, F.M., et al., Phytochemistry 34 (1993) 1337. — Downum, K.R., et al. (eds.), Phytochemical potential of tropical plants, vol. 27 (1993) of Recent Advances in Phytochemistry. — Edman, G., Svensk Bot. Tidskr. 30 (1936) 493. — Govindachari, T.R., Chemical and biological investigations on Azadirachta indica (the neem tree), Current Science 63 (1992) 117-122. — Govindachari, T.R., et al., Current Science 66 (1994) 295. — Gunning, P.J., et al., Phy- tochemistry 36 (1994) 1245. — Harborne, J.B., The flavonoids of the Rutales, pp. 147-173 in P.G. Waterman & M.F. Grundon (1983). — Hegnauer, R., Chemotaxonomie der Pflanzen 5 (1969) 54-71, 421-428, 456; 9 (1990) 31-49. — Hegnauer, R., Chemical characters and the classification of the Rutales, pp. 401—440 in P.G. Waterman & M.F. Grundon (1983). — Ishibashi, F., et al., Phytochem- istry 32 (1993) 307. — Ishida, M., et al., Biosci. Biotech. Biochem. 56 (1992) 1835. — Janprasert, J., et al., Phytochemistry 32 (1993) 67. — Kabaleeswaran, V., et al., Current Science 66 (1994) 362. — Kashiwada, Y., et al., J. Org. Chem. 57 (1992) 6946. — Kokpol, U., et al., J. Chem. Soc., Chem. Commun. (1994) 773. — Ley, S.V., et al., The chemistry of azadirachtin, Natural Product Reports 10 Mabberley et al. — Meliaceae 15 (1993) 109-157, 242 Ref. — Nakanishi, T., et al., Chem. Pharm. Bull. 36 (1988) 4148. — Naka- tani, M., et al., Phytochemistry 36 (1994) 39; Heterocycles 38 (1994) 327.— Nat, J.M. van der, et al., Ethnopharmacognostical survey of Azadirachta indica A. Juss. (Meliaceae), J. Ethnopharmacol. 35 (1991) 1-24. — Netolitzki, F., Anatomie der Angiospermen-Samen, Linsbauers Handbuch der Pflan- zenanatomie, Bd. 10, Borntraeger, Berlin (1926). — Ngnokam, D., et al., Phytochemistry 34 (1993) 1603. — Rembold, H., The azadirachtins — their potential for insect control, Economic Medicinal Plant Research 3 (1983) 57-72 (Academic Press). — Rembold, H., Azadirachtins. Their structure and mode of action, pp. 150-163 in J.T. Arnason et al. (1989). — Rembold, H. & I. Puhlman, Phytochemistry and biological activity of metabolites from tropical Meliaceae, pp. 153-165 in K.R. Downum et al. (1993). — Russell, G.B., et al., Phytochemistry 35 (1994) 1455. — Saifah, E. & J. Puripattanavong, J. Nat. Prod. 56 (1993) 473. — Satasook, et al., Insecticidal bioactivity of crude extracts of Aglaia species, Biochem. Syst. Ecol. 22 (1994) 121-127. — Savard, J., et al., Analyse chimique des bois tropicaux, Centre Technique Forestier Tropical, Nogent-sur-Marne, France (1954) 140, 166. — Saxena, R.C., Insecticides from neem, pp. 110-135 in J.T. Arnason et al. (1989). — Silva, M.F.das G.F. da, et al., Evolution of limonoids in the Meliaceae, Biochem. Syst. Ecol. 12 (1984) 299-310. — Silva, M.F.das G.F. da, et al., Chemosystematics of the Rutaceae: suggestions for a more natural taxonomy and evolutionary interpretation of the family, Pl. Syst. Evol. 161 (1988) 97-134. Compare Waterman (1990). — Stone, R., A biopesticidal tree begins to blossom, Science 255 (1992) 1070-1071. — Tang, W. & G. Eisenbrand, Chinese drugs of plant origin. Chemistry, pharmacology, and use in tradi- tional and modern medicine, Springer-Verlag, Berlin (1992), 647-658. — Taylor, D.A.H., Biogenesis, distribution and systematic significance of limonoids in the Meliaceae, Cneoraceae and allied taxa, pp. 353-375 in P.G. Waterman & M.F. Grundon (1983). — Triterpenes: Literature Reviews (incl. tetra- nortriterpenoids), Natural Product Reports | (1984) 53-65 (R.B. Boar); 2 (1985) 1-17; 3 (1986) 421- 442; 6 (1989) 475-501; 11 (1994) 91-117 (J.D. Connolly et al.). — Waterman, P.G., Chemosystem- atics of the Rutaceae: comments on the interpretation of da Silva et al., Pl. Syst. Evol. 173 (1990) 39— 48. Compare Silva, M.F.das G.F. da, et al. (1988). — Waterman, P.G., Phytochemical diversity in the order Rutales, pp. 203-233 in K.R. Downum et al. (1993). — Waterman, P.G. & M.F. Grundon, Chemistry and chemical taxonomy of Rutales, Academic Press, London (1983). — Welle, B.J.H. ter, Silica in woody plants of the Neotropics especially Surinam, pp. 107—142 in P. Baas et al. (eds.), Wood structure in biological and technical research, Leiden Bot. Series No. 3, Univ. Press, Leiden (1976). — Xie, Y.S., et al., Biochem. Syst. Ecol. 22 (1994) 129-136. — Xu, R.-S., et al., J. Chinese Pharm. Sci. 1 (2) (1992) 7; ex Natural Product Updates No. 16949 (1994). TAXONOMY Historical — This is summarized by Pennington & Styles (1975). As far as Malesia is concerned, the study of the family has been dominated by Miquel (1868), whose ‘Mon- ographia meliacearum archipelagi indici’ is a pioneering example of the monographic method he championed, and Harms (1896, 1940), who prepared accounts for both editions of Engler & Prantl’s Die Natiirlichen Pflanzenfamilien and described many dozens of Malesian species, particularly from the German possessions in New Guinea. His work gave a firm foundation for Pennington & Styles’s generic monograph of the family (1975), largely based on Pennington’s doctoral thesis of 1965 in the University of Oxford, which has been a centre of study for this family for several decades. Much remains to be studied, however, particularly in the fields of reproductive and dispersal biology (see above). Even at the alpha-taxonomic level, local workers can still contri- bute a great deal: collectors should be encouraged to collect flowers and ripe fruits in spirit, to note the largest size of leaves and the number of leaflets, to be sure to collect apical buds and to try to characterize the smell of broken branches. 16 Flora Malesiana ser. I, Vol. 12 (1) (1995) Present arrangement — The family is divisible into four subfamilies. A putative fifth, the monotypic Neomangenotioideae Leroy, from Madagascar, has been referred to Com- miphora (Burseraceae); see Cheek & Rakotozafy (1991). Two of the subfamilies, both monotypic, are restricted to Madagascar and to a certain extent provide links between the other two which are by far the largest: Melioideae and Swietenioideae, both pantropi- cal with the first much larger in terms of numbers of genera and species than the second. Subfamily Melioideae is divisible into seven tribes (Pennington in Pennington & Styles 1975) with 36—38 genera. All tribes are represented in Malesia, but only two (Guareeae and Trichilieae) are pantropical, while two more are restricted to the Old World (Turraeeae and Melieae), the remaining three being restricted to Indomalesia and the western Pacific (Vavaeeae, Aglaieae and Sandoriceae). The Trichilieae, Guareeae and Aglaieae are very closely allied and have been treated as one tribe by some workers, but the other four are more isolated around this core. With the possible exceptions of Lansium and Sandoricum, species of which are widely planted such that their natural distribution is blurred now, and maybe an undescribed genus here retained in Turraea, no genus is restricted to the Malesian region. Subfamily Swietenioideae is divisible into three tribes with 13 genera. All tribes are represented in Malesia, but by just one genus of each. Cedreleae are amphipacific in distribution; the other two tribes (Swietenieae and Xylocarpeae) are pantropical. At the generic level, the family demonstrates some remarkable transpacific affinities: Asiatic and Malesian Joona with neotropical Cedrela and Dysoxylum and Chisocheton of Indomalesia with Guarea, Ruagea and Cabralea of the Neotropics. Xylocarpus of East Africa to the Pacific is closely allied to Carapa of the Neotropics to West Africa. Types of species and their variation patterns — There are examples of the three major sorts of infraspecific variation in tropical trees pinpointed by Whitmore (1976), viz. the clear-cut, more-or-less uniform species, e.g. Dysoxylum arborescens, while ecogeo- graphically definable infraspecific taxa can be recognized in D. mollissimum and D. acu- tangulum, with more-or-less distinctive subspecies on each side of Wallace’s Line and in taxa treated as specifically distinct; there are similar affinities e.g. Dysoxylum cauli- florum and the closely allied D. cumingianum to the west, D. pettigrewianum to the east. The greatest difficulty is encountered with the third type, however, where there is in- sufficient ecogeographical differentiation yet there are sometimes recognizably distinct types at any one locality nevertheless: careful analysis of the massive amount of material now available shows that a number of species formerly recognized by earlier workers must be included in a complex around Chisocheton lasiocarpus and Aphanamixis poly- stachya for example. Such a variation pattern is not infrequently encountered in Melia- ceae, notably in New Guinea, where it is found in many other families too. Extending out into the Pacific, such is also seen in Vavaea amicorum (Pennington 1969). How- ever, A.C. Smith (1985) who resurrects several species in that complex for a local flora (Fiji), writes “It is not realistic to imply that overlapping characteristics in every case prohibit the recognition of taxa at the specific level. If one should propose that all spe- cies must be distinct from their congeners in all features, a species concept would be Mabberley et al. — Meliaceae 17 unattainable in most genera of flowering plants, and the genus would become the lowest category to mirror reality [sic!]. It would seem logical to take into consideration such dispersal barriers as distance, water expanses, topography, time, and many other isolat- ing factors in the recognition of taxa that, however superficially similar, are quite recog- nizable to many observers as ‘species’, albeit with occasional overlapping characteris- tics.” This is, then, an attempt to use a ‘biological species concept’ along the lines of the basic unit for classification being the breeding population. This is all fine theory but such theory has as its basis the results of experimental taxonomy and for the great majority of tropical woody plants such are neither available or even possible to obtain such that taxonomic work is inevitably based on museum studies: to pretend otherwise is to mis- lead. Moreover, what is the classification for? On one hand, it may be useful to give names of the ‘species’ or provenance at the local level, but, on the other, finely discrim- inated ‘species’ to which museum materials are difficult or impossible to assign with certainty scarcely encourage confidence in a monograph. If it suits local workers to resurrect species for local usage, then so be it: it may turn out that such prove to have biological significance (apomicts, jordanons, sibling species, yet they may represent infraspecific polymorphic variation), but as yet such ‘explana- tions’, like supposed introgression or other hybridity, must be considered with some caution. In my opinion, a discussion of variation within a complex, as untidy as it seems, is a fairer picture of the state-of-affairs than is forcing the facts into a false clarity exemplified by a linear series of neatly set-down binomials, the legacy of the essential- ists imbued with Linnean artificiality and the Great Chain of Being. It must be realized that there is no one classification in these difficult cases, nor indeed any other — a chemi- cal classification is a valid classification mirroring reality as perceived by creatures able only to sense chemical distinctions. So often, evidences from cytology, chemistry and other sources are considered ‘useful’ only if they bolster up those classifications based on the features of dried cellulose and lignin discernible optically and, as human beings perceive most of their world in this last way, such an approach is readily defended, though it does not make such classifications any better from a philosophical point of view. Relationships of the family — These are amply discussed by Pennington & Styles, who argue for the integrity of the family, which is, nevertheless, closely allied to Rutaceae, some of which have very similar limonoids, the oxidized terpenes which characterize the family (see ‘Phytochemistry’ ) and provide some of the most significant plant prod- ucts in terms of potential insecticides. The limonoids are recorded otherwise only from Cneoraceae and Harrisonia R.Br. ex A.Juss., currently placed in Simaroubaceae but perhaps better considered rutaceous. References: Cheek, M.R. & A. Rakotozafy, Taxon 40 (1991) 231—237. — Harms, H., in Engler & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 258—308; in ibid., ed. 2, 19b1 (1940) 1-172. — Miquel, F.A.W.., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 1-64. — Pennington, T.D., Blumea 17 (1969) 351-366. — Pen- nington, T.D. & B.T. Styles, Blumea 22 (1975) 419-540. — Smith, A.C., Flora Vitiensis Nova 3 (1985) 527-578. — Whitmore, T.C., Natural variation and its taxonomic treatment, in J. Burley & B.T. Styles (eds.), Tropical trees: variation, breeding and conservation (1976) 25—34. 18 Flora Malesiana ser. I, Vol. 12 (1) (1995) USES Timber — The timbers of certain Meliaceae are some of the most sought after in the world, such that natural stands have been much depleted and serious conservation mea- sures have been proposed for wild mahoganies (Swietenia spp.) in tropical America (see e.g. Knees & Gardner 1983; Newton et al. 1993b). The original ‘mahogany’ of Hepplewhite and Chippendale was S. mahagoni, allowing the construction of more graceful and woodworm-proof furniture than could the oak and walnut previously used in Europe: this species has suffered severe genetic erosion and most ‘mahogany’ (if meli- aceous at all!) seen today is derived from S. macrophylla introduced to the Old World, probably from Honduras, in 1876 and described from material cultivated in India. The other important timbers are also generally Swietenioideae, notably toon, Toona ciliata from India to Australia (‘red cedar’), where most of it has been long cut out, having been the most desirable timber in that continent. Others include the neotropical Cedrela odorata and species of the African genera Entandrophragma (E. sapele, E. utile), Khaya (African mahogany) and Lovoa (Nigerian golden walnut). Besides the Toona spp. and Chukrasia tabularis, the chickrassy wood of Malesia, indigenous species are not of commercial significance in the world timber market, despite their large boles, though the wild forms of Melia azedarach yield a second-quality timber. Locally important were several species of Dysoxylum, notably the gigantic species D. mollissimum (formerly the tallest trees of all in Java) and D. acutangulum. Locally, some Aglaia spp., Antho- carapa nitidula, Azadirachta excelsa, Dysoxylum spp., and Walsura spp. have been used for construction and furniture, while Xylocarpus wood is hard and used for boat- building. The standard name for Dysoxylum timber in Malaysia is jarum-jarum (a Sabah name, the ASEAN standard name being the Indonesian membalun). It and the timbers of Aglaia (incl. Amoora; bekak, pasak) and Azadirachta (sentang), Chukrasia (surian batu), Lansium (langsat), Sandoricum (sentul), Toona spp. (limpaga) and Xylocarpus (nyireh) are discussed by Wong (1982), but see “Vernacular names’ below. Silviculture — Most Meliaceae grown for timber in Malesia, then, are exotics, notably the neotropical Swietenia mahagoni, S. macrophylla and Cedrela odorata, which have all also been grown as shade or avenue trees though Swietenia is the only meliaceous genus treated in the PROSEA Handbook on Major commercial timbers (Prawirohatmadjo et al. 1993). The most serious problem besetting plantation forestry of Meliaceae is the attacks of the moths (Lepidoptera, Pyralidae) of the genus Hypsipyla, larvae of which burrow into young plants and seedlings, causing their collapse and death, though Melioideae seem to be rarely attacked — a good argument for their promotion as planta- tion trees. The shoot-borer is perhaps one of the most economically important insect pests in tropical forestry. In the Neotropics, the moths appear to be attracted by a chem- ical, possibly an alkaloid or limonoids, allied to substances which, in species from other parts of the world, are toxic to their local moths. Thus, Toona ciliata is resistant to attack in Costa Rica and this resistance may be transferred to susceptible Cedrela odor- ata if a scion of this is grafted on to a stock of Toona. Chukrasia tabularis, on the other hand, is not resistant in Costa Rica. There have been many attempts at biological control Mabberley et al. — Meliaceae 19 of the moths, for externally applied insecticides have little effect and systemic ones are expensive (Whitmore 1976). Mixed and enrichment planting with non-susceptible spe- cies has been shown to reduce damage and there are possible advances to be made in breeding resistance to attack. See Newton et al. (1993a) for further details. Other uses — The locally very important fruit trees, Lansium domesticum (langsat) and Sandoricum koetjape (sentul), exist in a number of forms, wild, cultivated and natural- ized, though they are not grown on a commercial plantation scale, those reaching markets being largely those selected from village trees. A good langsat may be worth more as a proposition than a good durian tree. Some species of Aglaia have acceptable fruits and one of these, A. korthalsti, is widely grown in villages in the north of the Malay Peninsula. Seeds of a number of species of Chisocheton and Aphanamixis yield an oil which has been used as an illuminant, while those of Lansium domesticum are used in arrow poisons, as is the bark. The bitterness (due to the limonoids) of the barks of Meliaceae has long been known and they have been used in medicine, some being those eagerly sought by Europeans in the eighteenth century. Soymida febrifuga (Roxb.) A. Juss. (Swietenioideae) from India was much prized but barks of other genera have been used, notably in the Malesian flora, species of Aphanamixis, Chukrasia, Lansium, Sandoricum, Toona, Trichilia and Xylocarpus, while the whole plant of Munronia pinnata is an important item of materia medica in southern Asia. Extracts from Walsura monophylla are being commercially tested for the treatment of cancer and AIDS in the U.S.A. Some others are poisonous in part, e.g. seeds of Dysoxylum spp., which contain the toxic dysoxylum acid. The bark and indeed, the leaves of the exotic Azadirachta indica, the neem, are powerful insec- ticides and this tree (q.v., and see also the paragraph on phytochemistry, p. 8) has a host of uses including planting in the reclamation of derelict land: it is perhaps one of the most all-round useful trees of Asia. The tetranortriterpenoids which are responsible for the insecticidal (and molluscicidal in e.g. Khaya spp. bark and seeds) properties have aroused considerable commercial interest and have been examined in a number of genera for their use as biological pesticides: this is dealt with genus by genus below. The biological activities of these compounds including insect antifeedant and growth- regulating properties, medicinal effects in humans and other animals, antifungal, bac- teriocidal and antiviral activity, are reviewed by Champagne et al. (1992). The leaves of Azadirachta excelsa are used as a green vegetable while the flowers of (male) Aglaia odorata, an introduced species, are used to flavour tea. For other local uses, especially in medicine, see Perry (1982). A number of indigenous species are highly ornamental but few are planted. Aglaia odorata makes a good, if slow growing, subject for hedging and topiary, which are practised in Java. An admirable example of the imaginative use of Malesian Meliaceae, mainly species of Aglaia, Aphanamixis and Dysoxylum is to be seen in Java around the Lady Raffles Memorial in the Kebun Raya, Bogor. A number of exotics, notably the Indian cultivars of Melia azedarach are planted for their elegant foliage and fruits, while the fruits of that species are imported into the Malay Peninsula from Sichuan and used as a febrifuge and vermifuge. 20 Flora Malesiana ser. I, Vol. 12 (1) (1995) References: Champagne, D.E., et al., Phytochem. 31 (1992) 377. — Knees, S.G. & M.F. Gardner, Oryx 17 (1983) 88. — Newton, A.C., et al., For. Ecol. Manag. 57 (1993a) 301; Biodiv. and Conserv. 2 (1993b) 114. — Perry, L.M., Medicinal plants East and Southeast Asia (1980) 260—264. — Prawiro- hatmadjo, S., et al. in I. Soerianegara & R.H.M.J. Lemmens (eds.), Pl. Res. SE Asia (PROSEA Handb.) 5 (1), Major commercial timbers (1993) 442—447. — Whitmore, J.L. (ed.), If C A Misc. Publ. 101 (1976). — Wong, T.M., Dict. Malays. Timbers, Mal. For. Rec. 30 (1982). VERNACULAR NAMES Because so many of the Malesian Meliaceae are not used by the indigenous peoples and because many species from different genera are superficially very similar, there is a great deal of confusion in the literature as well as in the field over the local names. The clearly obvious genera like Xylocarpus (nyireh), Toona (surian), Sandoricum (sentul) or with well known uses as Lansium (langsat etc.) have (usually) reliable names. The bulk of the species do not and many of these on herbarium labels and laboriously copied into manuals seem to have been made up on the spot or even to be those used for other trees with pinnate leaves in related families e.g. kedondong (Burseraceae). Lantupak is used in northern Borneo for almost any Meliacea, as is segera (Iban) in Sarawak, while variations of langsat indicate how the tree in question differs from Lansiwm domesti- cum. In the Malay Peninsula, bekak is used for some species of Dysoxylum and the larger species of Aglaia, while pasak lingga is used for species of both of these as well as of Chisocheton and Aphanamixis. Few vernacular names are indicated here unless there is evidence of use more than once and of their being of some discriminatory value. KEY TO SUBDIVISION OF THE FAMILY la. Buds naked; fruits capsules, berries or drupes; seeds neither corky nor winged .. . bgt Wie Te ge ing NE RE ding Sh bo Ste i AL RL MES Taped SN ate ARNON Subfam. Melioideae b. Buds with scale leaves; fruits woody septifragal capsules with central columella; SCCdS: WAN PEMIOM COLKY) eats ct od eme Cen e or Subfam. Swietenioideae The genera are arranged in the tribal system set out by Pennington & Styles (1975). The phytochemical studies since then have not upset it, though have shown how intimately allied are Trichilieae, Guareeae and Aglaieae and drawn attention to the apparently close relationship between Xylocarpeae and Melioideae. KEY TO GENERA NATIVE IN MALESIA (for either flowering or fruiting specimens) Mae NEC AVES DIUM AUC 512 slew ene ra set cio 8 ou at des 5 casserole uaplslevol by angie Melia (p. 329) DRIBCAVESHOIMIMALS CO yGTITNP LO aa asc ea Ree htc ee or eae cc ce a ete eo ea 2 Zawlceavessall simple: (Or wmilololate) ie es aie cpus ot coo sda ae ee a 3 Beavis qoinnmate to ttt Olt Lae pa fia olny De cases se klar a 8 Mabberley et al. — Meliaceae 21 Pid asap enly de. Sones uno Ruel! eon SiO ey 4 . Indumentum of at least some stellate hairs and/or stellate or peltate scales.... 7 MPCcianols Shrub; Inutacapsules:., soelysiss so ... YITBLE Nt | Turraea (p. 24) BE VGIOTeSD Ces mua DELLS) Siete tei ea Re eer ie CY 5 . Anthers in 2 whorls of 5 ....... Reinwardtiodendron celebicum (p. 328) Beemtnersin.| whorltsseioons) Seni Bente 1 jo zlvulw Sas eeutiod. . 6 . Flowers very small; small tree (Philippines) ... Walsura monophylla (p. 52) Flowers conspicuous; trees with Terminalia-branching ....... Vavaea (p. 34) . Hairs mixed simple and stellate, suckering shrublets to] m ................. pets. fat ts kis. ole ee ston meek tes ke eG ie: Munronia humilis (p. 32) (see also Turraea breviflora, p. 29) b. Indumentum of stellate hairs and/or stellate or peltate scales; suckers absent .... OE AT is oo 8 ce a ss , MR re Ee 3 Aglaia (p. 194) Seeeeavesiall tifoliolatehnisiar. (sto). ..0 > oats See. ile sings... 9 Pa Rs RULES 92592 os fo oucitete sk peed a) sow ee a eee Eels ee See gE 10 PeeeediTS Simple: ds a5 A! dca.) Saas Sandoricum (p. 344) b. Indumentum of stellate hairs and/or stellate or peltate scales ... Aglaia (p. 194) 10a. Leaves with pseudogemmula (apical bud) ........... Chisocheton (p. /36) Beeeavesumitnoutpseudocemmulanys 2.325282 2. ast Sel OY AE. 1] ieaseaveswuith'scalesiand/or stellate hairs: a2. 5 a. 12 Ee Rearesmumtth simple halts: e.c aid boa oes 2s secee so tee bo 15 12a. Shrublet with large white flowers and capsular fruits ...................20- Pe eC yr heh 7 2s Sle Sie a eo er e Siaes Munronia pinnata (p. 30) PeEGeeS: Oly ICC ICIS) Li thc... 2 Ie, ON ete es aces es Geis 13 13a. Disk present; capsule 5-locular, + persistent calyx of 5 free sepals; scales absent OL Se SANG = nee ers Dysoxylum stellatopuberulum (p. 73) Pubisksabsent; tnuitidifferent;scales.often:present —: ..42 - 44: «=. seetoeek: ok 14 14a. Female inflorescence and infructescence long spicate, scales absent ........... RPM eel AG tnt RO lae Oe Nis o cunie ae Aphanamixis polystachya (p. /S8) Bes Not this:combination of characters, s:\2o sso eee. Aglaia (p. 194) Pia. budseales present (Swietenioidede) isin, Palen waisjerg PREMERA ORE 16 Hs budscales absent... ij. 2. saeeentiis 2c. AO ee olis hn ie cow we 18 16a. Leaves (1—-)2—4(—5)-jugate, maritime ............... Xylocarpus (p. 37/) bs Leaves with mote leaflets; forestirees: 2 2... ao. SEO ee See 17 17a. Staminal tube present; capsule with 60—100 terminally winged seeds per locule cea AUST ranlretee lass Seer eee pesuseritet Chukrasia (p. 354) b. Stamens free; capsule with many fewer seeds with terminal wings or winged at BIOTEC” i. cieiescs dns sPakaghagle cies nk, eso wa heb Toona (p. 358) iSariLeaves patipinsates}.waioul soul are. “eel iee Folens eas 19 Ds LEAVES SUMP ALP INANE i585 a8 n ats 0500) 6, ayo cna ca terenais inteuee ci Faat cee, oy) Flora Malesiana ser. I, Vol. 12 (1) (1995) 19a. Stigma with conspicuous lobes; fruita drupe .......... Azadirachta (p. 337) b:. ,stiemaunlobed: iruitra capsule: or bemy sta ioe 24002 Seva. S0 se oe 20 20a. Diskvabsent; fruit aubeaby,, 2 4 52.34) a.ia vs SA: BE: Be ie eee 21 bi Disk presents fruit avcapsule hacen nueva ie ee I ee ee 22 21a. Anthers in 1 whorl of 10; berries on branches and trunk ... Lansium (p. 3/4) b. Anthers in 2 whorls of 5; berries on axillary infructescences ................ GY STSPOSPNEN SORES E8550 ch PE cba dere Reinwardtiodendron (p. 322) 22a a Stamimal tube cyathiiormiee aster. ia eerie Anthocarapa (p. /33) b. Stamuinal tube cylindrical:to patellifomi...02 7. 42 Da9 Dysoxylum (p. 6/) 23a. weal rachis;swollenat insertion of leaflets 45.22 S224 4e2 Walsura (p. 45) byl eatrachsinot'swollemthus tess isis Sleds hi carka oS wen aed 24 2Aa-. Diskrabsentiiruiiaberiyy aes ae eee ae ee Pseudoclausena (p. 55) bs eDiskspresent;iinut a.capsule ondrupes 52 ae a ee 25, 25a. Stigma with conspicuous lobes; fruit a 1- (or 2-)seeded drupe ............... PNM y wey nto D eh Sth tee MEE AE as TESS De, See Azadirachta (p. 337) b. Stigma without such lobes; fruit a capsule or drupe with 5 pyrenes......... 26 26a. Staminal tube deeply cleft; fruit a drupe or 1-seeded capsule .............. 4} b) Stamimalitube not deeply cleft; inmtacapsule. 7.28222 eee. 22 ee eee 28 27a. Corolla valvate; fruit a 5- or 6-pyrened drupe ............ Cipadessa (p. 57) be Corollaimbricate; fruit a, l-seeded capsuler):;aa- oe eee Heynea (p. 4/) 28d. EClAlSeosClISKZADSENE. 2272 sek b 2.2: MSE eNO e © sued ee Aphanamixis (p. 1/57) bs Petals 4 orn mores disk presenti a-toee ters eee Dysoxylum (p. 6/) SPOT CHARACTERS Unbranched or sparsely branched pachycaul treelets: Aglaia, Aphanamixis, Chisoche- ton, Dysoxylum Suckering shrublets: Munronia, Turraea breviflora Bushes: Turraea, Cipadessa Rheophytes: Aglaia, Dysoxylum angustifolium, Sandoricum borneense Halophytes: Xylocarpus Bark with white latex: Aglaia, Chisocheton, ?Dysoxylum Budscales present: Swietenioideae Budscales absent: Melioideae Indumentum of or with stellate hairs: Aglaia, Aphanamixis polystachya (sometimes), Chisocheton sect. Rhetinosperma, Dysoxylum stellatopuberulum, Melia Indumentum of stellate or peltate scales: Aglaia Leaves opposite: Dysoxylum Leaves simple or unifoliolate: Aglaia, Munronia, Turraea breviflora, Vavaea, Walsura monophylla Mabberley et al. — Meliaceae 23 Leaves trifoliolate: Aglaia, Sandoricum Leaves imparipinnate: Aglaia, Aphanamixis, Azadirachta, Chisocheton, Cipadessa, Dysoxylum, Heynea, Pseudoclausena, Walsura Leaves paripinnate (sometimes a lateral leaflet pseudoterminal): Anthocarapa, Azadi- rachta, Chisocheton, Chukrasia, Dysoxylum, Reinwardtiodendron, Toona, Xylo- carpus Leaves bipinnate: Melia Leaves with terminal bud (pseudogemmula): Chisocheton Inflorescences epiphyllous: Chisocheton Inflorescences cauliflorous: Chisocheton, Dysoxylum, Lansium Inflorescences ramiflorous: Aglaia, Chisocheton, Dysoxylum, Lansium Inflorescences flagelliflorous: Aphanamixis, Chisocheton, Dysoxylum Flowers yellow: Aglaia Sepals free, in a spiral: Dysoxylum Sepals leafy: Munronia Calyx deeply lobed with almost free orbicular sepal lobes: Aphanamixis, Lansium, Rein- wardtiodendron Calyx valvate: Xylocarpus Petals in a spiral, sometimes appearing as 2 whorls: Chisocheton Corolla valvate: Chisocheton, Cipadessa, Dysoxylum, Walsura Petals 3: Aglaia, Aphanamixis, Chisocheton (rare), Vavaea, Dysoxylum (rare) Filaments free: Toona, Walsura sect. Surwala Staminal tube globose: Aglaia, Aphanamixis, Lansium, Reinwardtiodendron Anthers in 2 or more whorls: Aglaia (rare), Reinwardtiodendron Anthers locellate: Chisocheton, Dysoxylum Disk absent: Aglaia, Aphanamixis, Lansium, Munronia, Reinwardtiodendron, Vavaea Disk tubular: Dysoxylum, Sandoricum Fruit a loculicidal capsule: Aglaia, Anthocarapa, Aphanamixis, Chisocheton, Dysoxy- lum, Heynea, Munronia, Turraea Fruit a septifragal capsule: Swietenioideae, Walsura dehiscens Fruit a berry: Aglaia, Lansium, Pseudoclausena, Reinwardtiodendron, Vavaea, Walsura Fruit a drupe: Azadirachta, Cipadessa, Melia, Sandoricum Seeds winged: Chukrasia, Toona Seeds corky: Xylocarpus 24 Flora Malesiana ser. I, Vol. 12 (1) (1995) SUBFAMILY MELIOIDEAE Melioideae;, T.D.Penn., Blumea 22 (1975) 451. Buds naked. Plants dioecious, polygamous or with hermaphrodite flowers entirely. Locules 1-, 2- or 3-ovular with vestigial third ovule, very rarely multiovulate; ovules collateral or superposed. Fruit a fleshy or leathery capsule, berry or drupe. Seeds un- winged, usually with a fleshy testa or aril or combination of these; endosperm present or absent. Seven tribes, pantropical. TRIBUS TURRAEEAE Turraeeae Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 267, 280; T.D. Penn., Blumea 22 (1975) 451. Aitonieae Harv., Gen. S. Afr. Pl. (1838) 46, sub Zygophyllaceae, nom. illeg.; M. Roem., Synops. Mon- ogr. 1 (1846) 78, 88; Baill., Hist. Pl. 5 (1874-75) 426, sub Sapindaceae. — Aitoniaceae (Harv.) Harv., FI. Cap. 1 (1860) 243 (‘Aitonieae’); Airy Shaw, Willis Dict. Flow. Pl. Ferns, ed. 7 (1966) xxi. Trees, shrubs or suffrutices. Indumentum of simple or stellate hairs. Leaves pinnate, trifoliolate or simple. Flowers hermaphrodite or very rarely apparently dioecious. Stami- nal tube complete, cylindrical or filaments united only in lower half (not in Malesia). Disk absent or feebly developed. Stylehead often modified to form ‘receptaculum polli- nis’. Fruit a capsule (Malesia); seed endospermous. Cotyledons flat, collateral; radicle superior, exserted. Distribution — Six or seven genera restricted to the Old World tropics. The largest is the viable Turraea, which is most widespread and the rest are satellite genera, one or two restricted to Indomalesia, two to Madagascar, one to southern Africa and one found in both India and southern Africa. TURRAEA Turraea L., Mant. Alt. (1771) 150; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 85, t. 20; T.D. Penn., Blumea 22 (1975) 455, f. 3 (see there for extra-Malesian synonyms); Mabb. & Cheek, Taxon 41 (1992) 541. Small trees or shrubs. Jndumentum of simple hairs. Leaves simple (pinnate in one African species), entire or (not in Malesia) sometimes sinuate or crenate. Inflorescences axillary, fasciculate, cymose or paniculate, or flowers solitary (terminal or ramiflorous outside Malesia). Flowers bisexual, greenish or white. Calyx (3—)4—5(—6)-lobed or -toothed. Petals (3) 4 or 5, free, imbricate or contorted. Staminal tube cylindrical or rarely (not in Malesia) cyathiform, complete or filaments at least 2/3 united; margin shallowly lobed or entire (not in Malesia) or terminated by simple or bilobed appendages, as many as or twice as many as the anthers; anthers (7—)8—10(—20). Disk small or absent. Ovary (3—)4—10(—20)-locular; locules biovulate; style usually expanded near apex forming a ‘receptaculum pollinis’ surmounted by a discoid stigmatic area. Fruit a (3—)4—10(—20)- Mabberley et al. — Meliaceae 25 valved, loculicidal capsule; locules 1- or 2-seeded. Seeds plano-convex, raphe-funicular- arillate; embryo embedded in thick endosperm, with rather narrow cotyledons. Distribution — About 30 species in Africa, perhaps 35 in Madagascar and the Mas- carenes with one in Indomalesia, extending from India, through Malesia to tropical Australia. Habitat & Ecology — The African species are found in a variety of forest types and bush vegetation, often markedly seasonal. Turraea pubescens Hell. Turraea pubescens Hell., Kon. Vet. Acad. Nya Handl. 9 (1788) 308, t. 10 f. 3; Willd., Sp. Pl. 2 (1799) 555; J.E. Smith in Rees, Cyclop. 36, 2 (1817) Turraea n. 2; DC., Prodr. 1 (1824) 620; G. Don, Gen. Syst. | (1831) 678; Benn., Pl. Jav. Rar. (1840) 181; M.Roem., Synops. Monogr. | (1846) 91; Miq., Fl. Ind. Bat. 1, 2 (1859) 534; F. Muell., Essay Pl. Fitz. (1860) 5; Fragm. Phyt. Austral. 5 (1866) 144; Descr. Not. Pap. Pl. 4 (1876) 53; Benth., Fl. Austral. | (1863) 379; C.DC. in DC., Monogr. Phan. | (1878) 446; Hance, J. Bot. 16 (1878) 9; Vidal, Rev. Vasc. Pl. Filip. (1886) 81; Forbes & Hemsl., J. Linn. Soc., Bot. 23 (1886) 113; F.M. Bailey, Proc. Roy. Soc. Queensl. | (1884) 63; Moore, Handb. Fl. NSW (1893) 37; F.M. Bailey, Queensl. Fl. 1 (1899) 226; Pellegr. in Fl. Indo-Chine | (1911) 735, t. 80, f. 5-12; Suppl. (1946) 687, incl. var. billardierei (DC.) Pellegr. (‘billardieri’); Koord., Exk. Fl. Java 2 (1912) 438; F.M. Bailey, Compreh. Catal. Queens]. PI. (1913) 86, t. 72; Craib, Fl. Siam. Enum. | (1926) 249, incl. var. billardierei; Merr., Lingn. Sc. J. 5 (1927) 103; How & Chen, Acta Phytotax. Sin. 4 (1955) 4; Backer & Bakh. f., Fl. Java 2 (1965) 191; Chun, Fl. Hain. 3 (1974) 60; T.D. Penn., Blumea 22 (1975) 456, f. 3b; Corner, Seeds Dicots (1976) 192, t. 392; Taylor & Harden, Flora NSW 2 (1991) 282; H6, Ill. Fl. Vietnam 2, 1 (1992) 484. Turraea virens L. var. billardierei DC., Prodr. 1 (1824) 620 (‘billardierii’); G. Don, Gen. Syst. 1 (1831) 678. — Turraea billardierei (DC.) A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 218 (‘bil- lardierii’); Benn., Pl. Jav. Rar. (1840) 181; M.Roem., Synops. Monogr. | (1846) 92; Migq., FI. Ind. Bat. 1, 2 (1859) 534; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 4; C.DC. in DC., Monogr. Phan. 1 (1878) 440 (‘billiardier’’); Backer, Schoolfl. (1911) 201; de Voogd, Trop. Nat. 25 (1927) 167, t. 5S. Turraea virens auct. non L.: J. Graham, Cat. Pl. Bombay (1839) 31; M.Roem., Synops. Monogr. | (1846) 92; Dalz. & Gibs., Bomb. Fl. (1861) 36; Drury, Handb. | (1864) 169; Hiern in Hook. f., Fl. Brit. India | (1875) 541 p.p.; Talbot, Trees Bomb. (1894) 38 & ed. 2 (1902) 72. Turraea villosa Benn., Pl. Jav. Rar. (1840) 182; M.Roem., Synops. Monogr. | (1846) 92; Wight, Ic. 4, 4 (1850) t. 1593; Drury, Handb. 1 (1864) 169; Hiern in Hook. f., Fl. Brit. India 1 (1875) 542; C.DC. in DC., Monogr. Phan. | (1878) 442; Talbot, Trees Bomb. (1894) 38 & ed. 2 (1902) 72; Cooke, Fl. Bomb. 2 (1902) 214; Brandis, Indian Trees (1906) 134; Talbot, For. Fl. Bomb. | (1909) 224, t. 134; Gamble, FI. Madras 1 (1915) 174; Basu, Ind. Med. PI. (1918) 298, t. 216; Anon., Wealth of India 10 (1976) 397. Turraea concinna Benn., Pl. Jav. Rar. (1840) 182; M. Roem., Synops. Monogr. | (1846) 92; Migq., FI. Ind. Bat. 1, 2 (1859) 534; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 4; C.DC. in DC., Monogr. Phan. | (1878) 441; Backer, Schoolfl. (1911) 201; Koord., Exk. Fl. Java 2 (1912) 438. Turraea zollingeri C.DC. in DC., Monogr. Phan. | (1878) 441. Turraea indica C.DC., op.cit. (1878) 439. Turraea brownti C.DC., op. cit. (1878) 442; Domin, Bibl. Bot. 22 (1927) 852; Brouwer, Fl. Males. Bull. 32 (1979) 3244; Stanley & Ross, Fl. SE Queensl. | (1983) 477. Turraea membranacea Merr., Philipp. J. Sc., Bot. 9 (1914) 306; Enum. Philipp. Flow. Pl. 2 (1923) 359. Deciduous shrub often of straggling habit, or, more rarely, a small tree to 6 m; trunk to 11 cm dbh. Bark silvery grey to reddish, sometimes with prominent lenticels; inner bark fawn, with sickly smell; sapwood creamy yellow. Leafy twigs 2—3 mm diam. 26 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 1. Turraea pubescens Hell. a, b. Habit, flowering; c. half flower; d. habit, fruiting; e. seeds (a—c: Womersley NGF 19133; d, e: Mabberley 1778). Drawing R. Wise. Mabberley et al. — Meliaceae 7 Leaves 1.5—10(—17) by 1—6 cm, often not completely expanded at anthesis, ovate to oblong, subglabrous or sparsely hairy on veins to subvelutinous adaxially; apex obtuse to acuminate; base rounded to cuneate; major veins c. 5 or 6 on each side, acute, arcu- ate, very occasionally with tufts of hairs in their axils; petiole S—9 mm. /nflorescences fascicles or condensed cymes of up to 6 sweetly scented flowers, sometimes produced on leafless branches; axis 2-5 mm long, pubescent; bracts 1-2 mm long, triangular, pubescent; pedicels 1.5—3 cm. Calyx 3—4 mm long, cupular, 5-lobed up to halfway down, pubescent. Petals 30—45 by 2.5—4 mm at widest, linear-spathulate, white be- coming yellowish with age. Staminal tube 20—30 by c. 3 mm at widest, cylindrical to weakly obconical, white, margin subentire or, usually, with ten appendages sometimes divided apically into 2(—4) lobes, reflexed at anthesis; anthers 10, c. | mm long, at apex of tube, glabrous, yellow, filament very short. Ovary 5-locular, subglabrous; style c. 30—45 mm, long-exserted, + pubescent in proximal region, stylehead expanded to form a ‘receptaculum pollinis’, surmounted by discoid stigmatic region pale yellow. Fruits up to 1.8 cm long, globose to subellipsoid, vermilion, 5-sutured, the 5 leathery valves reflexed at dehiscence starwise, 1-5 each bearing two seeds. Seeds c. 6 mm long, arillate; testa black, shiny; aril scarlet; endosperm slightly elongate. — Fig. 1. Distribution — Indomalesia from India and S China to tropical Australia, south to northern New South Wales; in Malesia: Java, Philippines (Luzon), Lesser Sunda Is- lands (Flores, Timor), New Guinea. Habitat — Semi-evergreen and deciduous forest to 1000 m altitude but usually much lower, and in coastal vegetation, behind mangrove, on dunes or rocks including lime- stone and basalt, as well as secondary vegetation at forest edges and on ridges, in vine thickets and Eucalyptus savanna. Ecology — The species flowers at the onset of the rains, when it may be leafless and then it is a conspicuous feature, as is Munronia pinnata of the spring flora of the teak forests of E Java. The scented white flowers with the long style suggest moth-pollina- tion which is recorded in allied species from Africa. Uses — The plant is of some slight medicinal value in India [see Wealth of India 10 (1976) 397]. There its pickled fruits are used as a dye. Corner (l.c.) notes that a red- dish-purple coloration is imparted to potash solution in which seeds have been steeped. Notes — |. Turraea pubescens is the only true Turraea in Asia and ts closely related to some of the East African species. A similar transoceanic distribution is seen in the allied Naregamia Wight & Arn. [see Cheek, Kew Bull. 45 (1990) 711], which com- prises one species in India and a very closely related one in Angola. 2. As Bentham (I.c.) noted, the staminal tube appendages, on which character several ‘species’ were segregated, vary greatly, even on one flowering shoot. The degree of pu- bescence also varies and may be associated with the level of exposure while shape of style- head varies with development and between flowers on the same plant. Immature fruits dehisce on drying so the ‘ripe’ fruit size appears to vary greatly in herbarium material. 3. Seeds of T. pubescens have been examined by Corner (I.c.), who regards them as very similar to those of Celastraceae, to which family many collectors have assigned fruiting material of this species in the field. 28 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 2. Turraea breviflora Ridley. a. Habit; b. half flower. Drawing R. Wise. Mabberley et al. — Meliaceae 29 SPECIES TO BE EXCLUDED FROM TURRAEA Turraea breviflora Ridley Turraea breviflora Ridley, Fl. Malay Penins. 1 (1922) 383; J. As. Soc. Str. Br. 86 (1922) 293: Hender- son, J. Roy. As. Soc. Mal. Br. 17 (1939) 40; T.D. Penn., Blumea 22 (1975) 458; Mabb. in Tree Fl. Malaya 4 (1989) 252. Suckering shrublet to | m tall, making stands of branching stems of 6 mm diam.; taproots long, tenacious. /ndumentum of stellate hairs with some simple ones. Leafy twigs 2—3 mm diam., densely pubescent. Leaves distichously arranged, 12—20 by 4—5 cm, elliptic to weakly obovate, somewhat bullate, subglabrous, pale abaxially; apex gradually long-acuminate; base acute to cuneate; venation prominent on both surfaces in sicco, midrib sparsely pubescent proximally, major veins 7-10 on each side, arcuate and looped at margin. Flowers in 3—6-flowered fascicles or cymes, in axils of leaves present or lost; branches 2—4 mm long, pubescent; pedicels 4-9 mm long, pubescent. Calyx c. 2—3 mm tall, very shallowly cupulate, 5-lobed, pubescent, green, the lobes triangular to 1.5 mm long. Petals 5, 6—7 by 2—3 mm, narrowly ovate, valvate, reflexed at anthesis, white. Staminal tube a little shorter than petals, white, bearing 10 filiform appendages alternating with anthers; anthers 10 with apical appendage. Disk cupuli- form, short, closely encircling ovary, margin somewhat erose. Ovary 5-locular, each locule 1-ovular, or, rarely, apparently 3-locular and some locules 2-ovular, pubescent with upward-pointing simple hairs; style c. 5 mm, pubescent; stylehead capitate, 5-lobed, orange. Fruit not observed. — Fig. 2. Distribution — Malesia: known only from a few localities in the Malay Peninsula (Selangor, Johore) and Singapore. Habitat & Ecology — Recorded from rather open hill Dipterocarp forest, often as- sociated with bertam, Eugeissona tristis, and from limestone rocks (Ridley, Hender- son). Note — This plant resembles no other Malesian Meliacea and, in the field, may be readily confused with /cacinaceae or other families. It is known from very well-worked sites such as Bukit Lagong and Ulu Gombak in Selangor but has never been found in fruit and the number of flowering collections made is small. No doubt it is frequently overlooked as it grows amongst tree seedlings and saplings which it greatly resembles. Ridley placed it in Turraea as it seemed the closest genus to him but he was hesitant and thought it might represent a new genus, though refrained from describing it as such un- til the fruit should be found. Harms placed it near the Turraea species of the Mascarenes formerly referred to the separate genus, Quivisia. It is to be excluded from Turraea be- cause it differs in its habit, indumentum, corolla aestivation, disk and the usual number of ovules in each locule, while it seems to be dioecious. In its indumentum, it approaches Munronia to which genus it seems on the whole more closely allied, but it differs from that genus, as presently understood, in its habit, valvate corolla, the usual number of ovules in each locule and apparent dioecy. In short, either it represents an undescribed genus or our concept of Munronia must be enlarged. 30 Flora Malesiana ser. I, Vol. 12 (1) (1995) MUNRONIA Munronia Wight, Ic. 1, 5 (1838) [1]; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 91; T.D. Penn., Blumea 22 (1975) 452. [Aengelenia Hassk., Cat. Bogor. Pl. (1844) 219, nom. in syn.] Philastrea Pierre, Bull. Mens. Soc. Linn. Paris 1 (1885) 475. Unbranched or sparsely branched shrublets, sometimes suckering with apparently short-lived shoots. /ndumentum a mixture of simple and bifid or stellate hairs. Leaves simple to imparipinnate, often crenate to serrate. Flowers hermaphrodite, solitary or in few-flowered thyrses, pseudopedicellate, usually white. Calyx 5-lobed to near base, lobes somewhat foliaceous. Petals 5, imbricate, adnate to staminal tube basally, rarely becoming free later. Staminal tube narrowly cylindrical or weakly obconical, the margin with 10 entire or bilobed appendages or, rarely, with 10 reflexed filiform appendages recurved some distance below margin; anthers 10, pubescent, inserted on tube rim, alternating with appendages, connective often produced apically forming an appendage, which is rarely filiform. Disk absent or tubular, free or united with base of staminal tube. Ovary 5-locular, each locule with two superposed ovules; stylehead capitate, some- times with 5 stigmatic lobes. Fruit a 5-valved loculicidal capsule, each locule with | or 2 seeds. Seeds plano-convex with a bony tegmen with an outgrowth enveloping a thick sarcotesta (Cheek); embryo embedded in thin endosperm. Distribution — About 6 species restricted to tropical Asia and subtropical China, two of them extending into Malesia, eastwards to Timor. Habitat — Drier forests up to about 900 m altitude (Lesser Sunda Islands). Note — Turraea breviflora Ridley (q.v.) may be referable here also. KEY, TO THE SPECIES |ReYondl Be eC ONC cig er ace Oke ERRORS OS aoe 6 oie es REISER 1. M. pinnata beebeavesssitmp lessee nears on oa anor oak eek Pca en RO Cak 2. M. humilis 1. Munronia pinnata (Wall.) Theob. Munronia pinnata (Wall.) Theob. in Mason, Burma, ed. 4, 2 (1883) 581; Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 78; Rao, Bull. Bot. Surv. India 5 (1963) 255, t. 1; Sald. & Nicols., Fl. Hassan Distr. (1976) 395; Whitmore, Enum. Flow. Pl. Nepal 2 (1979) 85; Stone, Malay. Nat. J. 37 (1984) 189; Mabb. in Tree Fl. Malaya 4 (1989) 202. — Turraea pinnata Wall., Pl. As. Rar. 2 (1830) 21, t. 19; Lindl., Bot. Reg. 17 (1831) t. 1413; G. Don, Gen. Syst. | (1831) 678; Span. — [Munronia wallichii Wight, Ill. 1 (1840) 147, nom. superfl.; Benn., Pl. Jav. Rar. (1840) 185; M.Roem., Synops. Monogr. 1 (1846) 91; Drury, Handb. 1 (1864) 168; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 5; Hiern in Hook. f., FI. Brit. India 1 (1875) 543; C.DC. in DC., Monogr. Phan. I (1878) 449; Gamble, FI. Madras | (1915) 175.] [Melia pumila Moon, Cat. (1824) 35, nom. nud.] — Munronia pumila Wight, Ic. (1839) t. 91; Illust. 1 (1840) 146; M. Roem., Synops. Monogr. | (1846) 91; Hiern in Hook. f., Fl. Brit. India 1 (1875) 543; C.DC. in DC., Monogr. Phan. | (1878) 448, t. 6, f. 8; Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 77. Turraea pinnata Span., Comp. Bot. Mag. | (1836) 345, nom. nud.; Linnaea 15 (1841) 183; M. Roem., Synops. Monogr. | (1846) 92; Miq., FI. Ind. Bat. 1, 2 (1859) 534; non Wall. (1830). Mabberley et al. — Meliaceae 3] Munronia neilgherrica Wight, Illust. 1 (1840) 147, t. 54; Drury, Handb. | (1864) 167; M. Roem., Synops.. Monogr. | (1846) 91. Munronia javanica Benn., Pl. Jav. Rar. (1840) 176, 185, t. 38; Hassk., Pl. Rar. Hort. Bogor., Pug. Nov. (1844) 11 (‘javana’); Pl. Jav. Rar. (1848) 281; M.Roem., Synops. Monogr. | (1846) 91; Moore, Gard. Comp. Flor. Guide (1852) 121 cum ic.; Lem., Jard. Fleur. 4 (1853) t. 360; Mig., FI. Ind. Bat. 1, 2 (1859) 535; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 4; C.DC. in DC., Monogr. Phan. | (1878) 448; Backer, Schoolfl. Java (1911) 201; Koord., Exk. Fl. Java 2 (1912) 438; Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 79; de Voogd, Trop. Nat. 1S (1927) 167, t. 3, 4; Backer & Bakh. f., Fl. Java 2 (1965) 119. [Ebermaiera pulchella Zipp. ex Span., |.c., nom. in syn. (“Ebermeyera’).] Didymocheton littorale Hassk., Tijds. Nat. Gesch. 10 (1843) 138; Cat. Pl. Bogor. (1844) 219; M. Roem., Synops. Monogr. | (1846) 105. [Didymocheton pumilum Blume ex Miq., |.c., nom. in syn.] [Trichilia humilis Zipp. ex Miq., |.c., nom. in syn.] Munronia timoriensis Baill., Adansonia 11 (1874) 266; C.DC. in DC., Monogr. Phan. | (1878) 499; Bot. Jahrb. 7 (1886) 461; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) t. 158 E; ed. 2, 19bI (1940) t. 21 E, F. Unbranched or sparsely branched shrublet to 65 cm tall. Stem to 7 mm diam., flexu- ous; rooting system with a taproot or up to five major tough roots. Bark finely fissured. Leafy shoots cicatrose, the leaves forming an apical rosette. Jndumentum of vegetative parts a mixture of simple and bifid or sometimes stellate distally-pointing hairs, most dense on innovations. Leaves imparipinnate, to 26 cm long; petiole 1.5—5.5 cm, + te- rete, pubescent. Leaflets (3, 5) 7 or 9 (11), the apical the largest, to 9 by 4 cm, ovate to obovate or subrhomboidal, the most proximal the smallest, to c. 4 by 2 cm, oval to or- bicular, scattered stellate-pubescent adaxially, with denser mixed simple and bifid hairs abaxially, esp. on veins, or both surfaces thus, apex acute to acuminate; base asymmet- ric, cuneate in apical leaflet to rounded or subcordate in most proximal; margin entire to + dentate, serrate or even lobed halfway to midrib in distal half; petiolule to 1.5 cm on apical leaflet, 0-4 mm elsewhere; nerves c. 4 on each side, arcuate, sometimes anasto- mosing well away from margin. Thyrses 2.5—15(—25) cm long, pubescent with up to 25 flowers, opening a few at a time, sometimes scented, pedicels 1-2 mm, densely hairy; bracts c. 2.5 mm long, subulate or narrowly triangular, densely hairy. Pseudo- pedicel c. 5 mm long, pubescent. Calyx lobes 5—7 mm long, spatulate, patent, pubes- cent on both surfaces. Corolla white, tube (10—)12—17 mm long, lobes 10-18 mm long, spatulate, 6—7 mm wide at broadest, acute, hairy abaxially; midvein conspicuous. Staminal tube free for apical 6—10 mm, simple hairy within (and sometimes without), margin with 10 entire appendages reflexed at anthesis; anthers yellow-brown, with short apical appendices. Disk 2—3 mm tall, cylindrical to conical, finely simple hairy. Ovary finely simple-hairy; style simple-hairy in proximal half, stylehead yellow-brown. Capsule c. 1—1.2 cm diam., depressed-globose, strongly 5-ribbed, sparsely stellate- hairy. Seeds brown. Distribution — Tropical Asia from India, Sri Lanka and eastern Himalaya to Thai- land and Malesia: Langkawi, Java, Lesser Sunda Islands (Bali, Sumba, Flores, Solor, Timor). Cultivated under glass in Europe. Habitat & Ecology — Altitude 0Q—900 m. Seasonal forests including those on lime- stone and just behind the mangrove as well as the Javanese teak forests, where, like 32 Flora Malesiana ser. I, Vol. 12 (1) (1995) Turraea pubescens, it is conspicuous when in flower in the spring, reminiscent of an Anemone (Ranunculaceae) in Europe according to De Voogd. In some parts of its range, notably Sri Lanka, it is heavily overcollected (see Uses below) and in that island, most remaining accessible specimens are small precociously flowering plants with few leaf- lets and flowers. Vernacular names — Godong lema (Jav.), singadepo laut (Sund.), Java; boan (Da- wan), Timor. Uses — Munronia pinnata is used in Ayurvedic medicine as a substitute for chiretta (Swertia spp., Gentianaceae) in the treatment of fevers and dysentery; of all native plant drugs in Sri Lanka it commands the highest price and has been used in the treatment of leprosy and other skin diseases. The whole plant is uprooted and dried to give the com- mercial materia medica. Andrographis paniculata (Burm. f.) Nees (Acanthaceae) 1s, in turn, used as a cheaper substitute for M. pinnata. Note — The description is that of Malesian material, in which there is very interest- ing variation, somewhat correlated with geography but not sufficiently clearly so for the recognition of formal infraspecific taxa based on the currently available evidence. Some plants from Timor are robust with 9 large entire or subentire leaflets and large flowers in short few-flowered inflorescences and thus resembling plants from India. Others are less robust, with more lobed leaflets and longer inflorescences, more closely resembling the bulk of the material from Java. Some specimens from that island, though, resemble the Indian plants while most of the specimens from Flores are rather delicate plants with much-lobed leaflets. The single specimen from Langkawi has the foliage of Indian spec- imens but the longer inflorescence more typical of the Javanese plants. 2. Munronia humilis (Blanco) Harms Munronia humilis (Blanco) Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 80; in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) t. 22; Pellegr. in Fl. Indo-Chine., Suppl. (1946) 688. — Plagian- thus humilis Blanco, Fl. Filip. (1837) 526; ed. 2 (1845) 367; ed. 3, 2 (1878) 315, t. 181. — Tur- raea humilis (Blanco) Merr., Philipp. Govt. Lab. Bur. Bull. 27 (1905) 30; FI. Manila (1912) 274; Philipp. J. Sc., Bot. 9 (1914) 308; Sp. Blanc. (1918) 208; Enum. Philipp. Flow. Pl. 2 (1923) 359; Craib, Enum. Fl. Siam. 1 (1926) 248; Adelb., Blumea 6 (1948) 315; Backer & Bakh. f., Fl. Java 2 (1965) 119! Turraea pumila Benn., Pl. Jav. Rar. (1840) 183; Miq., Fl. Ind. Bat. 1, 2 (1859) 534; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 4; C.DC. in DC., Monogr. Phan. | (1878) 440; Fern.-Vill., Fl. Filip. Nov. App. (1880) 41; Vidal, Rev. Pl. Vasc. Filip. (1886) 81; Backer, Schoolfl. Java (1911) 200; Koord., Exk. Fl. Java 2 (1912) 438. ?Turraea zollingeri C.DC. in DC., Monogr. Phan. | (1878) 441; ?Backer, Schoolfl. Java (1911) 200; ?Koord., Exk. Fl. Java 2 (1912) 438. Turraea palawanensis Merr., Philipp. J. Sc., Bot. 9 (1914) 307; Enum. Philipp. Flow. Pl. 2 (1923) 359; Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 80. — Munronia palawanensis (Merr.) Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 80; in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 93. Munronia pseudoturraea Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 79. Suffrutex with apparently short-lived semi-herbaceous shoots to 30 cm, but usually much less, and a woody stock to 8 mm diam. Aerial shoots unbranched or sparsely branched, 2—3 mm diam., leafy throughout. /ndumentum of simple and stellate hairs. Mabberley et al. — Meliaceae 33 Fig. 3. Munronia humilis (Blanco) Harms. a. Habit; b, c. half flower; d. detail androecium; e. fruit; f, g. seeds (a, e—g: Maxwell 75-239; b—d: Ramos 14994). Drawing R. van Crevel. 34 Flora Malesiana ser. I, Vol. 12 (1) (1995) Leaves simple, 3.5—8 by 1—4 cm, ovate or obovate, finely simply hairy especially on abaxial venation; apex acute or obtuse; base cuneate to decurrent; margin dentate or sinuately lobed, predominantly in more distal half; veins about 4 or 5 on each side, ascending, culminating in tips of major lobes; petiole 5-20 mm, pubescent. Flowers solitary or, more usually, in few-flowered thyrses up to 18 mm long, axillary to some- what supra-axillary, squarrose, pubescent; bracts c. 4 mm long, subulate, pubescent; bracteoles c. | mm long, long-triangular, pubescent. Pseudopedicels 6—9 mm long, pubescent. Calyx lobes 5-7 mm long, subulate, shortly hairy. Petals 2.75—4 cm long, 3-4 mm wide, linear-spatulate, white, cohering at base, apex acute. Staminal tube nar- rowly conical, white, with the filaments reflexed and curved outward at tips to form a subapical ruff; anthers at resultant apex, orange. Disk absent. Ovary subglabrous; style- head 5-lobed, orange-yellow. Fruit c. 1 cm, depressed globose, lobed, subglabrous. — Fig. 3. Distribution — Thailand and Malesia: Java, Philippines (Palawan, Luzon). Habitat — Seasonal forest including bamboo (Thailand) and that on limestone, thick- ets, open ravines and crevices in limestone rocks, 20—400 m altitude. Notes — 1. This species constitutes sect. Pseudoturraea Harms, characterized by the reflexed staminal tube appendages and the petals free from the tube. The third section, Philastrea (Pierre) Harms, is also monotypic but restricted to Indochina. The status of these sections cannot be evaluated in the absence of a modern revision of the genus. 2. The habit of Munronia humilis superficially resembles that of Hippobroma (Lau- rentia, Isotoma) longiflora (L.) G. Don (Campanulaceae). TRIBUS VAVAEEAE Vavaeeae Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 286; T.D. Penn., Blumea 22 (1975) 464. Trees or treelets with sympodial (‘Terminalia’-)branching. Indumentum of simple hairs. Leaves simple. Flowers mostly hermaphrodite. Staminal tube cylindrical or cya- thiform; filaments partly free. Disk obscure or absent. Ovary locules 1—multi-ovulate. Fruit a berry. Seeds with thin sarcotesta; thin endosperm sometimes present; cotyledons plano-convex, collateral. Monotypic. VAVAEA Vavaea Benth. in Hook. Lond. J. Bot. 2 (1843) 212; T.D. Penn., Blumea 17 (1969) 351; Blumea 22 (1975) 464. Lamiofrutex Laut., Nova Guinea 14 (1924) 147; Engl. in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19a (1931) 330 (sub Rutaceae). Calyx 4- or 5(—7)-lobed, with open rarely imbricate aestivation. Petals (3—)4—6, free, imbricate, rarely contorted. Filaments united in at least the proximal half; anthers 9—23, attached at ends of filaments. Disk patelliform or cyathiform, united to base of tube, or o>) Nn Mabberley et al. — Meliaceae forming androecial ribbing, or absent. Ovary 2—6-locular, locules with 1 or with 2 (3) collateral ovules or with 4—10 in 2 rows. Berry with fleshy to woody pericarp. Seeds 1—3(—7), ovoid or plano-convex; radicle superior, small, included or extending to the surface. Germination cryptocotylar, eophylls opposite, simple. Distribution — Four species from Sumatra eastwards through Malesia to tropical Australia, Micronesia, Melanesia and Polynesia, one restricted to Fiji and two to New Guinea. Divisible into two sections of two species each. KEY TO THE SPECIES la. Petals 4—9.5 mm; staminal tube 2—5.5 mm, barbate at throat; anthers 0.3—1.2 mm Poe OCUIES. L— OF 2—( OF 5-)OVULATE™ . saan he wos Be cic als ot l. V. amicorum b. Petals 35—80 mm, staminal tube 35—67 mm, not barbate at throat; anthers 2.5—6.5 mm long; locules 4—10-ovulate (sect. Grandiflorde) ...........0 000 c eens 2 2a. Calyx persistent in fruit; petals 4; staminal tube glabrous or rarely sparsely hairy without; filaments united to about halfway; ovary 4-locular; berry 12—18 mm, with EHEDESOHEPETIGALP) sic oe; vie Sepsis Mine a ees mee Ren... 19 19a. Costae c. S—8 on each side of midrib, leaves to 38 cm. — Malay Peninsula .... en At iaiee Cho cl 2s. eee. 21. C. sapindinus fateaves different, nut notrostrate:: 5.) Pataee ein Pena ee ise ede 32a. Petals c. 12 mm long, flowers crowded towards distal end of inflorescence, fruit Spuehical UR cen anls cos cea Nahm a ee 20. C. glirioides b. Petals up to 10 mm long, or, if slightly longer, then the tube villous within and the ledvesipilose abaxiallyiveswaiu ase ees cathe ctpoatelowwk meek OS 33a. Flowerbuds 9-10 mm, anthers 4—6 (7), style to0.15 mm ..... 17. C. sayeri 140 Flora Malesiana ser. I, Vol. 12 (1) (1995) b. Flowerbuds c. 11.5 mm, anthers 6—8, style 0.25 mm diam. .. 16. C. pilosus 34a. Inflorescence borne on supra-axillary branch resembling supra-axillary inflores- cence; petiolules fulvous tomentose, disk 1 mm .......... 22. C. laosensis b. Inflorescences: diterent-diskeabsenty Xen. tei eet 15. C. lasiocarpus 35a. Inflorescences borne on supra-axillary branch resembling supra-axillary inflores- cence; petiolules! fulvous:tomentose® =. 4232s a...) ae 22. C. laosensis b.. Inflorescences different we... AR. DER RRE 36 364), Hlaursisimaple stan os: ec We hab eee ATE, De RSA 21 22 ee 37 b. Hairs stellate, disk present, seeds sarcotestal. — Borneo eastwards......... 50 37a. ‘Corolla.aestivationambiicates.. .)..78.) 594s 20: oN. ee ee 4] 4la. Calyx 4—5-lobed, petals 5 or 6, anthers scarcely locellate ... 26. C. amabilis b. Calyx margin entire, petals (3) 4 (5), anthers locellate .. 27. C. cumingianus AD ae DiSkspreSemey (102%. cts'R Ge SASS 3 et ee SE ec 43 Dep Diskcalbsemt io etk mages sc chee ceese Oe delta. Se ey cece epi sacle ee a oh 44 43a. Venation very prominent on both sides (dried), seeds 3 cm diam. — Borneo .... URI Ee tte ar oe cae KGS GN IE Chega eg: 29. C. lansiifolius biaWenationvand scedsrdifferent 415612 2) eee noe tee 28. C. patens AAa. Petalselisammiulong.=—iPhilippimese.)\azieys kis oneal deen 35. C. mendozae brtRetals:Shorter cect use yew Rae A, 9 45 Adan. dubercrenate;antherssNainyccicters oe, aes ae iota te omen 24. C. sarawakanus b. Tube lobes long-triangular, anthers glabrous ............... 28. C. patens AG AIMDISKE PRESEN 3/5. = cede ov hares, wcedonct ck eo aE Rt ee See 36. C. vindictae bAeDiskiab sem terval se Se ete le FEN AS OM et 47 AT Mam ACS ISUb SlADTOUS! a4 27s elie ola cS oes ccaeeye sol ed Salts ee 48 by, Miwigs fawn PuUbeSCeMmte 224.622 sso i w xe tN RSE RE RTL erica 49 48a. Petals 13-19 by 2 mm, tube lobes + truncate, flowers in panicles, fruit 4.5 cm or TUN OLS Meas Solh kce TEINS ee ETN, EOE aa Lok seta Se Nak ane 38. C. ceramicus b. Petals 8—12(—19) mm, tube lobes + laciniate, flowers in panicles only if flowers Smallesinnititom. lem diam: az).in. nest ee Ree 40. C. pentandrus 49a., BetalsMlounmeilongs— Philippines, ......... -Se02@t. Ms ee 39. C. curranii bt Betalsio=iBammitlong ties: ahe.wteek beleeet ee 41. C. erythrocarpus 50a. Flowers 11-12 mm long. — Borneo, Celebes .......... 42. C. koordersii & PilowesstlessithanySimmnilon eps eee ORD. ae, eet a ee 51 51a. Inflorescence velutinous, style glabrous. — New Guinea ... 43. C. stellatus b. Inflorescence subglabrous, style densely hairy ...... 44. C. longistipitatus Mabberley et al. — Meliaceae 14] Section Clemensia Chisocheton sect. Clemensia (Merr.) Airy Shaw in Hook. Ic. Pl. 34 (1937) t. 3333; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 320. — Clemensia Merr. (genus), Philipp. J. Sc., Bot. 3 (1908) 143; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 155. Chisocheton § Graciles Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 295 (in sect. Euchiso- cheton); in ibid., ed. 2, 19b1 (1940) 153. Pachycaul trees and treelets to 28 m tall, unbranched or sparsely branched. Leaves to 220 cm long, imparipinnate or pseudogemmulate. /nflorescences unbranched or sparse- ly branched, to 7 m long, + flagelliform with arillate seeds, or shorter with sarcotestal seeds. Calyx + pubescent, margin truncate or variously 3- or 4-lobed. Petals (4—)5-—14, (16—)26—45 mm long, imbricate at apices. Staminal tube glabrous or sparsely pubes- cent without, with band of hairs below lobes and/or sparsely pubescent within up to apical quarter, margin entire to lobed; anthers 3—30, hairy or not, locellate or not. Disk flattened or annular, sometimes lobed. Ovary 4—6-locular; stylehead subdiscoid to capi- tate. Capsule to 13 cm diam., recurved, tomentose with stinging hairs. Seeds arillate or sarcotestal, never scutellar. Distribution — Seven species restricted to western Malesia. Note — The features which distinguish other sections intermingle here, so that sect. Clemensia may be considered a relict group from which the others have been derived. That is not to say, of course, that all features of this section are primitive. 1. Chisocheton macranthus (Merr.) Airy Shaw Chisocheton macranthus (Merr.) Airy Shaw in Hook. Ic. Pl. 34 (1937) t. 3333; Jacobs, Reinwardtia 3 (1955) 266: Meijer, Bot. News Bull. Sabah 8 (1967); Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 320, t. 1. — Clemensia macrantha Merr., Philipp. J. Sc., Bot. 3 (1908) 144; J. Str. Br. Roy. As. Soc., spec. no. (1921) 321; Enum. Philipp. Flow. Pl. 2 (1923) 371; Univ. Calif. Publ. Bot. 15 (1929) 122; Elmer, Leafl. Philipp. Bot. 9 (1937) 3349; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 155, t. 34; Heine, Feddes Rep. 54 (1951) 230. [Dysoxylum dehiscens Elmer, Leaf}. Philipp. Bot. (1937) 3349, nom. in syn.] Chisocheton medusae auct. non Airy Shaw: Heine, Mitt. Bot. Staatssamml. Miinchen 6 (1953) 233. Pachycaul tree to 13 m; bole to 22 cm diam., buttressed. Branching fastigiate, often with several limbs from near base. Bark smooth. Leafy twigs to 5 cm diam., blackish, with large scutellar cicatrices. Leaves to 220 cm long, pseudogemmulate, crowded in dense terminal spirals; petiole 5—20 cm, woody, dark-coloured, glabrescent to sparsely hairy. Leaflets in up to 16 pairs, sometimes + alternate at base or rachis, 20—45(—55) by (S—)8—12(—15) cm, oblong lanceolate or ovate when smaller, weakly bullate, gla- brous adaxially, + puberulous abaxially, base obtuse to subacute, apex acute to acumi- nate, costae c. 15—24 on each side, tertiary venation scalariform; petiolules to 8 mm. Inflorescence to 220 cm long, pendent; axis terete to weakly angled, weakly branched; branches crowded towards apex, pilose, with up to 12 flowers each; bracts c. 6 mm log, pubescent, caducous; pedicels c. 10 mm long, articulated with pseudopedicel. Calyx 14—20 mm long and across, cupular to cylindrical, pubescent, red-brown, mar- gin truncate to irregularly 3- or 4-lobed. Petals 6-10, 30—45 by 4—7(—12) mm, creamy 142 Flora Malesiana ser. I, Vol. 12 (1) (1995) pink. Staminal tube 25—40 by 6—7 mm, creamy white, glabrous without except on the lobes, pilose within at base, margin entire or with lobes 4—6 mm long; anthers 16—30, c. 5 mm long, rather recurved, connective somewhat pubescent. Disk flattened to weak- ly annular, glabrous. Ovary c. 5 mm diam., 5- or 6-locular, pilose; style pilose in proxi- mal half or glabrous; stylehead c. 2 mm diam., capitate. Infructescence borne in axils of last flush of leaves; axis to 3 m with terminal bunches of up to 60 fruits. Capsule to 12 cm diam., recurved, rostrate when immature, bright vermilion tomentose with irritant deciduous hairs. Seeds 2.5—3.3 cm long, triangular in cross section; aril reddish, cov- ering inner edges of black testa. Distribution — Malesia: northern Borneo (Sarawak, Sabah, E Kalimantan) and south- ern Philippines (Surigao, Mindanao). Habitat — Lowland rain forest including that on limestone, to 400 m altitude. Note — The stinging hairs and pachycaul habit are found in a number of Dysoxylum species, e.g. D. sessile (q.v.). 2. Chisocheton medusae Airy Shaw Chisocheton medusae Airy Shaw in Hook. Ic. Pl. 34 (1937) t. 3333; Jacobs, Reinwardtia 3 (1955) 264, incl. f. hiascens Jacobs; Meijer, Bot. News Bull. Sabah 8 (1967) 78; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 322. Megaphyllaea sp.: Merr., Univ. Calif. Publ. Bot. 15 (1929) 123. Pachycaul tree to 28 m; bole to 30 cm diam., sparsely branched, buttressed. Bark black with fine striations; inner bark dark brown; heartwood yellowish. Leafy twigs to 2.5 cm diam., fulvous tomentose. Leaves to 2 m long, pseudogemmulate (imparipin- nate and up to 4-jugate when young), bunched in terminal spirals; petiole 10—20 cm, terete or flattened adaxially, decurrent with twig and forming axillary cavity with it, + fulvous tomentose; rachis somewhat angular, glabrescent to fulvous tomentose. Leaf- lets in up to 14 pairs, opposite except for those near base of rachis, to 40 by 11 cm, lan- ceolate to elliptic-lanceolate, glabrous adaxially when mature, + densely fulvous pubes- cent abaxially, green when young, base narrowed into petiole, apex acute to acuminate when tip to 2 cm long, midrib stout, densely fulvous tomentose abaxially, costae 20—24 on each side, weakly arcuate near margin, subpubescent adaxially, prominent and pu- bescent abaxially. Inflorescences to 30 cm long, borne in axils of upper or undevel- oped leaves, weakly branched to narrowly paniculiform at base; axis flattened to angu- lar, shortly fulvous tomentose when young, glabrescent later; branches rather more densely pubescent, few-flowered with caducous bracts; pedicels 3—20 mm, somewhat angular, light-brown hirtellous, articulated with pseudopedicel, swollen at articulation. Calyx (10—)13—20(—23) mm long, 15—20 mm diam., cupular to subcylindrical, + den- sely ferrugineous-velutinous, margin truncate or irregularly split to halfway into 2 or 3 + triangular lobes. Petals 9-14, 35-40 by 2—6 mm, white. Staminal tube 27—32 mm, glabrous, thin proximally, margin truncate; anthers 15—20, slightly exserted or in- cluded, 3—4 mm long, glabrous. Disk glabrous. Ovary in female flowers 3—5 mm diam., 7- or 8-locular, glabrous to densely yellow hairy; the style + pubescent, especi- Mabberley et al. — Meliaceae 143 ally below the stylehead 2 mm diam., discoid to shal- lowly cylindrical, glabrous. Infructescence to 30 cm. Capsule to 13cm long, 10 cm diam., golden brown, densely hispid. Seeds to 5 cm long and orange-seg- ment-shaped; the sarcotes- ta densely vascularized. — Fig. 22. Distribution — Malesia: in northern Borneo (Sara- wak, Sabah, E Kaliman- tan). Habitat — Lowland rain forest to 300 m_ altitude, sometimes on limestone. Note — This species has a short infructescence with sarcotestal seeds whereas its apparently closest ally Chi- socheton macranthus has a flagelliform infructescence and arillate seeds. Chisoche- ton tomentosus (sarcotesta) and C. polyandrus (aril) are a similar pair but geographi- _ eaves. Sabah, Sepilok Forest Reserve (Mabberley 1680). Photo- cally vicarious. graph D. J. Mabberley, May 1974. 3. Chisocheton tomentosus (Roxb.) Mabb. Chisocheton tomentosus (Roxb.) Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 323, f. 1 (2); in Tree Fl. Malaya 4 (1989) 238. — Melia tomentosa Roxb., Hort. Beng. (1814) 90, nom. nud.; G. Don, Gen. Syst. 1 (1831) 681, nom. nud.; Roxb., Fl. Ind., ed. Carey, 1 (1832) 394; Walp., Rep. 1 (1842) 427; M. Roem., Synops. Monogr. | (1846) 96; Hiern in Hook. f., Fl. Brit. India 1 (1875) 543; C.DC. in DC., Monogr. Phan. | (1878) 458; Curtis, J. Str. Br. Roy. As. Soc. 25 (1894) 21. — Azedarach tomentosa (Roxb.) Kuntze, Rev. Gen. Pl. 1 (1891) 110. [Meliacea rugosa Wall., Cat. (1831/2) n. 4891, nom. nud.] Chisocheton princeps Hemsl. in Hook. Ic. Pl. 19 (1889) t. 1884; Curtis, J. Str. Br. Roy. As. Soc. 25 (1894) 22; King, J. As. Soc. Beng. 64, ii (1895) 29; Ridley, Fl. Malay Penins. | (1922) 388; Whit- more, Trop. Rain For. Far East (1975) t. 2.7. Chisocheton rubiginosus King, J. As. Soc. Beng. 64, ii (1895) 29; Ridley, Fl. Malay Penins. | (1922) 389; Burkill & Hend., Gard. Bull. Str. Settl. 3 (1925) 357. Chisocheton rugosus Pierre, Fl. For. Cochinch. 5 (1897) sub t. 347. 144 Flora Malesiana ser. I, Vol. 12 (1) (1995) Pachycaul tree to 21 m; bole to 20 cm diam., unbranched or sparsely and fastigiately branched, often from near base, sometimes slightly fluted below, knobbled or with small stiltroots, taprooted at least when young. Bark blackish brown, smooth to weak- ly fissured, with conspicuous scutellar cicatrices to 5 cm wide and long above; inner bark deep orange-yellow to brownish; sapwood ivory to fawn; pith wide, soft, white. Leafy twigs to 2 cm diam., all young parts brown-tomentose with irritant hairs, some- times inhabited by ants [Pannell, Kew Bull., Add. Ser. 16 (1992) 16]. Leaves to 2 m long, in terminal rosettes, maturing in flushes, pinkish red when young, imparipinnate, especially on saplings, or pseudogemmulate, pseudogemmula sometimes falling with- out further development, petiole 10—20 cm, terete, base swollen, massive, woody; rachis often angled, brown-tomentose. Leaflets 3—37 by 2—10 cm, the most proximal smallest, narrowly elliptic to oblong, sessile to subsessile, patent, rugose, shiny and glabrous adaxially except for brown-tomentose midrib, tomentose or tawny pubescent abaxially, surface strongly reticulate-areolate, costae 12—30 on each side, arcuate. /n- florescences borne in upper axils, appearing when fruit on old infructescences maturing (female trees), to 90 cm; axis massive, tough with flowers forming a terminal head to 45 by 10 cm, sometimes with short branches to 7 cm, composed of fascicles of sweetly scented pedicellate flowers, brownish pink in bud. Calyx (3—)4—8 mm tall and in diam., cupular, reddish brown, minutely puberulous, often warty, margin entire or obscurely 3- or 4-lobed, + setose. Petals 5 or 6(—10), 26—37 mm long, linear-spathulate, creamy red-striped without, white flushed pink within, waxy, concave distally, densely pilose without. Staminal tube cylindrical, slightly widened at mouth, villous with downward- pointing hairs within lower 3/4, sometimes sparsely hairy without especially just below lobes, lobes c. 10—15, often irregularly bifid, shorter than anthers; anthers 7—13(—15), 4.5—5 mm long, boat-shaped, locellate, glabrous, basifixed. Disk to 1 mm tall, annular, apically pilose. Ovary (4-) 5- (6-)locular; style cylindrical, sparsely pubescent; stylehead spherical. Capsule to 7 cm diam., subglobular, (4-) 5- (6-)locular, golden brown vel- vety with deciduous irritant hairs. Seeds 3—5, to 4 cm long; sarcotesta white. Distribution — Malesia: Sumatra (Barisan Range, De Vogel 2750, L), Malay Penin- sula. Habitat — Lowland and hill dipterocarp forest to 400 m altitude. Note — The type is a drawing prepared for Roxburgh when the tree was at Calcutta Botanic Garden and is reproduced in Ic. Roxb. Drawings Ind. Pl. 3 (1969) 16. 4. Chisocheton polyandrus Merr. Chisocheton polyandrus Merr., Philipp. J. Sc. 21 (1922) 520; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. Oi(1979) 3245 t. 2ise fo iC). Pachycaul tree to 15 m, unbranched or very sparsely branched, occasionally with stiltroots or small buttresses; inner bark pinkish; wood fawn. Leaves to 150 cm long, imparipinnate to 14-jugate or pseudogemmulate, when pseudogemmula densely long pubescent; petiole 1-5 cm, terete, woody; rachis terete. Leaflets 11-43 by 5-13 cm, oblong-lanceolate, opposite or subalternate at base of rachis, where they are often very Mabberley et al. — Meliaceae 145 small and even irregularly lobed, shiny, bullate at altitude, glabrous or veins + pubes- cent adaxially, appressed hirsute, particularly at altitude abaxially, base asymmetrical, cuneate to subcordate, apex somewhat acuminate, costae c. 15 on each side, often sunken adaxially; petiolules to 2 mm. /nflorescences borne in upper axils, up to 4 at a time, to 2 m long, unbranched or with a few squarrose branches to 13 cm long near tip, where flowers are condensed; axes ferruginous-pubescent when young. Calyx 5—8 mm long, 5—6 mm in diam., cupular to subcylindrical, densely ferruginous pubescent, green to deep red, margin truncate. Petals 5 or 6, 28—32 mm long, subspathulate, fleshy, creamy white with conspicuous pink or red tinge, densely pubescent without. Staminal tube cylindrical, white, subglabrous save conspicuous bands of hairs apically and basal- ly within, margin with c. 12—14 linear lobes, c. 3 mm long; anthers 12—14, c. 4 mm long, locellate, connective scattered ferruginous-pubescent. Disk c. 1 mm tall, thick, gla- brous, margin truncate. Ovary ?3—5-celled; style cylindrical glabrous; stylehead subcapi- tate, c. | mm diam. /nfructescence to 2 m long, pendent, with fruit aggregated at tip. Fruit spherical, covered with reddish irritant hairs, splitting into 3 or 4 valves. Seeds 3; testa black, covered on inner surface by orange-red aril. Distribution — Malesia: Borneo (northern Sarawak, Brunei, Sabah, S Kalimantan). Habitat & Ecology — Lowland and hill dipterocarp forest, 150—300 m altitude. Flow- ers are visited by spiderhunters. 5. Chisocheton penduliflorus Planch. ex Hiern Chisocheton penduliflorus Planch. ex Hiern in Hook. f., Fl. Brit. India 1 (1875) 550; C.DC. in DC., Monogr. Phan. | (1878) 536, t. 7 f. 4; Curtis, J. Str. Br. Roy. As. Soc. 25 (1894) 22; King, J. As. Soc. Beng. 64, ii (1895) 38; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 292, t. 162 f. E—G; ed. 2, 19b1 (1940) 139, t. 30 f. E—G; Ridley, Fl. Malay Penins. 1 (1922) 388, incl. var. kunstleri (King) Ridley; Burkill & Hend., Gard. Bull. Str. Settl. 3 (1925) 356; Briq., Mém. Inst. Nat. Genev. 24 (1935) 66; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 326; Mal. For. 45 (1982) 451; in Tree Fl. Malaya 4 (1989) 237. — [Melia penduliflora Wall., Cat. (1828) n. 1255, nom. nud.; M. Roem., Synops. Monogr. | (1846) 96, nom. nud.] Chisocheton kunstleri King, J. As. Soc. Beng. 64, ii (1895) 27. Pachycaul treelet or tree to 10 m; bole to 10 cm diam. Bark blackish; inner bark pale fawn. Leafy twigs 6-7 mm diam. (—15 mm in fruit), + densely to rusty tomentose. Leaves imparipinnate or pseudogemmulate, to 8-jugate; petiole to 22 cm, + channelled adaxially. Leaflets (8—)17—27.5 by (3—)8—11.5 cm, elliptic-ovate to elliptic-oblong, sub- coriaceous, subglabrous except for + fulvescent-tomentose on veins adaxially, + pubes- cent particularly on veins abaxially, base rounded to subcordate, sometimes asymmetri- cal, apex + gradually long-acuminate, costae c. 15—22 on each side with conspicuous secondaries, prominent abaxially; petiolules to 1 mm. /nflorescence to 7 m, supra-axil- lary, pendent, unbranched or with distal branches to 7 cm long, usually with flowers congested in subsessile cymes at distal end, like a bellrope; axis 3—5 mm diam., dense- ly rusty tomentose; bracts and bracteoles linear, hirsute. Calyx 3—4 mm tall, cupular to shortly cylindrical, pubescent, margin truncate to obscurely lobed. Corolla clavate in bud. Petals 4 or 5, 18—22 mm long, linear-spathulate, concave, thick, dull red, pubes- cent without, adnate to staminal tube below. Staminal tube pilose without below lobes, 146 Flora Malesiana ser. I, Vol. 12 (1) (1995) long-pilose with downward directed hairs in lower half within, margin with 3—5 irregu- larly bifid lobes; anthers 3—5, c. 3 mm long, locellate, glabrous, included or slightly exserted. Disk obscure to shallowly cupular, fleshy, glabrous. Ovary 3(?—5)-locular, densely long-pilose; style densely long-pilose in proximal 3/4; stylehead subdiscoid to capitate with + pubescent annulus. Capsules in infructescences of c. 10—15, to 5 cm long, recurved, rostrate when young, silky pubescent with (?)irritant hairs, splitting into 3 valves. Seeds 3; testa black, covered on inner surface by red-orange aril. Distribution — Lower Thailand and Malesia: Malay Peninsula, Riau/Lingga Archi- pelago. Habitat — Lowland rain forest to 900 m altitude. 6. Chisocheton crustularii Mabb. Chisocheton crustularii Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 327, f. 3. Pachycaul tree to 8 m; bole to 8 cm diam., sometimes with small rounded buttresses. Bark greyish, narrowly fissured. Leafy twigs c. 1.5 cm diam. Leaves to 135 cm long, imparipinnate, to 10-jugate; rachis terete, subglabrous, lateral leaflets opposite to 38 by 10 cm distally, 1.8 by 1 cm and pseudostipulate proximally, the terminal c. 45 by 13.5 cm, narrowly elliptic or oblong, glabrous adaxially, sparsely strigose, especially on venation abaxially, base shortly attenuate to subtruncate, apex acuminate, costae c. 24 on each side, prominent abaxially, tertiary venation conspicuous, somewhat scalari- form; petiolules 0-16 mm, swollen. /nflorescence (male only known) 38(—150) cm long, pendent, slender; axis 2-3 mm diam., sericeous, the flowers in fascicles in the distal half; pedicels 2—4.5 mm, recurved, hispid. Calyx 3 mm long, c. 5.5 mm diam., cupular, pubescent without, margin entire to weakly crenate. Petals 5, 16 by 4.5 mm, narrowly oblong, white, appressed pilose without, glabrous within. Staminal tube 14.5 mm long, subglabrous save broad band of adpressed hairs apically without, margin with 11 irregularly bifid lobes c. 3.5 mm long; anthers 11, c. 2 mm long, included, hardly locellate. Disk c. 0.5 mm tall, margin with recurved lobes. Style filiform with a tuft of long hairs at its base; stylehead c. 1 mm diam., spherical. Fruit unknown. Distribution — Malesia: Borneo, known from only two collections from Tinjar, N Sarawak. Habitat — Presumably rain forest. 7. Chisocheton setosus Ridley Chisocheton setosus Ridley, Bull. Misc. Info. Kew (1930) 366; Airy Shaw in Hook. Ic. Pl. 34 (1937) t. 3334; Meijer, Bot. News Bull. Sabah 8 (1967) 78; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 C1979) 327. Pachycaul treelet to 5 m, ?unbranched; bole c. 8 cm diam. Bark smooth; inner bark pale yellow. Leafy twigs c. 1 cm diam., densely ferruginous-setose. Leaves to 1 m long, imparipinnate, at least 6-jugate; petiole to 35 cm long, subterete, sometimes Mabberley et al. — Meliaceae 147 grooved adaxially, to 6 mm diam., ferruginous-setose with hairs 2—3 mm long, base swollen with conspicuous hollow at junction with shoot; rachis 1- or 2-sulcate, setose as petiole. Leaflets + densely ferruginous setose on both sides, setae tinkling when stroked (in sicco), pale when dry, proximal leaflets to 20 by 8.5 cm, elliptic oblong, distal to 36 by 10 cm, oblanceolate to oblong, base rounded to attenuate, apex acumi- nate, acumen 10—20 mm, costae 17—20 on each side, prominent abaxially, tertiary venation + prominently scalariform; petiolules 5-6 mm long, densely tomentose, that of terminal leaflet to 10 mm. /nflorescence to 2 m, pendulous, + densely setose, drying irregularly angled with flowers crowded in condensed cymes at distal end like a bell- rope; bracts c. 7 mm long, setose. Flowers recurved, shortly pedicellate. Calyx c. 3 mm long and 4 mm diam., cupular, somewhat elongated into a pseudopedicel, reddish, se- tose to pubescent, margin truncate to obscurely 3- or 4-lobed. Corolla 3—3.5 cm long, c. 3 mm diam., weakly clavate, glabrous, white or greenish; petals 4—6, to 4 mm wide, subspathulate, imbricate at spices. Staminal tube to 3.2 cm long, glabrous except for a band of hairs below lobes without, white, margin with 6—8 truncate or irregularly lobed lobes; anthers 6—8, c. 2 mm long, glabrous, scarcely locellate. Disk c. 1 mm tall, cupu- lar, glabrous, margin truncate to obscurely lobed. Ovary in female flowers unknown; style sparsely pilose proximally, glabrous distally; stylehead capitate, distinctly narrow- annular apically. Fruit (unripe) pale yellow, densely setose. Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, C Kalimantan). Collected only five times. Habitat — Presumably rain forest. Section Chisocheton Chisocheton sect. Chisocheton: Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 329. — Chisoche- ton sect. Euchisocheton Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 295, p.p. — Schi- zochiton Spreng. sect. Euschizochiton Kuntze in T. Post & Kuntze, Lex. Gen. Phan. (1903) 505, nom. superfl. Chisocheton sect. Tetrapetalum Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 26. — Chisocheton § Tetra- petali: (Miq.) Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 151. Chisocheton sect. Hexapetalum Miq., |.c. Trees or undergrowth treelets. /ndumentum of simple hairs. Leaves pseudogemmu- late, rarely imparipinnate (C. /asiogynus) or paripinnate (C. patens, p.p.). Inflorescences axillary, ramiflorous, cauliflorous or epiphyllous, or borne on supra-axillary branches. Corolla aestivation alternative, quincuncial or imbricate, white or cream to pink or red; petals 4—6. Staminal tube lobed or not; anthers (3—)5—10(—18), locellate. Disk some- times present. Ovary 2—8-merous; stylehead capitate. Fruit unarmed. Seeds 3—6, arillate. Distribution — 23 species throughout the range of the genus except Australia and New Hebrides. Notes — 1. The species are arranged in two series though these are linked by a small number of species which could be placed in either. 2. Allied to sect. Clemensia through arillate species of the latter. 148 Flora Malesiana ser. I, Vol. 12 (1) (1995) Series Schumannianti Chisocheton sect. Chisocheton series Schumanniani Harms ex Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 329. — Chisocheton § Schumanniani Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 153, nom. non rite publ. Leaves pseudogemmulate. Inflorescences (supra-)axillary or epiphyllous. Ovary 3—8- locular. Aril covering only half of testa. Distribution — 14 species from Celebes to the Solomon Islands, with 1 in Sumatra. 8. Chisocheton warburgii Harms Chisocheton warburgii Harms in Fedde, Rep. 42 (1937) 9; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 372; Mal. For. 45 (1982) 451. Tree to 7 m. Branchlets + densely hirsute-villous, leafy ones c. 1 cm diam. Leaves c. 1 m long, at least 15-jugate; petiole 7-15 cm, + villous. Leaflets 9-18 by 4—6 cm, oblong to oblanceolate or oblong-ovate, + pilose abaxially and sometimes on veins ad- axially, bases acute to obtuse, + asymmetric, apices narrowly acute, costae c. 10-16 on each side, arcuate, prominent abaxially in sicco; petiolules 5-10 mm, + pilose. Thyrse spiciform, to | m or more long, (to 280 cm in fruit) supra-axillary, the flowers on short branches or in fascicles forming a head in distal 40 cm; axis c. 5 mm diam., + pilose. Flowers subsessile. Calyx c. 5 mm long, 4 mm diam., cylindrical cupulate, densely vil- lous without, margin truncate. Petals 4 or 5, 10-15 by 2—3 mm, the innermost nar- rowest, ligulate, narrowly imbricate, adpressed pubescent without. Staminal tube sub- coriaceous, + with a sparse region of hairs outside near apex, glabrous within, margin truncate; anthers 8 or 9 (10), linear inserted c. 2 mm within throat, included, basifixed. Ovary densely villous; style glabrous apically; stylehead clavate-capitate. Capsule c. 5 cm diam., warty, + pubescent, recurved 4-valved, the valves woody, recurved, shortly and abruptly stipitate, apex weakly beaked. Seeds 4, c. 2.5 cm long, orange-segment- shaped. Distribution — Malesia: C and N Celebes. Habitat — Rain forest, 400—500 m altitude. Note — Flowering material was recollected by Kjellberg in Central Celebes in 1929, but the plant is known from only four surviving collections. 9. Chisocheton schoddei P.F. Stevens Chisocheton schoddei P.F. Stevens, Contr. Herb. Austral. 11 (1975) 38, t. 5; in Handb. Fl. Papua New Guinea | (1978) 161, t. 78; Mabb., Bull. Brit. Mus. Nat. Hist. 6 (1979) 329. Pachycaul tree to 12 m; bole to 15 cm diam. Bark brown to greenish grey, rather smooth; inner bark creamy white to fawn. Leafy twigs c. 7-8 mm diam., + terete, pu- bescent when young. Leaves to 1.3 m, bunched in terminal spirals; petiole 1O—15 cm; axis c. 3.5 mm diam., + terete. Leaflets in up to 10 pairs, 12—38(—50) by 6.5—14.5 (—20) cm, though most proximal much smaller, about 10 cm apart, ovate to oblong, Mabberley et al. — Meliaceae 149 sparsely pubescent adaxially, slightly more so abaxially, base + subcordate, apex acu- minate, costae 8—14 on each side, venation prominent abaxially. Thyrse to 1.2 m long, narrowly paniculate, borne in foliate axils; axis + pubescent, unbranched or with distal branches to 5 cm long; bracts c. 2 mm long, broadly ovate. Calyx 5—6.5 mm long, shal- lowly cupulate, pubescent without, margin truncate. Petals (4) 5 or 6, to 17 by 7 mm, ligulate, elliptic, red with- out, white to greenish with- in, + pubescent without. Staminal tube densely hairy within, less so without; mar- gin truncate to crenulate; the anthers 10-12, c. 3 mm long, locellate, connective + pubescent. Ovary c.4mm in diam., (4-) 5—8-locular, densely pubescent; the style pubescent for most of its length, occasionally gla- brous near apex; stylehead c. 1.5 mm diam., capitate. Infructescence to 65 cm; rachis c. 4 mm diam. Fruits crowded at apex, 6cm in diam., flattened globose, (4—)6(—8)-merous, densely tomentose, pericarp thick, creamy. Seeds up to 6, c. 18 mm long; testa black; aril orange-red covering adaxial half. — Fig. 23. Distribution — Malesia: southern Papua New Gui- OLAS ee iin Fig. 23. Chisocheton schoddei P.F. Stevens. P. Katik holding a Habitat — Rain foreststo fruiting twig. Papua New Guinea, Brown River (Mabberley 1773). 180 m altitude. Photograph D. J. Mabberley, July 1974. 10. Chisocheton tenuis P.F. Stevens Chisocheton tenuis P.F. Stevens, Contr. Herb. Austral. 11 (1975) 46, t. 7; P.F. Stevens in Handb. FI. Papua New Guinea | (1978) 168, t. 81; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 330; Fisher & Rutishauser, Can. J. Bot. 68 (1990) 2317. Chisocheton pohlianus auct. non Harms (1917): Harms, Bot. Jahrb. 72 (1942) 187, quoad spec. cit 150 Flora Malesiana ser. I, Vol. 12 (1) (1995) Tree to 8 m; bole to 7.5 cm diam. with ascending branches. Bark pale fawn; inner bark straw. Leafy twigs c. 4 mm diam. Leaves to 70 cm long, up to 7-jugate; petiole 5—10 cm; rachis 2—3 mm diam., terete with minute pseudogemmula and inflorescence scars and minute vegetative (Fisher & Rutishauser, I.c.) and inflorescence buds. Leaf- lets 7.5—25 by 3.3—9.5 cm, obovate to elliptic, + glabrescent except for hairier midrib, apex weakly acuminate, costae 6—12 on each side; petiolules (S—)10—23 mm long. Jn- florescence to 9 cm long, not or sparsely branched, epiphyllous and usually arising near petiolules; bracteoles to 1.5 mm long; pedicels 2-12 mm. Calyx 3—4 mm long, cupulate to cylindrical with pseudopedicel c. | mm long, pubescent, margin truncate, sometimes + split at one point. Petals 4, 7.5 mm long, + densely adpressed pubescent without, pinkish, alternative. Staminal tube c. 6.5 mm long, pubescent within and, except at top and bottom, without, margin + truncate; anthers 7 or 8, c. 2 mm long, locellate, insert- ed c. 3 mm within tube, connective pubescent. Disk small. Ovary pubescent (female flowers unknown); style with long ascending hairs except near apex; stylehead c. 0.5 mm diam. Fruits to 6 cm long, 2.3 cm diam., fusiform, 3- or 4-locular, + pubescent. Seeds 3 or 4, c. 18 mm long, + ellipsoid; testa black; aril orange-red, covering adaxial surface. Distribution — Malesia: New Guinea (W Sepik Province and highlands of Papua New Guinea). Habitat — Lowland and lower montane rane forest, to 1700 m altitude. 11. Chisocheton cauliflorus Merr. Chisocheton cauliflorus Merr., Philipp. J. Sc., Bot. 11 (1916) 188 (‘Chisochiton’); Enum. Philipp. Flow. Pl. 2 (1923) 366; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 330. Treelet to 3 m; bole to 4 cm diam.; young parts + fulvous villous or hirsute. Leafy twigs c. 5 mm diam., glabrous. Leaves to 50 cm long, to 6-jugate; petiole c. 10 cm, + conspicuously fulvescent like rachis. Leaflets 10-22 by 4—7 cm, oblong or (proximal ones) elliptic and weakly lobed, bases + rounded, apices acuminate, costae 8—15 on each side, venation + conspicuously fulvescent abaxially, prominent; petiolules 5—10 mm long. Inflorescences to 50 cm, narrowly paniculiform, borne on tubercles on trunk, branches, or axillary; axis 1.5 mm diam., + prominently fulvous villous, with few, few- flowered branches to 4 cm. Calyx 4—5 mm long, cupular to cylindrical, + densely ad- pressed fulvescent, margin truncate. Petals 4, to 18 mm long, spathulate, pinkish red, + subvillous without, alternative. Staminal tube c. 16 mm long, cylindrical, villous with- out, glabrous within, margin truncate; anthers 8, c. 2 mm long, locellate, glabrous, in- serted c. 1.5 mm within tube. Ovary densely villous; style adpressed pubescent except in distal third; stylehead subcapitate. Female flowers and fruits unknown. Distribution — Malesia: Philippines (Samar, Mindanao). Habitat — Lowland rain forest, to 470 m altitude. Note — The Mindanao material is somewhat less pubescent than the material from Samar. Mabberley et al. — Meliaceae 151 12. Chisocheton novobritannicus P.F. Stevens Chisocheton novobrittannicus P.F. Stevens, Contr. Herb. Austral. 11 (1975) 22, t. 3: in Handb. FI. Papua New Guinea | (1978) 151, t. 73; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 331. Tree to 13 m; bole to 20 cm diam. Leafy twigs c. 6 mm diam., terete, long villous when young. Leaves to 125 cm long, to 1 1-jugate; petiole 5—15 cm, rachis 5 mm diam. Leaflets 13—37 by 6.5—15 cm, ovate to oblong, subglabrous adaxially or venation + pu- bescent, abaxially glabrous save fine venation, bases rounded, sometimes asymmetric, apices acute to acuminate; costae 10—17 on each side; petiolules to 12 mm. /nflorescences to 80 cm long, axillary, narrowly paniculiform, 2-branched with branches to 2 cm, pa- tent or reflexed, the flowers + sessile. Calyx 1.5—2 mm deep, shallowly cupular, with pseudopedicel c. | mm long, pubescent without, margin truncate. Petals 4, c. 16 mm long, alternative, white, glabrous without except sometimes pubescent near apex. Sta- minal tube c. 14 mm long, glabrous or with spreading hairs near base within, margin truncate to very shallowly lobed; anthers 8 or 9, 1.5—2.5 mm long, locellate, glabrous. Disk c. 0.5 mm tall, glabrous. Ovary small (female flowers unknown); style densely pubescent in proximal half; stylehead c. 1 mm diam. Capsule c. 3.2 cm diam., 4-locu- lar, sparsely pubescent. Seeds c. 2 cm long, ellipsoid; aril on adaxial surface. Distribution — Bismarck Archipelago (New Britain). Habitat — Lowland rain forest, to 150 m altitude. 13. Chisocheton montanus P.F. Stevens Chisocheton montanus P.F. Stevens, Contr. Herb. Austral. 11 (1975) 18, t. 2; in Handb. Fl. Papua New Guinea | (1978) 147, t. 71; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 331; Fisher & Rutishauser, Can. J. Bot. 68 (1990) 2320. Tree to 8 m; bole to 24 cm diam. Bark dark brown, darker within; inner bark reddish. Leafy nvigs to 3 mm diam. Leaves to 75 cm long; petiole 5—15 cm long, 2.5 mm diam., terete. Leaflets in up to 13 pairs, 4.5—30 by 2.7—9 cm, obovate or elliptic to oblong, the proximal often conspicuously smaller than the distal, fulvous pubescent abaxially and on major venation adaxially, bases acute to cuneate; apices + acuminate, costae c. 7-15 on each side, venation weakly prominent abaxially; petiolules to 6.5 mm, pubescent. /n- florescence to 18 cm, weakly scented, paniculiform, borne in leafy axils, though subtend- ing leaf sometimes unexpanded, velutinous; axis unbranched (? female) or with branches to 2.5 cm (male); bracts to 3 mm long, subulate. Calyx 3—4 mm long, cupulate, pubes- cent without, margin truncate. Petals (3 or) 4, c. 7 by 2 mm, creamy white, pubescent without. Staminal tube weakly adnate to corolla, c. 6 mm long, sericeous without though sometimes glabrous at base, glabrous within though sometimes with a few hairs at base, margin + entire; anthers (5) 6, c. 1.5 mm long, locellate, inserted c. 3 mm within tube, connective glabrous to sparsely hairy. Ovary 4-locular; style to 6 mm long, gla- brous or with hairs in proximal half. Mature female flowers, fruits and seeds unknown. Distribution — Malesia: Papua New Guinea. Habitat — Lower montane forest (disturbed), 1450—1850 m altitude. 152 Flora Malesiana ser. I, Vol. 12 (1) (1995) 14. Chisocheton pohlianus Harms Chisocheton pohlianus Harms, Ber. Deutsch. Bot. Ges. 35 (1917) 341, t. 1; Bot. Jahrb. 72 (1942) 187, excl. spec. cit.; Hutch., Phylog. Flow. Pl. (1969) t. 342; P.F. Stevens, Contrib. Herb. Austral. 11 (1975) 28: Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 331, t. 3; Fisher & Rutishauser, Can. J. Bot. 68 (1990) 2320. Tree to 8 m; bole to 7 cm diam., sparsely branched. Bark greyish brown, scarcely cracking, cicatrose; inner bark claret. Leafy twigs to 4 mm diam. Leaves to 2 m long and to 28-jugate; petiole 5—10 cm, to 3.5(—6.5) mm diam., terete, with cicatrices of spent inflorescences. Leaflets 5—14.5(—22) by 3.2—5.5 (—7) cm, ovate to lanceolate or elliptic, pubescent on veins adaxially and abaxially, where sometimes whole surface thus, bases cuneate, apices acuminate; costae c. 14 on each side, venation weakly sunk- en adaxially, prominent abaxially; petiolules to 6 mm long. Inflorescences to 5 cm long, borne on concurrently flushing rachis, Cymbopogon-scented. Calyx c. 3.5 mm long, cupulate, sericeous, margin + entire. Petals (3 or) 4, to 10 by 2 mm, creamy green, sparsely hairy without. Staminal tube c. 9 mm long, subglabrous or with retrorse hairs within except at top and bottom, margin obscurely lobed; anthers (4—) 6 or 7, locellate, inserted up to 3 mm within tube. Ovary 3- or 4-locular; style to 8 mm long, glabrous or with ascending hairs throughout most of its length. Capsule to 4 cm long, 2.5 cm diam., subovovoid to ellipsoid. Seeds unknown. Distribution — Malesia: Papua New Guinea. Habitat — Lowland and lower montane forest, 200-1770 m altitude. 15. Chisocheton lasiocarpus (Miq.) Valeton Chisocheton lasiocarpus (Miq.) Valeton, Bull. Dép. Agr. Ind. Néerl. 10 (1907) 25; Steenis, Blumea 11 (1961) 132: P.F. Stevens, Contr. Herb. Austral. 11 (1975) 15; in Handb. Fl. Papua New Guinea 1 (1978) 144; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 333, f. 1 (4). — Dysoxylum lasiocar- pum Migq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 13; C.DC. in DC., Monogr. Phan. | (1878) 527; Bull. Herb. Boiss. II, 3 (1903) 168. — Alliaria lasiocarpa (Miq.) Kuntze, Rev. Gen. Pl. 1 (1891) 109. Melioschinzia macrophylla K.Schum. in K.Schum. & Hollr., Fl. Kaiser Wilh. Land (1889) 62; Warb., Bot. Jahrb. 13 (1891) 343. — Chisocheton macrophyllus (K.Schum.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 295; in K.Schum. & Laut., Fl. Deutsch. Schutzgeb. Siidsee (1901) 381, non King (1895). Chisocheton lauterbachii Harms in K.Schum. & Laut., Fl. Deutsch. Schutzgeb. Stidsee (1901) 382; C.DC., Bull. Herb. Boiss. H, 3 (1903) 168. Chisocheton pachyrachis Harms in K.Schum. & Laut., Fl. Deutsch. Schutzgeb. Siidsee (1901) 382; C.DC., Bull. Herb. Boiss. II, 3 (1903) 169; Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 314; Harms, Bot. Jahrb. 72 (1942) 187; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 27; in Handb. FI. Papua New Guinea | (1978) 153. Chisocheton novoguineensis C.DC., Bull. Herb. Boiss. II, 3 (1903) 169; Baker f., J. Bot. Lond. 61, Suppl. (1923) 8; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 25; in Handb. Fl. Papua New Guinea | (1978) 53. Chisocheton forbesii C.DC., Bull. Herb. Boiss. II, 3 (1903) 168; Nova Guinea, Bot. 8 (1910) 424; Baker f., J. Bot. Lond. 61, Suppl. (1923) 8. Chisocheton weinlandii Harms in K. Schum. & Laut., Nachtr. Fl. Deutsch. Schutzgeb. Siidsee 3 (1905) 283; Merr. & L.M. Perry, J. Arnold Arbor. 29 (1948) 157; Hartley et al., Lloydia 36 (1973) 261; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 50; in Handb. Fl. Papua New Guinea | (1978) 171, t. 82; Johns, Comm. For. Trees Papua New Guinea 5 (1976) 213. Mabberley et al. — Meliaceae 153 Chisocheton biroi Harms in K. Schum. & Laut., Nachtr. Fl. Deutsch. Schutzgeb. Siidsee 3 (1905) 283; C.T. White, Proc. Roy. Soc. Queensl. 34 (1922) 38; P.F. Stevens, Contr. Herb. Austral. 11 (1975) a3: Chisocheton multijugis C.DC., Bull. Herb. Boiss. II, 3 (1903) 168, incl. var. glabrior C.DC. Chisocheton schumannii C.DC., Nova Guinea, Bot. 8 (1910) 425; Harms, Bot. Jahrb. 72 (1942) 188: P.F. Stevens, Contr. Herb. Austral. 11 (1975) 40; in Handb. Fl. Papua New Guinea | (1978) 164, t. 79; Johns, Comm. For. Trees Papua New Guinea 5 (1976) 216. Chisocheton versteegii C.DC., Nova Guinea, Bot. 8 (1910) 424; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 49; in Handb. Fl. Papua New Guinea | (1978) 171. Chisocheton frutescens C.DC., Nova Guinea 8 (1914) 1013. Dasycoleum forbesii Baker f. & Norm., J. Bot. Lond. 61, Suppl. (1923) 8. Chisocheton lamekotensis Harms, Notizbl. Bot. Gart. Berlin 10 (1928) 276; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 53. [Chisocheton lamii Diels ex H.J. Lam, Nat. Tijds. Ned. Ind. 88 (1928) 216, nom. nud.; L.M. Perry, Sargentia 5 (1945) 59, nom. nud.]} Chisocheton sp. aff. schumannii C.DC.: C.T. White, J. Arnold Arbor. 10 (1929) 228. Chisocheton myrmecophilus Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 313. Chisocheton boridianus Harms, Bot. Jahrb. 72 (1942) 180. Chisocheton eurycalyx Harms, Bot. Jahrb. 72 (1942) 182. Chisocheton formicarum Harms, Bot. Jahrb. 72 (1942) 182; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 12; in Handb. Fl. Papua New Guinea | (1978) 141. Chisocheton gjellerupii Harms, Bot. Jahrb. 72 (1942) 183. Chisocheton caroli Harms, Bot. Jahrb. 72 (1942) 181; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 9; in Handb. Fl. Papua New Guinea | (1978) 137, t. 67. Chisocheton trichocladus Harms, Bot. Jahrb. 72 (1942) 189; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 48; in Handb. Fl. Papua New Guinea | (1978) 170. Chisocheton ledermannii Harms, Bot. Jahrb. 72 (1942) 184; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 53; in Handb. Fl. Papua New Guinea | (1978). Chisocheton schlechteri Harms, Bot. Jahrb. 72 (1942) 188; P.F. Stevens, Contr. Herb. Austral. 1 1 (1975) 37; in Handb. Fl. Papua New Guinea | (1978) 161, t. 77; Johns, Comm. For. Trees Papua New Guinea 5 (1976) 215. ?Chisocheton oreophilus Harms, Bot. Jahrb. 72 (1942) 185. Chisocheton torricelliensis Harms, Bot. Jahrb. 72 (1942) 188; P.F. Stevens, Contr. Herb. Austral. 11 tee) eee Chisocheton sp.: Menninger, Flow. Trees (1962) t. 228. Tree to 33 m; bole to 60 cm diam., fluted and with small buttresses to 1 m tall, when large. Bark blackish brown to red, sometimes cracking vertically and flaking; inner bark + red; wood pinkish straw to white. Leafy twigs (2—)4—9 mm diam., cicatrose, some- times myrmecophilous. Leaves to 150 cm; petiole 8—15 cm; rachis 2—4.5 mm diam., terete to + winged or rarely + flattened. Leaflets in up to 11 pairs, (7—)14—45 by (2.5-) 7—23 cm, ovate to elliptic or suboblong, indumentum of adpressed hairs usually rather inconspicuous or puberulous on veins adaxially and/or velutinous abaxially, base occa- sionally subcordate, fine venation slightly prominent especially abaxially; petiolules 3—8 (—12) mm. /nflorescences to 60 cm, but usually less, axillary or on short shoots in de- foliated axils of twigs to 2.5 cm diam., 0—2-branched, sweetly scented; branches to 20 cm; pedicels 0-5 mm. Calyx 2—4 mm long, margin truncate; pseudopedicel to 1.5 mm long. Petals (3) 4 or 5 (6), c. 7—16(—22) by 0.7—4.5 mm, white or flushed pink to claret, quincuncial, alternative or rarely imbricate. Staminal tube to 3 mm diam., pinkish, with- out glabrous or sparsely hairy in distal half, within retrorse hairy from (usually) just 154 Flora Malesiana ser. I, Vol. 12 (1) (1995) below anthers to base, very rarely glabrous, margin truncate to shallowly lobed; anthers (3—)5—10(—18), c. 1-3 mm long, locellate, inserted about 2—4 mm within tube. Ovary (3-) 4- or 5- (6-)locular; style 6-15 mm, hairy at least at base. Capsule to 4 cm diam., obovoid to + spherical, brownish red, hairs dense, sometimes of conspicuously differ- ent lengths; pericarp fibrous. Seeds up to 5; testa black, aril red surrounding hilum; co- tyledons superposed. 2n = 92. Distribution — Solomon Islands; Malesia: Moluccas (Ceram), New Guinea. Habitat & Ecology — Primary or secondary forest including riparian and submon- tane, persisting in logged and grazed-through forest, to 1525 m altitude. According to Beehler [Auk 100 (1983) 1], the fruits of at least one form of this species (‘Chisocheton weinlandil ) are taken exclusively by birds of paradise, nine species of which visited a tree over its three-month fruiting season, during which it produced some 4500 fruits. Uses — Leaflets used for wrapping sago and other food in cooking. Variation — Chisocheton lasiocarpus comprises a complex of forms, which together may be described as an ochlospecies in that in any particular locality, there may appear to be distinct entities co-existing, but intermediates are found elsewhere, often co-exist- ing with other apparently distinct entities, and so forth. Locally, nevertheless, it may be useful to recognize the major morphological entities, though there are many intermedi- ates. The following key (after Stevens 1975) may be useful, though the very nature of this complex denies the possibility of pigeonholing every specimen: KEY TO THE ENTITIES la. Leaf rachis prominently ridged or winged, adaxial surface flattened.......... 2 be Meat rachis channelledito terete: — 2.3. 4 ete asthe 2s On eee 3 2a. Inflorescence 3.5—12'cm long; flattened, petals 14—22 mmvlone —-) eee Bis NP ar ie ae oy aparece Rare REE ep AME? hope 0d 2 ee e. ‘pachyrhachis’ b. Inflorescence c. 8—45 cm long, + terete, petals 75-12 mm long .............. Ns Rl Aen HM SR AN i aie RN 5 a ere et A ch eae Re Pe Fg a. ‘novoguineensis’ 3a. Inflorescence to 8(—12) cm long, flowers dense; buds c. 4 mm across; indumentum never onlongserectihairsi ss 2 AONks, Rane at ee eS ee 4 b. Inflorescence usually more than 8 cm long; if less, flowers not dense, or buds c. 2 IMMMNACTOSS:OL both) W222 2.5 ae aE IN UC Sau es.). 2 Aga See 5 4a. Leaflets with dense + crisped short hairs on midrib adaxially; anthers c. 3 mm long; Styleswithvhams:formestof itsilengthi ii) sae ee ae CR Ee c. ‘lasiocarpus’ b. Leaflets with, at most, adpressed hairs adaxially; anthers less than 2 mm long; style AOA DLOUS MAEKAWA ce. tenet Ite ste ss of ARES ISOs OU. d. ‘formicarum’ 5a. Leaflets with erect or crisped hairs at least adaxially on midrib. ............. 6 b. Leaflets adaxially with adpressed hairs, often appearing glabrous ........... 8 6a. Leaflets without hairs abaxially; flowers always (?) S-merous ... g. ‘versteegii’ beiLeailetsiwithierecthawrsabaxially..-...2 80.) n2d02 . kde. 2 ee 7 ja eatletibasetshallowlyicordatenmauwy tore ok Aeeeee Se Ieee j. ‘schlechteri’ bi), Leatletibaseioundeditomeutes iow tied, Hepesa verse. vies i. ‘trichocladus’ Mabberley et al. — Meliaceae 155 8a. Leaflets subcoriaceous; inflorescences less than 12 cm long, branches narrowly as- g y BOOM GR TEI RIOWCRED trata eS. a! SEEDS a Dh. Garde igesh sible gins f. ‘caroli’ pee ogthic combination Of. characters ©. 2. sevem-ts -n dues OSE Pemsiee bee? 9 9a. No glabrous zone immediately below the anthers on inside of tube; flowers often STHEIRC TSG IND) eto) Pe A a eee ee Fen toe h. ‘schumannii’ b. Short glabrous zone immediately below anthers inside tube; flowers usually 4-me- Mats Calyx USUALLY == SPLCACING LO;SUDETECE ao so ae cach ls pon in hs lesion. sey in oye in = 10 10a. Style usually hairy for its entire length; fruit ovoid ..... a. ‘novoguineensis’ beste elabrous at apex; fruits spherical... Ao. ee b. ‘weinlandii’ a. ‘novoguineensis’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 334. — Chisocheton novoguineensis C.DC. — Chi- socheton forbesii C.DC. — Chisocheton biroi sensu C.T. White — Dasycoleum forbesii Baker f. & Norm. — Chisocheton myrmecophilus Merr. & L.M. Perry. Central East New Guinea, hill or submontane rain forest, (100—)600—1525 m alti- tude. b. ‘weinlandii’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 334. — Chisocheton weinlandii Harms — Chiso- cheton multijugis C.DC. incl. var. glabrior C.DC. — Chisocheton schumannii auct. non C.DC.: C.DC. (1910), quoad spec. cit. — Chisocheton frutescens C.DC. — Chisocheton sp. aff. schuman- nii C.DC.: C.T. White — Chisocheton boridianus Harms — Chisocheton eurycalyx Harms. Ceram, New Guinea, New Britain, primary and secondary rain forest to 1280 m alti- tude. Note — This is a very widespread form but tends to be difficult to separate from ‘novoguineensis’ on the one hand and, on the other, ‘/asiocarpus’ and ‘formicarum’ in its western range, as well as ‘schumannii’ in the east and ‘trichocladus’. Intermediate specimens are cited by Mabberley (l.c. and see p. 315), one of them, between ‘weinlan- div’ and ‘trichocladus’ being the type of Chisocheton biroi Harms. c. ‘lasiocarpus’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 335 — Chisocheton lasiocarpus (Miq.) Valeton, s. str. — Dysoxylum lasiocarpum Miq. — Alliaria lasiocarpa (Miq.) Kuntze. Western New Guinea, rain forest at low altitudes. d. ‘formicarum’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 335. — Chisocheton lamii Diels ex H.J. Lam — Chisocheton formicarum Harms. Northwest and central New Guinea, lowland rain forest. Note — This is very similar indeed to some forms of ‘pachyrachis’ and is connected by intermediates to ‘schumannit’. 156 Flora Malesiana ser. I, Vol. 12 (1) (1995) e. ‘pachyrhachis’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 335. — Chisocheton pachyrhachis Harms — Chiso- cheton gjellerupti Harms. New Guinea, primary rain forest to 1000 m altitude. f. ‘caroli’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 336. — Chisocheton caroli Harms. North-eastern New Guinea, primary rain forest to 1500 m altitude. g. ‘versteegii’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 336. — Chisocheton versteegii C.DC. West New Guinea, known only from the type. h. ‘schumannii’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 336. — Melioschinzia macrophylla K.Schum. — ?Chisocheton lauterbachii Harms — Chisocheton macrophyllus (K. Schum.) Harms — Chisocheton schumannii Harms — ?Chisocheton lamekotensis Harms — ?Chisocheton toricelliensis Harms. Northern New Guinea and (7?) New Ireland, in lowland rain forest. Notes — |. It intergrades with ‘trichocladus’ in the Solomons and is difficult to dis- tinguish from ‘weinlandii’ when in fruit. 2. Discovery of an isotype of Chisocheton toricelliensis (Schlechter 4402, BO) shows that it belongs 1n the Chisocheton lasiocarpus complex and possibly here. i. ‘trichocladus’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 336. — Chisocheton trichocladus Harms — Chiso- cheton ledermannii Harms. Northern New Guinea to the Solomon Islands, primary or secondary rain forest, to 150 m altitude. Note — This is linked to ‘schlechteri’ by the type of Chisocheton ledermannii and other specimens as well as to ‘schumanniv’ and ‘weinlandiv’. j. ‘schlechteri’ Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 337. — Chisocheton schlechteri Harms. North-east New Guinea, primary rain forest. Note — NGF 21573 from Morobe Province is a leptocaul treelet less than 3 m tall. The flowers are similar to those of the more glabrous forms of ‘weinlandii’, but the inflores- cence is 3-branched, the major branches being up to 16 cm long. In some respects it approaches the description of Chisocheton oreophilus Harms, which probably belongs in this complex, but has a smaller calyx and may be referable to a form of Chisocheton sayeri. Mabberley et al. — Meliaceae 157 16. Chisocheton pilosus C.DC. Chisocheton pilosus C.DC., Nova Guinea, Bot. 8 (1910) 423; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 337. — Chisocheton sayeri (C.DC.) P.F. Stevens var. pilosus (C.DC.) P.F. Stevens, Contr. Herb. Austral. 11 (1975) 36; in Handb. Fl. Papua New Guinea | (1978). Treelet 2.5 m tall. Leaves at least 25 cm; petiole c. 10 cm; rachis terete. Leaflets in up to at least 3 pairs, to 26 by 8 cm, oblong-ovate, glabrous adaxially, pilose abaxially, base cuneate, apex (?) acuminate, costae c. 13 on each side; petiolules c. 5 mm. /nflorescence to 12 cm, weakly paniculiform, axillary; branches to 3 cm long, pilose; bracts linear, pu- bescent; pedicels very short. Calyx campanulate, pubescent, margin truncate. Petals 4, c. 11.5 by 1.5 mm, puberulous without. Staminal tube villous within, margin truncate; anthers 6—8, c. 1.5 mm long, alocellate. Ovary and female flowers unknown. Distribution — Malesia: West New Guinea, collected once (a hybrid with C. sayeri?). 17. Chisocheton sayeri (C.DC.) P.F. Stevens Chisocheton sayeri (C.DC.) P.F. Stevens, Contr. Herb. Austral. 11 (1975) 32, excl. var. pilosus (C.DC.) P.F. Stevens; in Handb. Fl. Papua New Guinea | (1978) 158, t. 76, excl. var. pilosus; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 337. — Dasycoleum sayeri C.DC., Bull. Herb. Boiss. II, 3 (1903) 170. Chisecheton erythrocarpus auct. non Hiern: Lane-Poole, Rep. For. Res. Papua New Guinea (1925) 100. Chisocheton archboldianus Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 312. Chisocheton erythranthus Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 312. Chisocheton pohlianus auct. non Harms: Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 311. Chisocheton acartianthus Harms, Bot. Jahrb. 72 (1942) 180 (‘acariaeanthus’ ). Chisocheton graciliflorus Harms, Bot. Jahrb. 72 (1942) 183. ?Chisocheton leptopetalus Harms, Bot. Jahrb. 72 (1942) 184. Chisocheton schumannii auct. non C.DC.: Hartley et al., Lloydia 36 (1973) 261. Tree or treelet to 15 m; bole to 30 cm diam. Bark smooth or shallowly fissured, grey- green; inner bark pale brown. Leafy twigs to 3.5 mm diam. Leaves to 25(—75) cm; peti- ole 5—10 cm, 3.5 mm diam., terete. Leaflets in up to 9(—12) pairs, (S—)7.5—20(—43) by (2.5—)3.5—6.5(—9) cm, ovate to narrowly elliptic or lanceolate, subglabrous to veluti- nous On main veins adaxially and over whole abaxial surface; costae c. 9 on each side. Thyrses to 70 cm long, but usually much shorter, axillary, not or 1-branched (female) or 1—2-branched (male), sweetly scented; branches to 15 cm, patent or ascending, some- times with congested flowers; pedicels to 2.5 mm. Calyx 1—1.5 mm deep, reddish brown, obscurely lobed. Petals (3) 4 (5), 8—10 by 1—1.5 mm, pale cream, alternative to imbri- cate. Staminal tube pubescent without, glabrous to densely hairy within from just below anthers to base, white; anthers 4—6 (7), 0.7—1.2(—1.5) mm long, locellate or not, gla- brous. Ovary 3—5-locular; style to 9.5 mm, densely hairy in lower 2/3. Capsule to 2.8 cm long, 2 cm diam., obovoid to ellipsoid, obscurely stipitate, golden brown when young; pericarp fibrous with hairs of conspicuously different lengths. Seeds 3; aril cir- cumhilar; cotyledons collateral or superposed. Distribution — Malesia: New Guinea. Habitat — Lowland or submontane rain forest, to 1500(—1830) m altitude. 158 Flora Malesiana ser. I, Vol. 12 (1) (1995) 18. Chisocheton aenigmaticus Mabb. Chisocheton aenigmaticus Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 338, f. 4. Tree. Leafy twigs 8-14 mm diam., terete, lenticellate. Leaves to 78 cm, pseudogem- mulate; petiole 3—8 cm. Leaflets in up to 13 pairs, to 24 by 5.5 cm, elliptic oblong, sparsely adpressed hairy on veins adaxially, more so abaxially, bases obtuse to subcor- date; costae c. 10-14 on each side. Thyrses to 70 cm long, axillary or slightly supra- axillary, pyramidal, 3-branched; axis c. 6 mm diam.; most proximal branches to 25 cm long, ascending. Flowers subsessile. Calyx 2—2.5 mm deep, c. 3 mm diam., cupulate, shortly pubescent without, glabrous within, margin truncate or shallowly 4- or 5-lobed. Petals 5, 5-10 by 1.5 mm, narrowly spathulate, + adpressed pilose without, glabrous within, quincuncial or alternative. Staminal tube with long adpressed hairs without, + villous within, margin truncate or shallowly 5-lobed; anthers 5, c. 2.2 mm long, in- serted within tube or weakly exserted, basifixed, locellate. Style with ascending hairs; stylehead cylindric-capitate. Female flowers and fruits unknown. Distribution — Malesia: Sumatra, Simalur I. Habitat & Ecology — Unknown. 19. Chisocheton celebicus Koord. Chisocheton celebicus Koord., Meded. Lands Plant. 19 (1895/6) 385 & (1898) 636 (‘celebica’); Suppl. FI. NO Celebes 2 (1922) t. 42; 3 (1922) 22; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 340. Chisocheton glomeratus auct. non Hiern: Koord., Meded. Lands Plantent. 19 (1895/6) 385; Koord.- Schum., Syst. Verz. 3, 1 (1914) 63. Chisocheton sp. A: Koord.-Schum., Syst. Verz. 3, | (1914) 63. Tree. Leafy twigs c. 8 mm diam. Leaves to at least 30 cm, pseudogemmulate; petiole 3-10 cm. Leaflets in up to at least 5 pairs, to 32 by 14 cm, ovate to elliptic-oblong, ful- vous tomentose on venation adaxially and over whole abaxial surface, bases rounded, apices shortly and gradually acuminate, midrib sunken adaxially in sicco, costae in c. 19 subopposite pairs, tertiary venation distinct; petiolules to 8 mm long; pseudogemmula densely long fulvous tomentose. Thyrse to 85 cm long, 2-branched, supra-axillary; axis c. 3mm diam., branches to 9 cm long, + fulvous tomentose. Calyx c.2.5—3.5 mm long, 2-3 mm diam., tubular, long-pubescent without, margin truncate. Petals 5, 12-14 by 1.8 mm, narrowly spathulate, glabrous, alternative. Staminal tube densely sericeous without, especially in more distal half, up to anthers, glabrous or very sparsely villous within, margin truncate; anthers 6, 1.2—1.5 mm, narrowly oblong, locellate, glabrous, basifixed, included. Disk conspicuous. Style terete, long sericeous; stylehead cylindric- capitate. Fruits unknown. Distribution — Malesia: NE Celebes. Habitat — 100-700 m altitude. 20. Chisocheton glirioides P.F. Stevens Chisocheton glirioides P.F. Stevens, Contr. Herb. Austral. 11 (1975) 13, t. 1; in Handb. Fl. Papua New Guinea | (1978) 142, t. 69; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 340. Mabberley et al. — Meliaceae 159 Tree to 21 m; bole to 25 cm diam. Bark dark grey to grey-brown; inner bark dark straw. Leafy twigs to 4.5 mm diam. Leaves to 35 cm long, pseudogemmulate; petiole 7-10 cm, to 3.5 mm diam., terete. Leaflets 10—18(—24) by 3.7—7 cm, ovate-elliptic, subglabrous with adpressed hairs on venation, particularly abaxially, base acute, apex acute to weakly acuminate, costae up to 13 on each side; petiolules to 6.5(—12.5) mm. Thyrse to 40 cm, axillary, subglabrous, 1(—?2)-branched; branches to 4 cm, patent; bracts to | mm long, narrowly triangular. Calyx c. 2 mm long, cupulate, sessile, pubes- cent without, glabrous within, margin truncate. Petals 4, 12 by 1.7 mm, red, pubescent without, glabrous within, alternative to imbricate. Staminal tube c. 11 mm long, white, sparsely hairy in the middle on both surfaces, margin truncate or shallowly lobed; an- thers 6 or 7, c. 1.3 mm long, locellate, inserted c. 2.5 mm within the tube. Ovary 3- or 4-locular, c. 2 mm long (male flowers only known), style pubescent, stylehead c. 0.6 mm diam. Capsule c. 1.5 cm diam., spherical, shortly pubescent; pericarp fibrous. Seeds unknown. Distribution — Malesia: Papua New Guinea: Central Prov. Habitat — Lowland rain forest, to 240 m altitude. 21. Chisocheton sapindinus P.F. Stevens Chisocheton sapindinus P.F. Stevens, Contr. Herb. Austral. 11 (1975) 29, t. 4; in Handb. Fl. Papua New Guinea | (1978) 156, t. 75; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 340. Tree to 10 m, leptocaul, somewhat weeping, + riparian; bole to 10 cm diam. Bark greyish, smooth to finely fissured, reddish within; inner bark straw, sometimes un- pleasantly scented. Leafy twigs to 2 mm diam., thinly hairy when young, becoming glabrous, cicatrose. Leaves to 45 cm, with very small pseudogemmula; petiole 1—8 cm, 1—2 mm diam. Leaflets in up to 9 pairs, (4—)7.5—-15.5 by (2—)3.3—6.3 cm, ovate to el- liptic, glabrescent, coriaceous, bases acute to cuneate, + asymmetric, apices acuminate, venation impressed adaxially in vivo, prominent on both sides in sicco, costae 5—10 on each side; petiolules to 10 mm. /nflorescence racemose to thyrsoid, to 60 cm, but usu- ally very much less, unbranched in females, 0—2-branched in males; branches to 3 cm, patent, with scattered flowers, especially distally; bracts c. 1 mm long, subulate; pedi- cels to 2 mm long. Calyx 1.7—2.5 mm long, margin often with a split to 1 mm. Petals 4, c. 13.5 by | mm, though smaller in males, ligulate, white or pinkish green, glabrous or with a few hairs apically, alternative. Staminal tube glabrous, margin weakly lobed; anthers 4—6, c. 0.7 mm long, locellate, inserted c. 1.5 mm within tube, connective gla- brous or thinly pubescent. Style pubescent in proximal half; stylehead c. 0.5 mm diam. Capsule to 6.5 cm long, 2.25 cm diam., glabrous, red; valves 3 or 4. Seeds 2—4, to 3 cm long, 1.8 cm wide, 1.1 cm thick, ellipsoid to boatshaped; aril circumhilar, red; testa black; cotyledons superposed. Distribution — Malesia: very local in Papua New Guinea. Habitat — Primary rain forest, to 760 m altitude. 160 Flora Malesiana ser. I, Vol. 12 (1) (1995) Series Paniculati Chisocheton sect. Euchisocheton § Paniculati Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 295: Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 341 (as series in sect. Chisocheton). Chisocheton § Tetrapetali Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 151. Leaves pseudogemmulate, rarely imparipinnate or paripinnate. Inflorescences axil- lary, supra-axillary, cauliflorus, ramiflorous or borne on supra-axillary branches. Seeds arillate. White latex sometimes present in pith, pericarp etc. of some species. 22. Chisocheton laosensis Pellegr. Chisocheton laosensis Pellegr., Bull. Soc. Bot. Fr. 91 (1945) 178; in Fl. Indo-Chine, Suppl. (1946) 694; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 341. Tree to 20 m. All young parts densely fulvous tomentose. Leafy twigs 6—9 mm diam. Leaves to 50 cm, pseudogemmulate; petiole 5—8 cm. Leaflets in up to 8 pairs, to 22 by 10 cm, elliptic, coriaceous, glabrous adaxially save (usually) on venation, abaxially + tawny pubescent, apex abruptly and shortly acuminate, acumen 6—8 mm, costae c. 13 on each side, arcuate, prominent adaxially, tertiary venation conspicuous; petiolules c. 1 cm long, velutinous-fulvous. /nflorescences on supra-axillary branches to 90 cm long, with reduced caducous leaves, the whole resembling a supra-axillary thyrse, each to 25 cm long, not or sparsely branched with congested cymules of flowers; bracts lanceolate. Calyx to 3 mm long, cupular, pubescent without, glabrous within, margin + 4-lobed. Petals 4, 11-14 by 2.5 mm, linear-oblong, adpressed pubescent without, glabrous with- in, drying blackish, alternative to imbricate. Staminal tube glabrous except for a few hairs in notches of lobes without and towards base within, margin 7- or 8-crenulate or -lobed; anthers 6—8, oblong-elliptic, locellate, glabrous, included. Disk c. 1 mm tall, cylindrical, glabrous. Ovary apparently 2-locular; style filiform, papillose, pubescent in proximal half; stylehead subglobular, glabrous with an equatorial papillose band. Cap- sule to 3 cm long, 2.5 cm diam., rusty tomentose; valves 3 or 4. Seeds 1 or 2, c. 15 mm long, apparently arillate. Distribution — Malesia: Moluccas (Halmahera ?and Ceram). Habitat — Rain forest, to 600 m altitude. Note — The name derives from the type specimen which was believed to have been collected in Laos (Mabberley, |.c.) but, besides it, no non-Moluccan material is known. How this confusion came about is unclear. 23. Chisocheton ruber Ridley Chisocheton ruber Ridley, Bull. Misc. Info. Kew (1930) 365; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 342. Tree to 15 m; bole to 20 cm diam., fluted. Bark smooth to weakly and irregularly flaking, greenish grey or reddish, with conspicuous inflorescence bosses, sometimes bearing leafy shoots, arranged + spirally from ground level to 5 m; wood ivory. Leafy twigs c. 12-15 mm diam. Leaves to 1.5 m long, pseudogemmulate, subglabrous, in Mabberley et al. — Meliaceae 161 terminal spirals; petiole 8—20 cm; rachis + 3-ribbed. Leaflets in up to 15 pairs, to 42 by 10 cm, oblong, coriaceous, brilliant carmine when young and appearing in flushes of up to I1 pairs at once, very sparsely puberulous, bases subacute, asymmetric, apices acuminate, costae c. 12—14 on each side, tertiary venation conspicuous abaxially; petio- lules c. 6 mm. /nflorescences to 10 cm long, not or once branched, sweetly-scented, borne on bosses which produce them over several seasons; rachis pubescent; pedicels 1—3 mm long, pubescent, minutely bracteolate. Calyx c. 4 mm long, cupular, rugose, pubescent, red; margin + 4- or 5-lobed. Petals 5 or 6, 20—22 mm long, 4 mm wide at widest, 2.5 mm at narrowest, linear-oblong to spathulate, fleshy distally, pubescent without, imbricate to quincuncia!. Staminal tube pubescent distally without, villous within, white, adnate to corolla at base, the margin shallowly 6—8-lobed, each lobe praemorse or irregularly 2- or 3-fid; anthers 8—i0, 2 mm long, oblong, alocellate, yellow, sparsely hairy near connective, basifixed. Disk obscure. Ovary conical, 5- locular, appressed pubescent; style white, hairy in proximal 3/4 or throughout; style- head to 1.8 mm diam., very shortly cylindrical to subdiscoid. Capsule to 5 cm long, 3 cm diam., top-shaped, 5-merous, glabrous, reddish; pericarp with white latex. Seeds unknown. Distribution — Malesia: Borneo (Sarawak, Ist Division). Habitat & Ecology — Restricted to limestone formations, 80—250 m altitude. 24. Chisocheton sarawakanus (C.DC.) Harms Chisocheton sarawakanus (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 296; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 342; in Tree Fl. Malaya 4 (1989) 238. — Dasycoleum sarawakanum C.DC. in DC., Monogr. Phan. | (1878) 541. Chisocheton laxiflorus King, J. As. Soc. Beng. 64, ii (1895) 33, p. maj. p.; Ridley, Fl. Malay Penins. 1 (1922) 390; Craib, Fl. Siam. Enum. 1| (1926) 253. Chisocheton brachyanthus Merr., J. Str. Br. Roy. As. Soc. 86 (1922) 315; Univ. Calif. Publ. Bot. 15 (1929) 122; Meijer, Bot. News Bull. Sabah 8 (1967) 79. Chisocheton spec. A: Meijer, Bot. News Bull. Sabah 8 (1967) 79. Chisocheton glomeratus auct. non Hiern: Meijer, l.c. Tree 5—20 m high; bole to 30 cm diam., fluted below, with small buttresses to 2 m tall. Bark fawn to chocolate, smooth to weakly flaking; inner bark brownish yellow; wood white to pale fawn. Innovations + pale ferruginous pubescent. Twigs terete, dark brown, elenticellate, glabrous when leafless. Leafy twigs c. 5 mm diam., rarely myrme- cophilous. Leaves to 2 m long, pseudogemmulate; petiole 5S—12 cm; rachis brown, pu- bescent to ultimately glabrous. Leaflets in up to 26 pairs, flushing in up to 3 pairs at once, 8—29 by 4-8 cm, elliptic to elliptic-oblong, subcoriaceous, shiny and glabrous on both surfaces to ferruginous pubescent abaxially, particularly on veins, and on veins adaxial- ly, bases slightly narrowed or rounded, sometimes asymmetric, apices rather abruptly caudate-acuminate with acumen to 2 cm; costae 10—14 on each side, spreading, depressed adaxially and prominent abaxially in sicco; petiolules c. 6 mm, pubescent. Thyrses to 50 cm, narrowly paniculiform or subspiciform, supra-axillary; primary branches few, squarrose, bearing few secondary branches of cymules, the sweetly scented flowers 162 Flora Malesiana ser. I, Vol. 12 (1) (1995) usually borne in pairs, sessile. Calyx 2—3 mm long, c. 1.8 mm diam., cupular, gla- brous to puberulous without, glabrous within, margin truncate to obscurely crenate. Petals 4, c. 12 by 1.8 mm, linear, white, drying black, puberulous without, glabrous within, apex obtuse, slightly concave. Staminal tube cylindric, c. 2 mm diam., some- what appressed hairy distally, margin crenate; anthers (3—)4—6, 1-2 mm long, insert- ed just below rim, somewhat ciliate posteriorly. Disk absent. Ovary appressed pubes- cent; style pubescent proximally; stylehead subcapitate, c. 0.5 mm diam. /nfructescence with branches to 6 cm with up to 8 fruits on each. Capsule c.4 cm diam., depressed globose, shortly stipitate, crimson, obovoid and densely ferruginous pubescent when young; pericarp sometimes with white latex. Seeds 2; testa dark brown, partly envelop- ed by aril. Distribution — Malesia: Malay Peninsula, Bangka, Borneo. Habitat — Rain forest, to 250 m altitude. Note — There is a complete gradation between + glabrous and hairy forms, the latter being readily distinguished from hairy forms of Chisocheton patens in Borneo, where they occur together, by their prominent leaf venation. Elsewhere, however, sterile mate- rial of these two species 1s often very difficult to separate. 25. Chisocheton lasiogynus Boerl. & Koord. Chisocheton lasiogynus Boer]. & Koord. in Koord.-Schum., Syst. Verz. 2 (1910) 26; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 343. Treelet to 2 m. Leafy twigs 4—5 mm diam. Leaves 50—68 cm, imparipinnate, 3—5- jugate, subglabrous, drying pale brown; petioles 8—12 cm. Leaflets to 12 by 5 cm, ob- long to elliptic-ovate, sometimes subcrenulate opposite to subopposite, base cuneate, apex acuminate with acumen to 17 mm, costae c. 9 or 10 on each side, venation im- pressed adaxially, prominent abaxially. Raceme to 24 cm, bracteate, flowers aggregated near tip. Calyx 4.5—5 mm long, 4—4.5 mm diam., cupular to cylindrical, pubescent, margin truncate to obscurely lobed. Petals 4 or 5, 17 by 3 mm, creamy-white to pinkish or crimson, glabrous to sparsely pubescent proximally without. Staminal tube slightly expanded at mouth, white, densely pubescent in distal half, margin truncate to weak- ly crenulate; anthers 5—8, 1.5—2 mm long, narrowly oblong to boat-shaped, scarcely locellate, glabrous, basifixed, inserted. Ovary conical, 3-locular, densely hairy; style terete, sparsely pubescent in proximal half; stylehead subcylindrical, strongly exserted. Infructescences ramiflorous, sometimes apparently borne on reduced shoots, with fruits crowded at apex. Capsule strongly rostrate when immature, 2.5 cm long, subglobose- pyriform and red when mature. Seeds unknown. Distribution — Malesia: Sumatra and (?) Java but not seen in the latter for a century. Habitat & Ecology — Undergrowth treelet in rain forest, 40—450 m altitude. Note — The Java records are collections by Forbes and Junghuhn and may have come from Sumatra (Mabberley, op. cit. 344). Specimens from northern Sumatra (Aceh) are more pubescent and have rather larger flowers than do specimens from elsewhere. Mabberley et al. — Meliaceae 163 26. Chisocheton amabilis (Mig.) C.DC. Chisocheton amabilis (Mig.) C.DC. in DC., Monogr. Phan. | (1878) 537; Merr., J. Str. Br. Roy. As. Soc., spec. no. (1921) 319; Corner, Gard. Bull. Sing., Suppl. 1 (1978) 198; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 344; in Tree Fl. Malaya 4 (1989) 234. — Schizochiton amabile Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1968) 26, 27, incl. var. swmatranum Miq. Chisocheton illustris Ridley, Bull. Misc. Info. Kew (1930) 366. Chisocheton hackenbergii Harms, Notizbl. Bot. Gart. Berlin 15 (1941) 476. Chisocheton brachyanthus auct. non Merr.: Anderson, Gard. Bull. Sing. 20 (1963) 115. Tree 6—17 m; bole to 10 cm diam. Bark smooth to finely cracked or pustulate, grey- green; underbark orange-red to pinkish; inner bark cream, wood white. Leafy twigs 3-7 mm diam., elenticellate, reddish in sicco. Leaves 20—95 cm, pseudogemmulate; petiole 5—15 cm; rachis terete or laterally channelled in sicco; pseudogemmula fulvous tomentose. Leaflets in (4—)7—20 pairs, most proximal 2.2—11.5 by 1.9—4.8 cm, regu- larly elliptic, most distal 7.5—25.5 by 2.4—8.5 cm, elliptic-oblong, + asymmetric, cori- aceous, shiny adaxially, duller abaxially, glabrous on both sides, or midrib brown- tomentose adaxially and/or venation pubescent abaxially, bases subequally acute to obtuse, apices long-cuspidate, costae 5—14 on each side, ascending, prominulous to prominent abaxially, petiolules 2-5 mm long. Thyrses 8—45 cm, pendent, fragrant, borne in axils of youngest leaves, thus sometimes appearing terminal, often supra-axil- lary; axis glabrous to weakly pubescent, 3—5 mm diam., |—2-branched with pubes- cent pedicels articulated on slender branchlets c. 3—4 mm long arising from first-order branches to 9 cm long in male, unbranched, spiciform and minutely pedunculate with subsessile flowers condensed into short dense cymules, mostly at distal end of rachis in female. Calyx 3—4 mm long, cupular, 4- or 5-lobed, subglabrous to weakly pubescent, green. Petals 5 or 6, 15—25 by 2—3 mm, narrowly obovate, white or sometimes also tipped pink, sparsely hairy without or glabrous, drying reddish, alternative to quincun- cial. Staminal tube subglabrous to villous without, especially at base of lobes, villous within especially near base, margin 5—7-lobed, the lobes subentire to irregularly 2—3- fid; anthers 8—10, c. 1.5 mm long, scarcely locellate, long-pubescent dorsally, included within lobes. Disk c. 0.5—1 mm tall, subtubular, thick. Ovary 4-loculate; style pubes- cent, particularly in proximal half; stylehead subcylindric to spherical. /nfructescences borne on leafy twigs to 8 mm diam. Capsules c. 4 cm diam., spherical, long-stipitate, glabrous, pink ripening to bright rose-red, clustered in groups of 3—10 at end of rachis; stipe 1.7—2.2 cm; valves 3 or 4. Seeds 3 or 4, c. 9 mm long, with chestnut brown testa half covered in circumhilar yellow-orange aril. Distribution — Malesia: Sumatra, Malay Peninsula, Biliton, Borneo. Habitat & Ecology — Peat swamp forest and riparian forest (as at Sg. Sedili, Johore, Malay Peninsula, where it is a common tree), 0—20 m altitude. According to Corner (op. cit. 30, 198) it flowers gregariously in April-May at Sg. Sedili, where it is restrict- ed to the mempisang (Polyalthia sclerophylla Hook. f. & Thoms.) belt. Note — Specimens with inflorescences borne on reduced axillary branches are known from southern Borneo. 164 Flora Malesiana ser. I, Vol. 12 (1) (1995) 27. Chisocheton cumingianus (C.DC.) Harms Chisocheton cumingianus (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 296; Merr., Philipp. J. Sc. 1, Suppl. (1906) 72; West & Brown, Bull. Philipp. Dept. Agr. For. 20 (1920) 117, cum tab.; ibid. 22 (1921) 119; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 367; Briq., Mém. Inst. Nat. Genev. 24 (1935) 67; Elmer, Leafl. Philipp. Bot. 9 (1937) 3346; Mabb., Taxon 26 (1977) 528, incl. subsp. balansae (C.DC.) Mabb.; Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 347, incl. subsp. balansae & subsp. kinabaluensis (Merr.) Mabb.; Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 335, t. 255; H6, Ill. Fl. Vietnam 2, 1 (1992) 494, incl. subsp. balansae. — Dasycoleum cumingianum C.DC. in DC., Monogr. Phan. | (1878) 541; Fern.-Vill., Nov. App. (1880) 42; Vidal, Phan. Cuming. (1885) 84; Rev. Pl. Vasc. Filip. (1886) 84. Guarea gobara Buch.-Ham., Mem. Wern. Soc. 6 (1832) 306, p.min.p.; Wall., Cat. (1831/2) n. 4885, p.p. Guarea paniculata Roxb., Hort. Beng. (1814) 28, nom. nud.; FI. Ind., ed. Carey | (14 Jan 1832) 242; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 241; Wight, Ic. (1839) n. 146 (‘Guaria’); Walp., Rep. 1 (1842) 435, non Buch.-Ham. (1 Jan. 1832). — [Dysoxylum paniculatum Arn. ex Wight, Ic. (1839) sub n. 146, nom. in syn.] — Chisocheton paniculatus Hiern in Hook. f., Fl. Brit. India 1 (1875) 552, nom. superfl.; C.DC. in DC., Monogr. Phan. 1 (1878) 531; Gamble, Trees Beng., ed. 2 (1896) 16; Prain, Beng. Pl. 1 (1903) 315; Brandis, Indian Trees (1906) 139, 703; Pel- legr. in Fl. Indo-Chine 1 (1911) 736; ibid. Suppl. (1946) 692; Cowan & Cowan, Trees N Bengal (1929) 32; Kanj. et al., Fl. Assam 1, 2 (1936) 692; Chun, Sunyatsenia 4 (1940) 237; How & Chen, Acta Phytotax. Sin. 4 (1955) 18. — Schizochiton paniculatum (Hiern) Kurz, J. As. Soc. Beng. 44, ii (1875) 145; For. Fl. Burma (1877) 216. Dysoxylum multijugum Arn. in Wight & Arn., Prodr. (1834) 121; Steud., Nomencl., ed. 2, 1 (1840) 534; Voigt, Hort. Suburb. Calc. (1845) 135; M. Roem., Synops. Monogr. 1 (1846) 101; Drury, Handb. Ind. Fl. 1 (1864) 168. Cupania sp.: Wall., Cat. (1847) n. 4884B. Chisocheton balansae C.DC., Bull. Herb. Boiss. 2 (1894) 578; Pellegr. in Fl. Indo-Chine 1 (1911) 737, t. 81, f. 12-15; Lecomte, Bois Indoch. (1925) 134. Chisocheton coriaceus Pierre, Fl. For. Cochinch. 5 (1897) t. 346A; Pellegr. in Fl. Indo-Chine 1 (1911) (eiic Chisocheton thorelii Pierre, Fl. For. Cochinch. 5 (1897) sub t. 347; Pellegr. in Fl. Indo-Chine | (1911) 740 & Suppl. (1946) 696; H6, Ill. Fl. Vietnam 2, 1 (1992) 495. Chisocheton cochinchinensis Pierre, Fl. For. Cochinch. 5 (1897) t. 356B; Pellegr. in Fl. Indo-Chine | (1911) 739; H6, Ill. Fl. Vietnam 2, 1 (1992) 494. Chisocheton harmandianus Pierre, Fl. For. Cochinch. 5 (1897) t. 347; Pellegr. in Fl. Indo-Chine 1 (1911) 740. Chisocheton amboinensis Valeton in Hochr., Pl. Bogor. Exsicc. (1904) 67; Briq., Mém. Inst. Nat. Genev. 24 (1935) 66. Chisocheton chinensis Merr., Philipp. J. Sc. 21 (1922) 497. Chisocheton kinabaluensis Merr., J. Str. Br. Roy. As. Soc. 86 (1922) 316. Chisocheton sp.: Lane-Poole, Rep. For. Res. Terr. Papua New Guinea (1925) 100. Chisocheton siamensis Craib, Bull. Misc. Info. Kew (1926) 342; Fl. Siam. Enum. | (1926) 253; Pel- legr. in Fl. Indo-Chine, Suppl. (1946) 692. [Chisocheton benguetensis Elmer, Leafl. Philipp. Bot. 9 (1937) 3343, nom. non rite publ. [Dysoxylum sorsogonense Elmer, Leafl. Philipp. Bot. (1937) 3378, nom. non rite publ.] Chisocheton morobeanus Harms, Bot. Jahrb. 72 (1942) 185; P.F. Stevens, Contr. Herb. Austral. 11 (1975) 21; in Handb. Fl. Papua New Guinea | (1978) 147, t. 72; Johns, Comm. For. Trees Papua New Guinea 5 (1976) 214. Chisocheton cf. pachyrhachis: Hartley et al., Lloydia 36 (1973) 261. Tree to 37 m; bole to 14 m, 150 cm diam.; buttresses to 3 m tall, 2 m out or bole flut- ed to 10 m. Bark scaly, pale grey-brown; inner bark chestnut brown; sapwood straw. Limbs with conspicuous cicatrices. Leafy twigs 5—7 mm diam., dark brownish black, Mabberley et al. — Meliaceae 165 Fig. 24. Chisocheton cumingianus (C.DC.) Harms. Flowering shoot. Papua New Guinea, Gogol Valley (Mabberley 1757). Photograph D.J. Mabberley, June 1974. smooth but conspicuously lenticellate, sometimes with white latex, rarely myrmecophil- ous (and then sometimes the ant chamber housing scale insects, e.g. Coode 5454 in K). Innovations + rusty pubescent. Leaves to 120 cm, pseudogemmulate, crowded in dense terminal spirals; petiole 5-10 cm, 2.5—5 mm diam., terete or weakly flattened adaxially. Leaflets in up to 15 pairs, opposite or subopposite (proximally), (6—)10—42 by (2—)5— 14 cm, ovate to elliptic, papery to coriaceous, drying red-brown, glabrescent or hispid pubescent on the costae adaxially or, exceptionally, softly velutinous, bases asymmetric, acute, apices shortly cuspidate, costae 10—15 on each side, ascending, arcuate, + prom- inent abaxially, tertiary venation prominulous; petiolules (4—)6—12 mm, glabrescent to tawny tomentose. Thyrses to 50 cm, axillary, supra-axillary or borne on short shoots (3—8 thyrses per shoot) on defoliated twigs, branches or bole (Borneo), 2—3-branched; branches to 10 cm long, + pubescent; pedicels to 3(—4) mm; bracteoles 2 mm long, li- near; pseudopedicel to | mm long. Calyx 1-3 mm deep, campanulate, puberulous with- out, margin + entire. Petals (3) 4 (5), 12—20(—25) by 2.5 mm, spathulate, acute, pale yellow to white, reddish in sicco. Staminal tube c. | mm diam., + glabrous without, + pubescent within from just below anthers to base, margin 6—9-lobed, lobes to 2.5 mm long, entire to 2- or 3-fid; anthers 6—9, 1.5—2.2 mm long, elliptic-oblong, locel- late, glabrous to villous. Disk to 0.5 mm tall, annular, glabrous. Ovary in female (? and hermaphrodite) flowers 3- or 4-locular, each locule with | (2) ovule(s); style pubescent in proximal 3/4; stylehead disciform to capitate. /nfructescence to 30 cm, pendent. Cap- sule to 7 cm diam., globose to pyriform, occasionally weakly rostrate, orange-red, gla- 166 Flora Malesiana ser. I, Vol. 12 (1) (1995) brous to velutinous; stipe to 1.5 cm long; pericarp usually with white latex. Seeds 3 or 4; testa blackish brown; aril circumhilar, margin crenate, sometimes with extension to micropyle, orange-red; cotyledons superposed. 2n = 46, 92. — Fig. 24. Distribution — Continental Asia from Assam and tropical China through Indochina to eastern Malesia: Philippines, Celebes, Moluccas (Ambon, Ternate), New Guinea, Bismarck Archipelago (Manus, New Britain, New Ireland). Habitat — Rain forest, to 1300 m altitude. Uses — The wood of Indian specimens has been considered by the timber industry. In New Guinea used as a fish poison. Subdivision — Three subspecies are recognized, two of them occurring in and en- demic to Malesia. The third one, restricted to the Asiatic mainland, is subsp. balansae (C.DC.) Mabb. [Chisocheton balansae C.DC., Guarea paniculata Roxb., Dysoxylum multijugum Arn., Chisocheton paniculatus Hiern, Schizochiton paniculatum (Hiern) Kurz, C. coriaceus Pierre, C. thorelii Pierre, C. cochinchinensis Pierre, C. harmandia- nus Pierre, C. chinensis Merr., C. siamensis Craib]. It usually has rather pubescent leaves and axillary or supra-axillary thyrses, while 2n = 46. KEY TO MALESIAN SUBSPECIES la. Inflorescences borne on supra-axillary or ramiflorous short shoots, rarely supra- axallanvAanacSimplee ew, Marites. Actin cae 6 eee a. subsp. cumingianus be intlorescences bore On Dole. =. 2-4-2 ee eee b. subsp. kinabaluensis a. subsp. cumingianus Chisocheton cumingianus (C.DC.) Harms subsp. cumingianus; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 348. — Dasycoleum cumingianum C.DC. — Chisocheton cumingianus (C.DC.) Harms, s.str. — Chisocheton amboinensis Valeton — Chisocheton sp.: Lane-Poole — Chisocheton ben- guetensis Elmer — Dysoxylum sorsogonense Elmer — Chisocheton morobeanus Harms — Chiso- cheton cf. pachyrhachis: Hartley et al. Distribution — Malesian distribution as for species except Borneo. Vernacular name — Official common name in Philippines: balukanag. Note — There 1s a distinct trend from axillary inflorescences in the north-west (Philip- pines) to reduced branches bearing inflorescences in defoliated twigs in the southeast, though there are exceptions. 2n = 92. b. subsp. kinabaluensis (Merr.) Mabb. Chisocheton cumingianus (C.DC.) Harms subsp. kinabaluensis (Merr.) Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 349. — Chisocheton kinabaluensis Merr. Distribution — Mountains of northern Borneo: Kinabalu, Trusmadi. Note — The inflorescences are always borne on the bole, often very close to the ground indeed. It is noteworthy that, contrary to general expectation, it is the high altitude taxon which is the truly cauliflorous one in this species. Mabberley et al. — Meliaceae 167 28. Chisocheton patens Blume Chisocheton patens Blume, Bijdr. (1825) 169; Schultes & Schultes, Syst. Veg. 7 (1829) 83; (1830) 1626; G. Don, Gen. Syst. 1 (1831) 685; Migq., Fl. Ind. Bat. 1, 2 (1859) 537; C.DC. in DC., Mon- ogr. Phan. 1 (1878) 529, t. 7, f.5; King, J. As. Soc. Beng. 64, ii (1895) 34; Koord., Minah. (1898) 385; Ridley, J. Str. Br. Roy. As. Soc. 33 (1900) 591; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 350; in Tree Fl. Malaya 4 (1989) 235. — Schizochiton patens (Blume) Spreng., Syst. Veg. 4 (1827) 251; Walp., Rep. 1 (1842) 429 (‘Schizogiton’) in ind.; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 27, 29; M.Roem., Synops. Monogr. | (1846) 102. Chisocheton divergens Blume, Bijdr. (1825) 169; Schultes & Schultes, Syst. Veg. 7 (1829) 83; (1830) 1627; G. Don, Gen. Syst. 1 (1831) 685; Miq., Fl. Ind. Bat. 1, 2 (1859) 537; C.DC. in DC., Monogr. Phan. | (1878) 528; Curtis, J. Str. Br. Roy. As. Soc. 25 (1894) 22; King, J. As. Soc. Beng. 64, ii (1895) 35; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 292, t. 162, f. H; ibid., ed. 2, 19b1 (1940) 139, t. 30, f. h; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 99; Valeton in Hochr., Pl. Bogor. Exsicc. (1904) 68, incl. var. genuinus Valeton, var. minor Valeton, var. robustus Vale- ton; Brandis, Indian Trees (1906) 139; Backer, Schoolfl. Java (1911) 208; Koord., Exk. Fl. Java 2 (1912) 443; Craib, Aberd. Univ. Stud. 57 (1912) 36, incl. var. robustus Valeton; Koord., Atlas (1913) t. 165; Schmidt, Bot. Tidsskr. 32 (1916) 328, incl. var. robustus Valeton; Ridley, Fl. Malay Penins. 1 (1922) 390, incl. var. patens (Blume) Ridley, nom. illeg.; Brig., Mém. Inst. Nat. Genev. 24 (1935) 64; Backer & Bakh. f., Fl. Java 2 (1965) 124; T.D. Penn., Blumea 22 (1975) 496, t. 12, f. c & d; Corner, Seeds Dicots 2 (1976) & 2 (1976) t. 383 left. — Schizochiton divergens (Blume) Spreng., Syst. Veg. 4 (1827) 251; Walp., Rep. 1 (1842) 429; M. Roem., Synops. Monogr. | (1846) 102; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 26, 28. [Trichilia longissima Wall., Cat. (1847) n. 8069, nom. nud.] Cupania sp.: Wall., Cat. (1847) n. 8069. Schizochiton ?: Wall., Cat. (1847) n. 9040. Schizochiton tetrapetalum Turcz., Bull. Soc. Nat. Mosc. 1 (1858) 411. — Chisocheton tetrapetalus (Turez.) C.DC. in DC., Monogr. Phan. 1 (1878) 530; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 368; Briq., Mém. Inst. Nat. Genev. 24 (1935) 65; Elmer, Leafl. Philipp. Bot. 9 (1937) 3347. [Melia pendula Reinw. ex Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 29, nom. in syn.] [Trichilia hexandra Blume ex Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 29, nom. in syn.] Chisocheton fragrans Hiern in Hook. f., Fl. Brit. India 1 (1875) 551; C.DC. in DC., Monogr. Phan. 1 (1878) 529. Chisocheton glomeratus Hiern in Hook. f., Fl. Brit. India 1 (1875) 551 ; C.DC. in DC., Monogr. Phan. 1 (1878) 532; Curtis, J. Str. Br. Roy. As. Soc. 25 (1894) 22; King, J. As. Soc. Beng. 64, ii (1895) 30; Ridley, Fl. Malay Penins. | (1922) 389; Briq., Mém. Inst. Nat. Genev. 24 (1935) 66; H6, III. Fl. Vietnam 2, | (1992) 494. Chisocheton holocalyx Hiern in Hook. f., Fl. Brit. India 1 (1875) 551; C.DC. in DC., Monogr. Phan. 1 (1878) 530. Chisocheton vrieseanus C.DC. in DC., Monogr. Phan. | (1878) 536. Chisocheton barbatus C. DC., 1.c. Chisocheton laxiflorus King, J. As. Soc. Beng. 64, ii (1895) 33, p.p. Chisocheton fulvus Merr., Philipp. J. Sc., Bot. 3 (1908) 146; Enum. Philipp. Flow. Pl. 2 (1923) 367. [Chisocheton urdanetensis Elmer ex Merr., Enum. Philipp. Flow. Pl. 2 (1923) 368, nom. in syn.] [Chisocheton apoensis Elmer ex Merr., Enum. Philipp. Flow. Pl. 2 (1923) 368, nom. in syn.; Elmer, Leafl. Philipp. Bot. (1937) 3341, nom. non rite publ.] Tree to 35 m, but often flowering when 2—3 m tall; bole to 20 m and 70 cm diam., sometimes fluted or buttressed, buttresses to 2 m tall, 1 m out and 8 cm thick, concave. Bark pale greenish to black, smooth to faintly cracked, lenticellate, the lenticels in hori- zontal rows (Pennington); inner bark pale to dark brown; wood pale to dirty cream, often smelling of methyl mercaptan. Leafy shoots c. 6 mm diam., glabrous to decidu- ously tomentose, bark dark, cicatrices conspicuous. Leaves to 70 cm, paripinnate to 168 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 25. Chisocheton lansiifolius Mabb. a. Leaf and inflorescence; b. flower; c. inside staminal tube; d. half flower base; e. pistil; f. fruits (SAN 30737). Drawing Yap Pak Hau. © The Natural History Mu- seum, London. Bull. Brit. Mus. (Nat. Hist.), Series Botany, vol. 6 (4), 1979. Mabberley et al. — Meliaceae 169 pseudogemmulate, in terminal bunches; petiole 7-15 cm, glabrous to pubescent. Leaf- lets 6—28 by 2.5—10.5 cm, narrowly oblong to oblong- or elliptic-lanceolate, opposite to subopposite, often maturing all together, thinly coriaceous, often conspicuously paler abaxially, base + rounded or rarely subcordate, + unequal, shortly acuminate, adaxial surface glabrous or with tomentose midrib and pubescent veins, abaxial surface gla- brous or with tomentose midrib and pubescent veins, abaxial surface glabrous to softly tawny pubescent, midrib and veins tomentose, costae 9—14 on each side, + prominent abaxially, tertiary venation often conspicuous; petiolules 3—6 mm. Thyrse to 90 cm, borne in upper axils or supra-axillary, pendent, paniculiform; most proximal branches to 17 cm (10 cm in females), ultimate branchlets cymules of subsessile or shortly pedi- cellate fragrant flowers; axes glabrous to tomentose; bracteoles minute. Calyx 2.5—3 mm, cupular to shortly tubular, puberulous, margin subentire to minutely, irregularly toothed. Petals 4, 5—10 mm long, subspathulate-elliptic, glabrous to glabrescent. Sta- minal tube 5—7(—8) mm, glabrescent or minutely pubescent near mouth without, pubes- cent, tomentellous or very rarely villous within, margin with (S—)6—8 linear-triangular lobes a little shorter than anthers; anthers (5) 6 or 7 (8), basifixed, glabrous, locellate. Disk absent or very short, fleshy, glabrous, + lobed. Ovary pubescent; style glabrous to densely short pubescent; stylehead cylindric to clavate. Capsule to 5 cm long, 4.5 cm diam., subglobose, stipitate, glabrous to tomentose (especially when unripe), 2-locular; stipe to 2cm. Seeds 2, 5—11 mm long, to 8 mm wide, scutiform, half covered by an aril. Distribution — Peninsular Thailand; Malesia: Sumatra, Bangka, Malay Peninsula, Borneo, Java, Philippines, Celebes. Habitat — Common in lowland rain forest, to 500 m altitude. Notes — 1. This is an extremely variable species of which Chisocheton lansiifolius is a satellite. It cannot be satisfactorily subdivided throughout its range but in particular islands it may be possible and useful to recognize distinctive microtaxa. For example, in the Malay Peninsula, most herbarium specimens can be allotted to the almost glabrous Chisocheton patens s. str., the hairy ones to ‘Chisocheton glomeratus’, and, in the Phil- ippines they can be similarly divided between Chisocheton patens s. str. and ‘Chisoche- ton fulvus’. Even so, there are always a number of intermediates (Mabberley, |.c.). The variation pattern somewhat resembles that of Chisocheton lasiocarpus (q.v.) but is com- plicated by being spread over several islands and thus approaches the situation present- ed by Vavaea amicorum (q.V.). 2. Some specimens from Sarawak, particularly from Semengoh Forest Reserve near Kuching are curious for their large coriaceous leaflets, which resemble those of Chiso- cheton lansiifolius. These are treelets to 3 m tall and require further study (Mabberley, e:); 29. Chisocheton lansiifolius Mabb. Chisocheton lansiifolius Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 352, t. 5. Tree to 18 m; bole to 25 cm diam., sometimes fluted and buttressed, buttresses to 2 m tall, c. 5 cm wide, concave. Bark smooth, dark brown; inner bark yellow; sapwood 170 Flora Malesiana ser. I, Vol. 12 (1) (1995) pale brown. Twigs dark brown, lenticellate, cicatrose. Leafy twigs c. 8 mm diam. Leaves to 54 cm long, paripinnate, ?pseudogemmulate, to 5-jugate; petiole 8-15 cm. Leaflets to 42 by 10.5 cm, oblong-elliptic or -ovate, coriaceous, subglabrous, bases acute to ob- tuse, apices acuminate, acumen to 18 mm, costae c. 14 on each side, arcuate, venation prominent on both surfaces in sicco; petiolule 6—11 mm, drying blackish. Thyrse to 65 cm long, paniculiform; proximal branches to 18 cm, squarrose, branched, these branches passing interceptibly, like the major branches on the main axis, into cymose fascicles of 1—6 flowers. Calyx c. 1.5 mm deep, cupulate, rugose, margin obscurely 4-lobed. Petals 4, 8—9 by 1.5 mm, weakly pubescent without, glabrous within, creamy white to pinkish, connate at the base. Staminal tube glabrous without, cottony pubescent within, margin 6-lobed, the lobes entire, lanceolate, c. 2 mm long; anthers 6, c. 2.5 mm long, glabrous, weakly locellate. Disk annular, thick. Style terete, pilose in proximal 3/4; stylehead subcylindrical. /nfructescence to 85 cm long; axis c. 8 mm diam.; branches bearing | or 2 capsules. Capsule to 5 cm diam., subspherical, stipitate, red; valves 4. Seeds 2, scutiform, c. 3 cm diam. — Fig. 25. Distribution — Malesia: north and eastern Borneo. Habitat — Primary and secondary forest including peat swamp forest, to 100 m alti- tude. 30. Chisocheton granatum Mabb. Chisocheton granatum Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 835452 6: Tree to 12 m; bole to 8 m, 17 cm diam. Twigs cicatrose, lenticellate. Leafy twigs c. 6 mm diam. Leaves to 120 cm long, pseudogemmulate, in lax terminal spirals, + pubes- cent; petioles 5—10 cm long. Leaflets to 24 by 7.5 cm, in up to 12 pairs, oblong to ob- long-ovate, pale abaxially in sicco, bases cuneate, somewhat asymmetric, apices abrupt- ly shortly acuminate, costae c. 22 on each side, weakly arcuate, almost reaching margin, prominent abaxially in sicco; petiolules 8-12 mm; pseudogemmula markedly circinnate. Thyrse to 25 cm long, supra-axillary, often borne in axils of unexpanded leaves; branches to 6 cm or short-stalked cymules (female). Calyx 2.5—3 mm, campanulate, pubescent, margin truncate. Petals 4, c. 12-16 by 3.5 mm, linear-spathulate, creamy white, pubes- cent without, imbricate, forming a clavate corolla in male. Staminal tube pubescent in distal half without, glabrous within, inflated near anthers, margin obscurely crenate or truncate; anthers 6, 1.5 mm long, oblong, locellate, glabrous. Ovary 5-locular; style pilose in proximal 3/4, stylehead shortly cylindrical, glabrous, apically lobed. Capsule to 9 cm diam., flattened globose; valves 5, glabrous, tough, red brown without, white within, without latex. Seeds 4 or 5, c. 3 cm long, scutiform (where 4) or like orange- segments (when 5); cotyledons with white latex. Distribution — Malesia: Borneo (Mt Kinabalu area, and one sterile specimen from E Kalimantan). Habitat — Hill forest, (?300—)900—1500 m altitude. Note — This species is somewhat anomalous in this section in having exarillate seeds. Mabberley et al. — Meliaceae 171 Section Holopentas Chisocheton sect. Holopentas Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 27. Dasycoleum Turez. (genus). — Chisocheton sect. Dasycoleum (Turcz.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 296; Mabb., Bull. Brit. Mus. Nat. Hist., Bot. 6 (1979) 354. — Schizochi- ton Spreng. sect. Dasycoleum (Turcz.) Kuntze in T. Post & Kuntze, Lex. Gen. Phan. (1903) 505. Trees or treelets, pachycaul to slender leptocaul, usually with white latex. Jndumen- tum of simple hairs. Leaves pseudogemmulate. /nflorescences axillary or supra-axillary. Petals in 1-2 whorls, imbricate to valvate. Staminal tube lobed or not, anthers glabrous, locellate. Disk obscure to shallowly cupulate. Ovary 2—9(—11)-locular. Fruit laticiferous. Seeds sarcotestal. Distribution — 14 species, Indochina throughout Malesia to New Guinea. Notes — 1. Chisocheton macrophyllus [and its ally C. dysoxylifolius (Kurz) Hiern from mainland Asia] is better placed in this section than in the last one, where it is found in Mabberley, op. cit. 344. Miquel’s sectional name is employed here, as it is clear from his remarks under Dysoxylum setosum Miq. (op. cit. 14) that his § is meant to signify a section and is not rankless as hitherto held. (Type, here selected: Chisocheton ceramicus). 2. The section can be divided into two series, though C. mendozae and C. grandiflo- rus (Kurz) Hiern from Burma and Thailand are links. KEY TO THE SERIES la. Corolla enclosing at least one petal, imbricate; tube weakly lobed or unlobed, Sinlehead:discoid ito:subcapitate) iraitadehiscent yee ee cs See 2. A. flavida beliblowerm pentainerous; Inuit: mdehiscent, was not os 2 2 eee 3) 37a. Leaflets with a recurved margin, with numerous large dark orange-brown peltate scales on the lower surface and visible to the naked eye as evenly distributed spots PON cr thes eh tea Pe ER LAE AR Mah a PAO Tres eC En RR eT 27. A. variisquama b. Leaflet margin not recurved, with numerous small yellowish-brown or orange- brown peltate scales which have an entire or fimbriate margin or almost stellate scalesionmhe lowersuntacesa. ated Sate ae See ee 44. A. crassinervia Mabberley et al. — Meliaceae 205 38a. Fruit 2-locular, without prominent ribs ..:.......00.5.0.00.0. 28. A. rimosa . Fruit 10-locular, with 10 prominent longitudinal ribs ........ 29. A. costata PRIOR mScontnlt) xiii «keene eer aoat dam waite selax 13. A. lawii MEIEIMCID MCE ghee, 5, 2S Sap beers ste see edie wie oodas.e ese aes Se 40 . Leaflets with few to numerous small, reddish-brown, pale brown or orange-brown stellate hairs and scales or peltate scales on the midrib below; fruit large, up to 3.2 cm long and 3.8 cm in diameter, usually subglobose with an apical depression and 3-locular but sometimes the seed failing to develop in | or 2 of the locules; peri- Papubicknane Wweody asx). 34 oe Hanes Sok) leeme eats orfs dai. 48. A. edulis . Leaflets with large pale brown or orange-brown peltate scales which are entire or have a fimbriate margin, few to numerous on lower leaflet surface; fruits small, to 2.5 cm long and 1.5 cm in diameter, subglobose, ellipsoid or obovoid without an apical depression, with | or 2 locules, the pericarp thin . 25. A. elaeagnoidea . Staminal tube with a narrow pin-prick aperture c. 0.3 mm across, with an entire Midi. -AEREES CIRC LUGEE bh xs. 255/01 Sie See tacls en) ts oc. 38. A. silvestris . Staminal tube with the aperture wider than 0.3 mm, the margin shallowly lobed, amber prottudm:s through the apertures oa sedow. eden Qik i cea 42 . Leaflets usually markedly ovate, scales almost absent from the lower surface, veins often red when dry; fruit asymmetrically ellipsoid with one locule ....... PEE SN APS Sno, Sh eV Bde eda Sh paolo fees 39. A. perviridis . Leaflets elliptical, ovate or obovate, scales few to numerous on the lower leaflet surface, veins neither black nor red when dry; fruits symmetrical with 2 or 3 lo- RY hid o.oo, oh wm wy nn on, GEN UTIERENS OP aay. CNS CCD. ERR AIS ed 43 . Leaflets narrowly elliptical or narrowly obovate, scales numerous on lower sur- BAREPOMMCAOE 220 src aw ccm cnependas ea EE Le AEE 31. A. speciosa . Leaflets neither narrowly elliptical nor narrowly obovate, scales few to numerous Simewenleatletsnmace 00 Sie rabble A oe i eae Haren aa 44 . Lower surface of leaflet with few to numerous reddish-brown peltate scales which have a fimbriate margin and with paler scales interspersed; leaflet apex with a par- er SICOONHUISEraCUINEN (2.0.5 .442 54 55hcu as eee 41. A. cremea . Lower surface of leaflet with peltate scales all of one type, or if paler scales inter- Spersea then ‘acumen tiot parallel-sided) > iia iste See Sled ae Ce 45 B Beatles nsuallyOSiSio8 2c 25 IS Pe SER 30. A. agglomerata Malseaitens AEWeE tanlD cs. 4s /5:2.0.5.0 2d daetee eRe eee 32. A. korthalsii . Scales densely covering the midrib on lower surface of leaflet and immediately ad- jacent to the midrib, occasionally also on the lateral veins................. 47 / caies ty absentfronrlower:surface ofileafleh situ sia iyo. Ji a ie 54 . Scales large (many 0.2 mm across), orange-brown, reddish-brown or almost Wihite; witha tendency to flakeioffs ine neti cele cont ke: Bele 48 . Scales less than 0.2 mm across or if larger, then dark reddish-brown or purplish- brown and adhering closely: tothe leaflet oot. hen yo ih ot 50 . Leaflets with pale orange-brown or grey peltate scales few to numerous on the RuiciO.Ob the AOWEr SUDLACE ou, h2 anew ot a A 24. A. lepiorrhachis Flora Malesiana ser. I, Vol. 12 (1) (1995) Leaflets with numerous to densely covered with reddish-brown peltate scales on themidnbionithe: lowersurtace anise See A ee 49 . Peltate scales shiny orange-brown, often with a long fimbriate margin ........ SACRA IY, AST LUE ISR GALS EE SA EN Arctic Ook cone, OWE 35. A. flavescens Peltate scales large, shiny and reddish-brown, either entire or with a short fimbri- AT MAT SUA BAT ese CAL ee Fee 23. A. lancilimba . Anthers and/or staminal tube with simple white hairs.................... 5) Anthers and staminal tube without hairs :). 202.90) 22s. ee 59 . Flowers with the staminal tube appearing to be deeply lobed, the anthers continu- ous with the inner surface of the lobes, densely covered with simple yellow hairs on the margins of the lobes and the anthers ........... 56. A. euryanthera Staminal tube neither deeply lobed nor densely hairy, but with some simple white hareskoOnktheramthersi. Rees PAN ee ee ee eee 36. A. samoensis a. Leaflets with purplish-brown fimbriate peltate scales densely covering the midrib below and + absent from the rest of the lower surface of the leaflet ........... belie eb Bons ar nya eee Se kD re ek en ay Be, ©: 34. A. glabrata Leaflets with dark reddish-brown peltate scales numerous on the midrib below 53 a. Leaflets (7—-)9—13(—15), stellate scales absent ......... 33. A. scortechinii Leaflets 3—5(—7), some stellate scales interspersed among the peltate scales .. Tee Ne cay nl eae Se Weis hen HS chs ae Oe ee 50. A. odoratissima . Veins usually black when dry; petals 5; fruit indehiscent ... 40. A. leptantha Veins rarely black when dry; petals usually fewer than 5; fruit dehiscent ..... 25) a. Leaflets anarkedly curved and asymmietnical 22455 222.0. ...-: 1. A. cucullata lbeafletsonlyshghtlysasymmietrical)...\.csees ae) toe 2 eee 56 sRipeinuits up to. 3:Sicmiinidiameter 265,25 22 eieee seas 4 > > ola eee Si) Ripe drits more thants:s cnminidiametenr..n.ite ee ose oe a2 ee 58 . Leaflets brown when dry, with few pale brown peltate scales which have a fim- briate margin and compact reddish-brown stellate hairs interspersed on the lower surface; colour of fruits when fresh not known, dark reddish-brown when dry .. 2 BE RST OR Se a or MS EPPS Soc 4 cry Bree 9. A. lepidopetala Leaflets pale green or orange-brown when dry, with few to numerous almost white or pale orange peltate scales (rarely with a few hairs interspersed) on the lower surface; fruits pink, yellow, or carmine-red when fresh, pale brown or grey SA OCSTLAG lA ee Se aS onan, eR OES PORE RA Re ec al, Oe PER RARE Ce! 1 13. A. lawii . Leaflets 15—25, lanceolate, veins 20—50 on each side of the midrib, indistinct ... NUR, PE s.. Reet cares aah rg Los 8. A. multinervis Leaflets 7-15, ovate, obovate, elliptical or oblanceolate, lateral veins 6—20 on eachiside'ohthe midnribayii wee asses eens. Cee ee 59 . Leaflet surfaces reddish-brown or orange-brown and often rugulose when dry, with few reddish-brown or grey peltate scales which have a fimbriate margin on the midrib below and often interspersed with minute reddish-brown pits ....... SPST ETA RR FER SET RR, LES 3. A. macrocarpa Mabberley et al. — Meliaceae 207 60a. 61a. 66a. 67a. Leaflets pale brown and not rugulose when dry, with small orange, orange-brown or pale brown peltate scales which have a dark central spot numerous on the mid- rib below, sometimes numerous on lower surface of leaflet but usually absent . . . PE ER ON 6c Ae Ree RA whew ss wea eee Eee 2. A. flavida Leaflets with numerous or densely covered with dark reddish-brown peltate scales and sometimes paler stellate scales on the midrib below, few to numerous on the rest of lower surface of leaflet; the staminal tube appearing to be deeply lobed, the anthers continuous with the inner surface of the lobes, densely covered with simple yellow hairs on the margins of the lobes and the anthers . 56. A. euryanthera Leaflets without or with few dark reddish-brown peltate scales, but with numer- ous pale brown or orange-brown peltate and stellate scales on the midrib below, few to numerous on the rest of lower surface of leaflet; the staminal tube neither Mae ayeleveernor Uensely hairy Sis Gores ce ee eo ee ee ee OY 61 Leaflets with pale brown or reddish-brown peltate scales which have a fimbriate margin, numerous on or densely covering the midrib below, always with a few much darker peltate scales and dark reddish-brown pits interspersed, the inflores- Gener with reddish-brown stellate scales 22.22 iis: 5522/22 see ee Cees 62 Leaflets without dark reddish-brown peltate scales and dark reddish-brown pits, the indumentum usually brown or orange-brown, the pits, if present, orange- ERP 5. tone Gon. wenn = LTE SNA SAE SG Se te eset Ese Se 63 . Leaflets pale bluish-green or yellowish green on both surfaces when dry....... i a ano icer ss on.ca oi'% o'sk tet AtS Ls PORTO ND. EOE EE Sted Set OE) SELES 58. A. sapindina eotleismark¢brownowhen drys) iw. wae Mae ae aah Pte 57. A. polyneura . Leaflets pale yellowish-green when dry or with whitish-green or orange veins 64 eafletsnotpale yellowish-green: whem diy .s.. 15 tei ee hale. 67 . Fruit dehiscent (indicated in unripe fruits by the presence of three longitudinal MACESON tHE PELICATP) Jie m- wortcneinve nga Oese aos Ae Meson, 13. A. lawii Fruit indehiscent (any longitudinal markings on the fruits are slight depressions RMSE Tan TASES) este UN Le ed, CR AE Nae Nc wd, eh Re ies 65 a. Fruit subglobose with an apical depression or pyriform, pericarp thick (c. 3 mm) ATG 00000 | Ree PR ty 9 Ae tye RED rar ee iret yooh ee or 66 Fruits subglobose without an apical depression or obovoid, pericarp thin (c. 1 mm) Re ee ee ee ele tee eee re Scan ae ene caer ae ee 46. A. parviflora Leaflets pale brown or yellowish-brown when dry; fruit usually subglobose and 3-locular but sometimes the seed failing to develop in 1 or 2 of the locules ..... eS LA Cert As Sis ARE Pete SAS TR ee eee ae Meroe a ota 48. A. edulis Leaflets yellow or yellowish-green when dry; fruits usually pear-shaped, some- Gales subplobose ‘and 2:locularsy Sense. Ba 3 47. A. leucophylla Staminal tube obovoid with a narrow aperture, anthers included or just protruding err Serer) een Me. SS, Dales Ns AIT a eS 54. A. brassii Staminal tube cup-shaped, anthers protruding, ........ 2s 0G: Pes eee 68 Flora Malesiana ser. I, Vol. 12 (1) (1995) 70a. 76a. a. Inflorescences always axillary; fruit ellipsoid, up to2 cmlong ............... sity. a aR re Mere eee ae cease shee eet arte ot: 50. A. odoratissima Inflorescences ramiflorous and axillary; fruit pyriform, up to 4.5 cm long ...... WS APO oe ee Eee ee ere eo ea 49. A. macrostigma a. Leaflets with few to densely covered with stellate hairs or scales on the lower sur- face; when sparse, some hairs or scales occur evenly distributed between the veins and their presence visible withithe makedieye): 5.22254 =, ns52)-)-- 32) ee 70 Leaflets without or with few hairs on the lower surface, with scales visible only with a lens or densely covered with hairs on the midrib only, few and unevenly Scattered. on‘thejrest.ofithe lowersurface, ona net) ies oe. 3\-6-S ae 88 Reticulation subprominent on both surfaces of leaflet when dry; with some hairs (on the twigs and sometimes elsewhere) which have a central rachis and several whorlsjof anms radiating frombity 42-0. sa ree eee te 16. A. ramotricha Reticulation not subprominent; all hairs lacking a central rachis ............ 71 . Hairs pale yellowish-brown or if reddish-brown then flower trimerous...... 72 Hairs reddish-brown or brown, or if pale brown then flower pentamerous ... 74 we Weatlets Matai See oe ace SR ee PPI tate 10. A. densitricha eaflets!1=9, rately, li 2teds aie ate aes See ane oe See 73 suPeltate scales aDSentist.wee aioe aa ae. es eee cea ete 14, A. teysmanniana Peltate scales present, atleastion the'shootkapex 227.3... oar 13. A. lawii . Hairs on lower leaflet surface numerous and with the arms of adjacent hairs over- lapping, but leaving the surface of the leaflet visible..................... ip: Hairs on lower surface of leaflet few or at least with the arms not usually overlap- DING: ee%s ua {oGre ll sities eee ae) ee kere aoe ce Cr ee 79 . Hairs compact with arms all + equal in length c. 0.5 mm, brown, densely cover- ing the midrib, densely covering or scattered on the rest of the lower surface of LeatleE Di. Mee eos lene oe aerate eee eee tee ca LI ote 76 Hairs large and spreading, arms unequal in length up to 1 mm long and in some species (e.g. A. rufibarbis) up to 4 mm, usually reddish-brown and numerous on lowersustace. of leatlet, 05.2 screed feoienme soir s4ta-bo creak een 77 Lower surface of leaflet densely covered with persistent brown stellate hairs and SCANS ype wenesessyr eee: Gas oc ae eae ae Be he tant donee 45. A. sexipetala Lower surface of leaflet densely covered with brown stellate hairs which are de- ciduous and leave numerous pits and an uneven indumentum most dense near the OUTS er OM: 108 (=) 0 me ee ety Seg ee MCE an Rt ee en EO Spe me 42. A. forbesii . Stellate hairs with long arms, up to4mm .............. 84. A. rufibarbis Stellate hairs with arms not usually exceeding 1 mm in length ............. 78 . Lower surface of leaflet with numerous stellate hairs, the arms of adjacent hairs overlapping, interspersed with some pale brown hairs which have one or few ascending arms; fruit up to 5 cm long and 3.5 cm wide, with a hard woody pericarp. 2=4 mmsthicks, 0.6 ic pe I a ISS Se 82. A. hiernii Mabberley et al. — Meliaceae 209 b. Lower surface of leaflet with numerous stellate hairs, the arms of adjacent hairs 79a. 80a. 8la. 85a. S6a. 87a. overlapping, interspersed with pale brown stellate hairs which have several as- cending arms; fruit up to 2.5 cm long and 1.7 cm wide, with a thin brittle pericarp Fes Svea saat Keyser yh eS eA AE SD 79. A. tomentosa Leaflets with indumentum of few fimbriate peltate scales and stellate hairs on the lower surface; fruits small, up to 4mm in diameter, hairs few or absent ....... rR ACA re SESS DRT A Beg BS ean accede 77. A. palembanica Peltate scales absent from lower surface of leaflet; fruits | cm or more in diameter, Sante HSE HMUMICH TUITE ae: f2.ee Nee MRCS mee zens Soleo. Bales? 2 80 Leaflets usually more than !1, elliptical or oblong, the arms of adjacent hairs on PemeasacCe MOU OVERLAP PIS | GANS rei sane cvcce secs eteps yo Ne Meee. SD 81 Leaflets rarely more than 11, most obovate, if more than 11 then the indumentum dense and arms of adjacent hairs on lower surface overlapping ............ 83 Leaflet surfaces not rugulose and without pits, leaflets up to4 cm wide ........ rR HRA ANTS ors es ST tL O.. uk ee eeee, 76. A. exstipulata Upper surface of leaflet rugulose and pitted, lower surface with numerous pits, FcAbcEs tie Cl WAGE 96! ca2..2 SSR. PRE OL. 2B ls Fe Le se 82 Hairs and scales mostly reddish-brown ................ 75. A. rufinervis Hairs and scales mostly orange-brown ............... 47. A. leucophylla Pt aucun branched ree ay) 2.0 Lock aeisieits Mose Abe Mt DUZER 2GY eed. 84 Tree branching several or many times when mature ..................... 85 . Leaflets usually pale green when dry, with numerous reddish-brown stellate hairs on lower surface, not glabrescent; inflorescences axillary . 73. A. tenuicaulis Leaflets usually bluish-green when dry, with numerous to densely covered with orange-brown stellate hairs on the midrib below, interspersed on the lower sur- face with paler stellate hairs which have fewer arms, glabrescent; inflorescences STM OHOWS A MATL GINA ool eR oe ee ed EAE 59. A. ceramica Leaflets with 23—47 pairs of veins, with few stellate hairs on the lower surface . . TUS HI Ne ore all eee rr agTt sie bs ee ES 74. A. membranifolia Most leaflets with fewer than 23 lateral veins, with numerous hairs or scales on PET MIORVECRE SEUINACE 5.15 c, caseae proce See win & 00s 2a, bomwatanseewe woksgeie colle Neue 86 Leaflets with numerous pale stellate scales on the lower surface, interspersed with reddish-brown stellate hairs which have arms of different lengths ............ SMa AD SRSA, nes CE IL ls Be, ee Rae 79. A. tomentosa Leaflets with numerous pale brown or reddish-brown stellate hairs which have amms of similar lengths on the lowerisutiace: (ite Se Se aa eae Po 87 Leaflets with reddish-brown stellate hairs interspersed with paler brown stellate hairs on the lower surface, reticulation same colour as rest of lower surface of leaflets e orcreiiwth oad ee Ue a ae 73. A. tenuicaulis Leaflets with pale brown stellate hairs interspersed with pale brown stellate scales on the lower surface, reticulation white or pale brown on the lower surface when ETE i Us aes wei cis Wiad noosa cette hmm eR 78. A. brownii 210 Flora Malesiana ser. I, Vol. 12 (1) (1995) 88a. Lower surface of leaflet with numerous stellate or peltate scales .......... 89 b. Leaflets with hairs or scales few on the lower surface between the veins when mature,/but/sometimes densely covering the mudnibyi< <1: 24.5286 eee 93 89a. Upper and lower surfaces of mature leaflets with numerous stellate scales which havieva pale: niarsinsiers swam esc tt ee eee 68. A. pyriformis b. Scales stellate of uniform colour, scales usually absent from upper surface of ma- timmetoatletisns Aie.2isues Qs SRS BARES ae ora. 2 os arya renee ied oe 90 90a. Stellate scales interspersed with compact stellate hairs .... 45. A. sexipetala b. Scales peltate, often with a fimbriate margin, if scales stellate then stellate hairs BDSM Mersey eae creel 1 See eee, 0 ee ol Ola: PEruitidehiscenitr: ex isert as ea RAs, eee, ee 13. A. lawii bekinuiindehiscent (ey owas Ley Peis 2: 9 92 OQaascalesralliofroneity penta cues eee I. See eee 44. A. crassinervia b: “Stellate:andt peltatescalesspresent together’. se2 =) 47. A. leucophylla 93a. Stellate hairs or scales more than 0.15 mm in diam., numerous on or densely cov- ering the midrib, sometimes also on the lateral veins, almost absent elsewhere 94 b. Stellate hairs or scales either very small, less than 0.15 mm in diameter, or al- most totally absent from the midrib below and from the rest of the lower surface OPIEaH eH, PR oe ao rnta ite DEW oaks SENAY Pe a 109 94a. Leaves + sessile or with a short peduncle of not more than | cm; the basal part of leaflets much smaller than the rest and subrotund ....... 61. A. subsessilis b. Leaves not sessile, the basal leaflets only slightly smaller than the rest and of simlarshape’ .. Fie WA 22.1 k te A cee felts 2c er 05 95a. Reticulation subprominent on lower surface and often on upper surface of leaf- let). OE Ph AA AL eae Eee, GA EE es 96 b: Reticulation:may be visible, but not subprominent>. .:..... 225 Joe eee i) 96a. Leaflets 11-14, with pale brown stellate scales few to numerous on the midrib be- low, reticulation subprominent above and below ......... 67. A. laxiflora b. Leaflets 3-11, with pale yellowish-brown stellate hairs absent to numerous on the midrib below, reticulation visible on the upper surface, subprominent below Be bene Aira sane Sore eae RRS ODEN apr ey oh oy Meant reve A A 70. A. oligophylla 97a. Petals 3, densely covered with stellate scales on the outside; fruits dehiscent .. . We as aiosd SRE RA RINSE NS S115 AA ea ae eee eae 9. A. lepidopetala b. Petals 5, without scales on the outside, fruits indehiscent .............. 98 Osa ainecmnbranchedleatletsishinyoy). ese. a eee - 69. A. coriacea b-yiiree' branched; leaflets not: shinyaaa asa wee aca ene eae eee Be) 99a. Fruit c. 0.5 cm in diameter, with few stellate scales ...... 64. A. aherniana b. Ripe fruit 1 cm or more in diameter, with dense indumentum ........... 100 lOOatbeatletapexwith’ayparallel-sidediacumen: = 53... 26 as.aci.a ee eee 101 bvekKeatlenapexwithratapening acumen 11s | Sess Aaah ee ee 102 LOkapWeatletsrconmaceoust ways. uc baeed aa ee ere et 42. A. forbesii beleallets NOt iComaccOus. 6 %-. hee 5. R. celebicum bs akeaves pinmates3 nu ciemcden iis. Ste bo Bade See a se 2 2a. Secondary veins almost as prominent as primary, at least on adaxial surface ..... denne peck tater): Brtrdayhcs. seg? Biante. dc ote ed ae 4. R. humile b., Secondary veins, notso prominent <2 4. Sy. sad. ee: = st See eee 3 3a. Adaxial surface of leaflet midrib brown tomentose .......... 1. R. cinereum be Adaxial surface of-leafletssglabrous «.....53.0.2...%-. Jee. 120-2. 7 eee 4 4a. Peduncles pilose. Domatia frequently present ........... 2. R. kinabaluense b. Peduncles glabrous. Domatia unrecorded .............. 3. R. kostermansii 1. Reinwardtiodendron cinereum (Hiern) Mabb. Reinwardtiodendron cinereum (Hiern) Mabb., Malays. For. 45 (1982) 452; Blumea 31 (1985) 144; in Tree Fl. Malaya 4 (1989) 247, f. 7A. — Lansium cinereum Hiern in Hook. f., Fl. Brit: India 1 (1875)) 558; C.DC. in DC., Monogr. Phan. | (1878) 597; King, J. As. Soc. Beng. 64, ii (1895) 81; Ridley, Fl. Malay Penins. | (1922) 411. — Aglaia pseudolansium Kosterm., Rein- wardtia 7 (1966) 252, t. 6. Tree to 27 m, to 70 cm diam. and clear bole to 18 m. Buttresses to 1 m tall, extending out to 3 m, snake-like at extremities. Bark smooth with scaly patches and conspicuous knobbly tuber- cles; inner bark white; sapwood yellow. Twigs rather angular, with conspicuous leafscars and dense indumentum when young, aluminium grey later. Leaves 10-15 cm long; petiole 1.5—2 cm, peti- ole and rachis densely brown tomentose. Leaflets 2—4 (5) on each side, 4—5.5 (—8) by 1.5—2.5(—3.5) cm, the most Fig. 47. Reinwardtiodendron cinereum (Hiern) Mabb. Fluted bole. West Malaysia, Pahang. Photograph D. J. Mabberley, July 1981. oS) i) n Mabberley et al. — Meliaceae apical larger, up to 11.5 by 4.5 cm, elliptical, chartaceous to subcoriaceous, very sparse- ly pubescent abaxially, midrib bristly brown pilose on both surfaces, major veins thus abaxially; apex acuminate, acumen to 6—8 mm long; base weakly asymmetric, cuneate; major veins c. 6—8 on each side, arcuate, with pilose domatia in their axils, secondary veins less conspicuous; petiolules c. 5 mm, densely brown tomentose, weakly swollen at base. Spikes to 10 cm; peduncles densely pilose; bracts to 4 mm long, acute, often with smaller bracteoles. Calyx lobes c. | mm long, fawn sericeous, the margin ciliate. Petals 4 mm long, ovate. Fruit 1.5—2 cm diam., globose, glabrous, yellow. — Fig. 46, 47. Distribution — W Malesia: Sumatra, Malay Peninsula, and northern Borneo. Habitat — Rain forest to 700 m altitude. Rarely collected. Uses — The sarcotesta is edible. Note — The leaflets are very similar to those of Aglaia oligophylla (q.v.) but are more or less opposite and have no stellate indumentum. 2. Reinwardtiodendron kinabaluense (Kosterm.) Mabb. Reinwardtiodendron kinabaluense (Kosterm.) Mabb., Blumea 31 (1985) 145. — Aglaia kinabaluensis Kosterm., Reinwardtia 7 (1966) 253, t. 7. Tree to 20 m, to 25 cm diam. and clear buttressed bole to 15 m. Bark smooth; inner bark yellow to white; sapwood yellow. Leafy twigs soon glabrous, adpressed pilose at apices. Leaves to 27 cm; petiole 4—5 cm, sparsely pubescent to glabrous. Leaflets | or 2 on each side, the leaves thus largely trifoliolate, 8—15 by 3—6.5 cm, the most apical usually the largest (to 18 by 7 cm), oblong-elliptic, chartaceous to subcoriaceous, gla- brous; apex acuminate, acumen to 18 mm long; base cuneate; major veins c. 7 on each side, arcuate, frequently with small domatia in their axils, secondary veins less con- spicuous; petiolules c. 5 mm, sulcate, swollen at base. Spikes to 8 cm long or basally branched panicles with 2 or 3 branches to 8 cm long; peduncles more or less pilose especially in high altitude specimens; bracts | mm long, triangular, pilose. Calyx lobes c. 1.5 mm long, more or less pilose, margin ciliate. Petals 4-5 mm long, ovate. Fruits to 5 cm long, depressed-globose when young, obovoid with apical depression and 5 ribs when mature, sericeous, yellow. Distribution — W Malesia: northern Borneo. Habitat — Rain forest to 900 m altitude. Rarely collected. Note — This species is remarkably similar in overall facies to Aglaia oligophylla Mig. though the leaflets are larger in that species and the indumentum of stellate hairs, best seen in the fruit, distinguishes it at once. Furthermore, domatia are absent. 3. Reinwardtiodendron kostermansii (Prijanto) Mabb. Reinwardtiodendron kostermansii (Prijanto) Mabb., Blumea 31 (1985) 145. — Lansium kostermansii Prijanto, Reinwardtia 8 (1965) 63, t. la, b. — Aglaia kostermansii (Prijanto) Kosterm., Reinwardtia 8 (1966) 256, p.p., t. 9. 326 Flora Malesiana ser. I, Vol. 12 (1) (1995) Tree to 30 m, 40 cm diam. Bole fluted at base. Bark smooth, yellowish; inner bark yellowish. Leafy twigs soon glabrous, adpressed pilose at apices. Leaves 15—20 cm long; petiole 3—4.5 cm long, very sparsely and minutely puberulous. Leaflets 1-3 on each side, the leaves mostly 3- or 5-foliolate, 6.5—11 by 3—5 cm, oblong-elliptic, char- taceous to subcoriaceous, glabrous except for minutely puberulent abaxial surface of lamina and especially midribs and petiolules; apex acuminate, acumen to 9 mm, base acute to cuneate; major veins 7—10 on each side, arcuate; secondary veins less promi- nent; petiolule c. 5 mm, somewhat channelled when dry, weakly swollen at base. Jn- florescences spikes 11—18 cm long, axillary to slightly supra-axillary; rachis glabrous; bracts c. 0-5 mm long, triangular, pubescent. Calyx lobes c. 1 mm long, orbicular, glabrous, margin weakly ciliate. Petals c. 3.5 mm long, ovate. Fruits to 3 cm diam., globose, weakly dimpled at apex, obscurely 5-ribbed, velutinous, yellow. Seeds (fide Kostermans) 1—3 developing, 1.5—2.5 cm long, 1.5—2 cm diam. Distribution — E Malesia: W Sumbawa and W Flores. Habitat — Lowland to submontane rain forest, 200—500 m altitude; common in relict patches of valley forest. Vernacular names — Kaju (= tree) narab (Sumbawa); garu (Flores). Uses — The sarcotesta is edible. 4. Reinwardtiodendron humile (Hassk.) Mabb. Reinwardtiodendron humile (Hassk.) Mabb., Mal. For. 45 (1982) 452: Blumea 31 (1985) 145; in Tree Fl. Malaya 4 (1989) 249, f. 7B. — Lansium humile Hassk., Retzia, ed. nov. 1 (1858) 121; Miq., FI. Ind. Bat. 1, 2 (1859) 545; Suppl. (1861) 197; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 34; C.DC. in DC., Monogr. Phan. | (1878) 598; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 183; Hochr., Pl. Bogor. Exsicc. (1904) 74; Backer, Schoolfl. Java (1911) 215: Koord.-Schum., Syst. Verz. 1 (1911) 30: Koord., Exk. Fl. Java 2 (1912) 443; Backer & Bakh. f., Fl. Java 2 (1965) 125. — Apha- namixis humilis (Hassk.) Kosterm., Reinwardtia 7 (1966) 263 (‘humile’); Backer & Bakh. f., FI. Java 3 (1968) 654. Lansium dubium Merr., Philipp. Govt. Lab. Bur. Bull. 17 (1904) 23; Brown, Bull. Dept. Agr. Bur. For. Philipp. 21 (1920) 82, t. 40; ibid. 22 (1921) 304, t. 40; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 368; Webster, Bull. Dept. Agr. Bur. Agr. Philipp. 39 (1924) 125; Elmer, Leafl. Philipp. Bot. 9 (1937) 3384: Li, J. Arnold Arbor. 25 (1944) 208: How & Chen, Acta Phytotax. Sin. 4 (1955) 27; Anon., Fl. Hainan 3 (1974) 67, t. 569. — Aglaia dubia (Merr.) Kosterm., Reinwardtia 7 (1966) 254: Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 335. — Reinwardtiodendron dubium (Merr.) X.M. Chen, J. Wuhan Bot. Res. 4 (1986) 183. Lansium sp.: Merr., Lingn. Sc. J. 5 (1927) 104. Tree to 27 m, to 40 cm diam. Buttresses to 4 m tall, at apex running into fluted bole, at base running out to 5 m from tree (Kostermans). Bark smooth with scaly patches, fawn; inner bark yellow to reddish brown; sapwood yellowish. Twigs weakly cicatrose and subglabrous except minutely pubescent apices, drying dark brown. Leaves 15—20 cm long; petiole 2—4 cm long. Leaflets 2 or 3 on each side, (6—)8.5—10 by 1.8—3.5 cm, the most apical larger, up to 18 by 6.5 cm, elliptical to elliptical-obovate, chartace- ous to subcoriaceous, glossy on both surfaces, + glabrous; apex markedly acuminate, acumen to 1.5 cm long; base weakly asymmetric, cuneate; major veins not clearly dis- tinct from secondary ones, particularly adaxially, together some 20—30 on each side, Mabberley et al. — Meliaceae 327 parallel, straight, at about 60° to midrib, anastomosing at margin, prominent on both surfaces, rarely with domatia in axils adaxially; petiolules c. 4-6 mm long, weakly swollen and channelled. Spikes or panicles to 15 cm; peduncles very sparsely pubes- cent; bracts 2 mm long, acute, usually with a pair of smaller bracteoles, all sparsely pubescent. Calyx lobes 1-2 mm long, orbicular, margin ciliate. Petals to 2.5 mm long, Fig. 48. Reinwardtiodendron humile (Hassk.) Mabb. a. Flowering shoot; b. half flower; c. fruiting shoot. Drawing R. Wise. Reproduced (rearranged) with permission from Tree Flora of Malaya 4. 328 Flora Malesiana ser. I, Vol. 12 (1) (1995) yellow, ovate to obovate. Fruit to 5 cm diam., + fig-shaped or obovoid, especially when immature, with apical depression and five sutures, densely but very shortly pi- lose, yellow. — Fig. 48. Distribution — Hainan, Indochina; Malesia: Sumatra, Malay Peninsula, Borneo, Java, Philippines and SE Celebes. Habitat — Rain forest to 800 m altitude. Vernacular names — Tembangan, tipis kulit (Java); bintangor (Borneo); aragnan, ma- lakanasi (fide Merrill), uban-uban (Bik.), bulahan (P. Bis.), tulanan (S.L. Bis.), bisik, buahan (Sul.), official common name: lansones-bundok (Philippines). Note — The commonest and most variable species. 5. Reinwardtiodendron celebicum Koord. Reinwardtiodendron celebicum Koord., Minah.(1898) 389; Suppl. Fl. NO. Celebes | (1918) 23, t. 8a—b; Koord.-Schum., Syst. Verz. 3 (1914) 63; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 369; Steenis, Nova Guinea, n.s. 10 (1959) 210, t. 2; Mabb., Blumea 31 (1985) 145. — Aglaia reinwardtiana Kosterm., Reinwardtia 7 (1966) 230, t. 1. Reinwardtiodendron merrillii Perkins, Fragm. Fl. Philipp. (1904) 74; Merr., Philipp. J. Sc. 1, Suppl. (1906) 72; Elmer, Leafl. Philipp. Bot. 9 (1937) 3385. [Lansium monophyllum Merr. ex Perkins, Fragm. Fl. Philipp. (1904) 75, nom. in syn.] Tree to 25 m, to 40 cm diam., fluted. Buttresses to 1.75 m tall, extending out to 1.5 m. Bark smooth to scaling, light brown, thin; inner bark white; sapwood yellow; heart- wood yellow-brown. Leaves unifoliolate, 14—25.5 by 5—8 cm; elliptic or weakly ovate or obovate, subcoriaceous, almost glabrous; apex acuminate; base cuneate; major veins 5—9 on each side, arcuate usually with hollow setose domatia in axils, prominent like midrib abaxially, both sunken adaxially (in sicco); secondary venation reticulate, pro- minulous on both surfaces; petiole 1—4.5 cm, surmounted by swollen petiolule c. 3 mm. Spikes or basally branched panicles, sometimes on short shoots and superficially resem- bling larger panicles, to 12 cm long; peduncles subglabrous; bracts triangular, weakly pilose, c. | mm long; bracteoles c. 1 mm long, adpressed to calyx. Calyx lobes c. 1 mm long, + glabrous. Petals c. 3.5 mm long, ovate. Fruit to 3 cm diam., subglobose, den- sely shortly pilose with thin hard pericarp, without sutures, 1-seeded. Distribution — Malesia: E Borneo, Philippines, Celebes, Moluccas, W New Guinea. Habitat — Rain forest to 600 m altitude. Vernacular names — Balbisan, bianti, malakamanga (Philippines), sakogwa, tam (New Guinea). INSUFFICIENTLY KNOWN SPECIES One collection from northern Sumatra probably represents a sixth species in the region. See Mabberley, Blumea 31 (1985) 146. The flowers are unknown but the fruiting spec- imens differ from Reinwardtiodendron celebicum in having pinnate leaves, from R. ci- nereum and R. kinabaluense (and R. anaimalaiense) in having no domatia, from R. hu- mile in not having the very conspicuous secondary veins and from R. kostermansit, to which it seems most closely allied, in its conspicuously ribbed velutinous fruit. Mabberley et al. — Meliaceae 329 TRIBUS MELIEAE Melieae DC., Prodr. 1 (1824) 619; T.D. Penn., Blumea 22 (1975) 462. Trees. Indumentum of stellate and/or simple hairs. Leaves pinnate or bipinnate. Flow- ers hermaphrodite or male. Staminal tube cylindrical, lobed. Disk annular. Pistil with 3—8-lobed stylehead. Fruit a drupe. Seed exarillate; endosperm thin; cotyledons colla- teral; radicle superior, exserted. Two genera, in the wild state restricted to the Old World Tropics. MELIA Melia L., Sp. Pl. 1 (1753) 384; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 99, t. 26, 27; T.D. Penn., Blumea 22 (1975) 463, t. 4; Mabb., Gard. Bull. Sing. 37 (1984) 463. — [Zederachia Heist. ex Fabr., Enum. Meth. Pl. (1759) 221, nom. superfl. — Azedarac Adans., Fam. 2 (1763) 342, nom. superfl. — Azedara Raf., Fl. Ludov. (1817) 135, nom. superfl. — Azedaraca Raf., Med. Fl. 2 (1830) 199, nom. superfl.] Azedarach Mill., Gard. Dict., Abr. ed. 4 (1754) [170]. Antelaea Gaertn., Fruct. Sem. Pl. 1 (1788) 277. Trees, occasionally flowering precociously as shrublets. Jndumentum of simple and stellate-tufted hairs. Leaves 2(—3)-pinnate. Inflorescence thyrsoid, axillary. Flowers hermaphrodite and male on same tree (polygamous). Calyx 5- (or 6-)lobed to near base, lobes somewhat imbricate. Petals 5 (6), free, imbricate. Staminal tube narrowly cylin- drical, slightly expanded at mouth, 10- (or 12-)ribbed, with 10 or 12 truncate, bifid or 4-fid filiform lobes; anthers 10 (12), inserted at margin of or just within tube, alternat- ing with or opposite lobes. Disk small, surrounding base of ovary. Ovary 4—8-locular, each locule with 2 superposed ovules; stylehead capitate to coroniform with 4—8 short, erect or incurved stigmatic lobes. Drupe 3—8-locular; endocarp thick, bony, deeply dim- pled at base and apex; locules 1- (or 2-)seeded. Seed oblong, laterally compressed; testa leathery, sometimes slightly swollen and fleshy around hilum; embryo embedded in oily endosperm; cotyledons flat. Germination phanerocotylar; eophylls opposite, pinnatisect or trifoliolate. 2n = 28. Distribution — One species in Indomalesia with one or possibly two closely allied ones in south tropical Africa. Forms of the Indomalesian species are widely cultivated and naturalized throughout the warm parts of the world. Habitat — Forests of various types, the cultivated species also escaping in open sec- ondary vegetation. Notes — 1. As noted by Corner [Wayside Trees | (1940) 464], the inflorescences of cultivated Melia azedarach in Malaya are short shoots with terminal buds. The true inflo- rescences are borne in the axils of rudimentary leaves and after fruit abscission, the ter- minal bud may grow out into a leafy shoot. Similar short shoots are known to bear the inflorescences in Chisocheton (q.v.) while in South American species of Guarea, nota- bly G. kunthiana A. Juss. [T.D. Penn., Fl. Neotrop. 28 (1981) 8], there being no sign of rudimentary leaves, the shoots are termed indeterminate inflorescences which grow 330 Flora Malesiana ser. I, Vol. 12 (1) (1995) for some months, bearing ripe fruits at the base and young flowers at the apex, as fig- ured by Pennington (op.cit. t. 59) for G. pubescens (Rich.) A. Juss. 2. For details of embryology, see Nair, J. Ind. Bot. Soc. 38 (1959) 367 et seq. Melia azedarach L. Melia azedarach L., Sp. Pl. (1753) 384, incl. var. sempervirens L.; Burm. f., Fl. Ind. (1767) 101; DC., Prodr. 1 (1824) 621; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 219, t. 13, f. 4; Blanco, FI. Filip. (1837) 345 (‘acedarach’); Hassk., Cat. Pl. Bogor. (1844) 219; M. Roem., Synops. Monogr. (1846) 94; Mig., Fl. Ind. Bat. 1, 2 (1859) 533; Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 24, incl. var. subtripinnata Miq.; ibid. 4 (1868) 5, incl. var. incisa Miq., var. acuminatissima Miq., var. sambu- cina (Blume) Miq.; Hiern in Hook. f., Fl. Brit. India 1 (1875) 544; C.DC. in DC., Monogr. Phan. 1 (1878) 451, incl. var. glabrior C.DC. (nom. superfl.), var. australasica (A. Juss.) C.DC., var. squamulosa C.DC.; Morren, Belg. Hort. 30 (1880) 176, cum tab., incl. var. floribunda (Carr.) Morren; Fern.-Vill., Nov. App. (1880) 41; Vidal, Sin. Atl. (1883) 20, t. 29 f. E; Bisschop Grev., Pl. Ned. Ind. (1883) 490; C.DC., Bull. Herb. Boiss. 2 (1894) 577, incl. var. squamulosa C.DC.; Sarg., Gard. For. 7 (1894) 92, t. 20, incl. var. umbraculifera Knox ex Sarg.; King, J. As. Soc. Beng. 64, ii (1895) 20 (‘azidarach’); Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) t. 160 f. A-L; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 12, incl. var. javanica Koord. & Valeton, var. squa- mulosa C.DC.; Pierre, Fl. For. Cochinch. 5 (1897) t. 356B, incl. var. glandulosa Pierre (‘Melia composita var. biglandulosa’ in ic.); Koord., Minah. (1898) 390; Bailey, Cycl. Amer. Hort. 2 (1900) 1001, t. 1387, incl. var. umbraculiformis Hort. ex Berck. & Bail.; Backer, Fl. Batavia | (1907) 267; Schoolfl. Java (1911) 202; Pellegr. in Fl. Indo-Chine 1 (1911) 727, t. 79, f. 1-6, incl. var. cochin- chinensis (Pierre) Pellegr., var. biglandulosa Pierre; Merr., Fl. Manila (1912) 276; Koord., Exk. FI. Java 2 (1912) 439; Mak., Bot. Mag. Tokyo 28 (1914) 34, incl. var. japonica (G. Don) Mak., nom. superfl., subvar. semperflorens (Mak.) Mak., subvar. toosendan (Sieb. & Zucc.) Mak., subvar. in- termedia Mak. f. albiflora Mak.; Ridley, Fl. Malay Penins. | (1922) 384; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 360; Mak., J. Jap. Bot. 5 (1928) 20, incl. var. toosendan (Sieb. & Zucc.) Mak., var. intermedia (Mak.) Mak.; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1441; Briq., Mém. Inst. Nat. Genev. 24 (1935) 41, t. 2, f. 11, 12; Elmer, Leafl. Philipp. Bot. 9 (1937) 3384; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) t. 26, f. A—L, t. 27; Corner, Wayside Trees | (1940) 464, 2 (1940) t. 137; Adelb., Blumea 6 (1948) 315; Rehd., Bibl. Cult. Trees Shrubs (1949) 387, incl. f. wnbraculifera (Sarg.) Rehd.; Steenis, Fl. Scholen Indon. (1949) 233; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 889; Quisumb., Med. Pl. Philipp. (1951) 482; Nair, J. Ind. Bot. Soc. 38 (1959) 367; Backer & Bakh. f., Fl. Java 2 (1965) 120; Corner & Watanabe, III. Guide Trop. Pl. (1969) 404, cum tab.; Vaughan, Struct. Util. Oil Seeds (1970) 157, t. 82 A—D; T.D. Penn., Blumea 22 (1975) 461, t. 4, f. d; Corner, Seeds Dicots 1 (1976) 190; 2 (1976) t. 387; Henty, Bot. Bull. Lae 12 (1980) 103, t. 36; Mabb., Gard. Bull. Sing. 37 (1984) 55; Corner, Wayside Trees, ed. 3 (1988) 502, t. 149; Du Puy in Fl. Austral. 50 (1993) 298. — Azederach deleteria Medik., Geschl. Malv. (1787) 116. — [Melia florida Salisb., Prodr. (1796) 317, nom. superfl.] — Azedara speciosa Raf., Fl. Ludov. (1817) 35. — Azedaraca amena Raf., Med. Fl. 2 (1830) 199. — Azedarach vuigaris Gomez de la Maza, Rep. Méd.-farm. Havana 5 (1894) 296. Melia commelini Medik., Bot. Beob. 1782 (1783) 164. [Azedarach commelini Moench ex Steud., Nom. Bot., ed. 2, 1 (1840) 175, 2 (1841) 118 (‘Medik.’), nom. in syn.] Antelaea javanica Gaertn., Fruct. Sem. Pl. 1 (1788) 277, t. 58. Melia sempervirens (L.) Sw., Prodr. Veg. Ind. Occ. (1788) 67; Hassk., Cat. Pl. Bogor. (1844) 219; M. Roem., Synops. Monogr. | (1846) 95; Backer, Fl. Batavia | (1907) 268; Schoolfl. Java (1911) 202; Koord., Exk. Fl. Java 2 (1912) 439; Adelb., Blumea 6 (1948) 315; Backer & Bakh. f., Fl. Java 2 (1965) 120. — Azedarach sempervirens (L.) Kuntze, Rev. Gen. Pl. 1 (1891) 109, incl. var. gla- brior (C.DC.) Kuntze, var. australasica (A. Juss.) Kuntze & var. dubia (Cav.) Kuntze. Melia dubia Cav., Diss. 7 (1789) 364, ?nom. provis.; M. Roem., Synops. Monogr. | (1846) 96; Hiern in Hook. f., Fl. Brit. India 1 (1875) 545; C.DC. in DC., Monogr. Phan. 1 (1878) 453; Fern.-Vill., Nov. App. (1880) 41; Backer, Schoolfl. Java (1911) 201; Merr., Enum. Philipp. Flow. Pl. 2 (1923) Mabberley et al. — Meliaceae 331 360; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1443; Briq., Mém. Inst. Nat. Genev. 24 (1935) 44; Elmer, Leafl. Philipp. Bot. 9 (1937) 3384; Adelb., Blumea 6 (1948) 315; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 889; Quisumb., Med. Pl. Philipp. (1951) 485; Backer & Bakh. f., Fl. Java 2 (1965) 120; Corner, Seeds Dicots | (1976) 191, 2 (1976) t. 388; Guzman et al., Guide Philipp. FI. Fauna 3 (1986) 341, t. 260. [Azederach odorata Noronha, Verh. Bat. Genoot., ed. 5, 1 (1791) art. 4, 5, nom. nud.] Melia composita Willd., Sp. Pl. 2 (1799) 559; DC., Prodr. 1 (1824) 622; Blanco, Fl. Filip., ed. 2 (1845) 241; M. Roem. Synops. Monogr. | (1846) 94; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 5; Blanco, Fl. Filip., ed. 3, 2 (1878) 84, t. 420; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 9; Pierre, Fl. For. Cochinch. 5 (1897) t. 356A, incl. var. cochinchinensis Pierre; Koord., Exk. Fl. Java 2 (1912) 439; Ridley, Fl. Malay Penins. | (1922) 384. Melia robusta Roxb., Hort. Beng. (1814) 33, nom. nud.; G. Don, Gen. Syst. 1 (1831) 680. Melia superba Roxb., Hort. Beng. (1814) 33, nom. nud.; Fl. Ind., ed. Carey, 2 (1832) 396. — [Melia argentea Hiern in Hook. f., Fl. Brit. India 1 (1875) 545, sphalm. pro Melia superba. } Melia arguta DC., Prodr. 1 (1824) 622; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 220; M. Roem., Synops. Monogr. | (1846) 96; Miq., Fl. Ind. Bat. 1, 2 (1859) 532; Ann. Mus. Bot. Lugd.- Bat. 4 (1868) 5; C.DC. in DC., Monogr. Phan. | (1878) 454. Melia sambucina Blume, Bijdr. (1825) 162; Hassk., Cat. Pl. Bogor. (1844) 219; M. Roem., Synops. Monogr. | (1846) 95; Miq., Fl. Ind. Bat. 1, 2 (1859) 533; C.DC. in DC., Monogr. Phan. | (1878) 455; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 17. — Azedarach sambucina (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 110. Melia angustifolia Schum. & Thonn. in Schum., Beskr. Guin. Pl. (1827) 214. Melia guineensis G. Don in Loud., Hort. Brit. (1830) 168; Gen. Syst. 1 (1831) 681. Melia australis Sweet, Hort. Brit., ed. 2 (1830) 85; G. Don, Gen. Syst. 1 (1831) 680. Melia candollei A. Juss., Bull. Sc. Nat. Géol. 23 (1830) 239: Linnaea 6, lit. (1831) 111; Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 220, 258; M. Roem., Synops. Monogr. 1 (1846) 95; Migq., Fl. Ind. Bat. 1, 2 (1859) 532; Ann. Mus. Bot. Lugd.-Bat. 4 (1868)5; C.DC. in DC., Monogr. Phan. | (1878) 456; Fern.-Vill., Nov. App. (1880) 41; Perkins. Fragm. Fl. Philipp. (1904) 30; Merr., Fl. Manila (1912) 276; Sp. Blanc. (1918) 209. Melia australasica A. Juss., Bull. Sc. Nat. Géol. 23 (1830) 239; Linnaea 6, lit. (1831) 111; Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 220, 258. Melia japonica G. Don, Gen. Syst. 1 (1831) 680; C.DC. in DC., Monogr. Phan. 1 (1878) 457, incl. var. albicans C.DC.; Mak., Bot. Mag. Tokyo 18 (1904) 67, incl. var. semperflorens Mak. [Melia bukayun Royle, Ill. Bot. Himal. (1835) 141, nom. nud.] [Melia flaccida Zipp. ex Span., Linnaea 15 (1841) 182, nom. in syn.] Melia toosendan Sieb. & Zucc., Abh. Akad. Miinchen 4, 2 (1843) 159. Melia japonica Hassk., Cat. Pl. Bogor. (1844) 219, nom. illeg., non G. Don. — Melia hasskarlii K. Koch, Hort. Dendrol. (1853) 72. Melia javanica M. Roem., Synops. Monogr. | (1846) 96. Melia cochinchinensis M. Roem., op. cit. (1846) 95. Melia orientalis M. Roem., |.c. Melia tomentosa auct. non Roxb.: Migq., Fl. Ind. Bat. 1, 2 (1859) 532. [Melia chinensis Sieb. ex Miq., Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 23, nom. in syn.] Melia birmanica Kurz, J. As. Soc. Beng. 43, ii (1874) 183; C.DC. in DC., Monogr. Phan. | (1878) 458; Bull. Herb. Boiss. 2 (1894) 577. Melia bogoriensis Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 18; Hochr., Pl. Bogor. Exsicc. (1904) 74; Koord., Exk. Fl. Java 3 (1912) 359; Atlas 1 (1913) t. 81; Briq., Mém. Inst. Nat. Genev. 24 (1935) 43. [Zizipha candida Monspelliensium Lobel., Stirp. (1576) 546, cum tab.; Icones 2 (1581) 108, cum tab.] [Pseudosycomorus Camer., Pl. Epit. Util. Matth. (1586) 181, tab.; Besler, Hort. Eyst. 2 (1613) t. 2.) [Azedarach Dod., Stirp. Hist. Pempt. (1616) 848; Tourn., Inst. Rei Herb. (1700) 616; Burm., Thes. Zey]. (1737) 40.] [Arbor fraxinifolio flore caeruleo Bauhin, Pinax (1623) 415; Herm., Mus. Zeyl. (1717) 67.] [Arbor Azadirachta Syriaca, foliis ramosis, flore caeruleo & albo majore Breyne, Prod. Rar. Pl. 1 (1680) 3.) Lo Wo i) Flora Malesiana ser. I, Vol. 12 (1) (1995) [Azedarach floribus albis sempervirens Herm., Hort. Acad. Lugd.-Bat. Cat. (1687) 652, excl. syn. (= Aza- dirachta indica).] [Azedarach Indica foliis non deciduis, ossiculo polypyreno Herm., Parad. Bat. Prodr. (1689) 316.] [Azadirachta foliis ramosis minoribus Indica, flore albo subcaeruleo purpurascente majore Breyne, Prodr. Rar. Pl. 2 (1689) 21; Comm., Hort. Med. Amstel. (1697) 147, t. 76.] [Azadirachta foliis ramosis majoribus, Syriaca, sive vulgaris, flore caeruleo majore Breyne, 1.c.] [Azedarach sempervirens & florens Tourn., Inst. Rei Herb. (1700) 616, t. 387.] [Azedarach fructu polypyreno Herm., Mus. Zeyl. (1717) 3; Burm., Thes. Zeyl. (1737) 40.] [Melia foliis decompositis L., Hort. Cliff. (1738) 161; Royen, Fl. Leyd. Prodr. (1740) 462.] [Melia foliis duplicato-pinnatis L., Fl. Zeyl. (1747) 162.) Tree to 40 m, + deciduous; bole fluted below when old, to 60(—180, Elmer) cm diam. Bark grey-brown, smooth, lenticellate becoming lightly fissured or scaling with age; inner bark yellowish; sapwood whitish, soft; heartwood rusty brown. Crown of widely spread but sparsely branched limbs. Twigs upturned at ends of drooping branchlets, smooth, brown, lenticellate, with raised cicatrices. Leafy twigs c. 6—8 mm diam., + cloth- ed with fulvous stellate hairs. Leaves 15—80 cm, with 3-7 pairs of lateral rachides, each with 3-7 pairs of leaflets, the most proximal of which sometimes replaced with short rachides with a few pairs of leaflets, + weakly pubescent but usually subglabrous; peti- ole 8—30 cm, to 6 mm diam., terete, lenticellate, swollen at base; lateral rachides weakly ascendant, to 25 cm long, articulated with articulated main rachis and weakly swollen there. Leaflets 3—6(—10) by 1—2.5(—3) cm, ovate or oblong-lanceolate to elliptic, base acute to rounded, apex acuminate, margin entire to variously serrate, costae c. 7-10 on each side, subsquarrose to weakly ascendant and arcuate, looped at margin; petiolules 3-7 mm. Thyrses 10—22 cm, axillary or (see note | under the genus) on short shoots and in axils of rudimentary leaves thereon, primary branches c. 5—7.5 cm, weakly as- cendant, secondary to 2 cm, bearing fascicles of scented flowers; axes + mealy pubes- cent; bracts 3—10 mm long, filiform, pubescent caducous, bracteoles similar but smaller; pedicels c. 2-3 mm. Calyx c. 2 mm diam.; lobes c. 2 mm long, ovate, stellate- and simple-hairy without, margin ciliate. Petals 6-10 by c. 2 mm, narrowly oblong, white to lilac or bluish, stellate- and simple-hairy without, sometimes simple-hairy within, mid-vein conspicuous. Staminal tube subglabrous without, + densely simple hairy within, lobes bifid, or 4-fid, sometimes irregularly so; anthers c. 1.5 mm long, apicu- late, + hairy, inserted opposite lobes. Disk obscure and closely enveloping ovary. Pistil glabrous; stylehead c. 0.7 mm diam. Drupe 2—4 cm long, 1—2 cm diam., plum-shaped, glabrous, yellow-brown when ripe; endocarp very hard. Seed c. 3.5 by 1.6 mm, oblong, smooth, brown. — Fig. 49. Distribution — Wild trees are known from India and Nepal, Sri Lanka and tropical China, south and east through Malesia: Sumatra, Java, Philippines (Luzon, Negros, Mindanao), Lesser Sunda Islands (Flores, Timor, Wetar), New Guinea, to tropical Australia and the Solomon Islands. Habitat — Forests, particularly seasonal ones including bamboo thickets (Thailand) and those on limestone, Tamarindus woodland and Eucalyptus savanna, where it may coppice, to 1200 m altitude (to 1800 m in Himalayan tract). Cultivated forms persist and may become naturalized in secondary vegetation after clearance. Mabberley et al. — Meliaceae 333 Fig. 49. Melia azedarach L., wild form. a. Habit with inflorescence; a}. leaflet; a2. leaf; b. half flower; c. top of staminal tube; d. infructescence; e. fruit, cross section; f. seed (a—c, aj: de Wilde & de Wilde- Duyfjes 15884; a2, d—f: Metzner 9]). Drawing R. van Crevel. Taxonomy — The species is a complex of wild and cultivated forms discussed by Mabberley [Gard. Bull. Sing. 37 (1984) 53] of which the following is a condensation. Forms of this widespread species have been cultivated for over 2500 years and some of them look very different from the statuesque wild trees with pale flowers and mealy 334 Flora Malesiana ser. I, Vol. 12 (1) (1995) shoots. Plants resembling the wild forms were cultivated at Calcutta in the early 1800s and are grown under ‘stove’ conditions in Europe; they are rarely planted in the tropics, though they have been long established at Bogor in Java. By contrast, the widespread Persian Lilac 1s grown all over the warmer parts of the world and some forms of it can even be grown outside in the British Isles. Such forms are deciduous, almost glabrous and have rather large bluish or lilac, rarely white flowers. The plant was apparently introduced to Malesia by Europeans and, although now wide- spread, was not mentioned by Rumphius for example. The hardy forms reached Europe via the Persians and Arabs, who knew its medicinal properties but the species seems to have been domesticated long ago in India, where it is still grown for its flowers which are widely used in thank-offerings. The Arabs knew it by the time of Avicenna (sowing methods being discussed in Ibn Bassal’s Book of Agriculture [V, sect. 51] of c. 1080) and it was first mentioned in a European text by I’Obel (1576). About this time it seems to have been cultivated in Europe, where, particularly in the Mediterranean, it was soon widely planted and even naturalized. That the dried fruit had a natural channel through it, preadapted the tree for use in rosary-making, such that in Spain it became known as Arbor sancta. There was at least one more major introduction to Europe, from Sri Lanka, of these Indian cultivars. That form, named by Linné var. sempervirens, was being grown in the Netherlands at the end of the 1600s, and was noted for its being more evergreen and more tender than the old Persian Lilac. This seems to have been the form introduced into the West Indies, but it is very close to the original and it is not absolutely clear that it is the West Indian plant. Nevertheless the West Indian plant was taken thence to Africa and reintroduced to Europe as an African plant! Forms of these Indian cultivars with precocious flowering were selected and named as were ones with white flowers of deeply lobed leaflets some of which seem to have arisen in Java in the nineteenth century. Meantime, the Chinese seem to have been using the tree for patten making in Kwangtung and to have selected certain forms with large fruits and leaflets, with rather entire margins. These, like so many Chinese garden plants, were taken to Japan, where they were known as TO sendan, that is, Chinese Melia, but, again like many Chinese garden plants, Europeans named them ‘japonica’ (twice!). By the 1860s, these Chinese cultivars were well-established in cultivation in Europe and also in tropical botanic gardens, notably Bogor. Because these forms are different from the Indian ones and yet both seem to have been derived from populations of the wild tree, the synonyms relating to the two groups of cultivars are set out below. The very nature of these cultivars denies one hundred percent ‘pigeon-holing’ of specimens, however. ‘Wild’ trees Melia azedarach p.p., i.e. var. australasica, var. cochinchinensis, var. glandulosa, var. javanica, vat. squamulosa — ?Antelaea javanica — Melia dubia — Melia composita — Melia robusta — Melia superba — Melia australis — Melia candollei — Melia australasica — Melia azedarach sensu Blanco — Melia flaccida — Melia tomentosa sensu Mig. — Melia birmanica — Melia argentea — Azedarach sempervirens var. australasica & var. dubia — Melia bogoriensis. Mabberley et al. — Meliaceae 335 Tall forest trees to 40 m. Leaflets + entire, dark above, pale below, when young usu- ally densely stellate tomentose like young shoots. Flowers sweetly scented, scentless or malodorous (Roxburgh II.cc.). Petals white or pale mauve, often pubescent within. Sta- minal tube creamish or pale mauve darkening to purple with age. Drupe up to 4 cm long. Vernacular names — Marambung (Sumatra); gringging, mindi besar, mindi gedek (Java); mera (Flores); kamé, kamél (Timor); bagaluga (Visayan, Philippines). For fur- ther names see Merrill, Enum. Philipp. Flow. Pl. 2 (1923) 360. Note — This is the white cedar of commerce and it has been tried in plantations no- tably in the Philippines. Roxburgh (ll.cc.) reported that trees in the Calcutta Botanic Garden grew to some 12-15 m in 6 years and had boles over a metre in girth. Chinese cultivars Melia azedarach p.p., i.e. var. intermedia, var. japonica, var. toosendan, subvar. intermedia, subvar. sem- perflorens, subvar. toosendan, f. albiflora — Melia japonica G.Don — Melia toosendan — Melia japonica Hassk. — Melia javanica — Melia hasskarlii — Melia chinensis — cv. Toosendan: H6, Illus. FI. S. Viétnam 2, 1 (1992) 487. Small trees. Leaflets usually almost entire. Flowers sweetly scented. Petals mauve, pink or blue, rarely white. Staminal tube purple. Distribution — Cultivated. Rarely grown outside botanic gardens in Malesia. Note — It is the form with fruits larger than those of the Indian cultivars. Indian cultivars Melia azedarach s.s., incl. var. acuminatissima, var. floribunda, vat. glabrior, var. incisa, var. sambu- cina, var. sempervirens, var. subtripinnata, var. umbraculifera, var. umbraculiformis f. umbraculifera — Azederach deleteria — Melia sempervirens — Melia commelini — Azederach odorata — Melia florida — Azedara speciosa — Melia arguta — Melia sambucina — Melia angustifolia — Azedaraca amena — Melia guineensis — Melia bukayun — Azedarach commelini — Melia cochinchinensis — Melia orientalis — Melia floribunda — Azedarach sempervirens incl. var. glabrior — Melia sam- bucina — Azedarach vulgaris. Small trees. Leaflets irregularly serrate, pale green. Flowers sweetly scented. Petals mauve, pink or blue, rarely white. Staminal tube purple. Distribution — Cultivated throughout the warmer parts of the world, including Male- sia, where sometimes apparently naturalized. Two named cultivars are striking: ‘Flori- bunda’ (M. floribunda, M. azedarach var. floribunda) which flowers when a few dm tall and is widely used in bedding schemes, and ‘Umbraculifera’ (M. azedarach var. um- braculifera, f. umbraculifera, var. umbraculiformis, the Texas Umbrella Tree) which has a flattened crown of branches and arose as a mutant in Texas in the last century. The common form is the Persian Lilac, bead tree, Pride of India, Philippine Lilac, mindi kechil (Mal.), paraiso (Philippines). Uses — Persian Lilac is one of the most widely cultivated of all tropical trees and is so readily grown as to be despised by the discriminating. Good forms with dark heavily scented flowers are extremely desirable nevertheless. The almost continuous flowering of the species in some areas ensures an unbroken supply of the flowers and of the fruits 336 Flora Malesiana ser. I, Vol. 12 (1) (1995) which have been long used for beads [see Hoy & Catling, Davidsonia 12 (1981) 65—66 for illustration]. The fruits are toxic to man, some 6-8 considered a fatal dose, and also to pigs but apparently not to birds, sheep or goats; cows fed on the leaves yield tainted milk. The toxic principles are the limonoids, meliatoxins Al, A2, B1, B2 [Oerlichs et al., Phyto- chemistry 22 (1983) 531]. The wood of different forms of the species has been used for furniture and light con- struction, notably for ceilings, boats and tea-boxes, while, during the American Civil War, the trees were a commercial source of alcohol (10% by weight from the fruits). In tropical America it has been grown in plantations for the production of fibreboard [Pen- nington, Flora Neotropica 28 (1981) 24] and is important in the sports goods industries of Pakistan [Amjad & Mohammad, Pak. J. For. 1 (1980) 39]. It has been used as a fast-growing coffee-shade and it has been alleged that fruit trees grown under it remain relatively free from aphids. Indeed a decoction of the fruits has long been used as an insecticide for plants in India and fruits or leaves have been placed with dry fruit, clothing and in books to keep insects away. An ester of 2,3,16,20-tetra- hydroxypregnane (a C21-steroid), azedarachol, isolated from root-bark showed antifee- dant activity against the insect pest Ajrotis sejetum in Japan [Nakatani et al., Phytochem- istry 24 (1985) 1945]. In China, the seed-oil has been found to be quite effective as an antifeedant to yellow rice-borers, but not striped nor pink ones, effective against white- backed and brown plant-hoppers but not leaf-hoppers nor rice thrips [Hu et al., J. Agric. Res. China 1983 (1983) 63]. The oil is also effective in control of certain citrus pests, while powdered ripe seeds or lives are mixed with stored grain to preserve it from insect attack [W. & H. Olkowski, IPM Pract. 10, 9 (1988) 1]. Acetone extracts of the fruits are larvicidal but have no effect on the pupae of mosquitoes [Z. Al-Sharook et al., J. Appl. Entom. 111 (1991) 425]. Only trace amounts of the insecticidal salannin (see Azadirachta below) have been found in cultivars grown in the USA [Yamasaki et al., J. Chromatogr. 447 (1988) 277]. An extract has also been used as a fish poison. Although there has been much confusion with neem, Azadirachta indica A. Juss., Melia azedarach has been widely attributed with medicinal value. The root appears as Cortex Meliae azedarach in pharmacopeiae but it is generally held that the bark is most efficacious, particularly as a vermifuge. For details of its action and other medicinal uses, see Perry, Med. Pl. E & SE Asia (1980) 262. So valued are the fruits in the Malay Peninsula that they have been imported from Szechwan. A glycopeptide called meliacin (not the limonoid with that name), isolated from the leaves and roots is respon- sible for inhibition of in-vitro replication of various DNA and RNA viruses, e.g. polio- virus, herpes simplex Type I, and prevents development of encephalitis by Tacaribe virus in neonatal mice [Andrei et al., Antiviral Res. 9 (1988) 221]. It also triggers an antiviral state in cells prior to infection, but, although the mechanism is not like that in interferon, it is otherwise not understood. The seeds also yield an oil and the trunk a gum but these have been little utilized, though the first is suitable for soap and hair oil [Wealth of India 6 (1962) 323 & seqq., q.v. for further uses of the tree]. Mabberley et al. — Meliaceae 337 AZADIRACHTA Azadirachta A. Juss., Bull. Sc. Nat. Géol. 23 (1830) 236; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 102; Jacobs, Gard. Bull. Sing. 18 (1961) 71; T.D. Penn., Blumea 22 (1975) 464. Antelaea auct. non Gaertn.: Adelb., Blumea 6 (1948) 315. Trees. /ndumentum of simple hairs. Buds thinly encrusted with resin. Leaves pinnate with 2 pairs of glands at base of petiole. Flowers bisexual and male on same individual (polygamous). Calyx 5-lobed to proximal half, the lobes imbricate. Petals 5, free, im- bricate. Staminal tube cylindrical, slightly expanded at mouth, margin (8—)10-lobed, the lobes rounded, truncate, emarginate or bifid; anthers (8—)10, glabrous, inserted at base of and opposite lobes. Disk annular, united with base of ovary. Ovary 3-locular, each locule with 2 collateral ovules; stylehead with apical swollen torus with 3 acute, partially united papiliose stigmatic lobes. Fruit a 1- (or 2-)seeded drupe; endocarp thin, cartilagi- nous. Seed ovoid, distally pointed; testa thin, membraneous with small adaxial sarcotesta; cotyledons plano-convex, collateral; radicle superior, short projecting from cotyledons. Germination phanerocotylar; eophylls opposite, trifoliolate, leaflets deeply incised or pinnatifid. 2n = 28, 30. Distribution — Two species native to Indomalesia, of which one in Malesia proper. The other species, A. indica, the neem, is widely cultivated in warm countries through- out the world and is naturalized in some of them. Habitat — Lowland forest of various types, A. indica also colonizing deforested land. Note — Formerly confused with Melia, Azadirachta differs in its simple indumen- tum, pinnate leaves, collateral ovules, 3-lobed stylehead and 1- (or 2-)seeded drupes. Jacobs (1.c.) points out that sterile specimens may be separated in that in Melia there is a pair of orbicular glands at the base of the petiole, whereas in Azadirachta, there is, in addition, a pair of linear glands below them. Pennington (I.c.) notes that this is most readily seen in living material, where the leaf shape and indumentum are readily observ- ed anyway. The wood is readily distinguished in that that of Melia is ring-porous. KEY TO THE SPECIES Prmteatlets With CHLiTeMNAToIN 2.20 . huh Nees Slee b See eee 1. A. excelsa PECTS SEALS VA Pos. < 3.4-x ios, eae ae Ameena. Se 2. A. indica 1. Azadirachta excelsa (Jack) Jacobs Azadirachta excelsa (Jack) Jacobs, Gard. Bull. Sing. 18 (1961) 75; Wong, Mal. For. Rec. 28 (1976) 81, cum tab.; Mabb. in Tree Fl. Malaya 4 (1989) 233, f. 4. — Melia excelsa Jack, Mal. Misc. 11 (1820) 12; Corner, Gard. Bull. Str. Settl. 10 (1939) 263, t. 1, 2; Wayside Trees 1 (1940) 465, t. 155 & 2 (1940) t. 138; ibid., ed. 3 (1988) 504, t. 150; Merr., J. Arnold Arbor. 33 (1952) 235; Corner & Watanabe, Ill. Guide Trop. Pl. (1969) 404, cum tab. — Trichilia excelsa (Jack) Spreng., Syst. 4, 2 (1827) 252, excl. syn. Blume. — Azedarach excelsa (Jack) Kuntze, Rev. Gen. Pl. 1 (1891) 110. Azadirachta integrifoliola Merr., Philipp. J. Sc., Bot. 4 (1909) 272; Enum. Philipp. Flow. Pl. 2 (1923) 361; Elmer, Leafl. Philipp. Bot. 9 (1937) 3340. 338 Flora Malesiana ser. I, Vol. 12 (1) (1995) Rosemary AISLE. Fig. 50. Azadirachta excelsa (Jack) Jacobs. a. Flowering shoot; b. half flower; c. infructescences (a, b: Cuadra A 1278; c: Haniff 301). Drawing R. Wise. Reproduced with permission from Tree Flora of Malaya 4. Mabberley et al. — Meliaceae 339 Tree to 50 m; bole to 120 cm diam., regularly cylindric or rarely slightly buttressed over major roots. Bark smooth, pinkish grey or pinkish brown, in large trees becoming longitudinally fissured and scaling, the flakes oblong, greyish, breaking off at upper end and curling up from both ends before shedding, the bole appearing pale brownish or greyish buff and shaggy, sapwood white; heartwood light red. Crown rounded but rather open and uneven, deciduous for up to 3 months a year (Ng), the major branches ascending. Leafy twigs c. 8-12 mm diam., the pith pinkish but odourless or weakly onion scented. Young shoots puberulous, soon glabrous. Leaves 20—60(—90) cm, 7—11- jugate, pari- (or impari-)pinnate, tufted at ends of twigs; petiole 5—8 cm, + terete, swol- len at base; leaflets 4—12.5 by 2—3.5 cm, the largest near the middle, lanceolate-elliptic, asymmetric, + falcate, glabrous, pink when young, yellow when withering, bases un- equal, apices subacute to subacuminate, margin entire, costae c. 6—11 on each side, arcuate, tertiary venation laxly reticulate; petiolules c. 2 mm. Thyrses 20—45 cm, axil- lary, erect, sweetly scented; axes puberulous, green with 3 or 4 orders of branching, main proximal branches to 8 cm; bracts c. | mm long, narrowly triangular, adpressed; pedicels 1-3 mm, articulated with pseudopedicel of same length. Calyx c. 1 mm diam., puberulous without; lobes c. | mm long, rounded to subacute, pale green, margins ciliolate. Petals 5—6.5 by 1.5—2.2 mm, oblong-spathulate, puberulous without, pale creamy white. Staminal tube c.2—2.5 mm diam., glabrous without, sparsely hairy distally within, white or greenish, 10-ribbed, each rib terminating in a subbifid lobe; anthers (8—)10, c. 0.8 mm long, sessile, slightly exserted. Drupe 2.4—3.2 by 1.3—1.6 cm, ellipsoid, glabrous, green turning yellow at maturity; pericarp leathery; mesocarp soft, edible, with some white latex. Seed smelling of garlic when damaged. — Fig. 50:51: Distribution — Malesia: Sumatra (Bengkulu, East coast), Malay Peninsula (Penang, Perak, Selangor, Malacca, where also a village tree), Borneo (N & E), Philippines [Mindoro, Masbate, Samar, Palawan, Basilan (Elmer, |.c.)], Celebes, Moluccas (Aru Islands), W New Guinea. H6, Ill. Fl. Vietnam 2, | (1992) 488, records this tree from Vietnam. Habitat — Rain forest to 350 m altitude. Vernacular names — Kaju bawang, sentang (and variants), surian bawang (Mal.), bawang-bawang (Dugguh), ranggu (Iban, Dusun), turannggoh (Kwijan), danggo, maranggo (Tagb.); New Guinea: asrea (Kebar), bie (Mooi), dabiar (Biak), ibio (Amber- baken, Manikiong), marwa (Mejah), mekedjuf, numbwak (Sidei), nibwak, nubwak (Amberbaken). Uses — In the Malay Peninsula the timber is used in house-building and the young shoots are eaten as a vegetable (Corner, II.cc.). It coppices. The seeds yield azadirach- tin (see A. indica) and the more effective insect antifeedant, marrangin [Ermel et al., J. Appl. Entom. 112 (1991) 512]. Notes — 1. Though a conspicuous tree in the north of the Malay Peninsula and spe- cimens had been known for a long time, the identity of the tree was unknown or con- fused from the time of Jack until 1936 when Corner studied in Penang the trees which Jack himself may have known. Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 51. Azadirachta excelsa (Jack) Jacobs. Bole. West Malaysia, Selangor, Kepong. Photograph D.J. Mabberley, 1974. Mabberley et al. — Meliaceae 34] 2. Ata distance the tree may be confused with Ailanthus integrifolia Lam. (Simarou- baceae) but the bark of that tree seems always to be smooth and leaflets wither red and not yellow (Corner, Il.cc.). 2. Azadirachta indica A. Juss. Azadirachta indica A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 221, t. 13, f. 5; Miq., Fl. Ind. Bat. 1, 2 (1859) 533; Suppl. (1861) 502; C.DC. in DC., Monogr. Phan. 1 (1878) 459, t. 6, f. 10; Bisschop Grev., Pl. Ned. Ind. (1883) 491; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) t. 160, f. M—S; ibid., ed. 2, 19b1 (1940) t. 26, f. M—S; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 21; Prinsen Geerl., Teysmannia 13 (1902) 8, cum tab.; Valeton in Hochr., Pl. Bogor. Ex- sicc. (1904) 66, incl. var. minor Valeton & var. siamensis Valeton; Cat. Bogor. Nov. (1905) 21, 110, 131, incl. var. minor & var. siamensis; Backer, Fl. Batavia 1 (1907) 270; Schoolfl. Java (1911) 202; Koord., Exk. Fl. Java 2 (1912) 439; Atlas | (1913) t. 164; Brig., Mém. Inst. Nat. Genev. 24 (1935) 44, incl. var. minor & var. siamensis; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 889; Jacobs, Gard. Bull. Sing. 18 (1961) 74; Backer & Bakh. f., Fl. Java 2 (1965) 120; T.D. Penn., Blumea 22 (1975) 461, t. 4e; [Ruskin], Firew. Crops (1980) 114; Radwanski & Wickens, Econ. Bot. 35 (1981) 398; Ahmed & Grainge, Econ. Bot. 40 (1986) 201; Nicolson et al., Int. Hort. Malab. (1988) 190; Tewari, Monogr. Neem (1992). — Melia azadirachta L., Sp. Pl. (1753) 385; DC., Prodr. | (1824) 622; Blume, Bijdr. (1825) 161; Griff., Notul. 4 (1854) 500; Mig., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 5; Hiern in Hook. f., Fl. Brit. India 1 (1875) 544; Ridley, Fl. Malay Penins. | (1922) 384; Vaughan, Struct. Util. Oil Seeds (1970) 155, t. 82 E—F; Corner, Seeds Dicots 1 (1976) 191. — Antelaea azadirachta (L.) Adelb., Blumea 6 (1948) 315. — [Melia fraxinifolia Salisb., Prodr. (1796) 317, nom. superfl. — Azedarach fraxinifolia Moench, Meth. Suppl. (1802) 58, nom. superfl.] — [Melia pinnata Stokes, Bot. Mat. Med. 2 (1812) 482, nom. superfl.] — [Melia indica (A. Juss.) Brandis, For. Fl. NW & C India (1874) 67, nom. superfl.; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1443; Corner, Wayside Trees | (1940) 466, 2 (1940) t. 139; ibid., ed. 3 (1988) 504, t. 151; Corner & Watanabe, II]. Guide Trop. Pl. (1969) 405, cum tab.]} {Arbor indica, fraxino similis, oleae fructu Bauhin, Pinax (1623) 416.] [Arbor Azadirachta Indica foliis non ramosis, flore minore Breyne, Prodr. Rar. Pl. 1 (1680) 3.] [Aria Bepou Rheede, Hort. Mal. 4 (1683) 107, t. 52.] [Olea malabarica Nimbo dicta, fructu racemoso oblongo Ray, Hist. Pl. 2 (1688) 1545.] [Azadirachta Indica foliis Fraxini, sive non ramosis majoribus, flore minore albo Breyne, Prodr. Rar. PI. 2 (1689) 21.] [Olea malabarica, Fraxineo folio Pluk., Almag. (1696) 269, t. 247, f. 1.] [Azedarach altera foliis semilunatis Herm., Mus. Zeyl. (1717) 48.] [Azedarach foliis falcato-serratis Burm., Thes. Zeyl. (1737) 40, t. 15.] [Melia foliis pinnatis L., Hort. Cliff. (1738) 161; Fl. Zeyl. (1747) 161; Royen, Fl. Leyd. Prodr. (1740) 462.] Tree to 16 m; bole to 60 cm diam. Bark red-brown or greyish, fissured and flaking in old trees; inner bark red-brown with colourless, sticky foetid sap; sapwood whitish. Leafy twigs c. 4-8 mm, cicatrose, with pale vertical lenticels, garlic-scented when dam- aged. Leaves 15—35 cm, imparipinnate or paripinnate with terminal spike, 4—7-jugate, red when young, garlic-scented when damaged; petiole c. 3—7 cm, c. 1.5 mm diam., subglabrous, base weakly swollen. Leaflets opposite to subopposite, 5—9 by 1.5—3.5 cm, the subapical the largest, falcate-lanceolate, curved basipetally, subglabrous, bases very asymmetric, acute, apices long-acuminate, margin serrate, costae c. 15 on each side, obtuse, spreading; petiolules c. 1-2 mm. Thyrses to 30 cm, lax, paniculiform, axillary or in axils of fallen leaves, sweetly scented; axis c. 1.5 mm diam., subglabrous; 342 Flora Malesiana ser. I, Vol. 12 (1) (1995) branches to c. 15 cm, squarrose, bearing branchlets to 2 or 3 more orders, tipped with cymules of 1—3 flowers, finely sericeous; bracts and bracteoles c. 0.5—1 mm, lanceo- late, + pubescent; pedicels c. 2 mm, swollen at articulation with pseudopedicel, c. 2.5—4 mm, finely pubescent. Calyx c. 1 mm long, salveriform, the lobes rounded, pubescent, margins ciliate. Petals linear spathulate, 4-6 mm long, white, + pubescent on both sur- faces. Staminal tube glabrous to sparsely pubescent, 10-ribbed, margin with 10 rounded or somewhat laciniate lobes; anthers 10, c. 0.8 mm long, narrowly ellipsoid, basifixed, weakly exserted. Ovary glabrous to finely pubescent. Drupe c. 1-2 cm long, ellipsoid, green becoming yellow when ripe; mesocarp thin, fleshy. Distribution — Probably native in Burma but widely and long cultivated in tropical Asia and Africa, where it has become extensively naturalized. In Malesia planted in Sumatra, the Malay Peninsula, and West Java, also in East Java, and the Lesser Sunda Islands (Bali, Lombok, Sumbawa), where it is, like Tamarindus indica L. (Legumino- sae), so completely naturalized in the greatly modified seasonal forest areas there as to appear native. Habitat & Ecology — Where naturalized, in open thickets and savanna, to 225 m al- titude. In West Africa, where the tree has come to dominate large areas of the savanna, the seeds are dispersed by fruit bats and baboons, after passage through which their germination is enhanced [Lieberman et al., Ecology 60 (1979) 65-75]. Vernacular names — Neem or nim tree, mimba, mimbo, nimbo (from Sanskrit), mar- gosa tree (from Portuguese for bitter). Uses — See also the paragraph on Phytochemistry, p. 8. Held sacred by the Hindus, the neem is potentially one of the most important of all tropical seasonal forest trees, having proved to be very adaptable and able to withstand arid conditions. It can be grown in impoverished soil and is a fastgrowing source of fuelwood [Ruskin, I.c.; see also M. Jacobson (ed.), Focus on phytochemical pesticides, vol. 1 (1988); Ad hoc panel of Nat. Res. Counc., Neem — a tree for solving global problems (1992); Tewari, I.c.]. The form grown in West Africa is ignored by stock and is therefore readily establish- ed. Some forms in Asia, however, are used as fodder and in the Malay Peninsula the young leaves and flowers are boiled and eaten with rice (Corner, |.c.). In Central Amer- ica, it is now widely planted and recent trials have shown it to grow rapidly and produce fine timber which makes a good substitute for Swietenia mahogany [Pennington, FI. Neotropica 28 (1981) 25]. It is the most important plantation species in northern Nigeria and is used as poles or for fuel. It coppices well and the timber has a very high calorific value. Coppice shoots may reach 10 m height in two years. The wood is tougher than teak and resists decay. It is also widely planted as a windbreak, shade and avenue tree, the world’s biggest plantation being of 50,000 trees in the plains of Saudi Arabia, plant- ed to shade the two million or so Muslim pilgrims camping there annually for ‘Haj’ rites [Sallem Ahmed et al., Econ. Bot. 43 (1989) 35]. It is a soil ameliorant and is potential- ly a source of many valuable by-products (Radwanski & Wickens, I.c.). It has been an ingredient of soaps, toothpaste and lotions in commerce for some decades (Corner, lye): Mabberley et al. — Meliaceae 343 The seeds contain some 40% oil by weight and this bitter material has been used in lamps and as a lubricant and has potential as a fuel-source, the mesocarp being a prom- ising substrate for the production of methane gas. Neem cake is an excellent fertilizer and the leaves and twigs are used as a mulch in Asia. The bark produces a valuable gum and tannin worth exploiting. The pressed leaves have long been put in books to ward off insects, the repellent being the limonoid, aza- dirachtin, which is in the seeds as well as the leaves; house-sparrows in India incor- porate leaves into their nests, perhaps reducing parasite loads. Azadirachtin is absorbed by plants and acts as a systemic insecticide so efficient that Japanese beetles and other insects, even including the desert locust, will starve rather than eat plants treated with it. It has been alleged that the substance is repellent to nematodes as well (Ruskin, |.c.). Of the many limonoids known from the tree, deacetylaxadirachnol (salannin) is as potent as azadirachtin in inhibiting ecdysis in tobacco budworm [Kubo et al., Tetrahedron 42 (1986) 489]. Neem seed powder with carbofuran greatly reduces leaf-hoppers and rice tungro virus in rice [Kareem et al., Int. Rice Res. Newsl. 13 (1988) 35]. Reduction in the incidence of phloem-specific tungro viruses in neem-treated plants is attributed to a shift in insect-feeding from phloem to xylem vessels [Saxena et al., J. Econ. Entom. 80 (1987) 1079]. Neem is alleged to contain anti-culicid factors and may be effective in controlling the spread of malaria. Under the Sanskrit name nimba it is mentioned in the Ayurvedae (Systema Medicinae) of Susruta, one of the most ancient of the Hindu medical writings. The first European account was published by Garcia d’ Orta, physician to the Portuguese Viceroy at Goa in India, in 1563. The leaves, bark and seed oil have medicinal proper- ties summarized by Perry, Med. Pl. E & SE Asia (1980) 260, and have been used in the treatment of a wide range of ailments, including malaria, eczema, dysentery and ulcers, but is particularly effective as a parasiticide for skin diseases such as scabies. Nimbidin, a substance isolated from the seed oil, has been found to be an efficient anti-inflamma- tory agent in artificially induced arthritis and oedema in rats [Pillai & Santhakumari, PI. Medica 43 (1981) 59], while an aqueous extract of the bark has been shown to increase lymphocyte function such that production of MIF, a lymphokine, is increased, which may account for the generally stimulating and skin-healing properties long observed [Van der Nat et al., J. Ethnopharmacol. 19 (1987) 125]. Some of the triterpenoids have antibacterial activity [Siddiqui et al., J. Nat. Prod. 55 (1992) 303]. Neem oil also has significant post-coital contraceptive action [Sinha et al., Ind. J. Med. Res. 79 (1984) 131]. Notes — 1. Gametogenesis and embryogenesis are discussed by Garudamma, J. Ind. Bot. Soc. 35 (1956) 222 & 36 (1957) 227. 2. Neem can be an aggressively invasive tree and it needs careful control to prevent its swamping other crops (Ruskin, I.c.). 344 Flora Malesiana ser. I, Vol. 12 (1) (1995) TRIBUS SANDORICEAE Sandoriceae T.D. Penn., Blumea 22 (1975) 507. Trees. Jndumentum of simple hairs. Leaves trifoliolate. Flowers in axillary thyrses. Staminal tube cylindrical, ribbed distally; anthers 10, included. Disk tubular. Stylehead with 4- or 5-lobed stigma. Fruit a drupe. Seeds exarillate, pachychalazal; endosperm ab- sent; cotyledons thick, plano-convex, collateral. The following genus only. SANDORICUM Sandoricum Cav., Diss. 7 (1789) 359; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 170; T.D. Penn., Blumea 22 (1975) 507; in Fl. Neotrop. 28 (1981) 359; Mabb., Blumea 31 (1985) 146. Calyx + truncate to shallowly 4- or 5-lobed. Petals (4) 5, free, imbricate. Staminal tube margin with 5 or 10 short lobes; anthers glabrous. Disk free, margin coarsely toothed. Ovary slightly sunken in receptacle, 4- or 5-locular, locules each with 2 colla- teral ovules. Drupe 1—5-locular, pyrenes 1- (or 2-)seeded; outer mesocarp rather dry- fleshy or soft and fibrous, inner mesocarp fleshy or spongy-fibrous; endocarp thin, cartilaginous. Seeds kidney-shaped, laterally compressed with thin sarcotesta; radicle apical, extending to surface or slightly exserted. Germination phanerocotylar; eophylls trifoliolate, opposite. Distribution — Five species, all but one (S. koetjape) restricted to western Malesia, where the cultivated forms of S. koetjape may have arisen, though wild relations appear to be native as far east as New Guinea. All five are wild in Borneo, to which three spe- cies are restricted. Habitat — Dipterocarp, kerengas and other forest types. Sandoricum beccarianum is restricted to peat swamp forests, S. borneense to riparian ones. Uses — The fruit (where known as mature) of all species is edible and the timber of some value. KEY TO THE SPECIES la. Leaflets obovate (to elliptic), apex rounded or emarginate. Peat swamp forests ... Lae ane Sera Rie CE RA, 19, ROAR ETL SEER IN 2 5. S. beccarianum b. Leaflets lanceolate, ovate (or elliptic), apex + acuminate. Other habitats ....... 2 2a. Leaflets glabrous, lanceolate (or elliptic), base obtuse to rounded. — Borneo, river Wanlksy i252 Sree Se Oe Ses Res Se Nia Se de ot ein eee A 3. S. borneense b. Leaflets pubescent to subglabrous (or glabrous), ovate, base cuneate to rounded = 3 3a. Leaflets long acuminate (acumen to 24 mm), base cuneate. — Borneo .......... By a A ae eee ae ty Olan eee ne Se NRL Sh, Ges I 4. S. caudatum b. Leaflets acute to acuminate (acumen never more than 15 mm), base rounded, ob- tusestoracute (Or subcumeate) cc. eras) tatoo Sofie fe asics hbo See 4 Mabberley et al. — Meliaceae 345 4a. Calyx 4—4.5 mm long, dark brown; petals glabrous; staminal tube with 5 erose lobes; petiolule of apical leaflet 4-10 cm. — Borneo ...... 1. S. dasyneuron b. Calyx c. 3.5 mm, yellow-green; petals + pubescent; staminal tube with 10 bifid lobes; petiolule oftapical leaflet:3—5.5,cmay.i%,<)0:2.3.4 520 «eel e 2. S. koetjape 1. Sandoricum dasyneuron Baill. Sandoricum dasyneuron Baill., Adansonia 11 (1874) 265; C.DC. in DC., Monogr. Phan. | (1878) 462; Becc., For. Born. (1902) 602; Merr., J. Str. Br. Roy. As. Soc., spec. no. (1921) 319; Mabb., Blu- mea 31 (1985) 147. Tree 8—10(—25) m, bole 10—25 cm diam. Bark smooth; wood light red. Leafy twigs c. 6—8 mm diam. /ndumentum ferrugineous. Leaves 25—52 cm long; petiole 10—20.5 cm, + pilose, base + swollen. Leaflets broadly ovate, glabrous adaxially, + pilose ab- axially especially on veins, bases acute to subcuneate, apices acuminate, acumen to 15 mm, costae (10—)12 or 13 on each side, inarched only near margin, apical leaflet 17—24 by 12-16 cm, laterals 16—20 by 10—11 cm, petiolules 3—15 mm on lateral leaflets, 4—10 cm on apical. Thyrses 2—13 cm long, weakly sweetly scented, produced with new leaves over at least 6 nodes, sparsely branched often from very close to base, primary branches to 6 cm long, squarrose, bearing fascicles of 1-3 flowers; axes densely short-tomen- tose; bracts c. 7 mm long, narrowly triangular, densely pubescent, caducous; bracteoles c. 1-2 mm, narrowly triangular, densely pubescent, caducous; pedicels c. 5—7 mm, articulated with pseudopedicel, 1(—2) mm long, continuous with calyx. Calyx c. 4mm long, 4-5 mm diam., campanulate, glabrous, dark purplish brown, splitting into 5 ir- regular obtuse lobes to 1 mm deep, margin ciliate. Petals 5, 8—9 by c. 3 mm, spathu- late, glabrous, greenish-white, apices acute. Staminal tube fleshy, white, swollen at mouth, + pubescent without, pilose within, margin with 5 erose lobes; anthers 10, 1.5 mm, in two ranks, the more distal opposite lobes, the more proximal inserted between them, all weakly exserted. Disk c. 2 mm long, narrowing towards mouth, membrane- ous, glabrous, margin irregularly laciniate. Ovary and style glabrous; stigmatic lobes c. 1.5 mm long. Drupe globose-pyriform, densely and minutely fulvous tomentellous, when mature (non vidi) 10 cm diam., multi-ribbed longitudinally (Beccari, |.c.); py- renes at least 3, bean-shaped. Seeds (non vidi) 2.5 cm long (C.DC., l.c.). Distribution — Malesia: Borneo (Sarawak, Brunei, Kalimantan), rarely collected. Habitat — Hill-sides and ridges in mixed dipterocarp forest and ecotone to kerengas (Sarawak), Agathis forest on acid sands (W Kutai), 50—600 m altitude. Vernacular names — Atap bojig (Land Dayak), kelampu (Iban, Malay), krunpok (Iban). Uses — According to Beccari, the drupe has much edible sweet flesh but it leaves a garlicky taste in the mouth. 2. Sandoricum koetjape (Burm. f.) Merr. Sandoricum koetjape (Burm. f.) Merr., Philipp. J. Sc., Bot. 7 (1912) 237; Fl. Manila (1912) 274; Barrett, Philipp. Agr. Rev. 6 (1913) 498, t. 3a; Pratt et al., Philipp. J. Sc. Chem. Geol. 8 (1913) 80, t. 12, f. 1; Webster, Philipp. Agr. Rev. 8 (1915) 110, t. 8b; Merr., Interpr. Rumph. (1917) 346 Flora Malesiana ser. I, Vol. 12 (1) (1995) 308; Sp. Blanc. (1918) 209; Webster, Philipp. Agr. Rev. 13 (1920) 184, t. 17b; Popenoe, Man. Trop. Fr. (1920) 426; Brown, Bull. Philipp. Dept. Agr. Bur. For. 21 (1920) 86, t. 41; Merr., J. Str. Br. Roy. As. Soc., spec. no. (1921) 319; Philipp. J. Sc. 19 (1921) 329; Enum. Philipp. Flow. Pl. 2 (1923) 361; Mend., Man. Pl. Breed. Trop. (1926) 296; Ochse, Ind. Vrucht. (1927) 125, t. 61; Fr. Cult. Dutch E Ind. (1931) 65, t. 26; Juliano, Philipp. Agr. 23 (1934) 11, 253; Villegas, Univ. Philipp. Nat. Appl. Sc. Bull. 5 (1936) 293; Jimenez, Philipp. Agric. 26 (1937) 587; Corner, Way- side Trees | (1940) 466 & 2 (1940) t. 140, 141; Adelb., Blumea 6 (1948) 316; Steenis, Fl. Scholen Indon. (1949) 233; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 891; Quisumb., Med. PI. Philipp. (1951) 486; Intengan et al., Philipp. J. Sc. 84 (1955) 361; King & Morgan, J. Chem. Soc. (1960) 4738; Yiagan, Philipp. Agric. 44 (1961) 477; Ramirez, Philipp. Agric. 45 (1961) 275; Backer & Bakh. f., Fl. Java 2 (1965) 121; Molesw. Allen, Mal. Fr. (1967) 105, t. 38; Meijer, Bot. News Bull. Sabah 8 (1967) 81; Anon., Wealth India 9 (1972) 200; Whitman, Proc. Fla St. Hort. Soc. 87 (1974) 379, t. 3, 4; Corner, Seeds Dicots 1 (1976) 191 & 2 (1976) t. 375b; L.M. Perry, Med. Pl. E & SE Asia (1980) 262; Chin & Yong, Malays. Fr. Col. (1981) 29 cum tab.; T.D. Penn. in Fl. Neotrop. 28 (1981) 359; Mabb., Blumea 31 (1985) 147; Corner, Wayside Trees, ed. 3 (1988) 504, t. 152, 153; Phillips et al., Guide Market Fr. SE Asia (1985) 66; Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 342, t. 261; Mabb. in Tree Fl. Malaya 4 (1989) 249, f. 8B; de Vogel, Seedlings Dicots (1990) 355, f. 129; H6, Ill. Fl. Vietnam 2, 1 (1992) 489. — Melia koetjape Burm. f., Fl. Ind. (1768) 101. — [Trichilia nervosa Vahl, Symb. | (1790) 31, nom. superfl.; Willd., Sp. Pl. 2 (1799) 554; DC., Prodr. 1 (1824) 623; Spreng., Syst. 3 (1826) 68; G. Don, Gen. Syst. 1 (1831) 682. — San- doricum nervosum (Vahl) M. Roem., Synops. Monogr. | (1846) 108, non Blume.] Sandoricum indicum Cav., Diss. (1789) 359, t. 202, 203; Lam., Encycl. Méth. 3 (1789) 69; Ill. (1793) t. 350; Willd., Sp. Pl. 2 (1799) 556; Sm. in Rees, Cyclop. 31, 1 (1815) Sandoricum n. 1; Roxb., Pl. Coast Corom. 3 (1820) 58, t. 261; DC., Prodr. 1 (1824) 621; Blume, Bijdr. (1825) 164; G. Don, Gen. Syst. | (1831) 680; Roxb., Fl. Ind., ed. Carey, 2 (1832) 392; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 232, t. 16, f. 15; Wight & Arn., Prodr. (1834) 120; J. Graham, Cat. Pl. Bom- bay (1839) 31; Blanco, Fl. Filip., ed. 2 (1845) 242; M.Roem., Synops. Monogr. | (1846) 108; D. Dietr., Syn. 4 (1847) 789; Paxt., Mag. Bot. 15 (1849) 111, 116 cum tab.; Hassk., Retzia 1 (1855) 146; Hort. Bog. Descr. 1 (1858) 125; Miq., Fl. Ind. Bat. 1, 2 (1859) 541; Suppl. (1861) 196; Drury, Handb. Ind. Fl. 1 (1864) 165; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 32; Bedd., Fl. Sylv. (1872) iv, (1873) t. 319; Hiern in Hook. f., Fl. Brit. India 1 (1875) 553, incl. var. velutinum Hiern; Kurz, For. Fl. Burma | (1877) 217; C.DC. in DC., Monogr. Phan. | (1878) 461, t. 6, f. 11; Blanco, FI. Filip., ed. 3 (1878) 85, t. 127; Fern.-Vill., Nov. App. (1880) 42; Bisschop Grev., PI. Ned. Ind. (1883) 494; Laness., Pl. Util. Colon. Fran. (1886) 310; Vidal, Rev. Vasc. Pl. Fil. (1886) 82; Watt, Dict. Econ. Prod. Ind. 6, 2 (1893) 457; King, J. As. Soc. Beng. 64, ii (1895) 23; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) t. 161, f. A—E; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 27; Pierre, Fl. For. Cochinch. 5 (1897) t. 353A, incl. var. cochinchinense Pierre; Koord., Minah. (1898) 389; Ridley, Agr. Bull. Malay Penins. 8 (1898) 228; Agr. Bull. Str. Fed. Malay St. 1 (1902) 429; Becc., For. Born. (1902) 574; Gamble, Man. Ind. Timb., ed. 2 (1902) 149; Perkins, Fragm. Fl. Philipp. (1904) 31; Brandis, Indian Trees (1906) 137; Merr., Philipp. J. Sc. 1, Suppl. (1906) 71; Backer, Fl. Batavia (1907) 274; Guillaumin, Rev. Gén. Bot. 22 (1910) 468, t. 12, f. 3; Backer, Schoolfl. Java (1911) 210; Pellegr. in Fl. Indo-Chine 1 (1911) 732, cum tab., incl. var. cochinchinense; Whitford, For. Philipp. 2 (1911) 46, t. 34; Koord., Exk. Fl. Java 2 (1912) 440; Atlas 1 (1913) t. 182; Crev. & Lem., Cat. Prod. Indoch. 1 (1917) 227; Basu, Ind. Med. PI. (1918) 313, t. 221; Troup, Silv. Ind. Trees (1921) 204; Ridley, Fl. Malay Penins. 1 (1922) 385; C.E. Parkinson, For. Fl. Andam. (1923) 117; Craib, Enum. Pl. Siam. 1 (1926) 353; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1946; Brig., Mém. Inst. Nat. Genev. 24 (1935) 45; Rodger, Handb. For. Prod. Burma (1936) 64; Elmer, Leafl. Philipp. Bot. 9 (1937) 3386; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) t. 35, f. A—E; Pellegr. in Fl. Indo-Chine, Suppl. (1946) 687; Tix., J. Agr. Trop. Bot. Appl. 5 (1950) 596; Chittenden, Dict. Gard. 4 (1951) 1863; Nair, Phyton 10 (1958) 145; Worth., Ceyl. Trees (1959) t. 129; H6 & Duong, Fl. Vietnam (1960) 248, t. 90OB. [Azedarach edulis Noronha, Verh. Batav. Gen. 5, ed. | (1791) art. 4, 5, nomen. ] Mabberley et al. — Meliaceae 347 Sandoricum nervosum Blume, Bijdr. (1825) 163; Hassk., Retzia 1 (1855) 146; Hort. Bot. Descr. (1858) 125; Miq., Fl. Ind. Bat. 1, 2 (1859) 541; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 32; Koord. & Vale- ton, Bijdr. Booms. Java 3 (1896) 30; Backer, Schoolfl. Java (1911) 210; Koord., Exk. Fl. Java 2 (1912) 440; Koord., Atlas (1913) t. 183; Ridley, Fl. Malay Penins. 1 (1922) 385; Craib, Enum. PI. Siam. 1 (1926) 254; Burkill & Haniff, Gard. Bull. Str. Settl. 6 (1930) 183; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1948. Trichilia venosa Spreng., Syst. 3 (1826) 68; G. Don, Gen. Syst. 1 (1831) 682. — Sandoricum veno- sum (Spreng.) M. Roem., Synops. Monogr. | (1846) 109. ?Sandoricum serratum G. Don, Gen. Syst. 1 (1831) 680; M. Roem., Synops. Monogr. | (1846) 109; D. Dietr., Syn. 4 (1847) 789. Sandoricum ternatum Blanco, FI. Filip. (1837) 346; M. Roem., Synops. Monogr. | (1846) 109. Sandoricum glaberrimum Hassk., Retzia 1 (1855) 145; Hort. Bogor. Descr. (1858) 124; Miq., Fl. Ind. Bat. 1, 2 (1859) 541. Sandoricum maingayi Hiern in Hook. f., Fl. Brit. India 1 (1875) 554; C.DC. in DC., Monogr. Phan. 1 (1878) 462, incl. var. guadripetalum C.DC.; King, J. As. Soc. Beng. 64, ii (1895) 22; Becc., For. Born. (1902) 201, 599; Merr., J. Str. Br. Roy. As. Soc., spec. no. (1921) 319, incl. var. quadripeta- lum; Ridley, Fl. Malay Penins. | (1922) 385; Meijer, Bot. News Bull. Sabah 8 (1967) 81. Sandoricum borneense auct. non Miq.: Fern.-Vill., Nov. App. (1880) 43; Vidal, Cat. Pl. Prov. Man. (1880) 22. [Sandoricum harmandianum Pierre ex Laness., Pl. Util. Colon. Fran. (1886) 310, nomen.) Sandoricum radiatum King, J. As. Soc. Beng. 64, ii (1895) 21; Ridley, Agr. Bull. Str. Fed. Malay St. 1 (1902) 429. Sandoricum harmandii Pierre, Fl. For. Cochinch. 5 (1897) t. 353B, excl. fol. (= Aglaia sp.); Pellegr. in FI. Indo-Chine | (1911) 733. Sandoricum vidalii Merr., Philipp. Govt. Lab. Bur. Bull. 6 (1904) 8; Whitford, For. Philipp. 2 (1911) 46, t. 35; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 361; Elmer, Leafl. Philipp. Bot. 9 (1937) 3387. Sandoricum harmsianum Perkins, Fragm. Fl. Philipp. (1904) 31. Sandoricum ledermannii Harms in Eng]. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 172, 177, no- men; Bot. Jahrb. 72 (1942) 204. [Hantol Kamel, App. Herb. Luz. Philipp. (1704) 54.] [Sandoricum domesticum Rumph., Herb. Amb. | (1741) 167, t. 64.] Tree to 45(—50) m; bole to 1 m diam., fluted and sometimes with buttresses to 3 m tall. Bark pale pinkish brown, smooth, lenticellate to peeling with round flakes; inner bark pink; sapwood pale yellow; heartwood pink or reddish. Twigs cicatrose, lenticel- late, rough, grey-brown. Crown domed. Leafy twigs 4—7 mm diam., subglabrous to fulvous-tomentose. Leaves 18—40 cm long; petiole 7.5—16 cm, flattened (or even wing- ed in sicco) adaxially towards the + swollen base, subglabrous to fulvous pubescent. Leaflets ovate, glabrous or with a few brown hairs on midrib adaxially, subglabrous to densely brown pubescent abaxially, pink when young, withering yellow or reddish, bases acute to rounded, + asymmetric on lateral leaflets, apices acuminate, costae 7—14 (—20 in cuitivated forms) on each side, looped near margin, apical leaflet 8—20 by 5—14 cm, laterals usually smaller and narrower; petiolules 4—9 mm on lateral leaflets, 3—5.5 cm on apical. Thyrses 2.5—24 cm long, fragrant, produced in up to 8 of the most apical axils, erect or weakly drooping; axes + fulvous pubescent; primary branches to 8 cm long, squarrose, bearing secondary branches of glomerules of 1—5 flowers; bracts c. 7 mm long, narrowly triangular, densely pubescent, caducous, bracteoles somewhat smaller, subopposite; pedicels c. 3—5 mm, articulated with pseudopedicels, ¢. |-1.5 cm long, continuous with calyx. Calyx c. 3.5 mm long, campanulate to cupular, splitting 348 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 52. Sandoricum koetjape (Burm. f.) Merr. a. Leaf and inflorescences; b. half flower; c. fruit; d. seed- ling. — S. beccarianum Baill. e. leaf and inflorescence; f. half flower; g. fruit. Drawing R. Wise. Repro- duced with permission from Tree Flora of Malaya 4. Mabberley et al. — Meliaceae 349 into 5 irregular obtuse or rounded lobes to 0.75 mm deep, pubescent, margin ciliate, yellow-green. Petals (4 or) 5,6—9 mm long, linear-lanceolate to oblanceolate, yellowish- green or pinkish, + pubescent without, reflexed at anthesis, apices rounded to emargi- nate. Staminal tube + pubescent without, pilose within, pale yellow to orangeish, mar- gin with 10 lobes acute to bifid, somewhat reflexed at anthesis; anthers (8 or) 10, 1-1.5 mm long, narrowly oblong, apiculate, + in 2 ranks, weakly exserted. Disk c. 1.5 mm long, membraneous, glabrous, margin irregularly laciniate. Ovary and style glabrous; stigmatic lobes c. 1.5 mm long. Drupe 5—8 cm diam., flattened-globose, velvety, yel- low or brownish when ripe, smooth to longitudinally wrinkled; pericarp with milky latex; mesocarp white, translucent, juicy sweet to very sour; endocarp tough, with | or 2 seeds. Seeds 20—35 by 12—21 mm, 9-16 mm thick, germinating in rotting fruit; co- tyledons pink inside. — Fig. 52a—d. Distribution — Planted widely in the Asiatic tropics but probably native only in Malesia, the wild form (see below) extending from the Malay Peninsula and Sumatra throughout the archipelago to New Guinea (Madang). The pubescent cultivated form with leaves withering red (Parkinson, I.c.) is that planted in India including the Anda- mans, Burma, Indochina, the Mascarenes and the New World as well as grown under glass in Europe. Habitat & Ecology — Native and naturalized in primary and secondary forests to 1200 m altitude or more. Wild trees are very scattered in the forest, Whitmore [Trop. Rain For. Far East (1975) 237] recording them at a density of 0.8 trees per 40 ha in Ulu Kelantan, Malay Peninsula. In Singapore the leaves are dropped biannually, leaving the crown almost bare, but the new leaves appear within a week (Corner, |.c. 1940). The fruits mature in 2—3 months and may lie carpeting the ground under the tree thereafter. The flowering in the Malay Peninsula is so regular that formerly it was recognized as the cue for rice-planting (Burkill, |.c.). Morphology — Juliano (Il.cc.) has shown that the sequence of development of the floral parts is: sepals, petals, androecium, carpels and lastly disk. He also gives details of micro- and macrosporogenesis. Villegas (1.c.) reports that the mesocarp is derived from the inner pericarp walls and as outgrowths from the endocarp, and that the indu- mentum of the drupe develops after fertilization, the ovary being glabrous. Similar is reported by Pennington [Flora Neotropica 28 (1981) 11] for two neotropical species of Trichilia. Cytology — The form known as ‘Native’ has 2n = 22, whereas the robust cultivated ‘Bangkok’ (= ‘Manila’ ) has 2n = 44 and is a tetraploid (Ramirez, |.c.) and 2n = 16, 28, 32 have also been recorded by Pennington [Blumea 22 (1975) 508]. There is no evidence for apomixis and records of polyembryony probably rest on a confusion of the |- or 2- seeded ‘stones’ with seeds (Juliano, Il.cc.). Juliano also recorded a case of one tree in Manila which never produced fruit. This was due to the failure of the corolla to open and no self-pollination took place even though mega- and microsporogenesis were normal. Systematics — This species embraces the cultivated fruit trees known as sentul (and variants) and kechapi (and variants), grown throughout western Malesia, largely as vil- lage trees, for shade as well as fruit. The form known as kechapi in the narrow sense, 350 Flora Malesiana ser. I, Vol. 12 (1) (1995) i.e. that with robust pubescent twigs, a brown pubescent adaxial surface to the large leaves, and with many costae in the leaflets is that most frequently encountered in the Philippines and probably the only form cultivated outside Malesia, being early carried to Indochina, the Mascarenes and now increasingly in the New World, notably Costa Rica (Pennington, |.c.) and especially Florida (Whitman, I.c.), where a robust form of it is known as ‘Manila’ which may serve as a cultivar name. The robust form introduced to the Philippines in 1949 from Thailand was called ‘Bangkok’, compared with the original ‘Native’ (Ramirez, |.c.). This latter is probably the Red Sentol of Corner (1940) who notes that the tree has leaves which wither red, pale green petals and a sweet or sour fruit with a thick generally wrinkled or uneven pericarp and often rotting on the tree. It seems to be the form originally described by N. Burman and Cavanilles, though there is considerable variation in the cultivated trees in terms of pubescence and the form and taste of the fruit. A specimen from J. Burman’s herbarium and now preserved at L has much less pubescent leaflets, for example. Wild trees in the Malay Peninsula and eastwards to New Guinea, beyond the range of cultivation of the Red Sentol, have more delicate twigs, smaller subglabrous leaves, withering yellow, the leaflets with fewer costae and pinkish petals in rather shorter inflorescences, the fruits sweet-tasting with thinner smoother pericarp and falling when ripe. In the Philippines and from Celebes eastwards, the two forms seem perfectly distinct but in western Malesia it becomes impossible to draw a clear line between them. Indeed, it is tempting to speculate that it is here, where all the other species in the genus are also native, different forms have been selected from the variable wild populations. It is interesting to compare the origin of ‘Manila’ with the robust form of the Lansium domesticum aggregate known as ‘Kokossan’ (q.V.). Vernacular names — Throughout the archipelago and beyond, different forms are known variously as sentul (or variants) and kechapi (or variants) though apparently, as in the case of Lansium domesticum, not consistently, even within islands (see Corner 1940). It is notable that the scientific name is compounded of both names, Sandoricum being Rumpf’s latinization of the first and koetjape being Burman’s hollandicization of the second. Variants are listed by K. Heyne (I.c.) and Merrill (1923). The wild form in the Philippines is known as malasantol or bago-santol or variants (Merrill, |.c.). Uses — The mesocarp is the part eaten. In some forms it is exceedingly sweet and the sour ones may be ‘excruciating’ (Corner 1940). Nevertheless, some trees have ex- cellent flavour and Popenoe (lI.c., quoting Webster) remarks that should seedless or semi-seedless forms be found, the sentul could “become one of the most popular of the tropical fruits.” An excellent jam has been prepared from the fruits (Barrett, I.c.) which in some forms smell like ripe peaches. Rumpf notes that the fruit may be eaten raw or cooked, like lemons, with fish in Ambon. It may also be candied or fermented with rice in the production of an alcoholic drink. Although not particularly high in Vitamin C, it is a good source of Vitamin B [Hermano & Sepulveda, Philipp. J. Soc. 54 (1934) 70]. See also Sotto in E.W.M. Verheij & R.E. Coronel (eds.), Edible fruits and nuts, Plant Res. SE Asia (PROSEA Handbook) 2 (1991) 284. The tree is fast growing when young, Roxburgh (l.c.) recording that a 24-year-old tree in the Calcutta Botanic Garden had a bole with a circumference of over 2 m. The Mabberley et al. — Meliaceae 35] tree has been widely planted for shade: it is recommended as an avenue species. The timber is red, moderately hard and takes a fine polish. Rumpf records that it was a dur- able timber for house construction in Ambon but Anon. (1972) reported that it perishes on exposure to water and borers. Nevertheless, it is in much demand in Burma for the manufacture of sandals and has been used in the construction of barrels, boats, carts and butchers’ blocks. The bark has been used in tanning fishing nets. It, and also par- ticularly the roots, are claimed as efficacious in the treatment of a number of medical conditions (see Perry for digest), its value being known in the seventeenth century and recorded by Mercado [see Fern.-Vill., Nov. App. (1880) 42]. Stems yield the triterpen- oids 3-oxo-olean-12-en-29 oic acid, its 3-dihydroderivative katonic acid and an A-ring seco derivative of katonic acid called koetjapic acid, C39H46O4; the first-named two components have significant cancerostatic activities in cell cultures [Kaneda et al., J. Nat. Prod. 55 (1992) 654]. Two new limonoids, sandoricin and 6-hydroxysandoricin, have been isolated from the seeds; both have been tested for insect-antifeedant activity with larvae of two lepidopteran species and found to be active [Powell et al., J. Nat. Prod. 54 (1991) 241]. 3. Sandoricum borneense Mig. Sandoricum borneense Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 33; C.DC. in DC., Monogr. Phan. 1 (1878) 463; Valeton, Ic. Bogor. | (1897) t. 10; Merr., J. Str. Br. Roy. As. Soc., spec. no. (1921) 319; Schwartz, Mitt. Inst. Allg. Bot. Hamb. 7 (1931) 235; Meijer, Bot. News Bull. Sabah 8 (1967) 81; T.D. Penn., Blumea 22 (1975) 506, t. 14e; Ashton in UNESCO, Trop. For. Ecosyst. (1978) 92; Mabb., Blumea 31 (1985) 150. Tree 7—20 m:; bole to 15 m, 20—40(—60) cm diam., unbuttressed. Bark smooth, pale brown, with small lenticels; inner bark light brown or pinkish; sapwood white. Leafy twigs 3—4(—5) mm diam., smooth, often pale in sicco. /ndumentum restricted to inno- vations. Leaves 22—38 cm long, red when young; petiole 4—10 cm, terete to somewhat flattened adaxially near base. Leaflets elliptic to lanceolate-ovate, bases obtuse to round- ed, those of laterals often markedly asymmetric, apices acuminate, costae about 9—12 on each side, arising almost at right angles from midrib, very weakly arcuate and looped at margin; apical leaflet not conspicuously larger than laterals, 8-22 by 3.5—8.5 cm; petiolules 3—5(—10) mm on lateral leaflets, 20—45 mm on apical. Thyrses 10-16 cm long, + erect, narrow, fragrant; primary branches to 6 cm long, subsquarrose to weakly ascending, the secondaries c. 1.5 cm long, bearing cymules of 1—5 flowers; bracts nar- rowly triangular, c. 8 mm long, to foliaceous, when lanceolate to oblanceolate, 2 cm long and petiolate, caducous; bracteoles narrowly triangular, c. 4 mm long, caducous; pedicels 3—7 mm long, glabrous to subpuberulous. Calyx 1.5—2 mm long, shallowly cupular, + puberulous, pale green, splitting irregularly into 5 obtuse lobes c. 0.5 mm deep. Petals 5, 6—8 mm long, linear-oblong, glabrous, creamy green to pinkish, apices acute. Staminal tube creamy green to pinkish, glabrous and with c. 20 ribs without, vil- lous within, margin 10-lobed, each lobe + bifid; anthers c. 0.5 mm long, in two ranks, alternating with the lobes, apiculate, weakly exserted. Disk c. 2 mm long, glabrous, margin 5-lobed, the lobes + bifid. Ovary and style glabrous; stigmatic lobes c. | mm 352 Flora Malesiana ser. I, Vol. 12 (1) (1995) long. Drupe 2.5—4 cm diam., subglobose to pyriform, velutinous, buff-yellow to orangeish, obscurely longitudinally ribbed, usually solitary; pericarp with milky latex; mesocarp sweet but mealy; endocarp tough; pyrenes 2—5. Seeds c. 14 by 9 mm (Vale- ton, I.c.); cotyledons pink within. Distribution — Malesia: Borneo (Sarawak, southern Sabah, northern Kalimantan). Habitat & Ecology — Riverbanks, subject to inundation down to just above tidal influence. According to Ashton (l.c. and in litt. 29 xii 1982), it is a common river- side tree in Brunei, overhanging the water like Dipterocarpus oblongifolius (neram). The fruits are dropped into the water and devoured by fish which appear to spit out the pyrenes. Whether such leads to dispersal is unknown (cf. Dysoxylum angusti- folium). Vernacular names — Apoh (Kaya), apok (Kenyah), kelampu (Iban). Uses — The sparsely produced fruits are edible and the timber is good for “sapé’- making. 4. Sandoricum caudatum Mabb. Sandoricum caudatum Mabb., Blumea 31 (1985) 150, f. 1. Small tree to 10 m and bole to 15 cm diam. Bark smooth, grey-green. Leafy twigs c. 3—4 mm diam. /ndumentum fulvous, restricted to innovations. Leaves 20—25 cm long; petiole 5—9 cm, wrinkled in sicco, base swollen and flattened adaxially. Leaflets ovate, bases cuneate, + asymmetric on lateral leaflets, apices long acuminate, acumen to 24 mm, costae 8—10 on each side, arcuate, looped well clear of margin; apical leaflet 14— 15.5 by 6-7 cm, laterals 10-13 by 4—6 cm; petiolules wrinkled in sicco, 6—9 mm on lateral leaflets, 4-5 cm on apical. Thyrses 4—7 cm long, sparsely branched; borne in axils of undeveloped leaves, primary branches to 2 cm long, squarrose, bearing fasci- cles of 2 or 3 flowers; axes minutely puberulous; bracts subtending primary branches 5—7 mm long, narrowly lanceolate, puberulous, caducous; bracteoles at base of pedicels smaller, often with | or 2 more, smaller ones half way to articulation with pseudopedi- cel; pedicels c. 5—6 mm; pseudopedicels c. 1-2 mm continuous with calyx. Calyx c. 2.5 mm long, shallowly campanulate, + puberulous, pale green, splitting into 5 irregular obtuse lobes c. 0.75 mm deep, margin ciliate. Petals 5, c. 4.5 mm long (immature), c. 2 mm wide, elliptic, apex rounded, creamy white. Staminal tube glabrous without, villous within, margin with 10 irregular lobes, creamy white; anthers 10, + in two ranks, alter- nating with lobes, c. 0.75 mm long, oblong, weakly exserted. Disk c. 1.5 mm long, glabrous, membraneous, clasping ovary, margin laciniate. Ovary and style glabrous; stigmatic lobes c. 1 mm long. Drupe solitary, at least 5 cm long, 3.5 cm diam., stipitate, rostrate, stipe to | cm, beak to 6 mm, densely yellow-brown, velutinous, + ribbed lon- gitudinally, calyx marcescent, pyrenes 3 or 4. Seeds c. 16 mm long, 9 mm wide (imma- ture); cotyledons reddish in sicco. Distribution — Malesia: Borneo (Sarawak, Ist Div.: 8 collections only). Habitat — Lowland dipterocarp forest and kerangas to 350 m altitude. Note — Neither mature flowers nor mature fruits are known. Mabberley et al. — Meliaceae 353 5. Sandoricum beccarianum Baill. Sandoricum beccarianum Baill., Adansonia 11 (1874) 264; Mabb., Blumea 31 (1985) 151; in Tree FI. Malaya 4 (1989) 249. Sandoricum emarginatum Hiern in Hook. f., Fl. Brit. India 1 (1875) 264; C.DC. in DC., Monogr. Phan. | (1878) 461; King, J. As. Soc. Beng. 64, ii (1895) 22; Becc., For. Born. (1902) 574; Merr., J. Str. Br. Roy. As. Soc., spec. no. (1921) 319; Ridley, Fl. Malay Penins. 1 (1922) 385; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1946; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 890; Ander- son, Gard. Bull. Sing. 20 (1963) 165; Meijer, Bot. News Bull. Sabah 8 (1967) 81; Corner, Gard. Bull. Sing., Suppl. 1 (1978) 77, 86, 89. Tree to 35 m with bole to 25 m and 70 cm diam. Bark smooth with minute cracks to deeply fissured; inner bark red-brown, to 10 mm thick; sapwood whitish to pale brown; heartwood pink to red-brown. Leafy twigs c. 3-5 mm diam. /ndumentum restricted to innovations. Leaves 11—25 cm long; petiole 3—7 cm, weakly swollen and flattened ad- axially at base. Leaflets obovate (to elliptic), bases acute to subcuneate; apices emargi- nate or rounded (to obtuse), sometimes mucronate, costae 6—8 on each side, weakly arcuate and rather obscurely looped near margin; apical leaflet 5.5—14 by 3.5—9 cm, laterals 4-12 by 2.5—7 cm; petiolules 3.5—10 mm on lateral leaflets, 4-6 cm on apical, all somewhat swollen at junctions with laminae and grooved adaxially in sicco. Thyrses 1—5.5 cm long, produced with new leaves and from axils of fallen ones up to at least ten nodes from apex, fasciculate, apparently arising in the axils of undeveloped leaves in axillary buds; primary branches to 2 cm long, bearing fascicles of 1—4 flowers; axes minutely puberulous to glabrous; bracts c. 1 mm long, narrowly triangular, pubescent, caducous; bracteoles 1—3, c. 0.5 mm long, narrowly triangular, pubescent, borne half way up pedicel to articulation with pseudopedicel; pedicels c. 4-6 mm long, conspic- uously articulated with pseudopedicel, c. 1 mm long, continuous with calyx. Calyx c. 2—2.5 mm long, shallowly cupular, + puberulous, red-brown, margin truncate to irregularly (4- or) 5S-lobed, the lobes up to 1 mm deep, obtuse, margin + ciliate. Petals (4) 5, 6-7 by 2.5 mm, oblanceolate, yellow-green to white, glabrous. Staminal tube fleshy, deeply (16- or) 20-ribbed, cream, + pilose within, margin with (8) 10 emargi- nate lobes; anthers (8) 10, 1 mm long, ovate, apiculate, in one rank, inserted opposite lobes, very weakly exserted. Disk c. 2 mm long, membraneous, glabrous, margin irregularly laciniate. Ovary and style glabrous; stigmatic lobes c. 1 mm long. Drupe (immature) c. 3 cm long, 2.5 cm diam., subglobose, stipitate, stipe to 5 mm, densely minutely tomentellous, orange-red or pinkish-yellow, pericarp with white latex. Seeds 2, c. 2 cm long, 1 cm diam. — Fig. 52e-f. Distribution — Thailand and Malesia: coastal regions of Sumatra, Malay Peninsula, Borneo. Habitat — Peatswamp forests; it may be codominant locally (Briinig), to 30 m altitude. Vernacular names — Apau, kelampu apau (Sarawak), gapas-gapas, kapas-kapas (Sa- bah), kechapi or sentul kera or kerok (i.e. baboon’s sentul) (Sumatra). Uses — The fruit is edible and also eaten by Siamang (Chivers). The timber is yel- lowish-red and durable, being used as planks in ship- and house-building in Sumatra. Burkill (1.c.) reports that it reached the Singapore sawyers. oe) Nn TS Flora Malesiana ser. I, Vol. 12 (1) (1995) SUBFAMILY SWIETENIOIDEAE Swietenioideae Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 267, 270; T.D. Penn. & Styles, Blumea 22 (1975) 511. Monoecious trees. Buds nearly always protected by scale leaves. Leaves pinnate, spi- rally arranged. Locules nearly always with 3 or more ovules, these biseriate; stylehead discoid or very rarely capitate. Fruit a septifragal capsule with a central columella and seeds winged, or capsule subwoody or leathery with rudimentary columella and seeds unwinged but with a woody or corky outer layer. TRIBUS SWIETENIEAE Swietenieae A. Juss., Bull. Sci. Nat. Géol. 23 (1830) 236 (tribus Cedrelacearum); T.D. Penn. & Styles, Blumea 22 (1975) 513. Flowers with or without a gynophore. Petals free. Stamens 8—10 (11), filaments part- ly or completely united into a cylindrical, urceolate or bowl-shaped tube, margin lobed or not. Capsule woody, nearly always with a well-developed columella, the valves sometimes splitting into an outer woody and an inner membraneous layer. Seeds vari- ously winged; cotyledons collateral, flattened; (residual) endosperm present. Distribution — Pantropical. Nine genera, five restricted to Africa and Madagascar, one to India and Sri Lanka, two to tropical America and the following. CHUKRASIA Chukrasia A. Juss., Bull. Sci. Nat. Géol. 23 (1830) 239; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 65; T.D. Penn. & Styles, Blumea 22 (1975) 519. — [Chickrassia Wight & Arn., Prodr. (1834) 122, nom. superfl.] — [Plagiotaxis Wall., List (1829) n. 1269, 1270, nom. nud., ex Kuntze, Rev. Gen. Pl. | (1891) 110, nom. superfl.] Deciduous trees. /ndumentum of simple hairs. Leaves paripinnate with terminal spike, imparipinnate and bipinnate with incised or lobed leaflets in juveniles, rarely retained at maturity. Thyrses axillary, often subterminal appearing terminal. Calyx 4- or 5-lobed. Petals 4 or 5, free, contorted and much longer than calyx in bud. Staminal tube cylindri- cal, somewhat narrowing distally, margin entire to crenulate; anthers attached to margin. Disk obscure to narrowly cushion-shaped. Ovary flask-shaped, 3—5-locular, each lo- cule with numerous ovules; stylehead capitate with 3—5 stigmatic ridges. Capsule ovoid or ellipsoid, woody, opening by 3—5 valves from the apex, the valves splitting into an outer and inner bifid layer; columella with 3—5 sharply angled ridges, extending to apex of capsule; seed scars conspicuous. Seeds 60-100 per locule, wings terminal, arranged laterally in tiers in two ranks; endosperm present; cotyledons subcircular; radicle oblique- ly exserted. Germination phanerocotylar; cotyledons unequal-sided; eophylls opposite, irregularly imparipinnate, the leaflets lobed or irregularly toothed. 2n = 26. Distribution — One somewhat variable species from India and Sri Lanka, eastwards through tropical Asia to western Malesia. Mabberley et al. — Meliaceae 355 Chukrasia tabularis A. Juss Chukrasia tabularis A. Juss., Bull. Sci. Nat. Géol. 23 (1830) 241; Linnaea 6 lit. (1831) 115; Mém. Mus. Nat. Hist. Nat. 19 (1832) 251, t. 22, f. 27; C.DC. in DC., Monogr. Phan. 1 (1878) 726, t. 8, f. 9, incl. var. malaccana C.DC.; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 271, t. 152, f. S, T; Pierre, Fl. For. Cochinch. 5 (1897) t. 357C, incl. var. attopeuensis Pierre; Cooke, Fl. Bomb. (1902) 216; Brandis, Indian Trees (1906) 144; Pellegr. in Fl. Indo-Chine 1 (1911) 780, t. 84, f. 5-11, incl. var. attopeuensis, var. dongnaiensis (Pierre) Pellegr., var. microcarpa (Pierre) Pellegr., var. ve- lutina (M. Roem.) Pellegr.; Gamble, Fl. Madras 1 (1915) 186; C.E. Parkinson, For. Fl. Andam. (1923) 119, t. 2, f. 37; Livera, Ann. Roy. Bot. Gard. Perad. 9 (1925) 308; Merr. & Chun, Sunyat- senia | (1930) 61, incl. var. velutina; Alston in Trim., Handb. Fl. Ceylon 6 (1931) 46; Merr. & Chun, Sunyatsenia 2 (1934) 32; Briq., Mém. Inst. Nat. Geney. 24 (1935) 34, t. I, f. 9; Harms, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 50, t. 4, f. S, T; Pellegr. in Fl. Indo-Chine, Suppl. (1946) 721, incl. var. quadrivalvis Pellegr.; Anon., Wealth of India 2 (1950) 152, t. 66; How & Chen, Acta Phytotax. Sin. 4 (1955) 32, incl. var. velutina; Worth., Ceylon Trees (1959) t. 124; Anon., Icon. Corm. Sin. 2 (1972) 571, t. 2872, incl. var. velutina; Bull. Dept. Med. Pl. Nepal 4 (1973) 12; FI. Hainan 3 (1974) 70, t. 572; T.D. Penn. & Styles, Blumea 22 (1975) 522, t. 18c, d; Wu, Fl. Yun- nan. 1 (1977) 211, t. 49, f. 1—4, incl. var. velutina; Whitmore, Enum. Flow. Pl. Nepal 2 (1979) 85; Mabb. in Tree Fl. Malaya 4 (1989) 256, f. 10. — Chickrassia tabularis (A. Juss.) Wight & Arn., Prodr. (1834) 123; J. Graham, Cat. Pl. Bombay (1839) 32 (‘tubularis’); Wight, Il. 1 (1840) t. 56 (‘mbularis’); M. Roem., Synops. Monogr. | (1846) 135; Thwaites, Enum. (1858) 61; Drury, Handb. | (1864) 171; Bedd., Fl. Sylv. (1869) 9, t. 9; Kurz, Rep. Veg. Andam. (1870) 33; Hiern in Hook. f., Fl. Brit. India 1 (1875) 568: Kurz, For. Fl. Burma | (1877) 227: Theob. in Mason, Burma, ed. 3, 2 (1883) 586, incl. var. genuina Theob., nom. superfl. & var. velutina (M.Roem.) Theob.; Laness., Pl. Util. Colon. Fr. (1886) 312; Watt, Dict. Econ. Prod. India 2 (1889) 268; Trimen, Handb. Fl. Ceylon 1 (1893) 252; Talbot, Trees Bomb., ed. 1 (1894) 43; ed. 2 (1902) 117; King, J. As. Soc. Beng. 64, ii (1895) 88, incl. var. velutina (M. Roem.) Theob.; Gamble, Man. Ind. Timb., ed. 2 (1902) 156; Prain, Beng. PI. (1903) 223; Talbot, For. Fl. Bomb. 1 (1909) 248, t. 147; Basu, Ind. Med. Pl. (1918) 320, t. 229; Troup, Sylv. Ind. Trees (1921) 191, t. 79; Ridley, Fl. Malay Penins. | (1922) 415; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 524; Kanjilal et al., FI. Assam | (1936) 241; Chittenden, Dict. Gard. 1 (1951) 459. Swietenia trilocularis [Roxb. ex Buch.-Ham., Journey Madras | (1807) 184, nom. nud., ex] G. Don, Gen. Syst. | (1831) 688. — Chickrassia trilocularis (G. Don) M. Roem., Synops. Monogr. | (1846) iS: Swietenia chickrassa Roxb., Hort. Beng. (1814) 33, nom. nud.; Fl. Ind., ed. Carey, 2 (1832) 399. — Chukrasia chickrassa (Roxb.) Schultze-Motel, Kulturpfl., Beth. 4 (1966) 209. — [Plagiotaxis chic- krassa Wall., Cat. (1829) n. 1269, nom. nud.; Kuntze, Rev. Gen. PI. 1 (1891) 110, nom. superfl.] [Plagiotaxis velutina Wall., Cat. (1829) n. 1270, nom. nud.; Kuntze, Rev. Gen. Pl. 1 (1891) 110, nom. superfl.] — Chickrassia velutina M. Roem., Synops. Monogr. | (1846) 135; Kurz, For. Fl. Burma 1 (1877) 227. — Chukrasia velutina (M. Roem.) C.DC. in DC., Monogr. Phan. | (1878) 727, incl. var. macrocarpa C.DC.; Pierre, Fl. For. Cochinch. 5 (1897) t. 357, incl. var. dongnaiensis Pierre & var. microcarpa Pierre; Cooke, Fl. Bomb. (1902) 216; Brandis, Indian Trees (1906) 145; Livera, Ann. Roy. Bot. Gard. Perad. 9 (1925) 308; Alston in Trim., Handb. Fl. Ceylon 6 (1931) 46; Brig,, Mém. Inst. Nat. Genev. 24 (1935) 34; Worth., Ceylon Trees (1959) t. 125. — [Swietenia velutina Wall. ex Kurz, J. As. Soc. Beng. 42, ii (1873) 65, nom. in syn.]} [Swietenia sotrophola Buch.-Ham. ex Wall., Cat. (1829) n. 1269, nom. in syn.] Cedrela sp.: Wall., Cat. (1831/32) n. 4892. Chickrassia nimmonii R. Graham ex Wight, Ill. 1 (1840) 148; Dalz. & Gibs., Bomb. Fl. (1861) 38; Drury, Handb. 1 (1864) 171. — Chukrasia nimmonii (R. Graham ex Wight) Merr. & Chun, Sun- yatsenia | (1930) 61, nom. provis. Sapindus multijugus Wall., Cat. (1847) n. 8099. Melia tomentosa auct. non Roxb.: Kurz, Rep. Andam. (1867) iv. [Swietenia villosa Wall. ex Kurz, J. As. Soc. Beng. 42, ii (1873) 65, nom. in syn.} Dysoxylum esquirolii Lév1., Cat. Pl. Yunnan (1916) 176; China Rev. Ann. (1916) 23. 356 Flora Malesiana ser. I, Vol. 12 (1) (1995) - . : bt S 8 a — ar es une MAK - Fig. 53. Chukrasia tabularis A. Juss. a. Leaf and inflorescence; b. seedling; c. young leaf; d. half flower; e. fruit, closed and dehisced; f. seed. Drawing R. Wise. Reproduced with permission from Tree Flora of Malaya 4. Mabberley et al. — Meliaceae 357 Tree to 40 m; bole to 25 m, fluted below, to 120 cm diam. Buttresses convex, to 150 cm tall. Bark dark brown, fissured vertically and scaling or cracking into rectangular blocks; inner bark red brown or pinkish; sapwood straw, heartwood yellow to reddish- brown. Twigs grey, bark cracking horizontally. Leafy twigs 4—6 mm diam., lenticel- late. Leaves 30—50 cm, with 6—12 leaflets on each side, + short-pubescent; petiole 4—9 cm, + terete, swollen at base. Leaflets ovate to oblong, + asymmetric or even falcate, the subapical the largest, 10—-17.5 by 3.5—6.5 cm, the most proximal as small as 4 by 2.2 cm, bases obtuse to rounded distally, acute to cuneate proximally, apices acute to acuminate, subglabrous to finely scattered short pubescent adaxially, subglabrous ex- cept for short hairs on veins and domatia in axils of costae to densely velutinous abaxi- ally, chartaceous to subcoriaceous, costae c. 9-11 on each side of largest leaflets, arcu- ate, + bifurcating near margin; petiolules 2-6 mm. Thyrses 10—30 cm long, primary branches to 16 cm, squarrose to ascending, secondary to 4 cm, bearing fascicles of sweetly-scented (Muscari, Mabberley) flowers; axes short-pubescent; bracts 2—7(—10) mm, narrowly triangular, often caducous, bracteoles similar but smaller; pedicels c. 3—4 mm, articulated with pseudopedicels c. 2 mm long, continuous with calyx. Calyx c. 2.5— 3.5 mm diam., shallowly cupular, + pubescent without lobes obtuse, c. 1/3 length of calyx. Petals c. 12-16 mm long, narrowly oblong to subspathulate, creamy green or yellowish, often tinged pink, subglabrous or puberulent (especially on sectors exposed before anthesis). Staminal tube glabrous, colour as petals, anthers c. 1 mm long, ob- long. Ovary densely adpressed pubescent. /nfructescence pendulous with up to 6 fruits. Capsule (2.5—)3.5—5 cm long, 2.5—4 cm diam., dark brown, lenticellate. Seeds c. 1.2 cm long. — Fig. 53. Distribution — India and Nepal, east- and southeastwards from southern China and Hainan, Sri Lanka and the Andamans to western Malesia: Sumatra (north but rare), Malay Peninsula (but not in the south), Borneo (limestones of Ist Div., Sarawak). Habitat — Rain forest and semi-deciduous forest, including bamboo forests and those on limestone, to 900 m altitude. It is a colonist of bare ground, including road cut- tings in the Malay Peninsula. Vernacular names — Suntang (Mal.); ranggu (Sarawak). Uses — The timber is an export of Burma and India, where boles to 6 m girth have been recorded, and is known in world commerce as chickrassy, yinma or Chittagong wood. On seasoning, it tones down to a golden mahogany shade with a beautiful satiny sheen and is often figured handsomely. In China it is favoured for carving as well as construction work. The tree may be coppiced and it has been tried in tropical Australian plantations, but it is susceptible to the depredations of Hypsipela moths. The tree yields a gum which is marketed mixed with others in India, while the flow- ers are the source of a dye. An extract from the twigs has proved an efficacious anti- feedant against Pieris rapae in southern China [W. & H. Olkowski, IPM Pract. 10, 9 (1988) 3]. The seeds contain some 50% oil but how this is utilized is unrecorded. The bark is astringent. In Europe, the plant has been cultivated as a greenhouse ornamental. Note — The trees from Sarawak and the Malay Peninsula are almost glabrous, whilst in Sumatra both this form and a densely pubescent one have been collected. The 358 Flora Malesiana ser. I, Vol. 12 (1) (1995) latter is paralleled by ‘Chukrasia velutina’ in India and Sri Lanka, where Livera [Ann. Roy. Bot. Gard. Perad. 9 (1925) 308] has argued that it is possible to distinguish two species on the basis of pubescence, the number of anthers and of leaflets, the shape of the latter and so forth. Examination of more copious materials break down these dis- tinctions and there are, in any case, a number of specimens intermediate in pubescence between these extremes. Both the most pubescent and the subglabrous are recorded from India, Burma, Thailand and southern China as well, again with intermediates. Those concerned with provenances for forestry may find it useful to use an informal classification for such forms, the pubescent of which may prove to be ecotypes of more seasonal forests (N.B.: If these forms were to be recognized at the species level, they would have to be known as C. nimmonii, not C. velutina). Moreover, bracts from pop- ulations in the west of the range appear, on the whole, to be longer, while, in Indo- china, there are forms with smaller-than-average fruits and one with juvenile foliage apparently retained at maturity [var. dongnaiensis (Pierre) Pellegr. ]. TRIBUS CEDRELEAE Cedreleae (R.Br.) DC., Prodr. 1 (1824) 624; T.D. Penn. & Styles, Blumea 22 (1975) 511. — Cedrela- ceae R.Br. in Flinders, Voy. Terra Austr. 2, App. (1814) 64. Trees. Flowers with an androgynophore, petals adnate to it. Stamens 5, free, some- times with a whorl of staminodes. Capsule woody or membraneous, with a soft colu- mella. Seeds winged at one or both ends; cotyledons flat, leaf-like; residual endosperm present. Distribution — Two genera, Cedrela L. in tropical America and the following. TOONA (J. M. Edmonds) Toona (Endl.) M. Roem., Synops. Monogr. | (1846) 131, 139; Pellegrin in Fl. Indo-Chine | (1911) 792; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 269; ed. 2, 19b1 (1940) 44, t. 2.; T.D. Penn. & Styles, Blumea 22 (1975) 512; Bahadur, Monogr. Toona (1988) 1-251. — Cedrela L. sect. Toona Endl., Gen. Pl. 2 (1840) 1055. — [Surenus Rumph., Herb. Amb. 3 (1743) 66, t. 39, ex Kuntze, Rev. Gen. Pl. 1 (1891) 110, nom. superfl. pro Toona; T. Post & Kuntze, Lex. Gen. Phan. (1903) 543.] [Cuveraca Jones, Asiat. Res. 4 (1790) 281, nom. inval.] {Mioptrila Raf., Am. Man. Mulb. Trees (1839) 37, nom. provis.] Cedrela L., p.p.: Ridley, Fl. Malay Penins. | (1922) 415; Burkill, Gard. Bull. Str. Settl. 5 (1930) 120; Dict. Econ. Pl. Malay Penins. (1935) 499; Symington, Malay For. 4 (1935) 119. Deciduous or semi-evergreen trees up to 50 m tall, monoecious. Bark fissured, some- times flaking irregularly, grey-brown; inner bark pink to red; sapwood cream. Leaves paripinnate, occasionally imparipinnate. Leaflets entire, serrate or dentate; glabrous or pubescent with simple hairs; club-shaped glands often associated with the vascular sys- tem; domatia (small deltate axillary pockets) usually present on proximal lateral veins of lower surfaces, often bordered with simple hairs. Inflorescences much-branched pen- Mabberley et al. — Meliaceae 359 dent thyrses, often exceeding a metre in length. Flowers 5-merous, unisexual with well- developed vestiges of the opposite sex present, rarely hermaphrodite, small. Calyx 5- (rarely 6-)lobed or of 5 (rarely 6) free sepals, which are imbricate or cupulate in bud; sepal margins always ciliate. Petals 5 (rarely 6), free, longer than the calyx in bud, im- bricate (quincuncial), basally adnate to a short pulvinate (cushion-shaped) androgyno- phore (disk); white, cream or pink. Stamens 5 (rarely 6), free, arising from the andro- gynophore, sometimes alternating with 1—5 filamentous staminodes; anthers in male flowers yellow, dehiscing laterally; antherodes in female flowers often sagittate, brown with abortive pollen. Ovary 5-locular, each locule with 6—10 ovules, vestigial in male flowers; styles short in female flowers, pistillodes long and slender in male flowers; stylehead discoid with stigmatic papillae, usually 5-rayed. Fruit a pendulous thinly woody, ellipsoidal or obovoidal, septifragal capsule; valves 5, brown, smooth to ver- rucose, opening from the apex. Columella softly woody, concavely or convexly 5- angled, extending to the capsule apex. Seeds winged either at both ends when attached towards the distal end of the columella, or at one end when attached by the seed-end to the proximal part of the columella; wings membraneous, seeds with residual endosperm. Cotyledons collateral, flattened, leaf-like; radicle laterally exserted. Germination phane- rocotylar; first true leaves opposite, trifoliolate with deeply lobed or dentate leaflets. Distribution — An Old World genus of 4 or possibly 5 species, extending from east- ern Pakistan through SE Asia and southern China to eastern Australia. Three species occur in Malesia. Habitat — Primary and secondary rain forest, low- to mid-montane, up to 2900 m altitude. Uses — The timbers of Toona spp. are highly prized but generally scarce. Toona ciliata was the most important native timber in Australia but is now largely cut out; the Malesian species seem to be undergoing a similar fate. Extracts from the bark, heart- wood and leaves apparently have insecticidal qualities. Systematics — The genus is closely allied to the neotropical Cedrela with which it has been repeatedly united and separated from since 1840. Pennington & Styles (1975) proposed clear arguments for the recognition of two distinct genera, with Ce- drela being differentiated from Toona by a columnar androgynophore longer than the ovary, and seedlings having entire leaflets. The genera are also distinguishable by various other characters, with chromosomal evidence suggesting different evolution- ary histories. Species of the genus Toona are extremely variable, especially in their vegetative morphology; average parameters have not been included in all of the spe- cies descriptions. Monographic study of this genus has demonstrated that it is composed of very few species, and that phenotypic plasticity and genetic variation are responsible for much of the taxonomic complexity reflected in the literature. The species exhibit a phenomenal range of morphological variation, both within and between trees of the same population, and many of the features used by earlier workers to define their taxa, have proved to be only slight morphological variants. Such vegetative characters include leaf and leaflet 360 Flora Malesiana ser. I, Vol. 12 (1) (1995) size; leaflet and leaflet-margin shape; indumentum type and hair density. In particular, the velutinous pubescence, on which a number of Toona taxa have been based, occurs throughout the genus with the exception of 7. sinensis, both inter- and infra-specifi- cally, and even between seedlings of the same population. Though the monographic revision of these species at the specific level is nearing completion, the floral and vegetative variation observed in the species could represent extreme ranges of polymorphic species, or be genetically stable and eco-geographically correlated, and therefore be given formal recognition. The taxonomy at the infraspecific level is therefore complex and still under review; future conclusions could necessitate some changes in the taxonomy of this genus. KEY -1O THE’ SPECIES la. Bark obnoxiously pungent. Leaflets serrate or dentate. Petal margins, ovary and disk glabrous. Capsule columellas convex without apical scarring. Seeds winged at 059 (cae! 010 ara nd oe Ea nr an SR a, acer a, 1. T. sinensis b. Bark sweetly aromatic. Leaflets entire. Petal margins ciliate; ovary and disk pilose. Capsule columellas concave with apical scarring. Seeds winged at bothends... 2 2a. Leaflets usually conspicuously pilose on upper midribs. Flower bud petals usually with dense central ciliate bands externally; styles pilose. Capsule valves usually verrucose with conspicuous tusty lenticels’ 4. aaeee os ere ae 2. T. sureni b. Leaflets usually glabrescent on upper midribs. Flower bud petals glabrescent to sparsely pilose externally; styles glabrous. Capsule valves usually smooth to mi- nutely lenticellate with small inconspicuous lenticels ...................-2-- Yrs Ue TIE ee ee SU Sere ities aa 3. T. ciliata (and see 3a. T. calantas) 1. Toona sinensis (A. Juss.) M.Roem. Toona sinensis (A. Juss.) M.Roem., Synops. Monogr. | (1846) 139; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 269; Diels, Bot. Jahrb. 29 (1901) 425; Schneider, Illus. Handb. Laubh. 2 (1907) 132; Koord., Exk. Fl. Java 2 (1912) 437; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46; Mitt. Deutsch. Dendrol. Ges. 53 (1941) 183; A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 156; Li, Trop. Woods 79 (1944) 19; Adelb., Blumea 6 (1948) 313; How & Chen, Acta Phytotax. Sin. 4 (1955) 41; Backer & Bakh. f., Fl. Java 2 (1965) 117; Wu, Fl. Yunnan. 1 (1977) 210; L.M. Perry, Med. Plants E & SE Asia (1980) 263; Chen, J. Wuhan Bot. Res. 4 (1986) 186; Bahadur, Monogr. Toona (1988) 115, 124; Mabb. in Tree Fl. Malaya 4 (1989) 256. — Cedrela sinensis A. Juss., Bull. Sci. Nat. Géol. 23 (1830) 241; Linnaea 6 (1831) 115; Mém. Mus. Nat. Hist. Nat. 19 (1832) 255; Carriére, Rev. Hort. (1875) 86; C.DC. in DC., Monogr. Phan. | (1878) 743; Franchet, Nouv. Arch. Mus. Paris 2, 5 (1884) 220 (Plantae David. 1: 68) (‘chinensis’); Forbes & Hemsley, J. Linn. Soc. 23 (1886) 114; André, Rev. Hort. (1891) 573; Kuntze, Rev. Gen. Pl. 1 (1891) 111; Backer, Schoolfl. Java (1911) 219; C.DC., Rec. Bot. Surv. India 3 (1908) 360; Shira- sawa, Icon. Ess. Flor. Japon. 2 (1908) t. 35, f. 1-13 (‘chinensis’); Dunn & Tutcher, Bull. Misc. Info. Kew, Add. Ser. 10 (1912) 58; Sargent, Pl. Wilson. 2 (1916) 156; K. Heyne, Nutt. Pl. Ned.- Indié, ed. 1, 3 (1917) 42; Rehder, J. Arnold Arbor. 7 (1926) 189; Burkill, Gard. Bull. Str. Settl. 5 (1930) 121; Makino, J. Jap. Bot. 8 (1932/33) 341; Troup, Exotic Trees (1932) 64; Li, Trop. Woods 79 (1944) 18; Bean, Trees & Shrubs, ed. 8, 1 (1970) 557. — Surenus sinensis (A. Juss.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. Mabberley et al. — Meliaceae 361 Cedrela serrata Royle, Ill. Bot. Him. (1839) 144, t. 25; Brandis, Forest Fl. NW & C India (1874) 73; Kurz, J. As. Soc. Beng. 44, 1i (1875) 150; For. Fl. Brit. Burma | (1877) 229; C.DC. in DC., Monogr. Phan. | (1878) 742; Gamble, Man. Indian Trees (1880) 79; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 204; Kanjilal, Forest Fl. School Circle, NWP (1901) 60; Collett, Fl. Sim- lensis (1902) 83; Brandis, Indian Trees (1906) 145; C.DC., Rec. Bot. Surv. India 3 (1908) 361; Lace, List Trees Burma (1922) 31; Burkill, Gard. Bull. Str. Settl. 5 (1930) 121; Troup, Exotic Trees (1932) 64; Midon, Malays. For. Serv. Trade Leaflet 93 (1985). — Toona serrata (Royle) M. Roem., Synops. Monogr. | (1846) 139; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 269; Briq., Mém. Inst. Nat. Genev. 24 (1935) 31; Harms in Engl. & Prantl, Nat. Pflanzen- fam., ed. 2, 19b1 (1940) 47; A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 158; Hara & Williams, Enum. Flow. Pl. Nepal 2 (1979) 86; Stainton, Flowers Himal. (1988) 11; Styles & White, Flora Trop. East Africa, Meliaceae (1991) 46. — Surenus serrata (Royle) Kuntze, Rev. Gen. Pl. 1 (1891) BLE. Mioptrila odorata Raf., Am. Man. Mulb. Trees (1839) 37, nom. provis., excl. fol. (= Zanthoxylum sp.). Toona longifolia (Wall. ex M. Roem., Synops. Monogr. | (1846) 140, nom. nud., ex] Harms in Engl. & Prantl, Nat. Pflanzenfam., ed 2, 19b1 (1940) 46. Cedrela serrulata Miq., Fl. Ind. Bat., Suppl. (1861) 508; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 64; C.DC. in DC., Monogr. Phan. | (1878) 746. — Surenus serrulata (Miq.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. — Toona serrulata (Miq.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 269; ibid., ed. 2, 19b1 (1940) 46. Ailanthus flavescens Carriere, Rev. Hort. (1865) 366 (‘Ailantus’). Cedrela glabra C.DC. in DC., Monogr. Phan. | (1878) 742; Blanco, Fl. Filip. 4, Nov. App. (1880) 45. — Surenus glabra (C.DC.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. — Toona glabra (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 269; Panigrahi, Bangl. J. Bot. 3 (1974) 54. — [Cedrela longifolia Wall., Cat. (1829) n. 1273, nom. nud. ex C.DC., Rec. Bot. Surv. India 3 (1908) 375, nom. superfl. pro Cedrela glabra.) — Toona sinensis (A. Juss.) M. Roem. var. glabra (C.DC.) Bahadur, Monogr. Toona (1988) 120. Cedrela longifolia Wall. ex C.DC. var. kumaona C.DC., Rec. Bot. Surv. India 3 (1908) 376. Cedrela serrata Royle var. puberula C.DC., Rec. Bot. Surv. India 3 (1908) 363. — Toona sinensis (A. Juss.) M. Roem. var. puberula (C.DC.) Bahadur, Monogr. Toona (1988) 122. Cedrela sinensis A. Juss. var. hupehana C.DC., Rec. Bot. Surv. India 3 (1908) 361. — Toona sinensis (A. Juss.) M. Roem. var. hupehana (C.DC.) A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 156. Cedrela sinensis A. Juss. var. schensiana C.DC., Rec. Bot. Surv. India 3 (1908) 361; Li, Trop. Woods 79 (1944) 19. — Toona sinensis (A. Juss.) M. Roem. var. schensiana (C.DC.) Wu, Fl. Yunnan. | (1977) 210, comb. invalid.?; X.M. Chen, J. Wuhan Bot. Res. 4 (1986) 187. Toona sinensis (A. Juss.) M. Roem. var. grandis Pamp., Nuov. Giorn. Bot. Ital. 18 (1911) 171; A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 156. Cedrela sinensis A. Juss. var. lanceolata Li, Trop. Woods 79 (1944) 19. Toona microcarpa (C.DC.) Harms var. denticulata A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 158. — Toona sinensis (A. Juss.) M. Roem. var. denticulata (A. Chev.) Bahadur, Monogr. Toona (1988) 118. Toona microcarpa (C.DC.) Harms var. grandifolia A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 158. — Toona sinensis (A. Juss.) M. Roem. var. grandifolia (A. Chev.) Bahadur, Monogr. Toona (1988) 121. Toona sinensis (A. Juss.) M. Roem. var. incarvillei A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 156. Tree to 40 m; bole to 20 m and girth to 150 cm, buttressed. Bark grey to dark brown, fissured; inner bark pink to red, fibrous; sapwood cream, fibrous; smelling strongly of garlic and pepper when cut. Leaves 32—120 cm long, usually 8—20-jugate; rachides gla- brous to pilose, often reddish; petioles 5.5—20 cm long, glabrous to pilose. Leaflets narrowly lanceolate to linear-lanceolate, serrate or serrulate, rarely entire, 1 1—22 by 3— 5.6 cm, glabrous to pilose, especially on the midribs and veins, club-shaped glandular 362 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 54. Toona sinensis (A. Juss.) M. Roem. a. Leaf and inflorescence; b. female flower; c. ditto with ovules; d. male flower; e. ditto with vestigeal ovules; c & e in longitudinal section; f. infructescence with open capsules; g. seed (a: Lace 961; b, c: Symington 30980; d, e: Whitmore 20098; f, g: Clemens 6218). — T. sureni (Blume) Merr. h. Open capsule; i. seed (h, i: Boschproefstation 11976). Drawing R. Wise. Mabberley et al. — Meliaceae 363 hairs conspicuous on upper midrib areas and junctions between leaf rachides and petio- lules, apex acuminate, base asymmetric; petiolules 3—9 mm long, glabrescent. /nflores- cences up to | m, pendent, rachides glabrescent, pilose or villous with short appressed or spreading simple hairs; pedicels c. 0.5 mm, glabrescent to pilose. Flowers 3.5—4.5 mm long. Calyx cupulate, 1—1.5 mm long, glabrous externally; sepals 0.5—1.1 by 0.6— 1.75 mm, margins ciliate. Petals white or flushed pink, 2.8—4.2 by 1.1—2.9 mm, gla- brous externally, margins non-ciliate. Androgynophore 2.5—4 mm long, glabrous. Sta- minodes always present, usually 5, sometimes 1—4, 0.75—1.75 mm long, glabrous. Fi- laments 1.3—1.75 mm long (male flowers), 1—1.5 mm long (female flowers), glabrous. Anthers 0.8—1.2 mm long, 0.5—0.75 mm broad. Antherodes 0.3—0.8 mm long, 0.3— 0.5 mm broad, apices apiculate, sometimes extended. Disk 1—1.5 mm diam., orange, glabrous. Ovary 1.6—2.25 mm diam., glabrous; locules to 6-ovulate. Style 1.1—1.5 mm long, 0.5 mm broad (male), 0.5—0.8 mm long, 0.3—0.4 mm broad (female), glabrous; stylehead 0.5—1 mm diam. Capsule 15—30 mm long; columella 12—29 mm long, 6—11 mm broad, convex; valves reddish to dark brown, 15—30 by 4—7 mm, elenticellate and smooth to punctate with small (0.3—0.5 mm diam.) scattered lenticels. Seeds winged at one end, 8—16 by 3.5—6.2 mm; seed body 8-10 by 1.7—4 mm. — Fig. 54a—g. Distribution — Highlands of tropical Asia from India, Nepal and China through Bur- ma, Thailand to Malesia: Sumatra (W & E Coast, Palembang, Tapanuli), Peninsular Malaysia (Perak, Pahang), Borneo (Sabah), Java (Preanger, Cibodas, Mt Merbabu). It is planted in India and Sri Lanka for shade, and in various European cities (e.g. Paris, where introduced via Japan in 1862, the original tree in the rue Buffon being felled in the 1970s) as an avenue tree. Habitat — An upland species (350—2000 m altitude), usually confined to primary montane forests, where it is found on light, steep hillside slopes often near streams; also occurring in secondary forests and disturbed sites. Vernacular names — Peninsular Malaysia: surian bawang, surian wangi; Borneo (Sabah): rangau (Dusun), limpaga (Malay); Sumatra: ingaal; Java & Sumatra: soeren, soerian, suren. Uses — The timber is used for furniture and sieve hoop making, and in bridge con- struction. The leaves are used as food in China and Malaysia, and as animal fodder in India (Burkill 1930). The trees are widely used medicinally, with the bark being used as an astringent and depurative; powdered root as a refreshment and a diuretic, and tender leaves as a carminative (Perry 1980). Cultivated plants in Europe include cy. Flamingo with brightly coloured young leaves. Note — The leaf morphology and indumentum types and density are particularly variable in this species, with capsule valve form also showing some variability. On the basis of such features, eight infraspecific variants have been recognized by other authors. 2. Toona sureni (Blume) Merr. Toona sureni (Blume) Merr., Interpr. Rumph. Herb. Amb. (1917) 305; Enum. Philipp. Flow. Pl. 2 (1923) 357; Merr. & Chun, Sunyatsenia | (1930) 62; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 45; Adelb., Blumea 6 (1948) 313; How & Chen, Acta Phytotax. Sin. 4 (1955) 364 Flora Malesiana ser. I, Vol. 12 (1) (1995) 39; Backer & Bakh.f., Fl. Java 2 (1965) 117; Hara & Williams, Enum. Flow. Pl. Nepal (1979) 86; L.M. Perry, Medic. Pl. E & SE Asia (1980) 263; Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 345, t. 264; Bahadur, Monogr. Toona (1988) 129; Mabb. in Tree Fl. Malaya 4 (1989) 258. — Swietenia sureni Blume, Cat. Gew. Buitenzorg (1823) 72. — Cedrela sureni (Blume) Burkill, Gard. Bull. Str. Settl. 5 (1930) 121; Dict. Econ. Prod. Malay Penins. (1935) 499. [Surenus rubra Rumph., Herb. Amb. 3 (1750) t. 39; cf. Hassk., Neue Schluss. Rumph. Herb. Amb. (1886) 51.] [Azederach recisa Noronha, Verh. Batav. Gen., ed. 5, | (1790/91) art. 4, 5 (‘recisum’), nom. nud.; cf. Mabb., Gard. Bull. Sing. 37 (1984) 62.] Cedrela febrifuga Blume, Verh. Batav. Gen. 9 (1823) 135; Bijdr. (1825) 180; Nees, J. Pharm. 11 (1825) 518; Sprengel, Syst. Veg., ed. 16, 4 (1827) 95; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris (1830) 255; G. Don, Gen. Syst. 1 (1831) 687; Forsten, Diss. Cedrela febrifuga, Univ. Leiden (1836) 18; Hassk., Hort. Bogor. Descr. (1858) 129; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 63; C.DC. in DC., Monogr. Phan. | (1878) 744; King, J. As. Soc. Beng. 64, i1 (1895) 89; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 197; Brandis, Indian Trees (1906) 146; C.DC., Rec. Bot. Surv. India 3 (1908) 372; Backer, Schoolfl. Java (1911) 219; K. Heyne, Nutt. Pl. Ned.-Indié, ed. 1, 3 (1917) 40; Lace, Trees Shrubs Burma (1922) 31; Ridley, Fl. Malay Penins. 1 (1922) 415. — Toona febrifuga (Blume) M.Roem., Synops. Monogr. | (1846) 139; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 269; Pierre, Fl. For. Cochinch. 5, fasc. 23 (1897) t. 358; Pellegr. in Fl. Indo-Chine | (1911) 793; Koord., Exk. Fl. Java 2 (1912) 437; Atlas 1 (1913) f. 168; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1l (1940) 46. — Surenus febrifuga (Blume) Kuntze, Rev. Gen. PI. 1 (S91) 11 Te Cedrela febrifuga Blume var. assamensis C.DC., Rec. Bot. Surv. India 3 (1908) 373; Burkill, Rec. Bot. Surv. India 10 (1925) 257. Cedrela febrifuga Blume var. glabrior C.DC. in DC., Monogr. Phan. | (1878) 744, p.p.; Rec. Bot. Surv. India 3 (1908) 373; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 199. — ?Toona febrifuga M. Roem. var. glabrior (C.DC.) Pierre, Fl. For. Cochinch. 5, fasc. 23 (1897) t. 358, p.p.; Pellegr. in Fl. Indo-Chine | (1911) 794. Cedrela febrifuga Blume var. pealii C.DC., Rec. Bot. Surv. India 3 (1908) 374. ?Cedrela febrifuga Blume var. verrucosa C. DC., l.c. ?Cedrela toona Roxb. ex Rottl. & Willd. var. pilistila C.DC., Rec. Bot. Surv. India 3 (1908) 365. Cedrela toona Roxb. ex Rottl. & Willd. var. pilistaminea C.DC., op. cit. (1908) 366, p.p. Cedrela toona Roxb. ex Rottl. & Willd. var. yunnanensis C. DC., l.c. — Toona ciliata M. Roem. var. yunnanensis (C.DC.) C.Y. Wu, Fl. Yunnan. | (1977) 207, ?nom. superfl.; Chen, J. Wuhan Bot. Res. 4 (1986) 187. ?Cedrela toona Roxb. ex Rottl. & Willd. var. henryi C.DC., Rec. Bot. Surv. India 3 (1908) 369. — Toona ciliata M. Roem. var. henryi (C.DC.) C.Y. Wu, Fl. Yunnan. | (1977) 209, ?nom. superfl. ?Cedrela toona Roxb. ex Rottl. & Willd. var. warburgii C.DC., Rec. Bot. Surv. India 3 (1908) 370. Cedrela microcarpa C.DC. var. grandifoliola C.DC., op. cit. (1908) 371. — Toona ciliata M. Roem. var. grandifoliola (C.DC.) Bahadur, Monogr. Toona (1988) 91. ?Toona ciliata M. Roem. var. candollei Bahadur, Monogr. Toona (1988) 99. Toona ciliata M. Roem. var. hainesii (C.DC.) Panigrahi & Mishra, Ind. J. For. 11 (1988) 141, p.p. Toona microcarpa (C.DC.) Harms var. pilipetala (C.DC.) Bahadur, Monogr. Toona (1988) 113, p.p. ?Toona microcarpa (C.DC.) Harms var. sahnii Bahadur, Monogr. Toona (1988) 114. Medium- to large-sized tree to 40 m tall and > 3 m girth; bole to 25 m, with or with- out buttresses (to 2 m); crown fairly wide, spreading, occasionally dense. Bark whitish, grey-brown, grey or light-brown, usually vertically fissured and flaking; inner bark pinkish-white, -brown, reddish-brown or -orange, fibrous; sapwood white, pink or pale red; sweetly aromatic when cut. Twigs often prominently lenticellate with verru- cose lenticels. Leaves 29—84 cm long, usually 6—9(—12)-jugate; rachides glabrescent to moderately pilose, occasionally velutinous; petioles 7-12 cm long, glabrescent to pilose, Mabberley et al. — Meliaceae 365 Fig. 55. Toona trees, probably T. sureni (Blume) Merr., planted for shade and roadside boundary in Cen- tral Java. Provenance of photograph not known. often lenticellate. Leaflets lanceolate to ovate-lanceolate, margins entire, 7—14(—19.5) by 3—6(—7) cm, glabrescent to moderately pilose, usually with short hairs and club-glands on the upper midribs, lower vascular system pilose, apices acuminate, occasionally acute, bases symmetrical to asymmetrical; petiolules (0.2—)0.4—1.2 cm long, glabrescent to pilose/villous. Inflorescences up to 40 cm long, pendent, sweetly aromatic; rachides pilose to villous with medium to long spreading hairs, occasionally glabrescent; pedicels (0.3—)0.75—1.25 mm long, pilose to villous. Flowers 4—5 mm long. Calyx 1—1.5 mm long, glabrescent to pilose externally, lobes imbricate; sepals usually triangular, espe- cially in bud, 0.6—1 by 0.8—1.5 mm, glabrescent to villous externally, apices usually acute, margins ciliate. Petals white, creamy-white or pinkish, 3.5—5 by 1.6—3.2 mm, glabrescent to villous, but usually with conspicuous central bands of long appressed hairs in bud, margins ciliate with long hairs. Androgynophore 2.5—4.7 mm long. Fila- ments 1.25—2.5 mm long (male flowers), 1—1.3 mm long (female flowers), pilose to villous with scattered to dense long hairs. Anthers 0.75-1.25 mm long, 0.3—0.8 mm broad, apices usually apiculate. Antherodes 0.5—0.9 mm long, 0.25—0.6 mm broad, sagittate. Disk 1.25—2.5 mm diam., orange to red, densely pilose. Ovary 1.6—2.75 mm diam., moderately to densely pilose; locules to 6-ovulate. Style 1.25—3 mm long, 0.25— 0.5 mm broad (male), 0.5—1 mm long, c. 0.3 mm broad (female), pilose with scattered 366 Flora Malesiana ser. I, Vol. 12 (1) (1995) usually appressed hairs especially on the lower half; stylehead 0.75—1.25 mm diam. Capsule 14—20(—24) mm long; columella 14—20(—24) mm long, 5—8(—10) mm broad, concave with apical scarring; valves dark- to blackish-brown, rough, verrucose with conspicuous, often ovate rusty lenticels, 0.3—2 by 0.4—1.25 mm. Seeds winged at both ends, 11—20(—22) by (3—)4—4.8 mm, wings unequal with broadly obtuse apices; seed body 5—8 by 1.5—2 mm. — Fig. 54h, i, 55. Distribution — From India, through Nepal, Bhutan, Burma, S China, Thailand to Malesia: Sumatra (W Coast, Tapanuli, Lampongs, Madailing, Karolands), Peninsular Malaysia (Selangor, Pahang, Kelantan, Kedah, Perlis, Penang, Langkawi, Mersing, Aur), Borneo (Sabah, Sarawak, Kalimantan), Java (W & E), Lesser Sunda Islands (Bali, Sumba, Timor), Moluccas (Halmahera, Tanimbar), Celebes (N, Kawata, Amu- rang), New Guinea (Irian Jaya: Kebar Valley; Papua New Guinea: Chimbu Valley). Habitat — Common to rare, in primary and secondary forests, often in logged and disturbed areas, usually on riparian hillsides or slopes, from sea-level to 1700 m alti- tude. Vernacular names — Sumatra: serijan (Lampongs), ?para, soerian amba, ingoe; Ma- lay Peninsula: soren, surian, surian wangi, ?tangipan burang; Borneo (Kalimantan): suren, ?mesal; Java: kiketaro, odjang, soren, soeren, soeren keling, suren (Madura), redani; New Guinea: boerwam, moeroewan (Kebar), ongguno (Kuman), muwa (Ma- ring); Philippines, official common name: danupra. Uses — The timber is said to be ‘excellent’ (Don 1832). Medicinally, the bark is used as a powerful astringent and a purative throughout its range [see Horsfield, Verh. Batav. Gen. Wet. 8 (1816) 125]; in Indochina it is considered to be a tonic, an antiperi- odic and an antirheumatic, while in Indonesia it is used as an astringent and a tonic for treating diarrhoea, dysentery and other intestinal infections. Leaf extracts apparently have an antibiotic activity against Staphylococcus, with leaf tip concoctions being ap- plied to swellings (Perry 1980). Note — There are several large-fruited variants of this species, whose taxonomic status has yet to be determined. The most striking was described as Cedrela celebica Koord. [Minah. (1898) 636] from NE Celebes, where it is known as ‘lalumpehe’. Flowers of this variant are rarely collected, but its capsules are morphological enlarge- ments of those found in T. sureni. They exhibit the following characteristics: 35—42 mm long; columellas 29—38 mm long, 13—18 mm broad, valves verrucose, 36—41 by 9-11 mm, with large rusty lenticels 0.75—1.25 mm diam.; seeds 18—29 by 6—9 mm. The tree is used for furniture manufacture, and in house and boat construction. Whether it is merely a large-fruited variant of 7. swreni or a distinct geographical taxon remains to be resolved. 3. Toona ciliata M. Roem. Toona ciliata M. Roem., Synops. Monogr. | (1846) 139; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 270; Merr., Dept. Int. For. Bur. Bull. 1 (1902) 27; Bull. Dept. Agric. Nat. Res. Bur. For. Philipp. | (1903) 27; Brig., Mém. Inst. Nat. Genev. 24 (1935) 32; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 45; Kitamura in Kihara, Fauna & Flora Nepal Himal. | Mabberley et al. — Meliaceae 367 (1955) 170; Wu, Fl. Yunnan. | (1977) 207; Hara & Williams, Enum. Flow. Pl. Nepal 2 (1979) 85; Chen, J. Wuhan Bot. Res. 4 (1986) 187; Panigrahi & Mishra, Ind. J. For. 11 (1988) 140; Baha- dur, Monogr. Toona (1988) 64; Styles & White, Fl. Trop. E Afr., Meliaceae (1991) 45; Taylor & Harden in Harden (ed.), Fl. NSW 2 (1991) 279. — Cedrela toona Roxb. ex Rottl. & Willd., Neue Schr. Naturf. Freunde Berlin 4 (1803) 198; Roxb., Hort. Bengal. (1814) 18; Plant. Coromand. 3 (1819) 33; Roem. & Schultes, Syst. Veg. 5 (1819) 464; DC., Prodr. 1 (1824) 624; Roxb., FI. Ind. 2 (1824) 423; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 255; G. Don, Gen. Syst. | (1831) 687; Wight & Arn., Prodr. 1 (1834) 124; Wight, Ic. 1 (1840) pl. 161; Hassk., Hort. Bogor. Descr. (1858) 127; Miq., Fl. Ind. Bat. 1 (1859) 548; Suppl. (1861) 197; Dalzell & Gibson, Bom- bay Fl. (1861) 38; Benth., Fl. Austral. 1 (1863) 387; Bedd., Fl. Sylv. S Ind. 1 (1869) 10; Hiern in Hook. f., Fl. Brit. India 1 (1872) 568; Brandis, For. Fl. NW & C India (1874) 72; Kurz, For. FI. Brit. Burma | (1877) 228; C.DC. in DC., Monogr. Phan. | (1878) 745; Gamble, Man. Ind. Trees (1880) 77; Blanco, Fl. Filip. 4, Nov. App. (1880); Franchet, Bull. Soc. Bot. Fr. 33 (1886) 452; Talbot, Trees Shrubs Bombay (1894) 42; ed. 2 (1902) 80; Kanjilal, For. Fl. School Circle, NWP (1901) 60; Collett, Fl. Simlensis (1902) 83; Cooke, Fl. Pres. Bombay | (1903) 217; Duthie, FI. Upper Gang. Plain 1 (1903-05) 153; Brandis, Indian Trees (1906) 145; C.DC., Rec. Bot. Surv. India 3 (1908) 364; Haines, For. Fl. Chota Nagpur (1910) 249; Burkill, Rec. Bot. Surv. India 4 (1913) 102; Haines, Bot. Bihar & Orissa 2 (1921) 174; Lace, Trees Shrubs Burma (1922) 32; Craib, Fl. Siam. Enum. | (1926) 267; Burkill, Gard. Bull. Str. Settl. 5 (1930) 121; Troup, Exotic Trees (1932) 65; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 500; Li, Trop. Woods 79 (1944) 22. — [Swietenia toona Stokes, Bot. Mat. Med. 2 (1812) 480, nom. nud., ‘Suitenia toon’.| — Cedrela australis Mudie, Pict. Austral. (1829) 147, nom. superfl. pro C. toona; cf. Mabb., Austral. Syst. Bot. Soc. Newsl. 70 (1992) 14. — Surenus toona (Rottl. & Willd.) Kuntze, Rev. Gen. Pl. 1 (1891) 1 fu ?Cedrela velutina DC., Prodr. 1 (1824) 625; G. Don, Gen. Syst. 1 (1831) 687; C.DC. in DC., Monogr. Phan. 1 (1878) 747. — ?Toona velutina (DC.) M. Roem., Synops. Monogr. | (1846) 139. — ?Su- renus velutina (DC.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. — ?Toona ciliata M. Roem. subsp. velu- tina (DC.) Bahadur, Monogr. Toona (1988) 97. Cedrela hexandra Wall. in Roxb., Fl. Ind. (1824) 425, nom. in obs.; Aitken, Trans. Soc. Arts 48 (1833) 448. — Toona hexandra (Wall.) M. Roem., Synops. Monogr. | (1846) 139. — Toona ciliata M.Roem. var. hexandra (Wall.) Bahadur, Monogr. Toona (1988) 93. Cedrela australis F. Muell., Fragm. Phyt. Austral. 1 (1858) 4, non Mudie; C.DC. in DC., Monogr. Phan. 1 (1878) 743. — Surenus australis Kuntze, Rev. Gen. Pl. 1 (1891) 111. — Toona australis Harms in Engl. & Prantl., Nat. Pflanzenfam. 3, 4 (1896) 270; Briq., Mém. Inst. Nat. Genev. 24 (1935) 31. ?Cedrela teysmannii Hassk., Hort. Bogor. 1 (1858) 133; Miq., Fl. Ind. Bat. 1, 2 (1859) 549; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 63; C.DC. in DC., Monogr. Phan. 1 (1878) 747 — Surenus teysmannii (Hassk.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. — ?Toona sureni (Blume) Merr. var. teysmannii (Hassk.) Bahadur, Monogr. Toona (1988) 143. Cedrela toona Roxb. ex Rottl. & Willd. var. parviflora Benth., Fl. Austral. 1 (1863) 387; C.DC., Rec. Bot. Surv. India 3 (1908) 368. — Toona ciliata M. Roem. var. parviflora (Benth.) Bahadur, Monogr. Toona (1988) 94. Cedrela febrifuga Blume f. ternatensis Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 63. — Toona terna- tensis (Miq.) Bahadur, Monogr. Toona (1988) 144. Cedrela febrifuga Blume f. timorensis Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 63. Cedrela microcarpa C.DC. in DC., Monogr. Phan. | (1878) 745; Rec. Bot. Surv. India 3 (1908) 370; Brandis, Indian Trees (1906) 145; Lace, Trees Shrubs Burma (1922) 31; Craib, Fl. Siam. Enum. | (1926) 267; Chun, Fl. Kwantung (1934) 257; Merr. & Metcalfe, Lingn. Sc. J. 16 (1937) 196; Li, Trop. Woods 79 (1944) 20. — Surenus microcarpa (C.DC.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. — Toona microcarpa (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 270; Pellegr. in Fl. Indo-Chine 1 (1911) 795; Brig., Mém. Inst. Nat. Genev. 24 (1935) 32; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46; How & Chen, Acta Phytotax. Sin. 4 (1955) 40; Wu, Fl. Yunnan. 1 (1977) 209; Hara & Williams, Enum. Flow. Pl. Nepal (1979) 86; Chen, J. Wuhan Bot. Res. 4 (1986) 188; Bahadur, Monogr. Toona (1988) 109. 368 Flora Malesiana ser. I, Vol. 12 (1) (1995) Cedrela toona Roxb. ex Rottl. & Willd. var. pubescens Franch., Bull. Soc. Bot. Fr. 33 (1886) 452; Pl. Delav. (1889) 126; C.DC., Rec. Bot. Surv. India 3 (1908) 369; Li, Trop. Woods 79 (1944) 24. — Toona ciliata M. Roem. var. pubescens (Franch.) Hand.-Mazz., Symb. Sin. 7 (1933) 631; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 45; Wu, Fl. Yunnan. | (1977) 207; Chen, J. Wuhan Bot. Res. 4 (1986) 187. — Toona sureni (Blume) Merr. var. pubescens (Franch.) Chun in How & Chen, Acta Phytotax. Sin. 4 (1955) 40. Toona febrifuga (Blume) M. Roem. var. cochinchinensis Pierre, Fl. For. Cochinch. 5, 23 (1897) t. 358; Pellegr. in Fl. Indo-Chine 1 (1911) 795. — Cedrela febrifuga Blume var. cochinchinensis (Pierre) C.DC., Rec. Bot. Surv. India 3 (1908) 374. — Toona sureni (Blume) Merr. var. cochinchinensis (Pierre) Bahadur, Monogr. Toona (1988) 135. Toona febrifuga (Blume) M. Roem. var. griffithiana Pierre, Fl. For. Cochinch. 5 (1897) 23, t. 358. Toona febrifuga (Blume) M. Roem. var. ternatensis Pierre, Fl. For. Cochinch. 5 (1897) 23. ? Cedrela hainesii C.DC., Rec. Bot. Surv. India 3 (1908) 363, p.p. — ?Toona hainesii (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 47, p.p.; Bahadur, Monogr. Toona (1988) 108. — ?Cedrela toona Roxb. ex Rottl. & Willd. var. hainesii (C.DC.) Haines, Bot. Bihar & Oris- sa 2 (1921) 174. — Toona ciliata M. Roem. var. hainesii (C.DC.) Panigrahi & Mishra, Ind. J. For. 11 (1988) 141, p.p. Cedrela toona Roxb. ex Rottl. & Willd. var. latifolia Miq. ex C.DC., Rec. Bot. Surv. India 3 (1908) 365. — Toona ciliata M. Roem. var. latifolia (C.DC.) Chandr., Biol. Mem. 2 (1977) 28. Cedrela toona Roxb. ex Rottl. & Willd. var. nepalensis C.DC., Rec. Bot. Surv. India 3 (1908) 365. — Toona ciliata M. Roem. subsp. nepalensis (C.DC.) Panigrahi, Bangl. J. Bot. 3 (1974) 54. Cedrela toona Roxb. ex Rottl. & Willd. var. cuspidata C.DC., Rec. Bot. Surv. India 3 (1908) 366, p.p. Cedrela toona Roxb. ex Rottl. & Willd. var. pilistaminea C.DC., l.c., p.p. Cedrela toona Roxb. ex Rottl. & Willd. var. gamblei C.DC., Rec. Bot. Surv. India 3 (1908) 367. Cedrela toona Roxb. ex Rottl. & Willd. var. pilipetala C.DC., |.c., p.p. — Toona microcarpa (C.DC.) Harms var. pilipetala (C.DC.) Bahadur, Monogr. Toona (1988) 113, p.p. Cedrela toona Roxb. ex Rottl. & Willd. var. stracheyi C.DC., Rec. Bot. Sury. India 3 (1908) 367. Cedrela toona Roxb. ex Rottl. & Willd. var. talbotii C.DC., l.c. Cedrela toona Roxb. ex Rottl. & Willd. var. australis C.DC., Rec. Bot. Surv. India 3 (1908) 368; Lane- Poole, For. Res. Papua New Guinea (1925) 100 (‘Cedrella’). — Toona ciliata M. Roem. vat. austra- lis (C.DC.) Bahadur, Monogr. Toona (1988) 78, nom. invalid. Cedrela toona Roxb. ex Rottl. & Willd. var. pubinervis C.DC., Rec. Bot. Surv. India 3 (1908) 368. — Toona ciliata M. Roem. var. pubinervis (C.DC.) Bahadur, Monogr. Toona (1988) 96. Cedrela toona Roxb. ex Rottl. & Willd. var. puberula C.DC., Rec. Bot. Surv. India 3 (1908) 369. Cedrela toona Roxb. ex Rottl. & Willd. var. sublaxiflora C.DC., 1.c. — Toona ciliata M. Roem. var. sublaxiflora (C.DC.) C.Y. Wu, Fl. Yunnan. | (1977) 209. Cedrela kingii C.DC., Rec. Bot. Surv. India 3 (1908) 371. — Toona kingii (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46. — Toona ciliata M. Roem. var. kingii (C.DC.) Bahadur, Monogr. Toona (1988) 100. Cedrela kingii C.DC. var. birmanica C.DC., Rec. Bot. Surv. India 3 (1908) 372. ?Cedrela mannii C.DC., Rec. Bot. Sury. India 3 (1908) 374. — ?Toona mannii (C.DC.) Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46. Cedrela toona Roxb. ex Rottl. & Willd. var. hasletii Haines, For. Fl. Chota Nagpur (1910) 250; Bot. Bihar & Orissa 2 (1921) 174. — Toona ciliata M. Roem. var. hasletii (Haines) Panigrahi & Mishra, Ind. J. For. 11 (1988) 141. — Toona ciliata M. Roem. var. hasletii (Haines) Bahadur, Monogr. Toona (1988) 92 (‘haslettii’ ). Cedrela toona Roxb. ex Rottl. & Willd. var. vestita C.T. White, Queens]. Agr. J. 13 (1920) 66. — Toona ciliata M. Roem. var. vestita (C.T. White) Bahadur, Monogr. Toona (1988) 106. Cedrela mollis Hand.-Mazz., Anzeig. Akad. Wiss. Wien 57 (1920) 266. — Toona mollis (Hand.-Mazz.) A. Chev., Rev. Bot. Appl. Agr. Trop. 24 (1944) 157. — Toona ciliata M. Roem. var. mollis (Hand.- Mazz.) Bahadur, Monogr. Toona (1988) 101. Cedrela toona Roxb. ex Rottl. & Willd. var. multijuga Haines, Bot. Bihar & Orissa 2 (1921) 174. Cedrela brevipetiolulata Haines, Bot. Bihar & Orissa 1 (1921) 174, nom. provis. — Toona ciliata M. Roem. var. brevipetiolulata (Haines) Mishra & Panigrahi, Ind. J. For. 11 (1988) 140. Mabberley et al. — Meliaceae 369 Medium- to large-sized tree to 35 m tall and 152 cm girth; bole to 22 m, with or with- out buttresses (to 3.5 m); crown usually rounded and spreading, occasionally dense. Bark greyish-white to brown, usually fissured and flaking; inner bark brown to reddish, fibrous; sapwood white to pink or red; smelling strongly of cedar when cut. Leaves (15—)26—69 cm long, usually (5—)9—15-jugate; rachides glabrous to sparsely pilose, often reddish; petioles 6—11 cm long, glabrous to pilose. Leaflets lanceolate to ovate- lanceolate, margins entire, (7—)9—12.8(—16) by (2.2—)3.2—5(—6) cm, glabrescent with hairs on upper midribs rare, apex acute to acuminate, base usually asymmetric; petio- lules 0.2—1(—1.4) cm long, glabrescent. /nflorescences up to 55 cm, pendent, fragrant; rachides pilose to pilose-villous with short to long, spreading or appressed hairs; pedicels 0.5—1 mm, usually pilose, occasionally villous. Flowers 3.5—5(—6) mm long. Calyx 0.75—1.25 mm long, usually glabrescent externally, lobes imbricate; sepals spathulate, (0.4—)0.75—1 by (0.5—)0.75—1.25 mm, margins ciliate. Petals white to creamy-white, 3.5—5.8 by 1.3—3.1 mm, usually glabrescent occasionally pilose externally, margins ciliate with long hairs. Androgynophore (1.75—)3—4.9(—5.5) mm long. Filaments 1.25—2.5 mm long (male flowers), 0.75—1.75 mm (female flowers), glabrous to pilose/ villous. Anthers 0.6—1.1 mm long, 0.4—0.9 mm broad, apices usually apiculate, often with long appendage. Antherodes 0.5—0.9 mm long, 0.3—0.6 mm broad, usually sagit- tate, often with long apiculate appendage. Disk 1.25—2.5 mm diam., reddish-orange, densely pilose. Ovary 1.25—1.8 mm diam., moderately pilose; locules to 8-ovulate. Style 1.2—3 mm long, 0.2—0.4 mm broad (male), 0.3—1.5 mm long, c. 0.3 mm broad (female), glabrous; stylehead 0.75—1.25 mm diam. Capsule 15—20(—25) mm long; columella 15—20(—24) mm long, 5—7(—10) mm broad, concave with apical scarring; valves reddish-brown, smooth to lenticellate with small (0.1—0.5 mm diam.) scattered lenticels. Seeds winged at both ends, 11-19 by 2.5—4(—5.8) mm, wings unequal, api- ces narrowly obtuse; seed body 5—7 by 1.2—3 mm. Distribution — Widely distributed within the generic range, from Pakistan through India, Bangladesh, South China, Burma, Thailand to Malesia: Sumatra (East & West Coast, Lampongs), Bangka Islands, Peninsular Malaysia (Perak, Kedah, Langkawi), Borneo (Sabah), Java (W, C & E), Philippines (Luzon, Mindoro), Celebes, Lesser Sunda Islands (Flores, Timor), Moluccas (Ternate, Tanimbar), New Guinea (Irian Jaya: Manokwari; Papua New Guinea: Morobe Prov., Koitani, Yalu, Moresby Distr.). Also in New Britain and eastern Australia. Commonly cultivated as an avenue tree in India. Habitat — Common to scarce, in primary and disturbed, often riparian, rain forests from sealevel to 1500 m altitude, occasionally higher. Vernacular names — Sumatra: ingoe (?), serian (Lampongs), limprah, taal-batak (Karo), gerpa, ? keritang; Malaya: suryan, surian; Java: soeren, soeren-kapar, ?ki bewan, suren batu (Djasilin), soeren-mal, beroeh; Timor: hoerenii, surian (Dawan), ai seriq (Tetum), hao lonis; Sabah: surian-limpaga, limpaga (Kwijau), ranggoh (Dusun); Celebes: malapoga (Baree lang.), mapala (Koemea); Moluccas: kukoru, maroa, ladeje (Otimmer); Philippines: danupra; New Guinea: boerwaan (Kebar), djoer-er (Tor), ibio (Manikiong), seba (Mooi), maroa, mafus (Yalu), soi (Waria). 370 Flora Malesiana ser. I, Vol. 12 (1) (1995) Uses — The timber is highly valued, especially in India and Australia; it is used in house and boat construction, for high grade furniture and carvings, and to make tea- chests, oil casks, pencils, musical instruments. The flowers are used as a source of red and yellow dyes for silk, cotton and wool fabrics and as an emmenagogue. The leaves and young shoots are lopped for cattle fodder in India. Various parts are used medici- nally throughout its geographical range; the bark is a powerful astringent, a tonic and an antiperiodic, and used to treat dysentry and wounds (Brandis 1874; Burkill 1935). The limonoid, cedrelone, exhibits antifeedant properties against a variety of insects, e.g. in- hibiting noctuid larval growth but is believed not to affect the insect’s endocrine system [Koul & Isman, Entom. Exp. Appl. 64 (1992) 281]. Notes — 1. This is the most wide-ranging Toona species, which exhibits consider- able variation in filament pubescence. It was first described from India, where it is the dominant Joona and is characterized by glabrous filaments. This variant extends east- wards to Hainan. Showing a more restricted distribution within this range, are plants with glabrescent or sparsely pilose/villous filaments, while extending the range as far as eastern Australia are plants with conspicuously villous filaments. Whether the eastern variants should be recognized infraspecifically has yet to be determined. Cedrela velutina is included here with some hesitation as it is based on a scrappy sterile specimen, which was included in Chukrasia by e.g. Hiern in Hook. f., Fl. Brit. India 1 (1875) 568. 2. The correct status of the following species has yet to be determined; it may be a good morphologically distinct species, or simply a large-fruited geographical variant of Toona ciliata. 3a. Toona calantas Merr. & Rolfe Toona calantas Merr. & Rolfe, Philipp. J. Sc., Bot. 3 (1908) 105; Whitford, For. Philipp. 2 (1911) 45; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 357; Elmer, Leafl. Philipp. Bot. 9 (1937) 3388; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46; Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 344, t. 263. Cedrela odorata Blanco, FI. Filip., ed. 1 (1837) 184, non L.; Merr., Sp. Blanc. (1918) 31. ?Cedrela inodora Hassk., Hort. Bogor. 1 (1858) 131; Miq., Fl. Ind. Bat. 1, 2 (1859) 549; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 64; C.DC. in DC., Monogr. Phan. | (1878) 747. — ?Surenus inodora (Hassk.) Kuntze, Rev. Gen. Pl. 1 (1891) 111. — ?Toona inodora (Hassk.) Hochr. in Brig., Mém. Inst. Nat. Genev. 24 (1935) 32. — ?Cedrela febrifuga Blume var. inodora (Hassk.) C.DC., Rec. Bot. Surv. India 3 (1908) 373. — ?Toona sureni (Blume) Merr. var. inodora (Hassk.) Bahadur, Monogr. Toona (1988) 136, nom. illeg. Toona paucijuga Merr., Philipp. J. Sc., Bot. 7 (1912) 279; Enum. Philipp. Flow. Pl. 2 (1923) 357; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46. [?Toona philippinensis Elmer, Leafl. Philipp. Bot. 9 (1937) 3388, nom. non rite publ., anglice (‘philip- pinense’); Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 46. — ?Toona sureni (Blume) Merr. var. philippinensis Bahadur, Monogr. Toona (1988) 138, nom. non rite publ., anglice (‘philippinense’).| See Note 2 to the previous species. This is an important timber tree in the Philippines, where it is commonly known as ‘kalantas’ or ‘calantas’; it is reported to occur at low and medium altitudes in primary for- ests in the Batan Islands, Luzon, Mindoro, Samar, Negros, Leyte, Cebu and Mindanao. Mabberley et al. — Meliaceae 371 Flowering specimens are rare, but the fruits are characterized by the following fea- tures: Capsules large, (20—)28—40 mm long; columella 24—40 mm long, 7-15 mm broad; valves dark red to red-brown, smooth, 24—41 by 4—14 mm, lenticellate with numerous lenticels (0.1—1 mm diam.), which are often smaller and denser towards the base of the capsule. Seeds (14—)20—32 by 3.2—6 mm, wings unequal with broadly ob- tuse apices; seed body 4.2—10 by 1.5—3 mm. Specimens which might also belong to this taxon occur in Sumatra (Lampong), Penin- sular Malaysia (Selangor, Pahang), Borneo (Sabah), Java (Madiun, Besuki), and New Guinea (Manokwari, Port Moresby, Manus Island). Vernacular names — Philippines: anipla (Iv.); danigga (Ibn.); danigpa, danupra (IIk.); kalantas (Pang., Tag.); kantingen (IIk., Sbl.); lanigda, lanigpa (Bik.); lanipga (Bik., S.L. Bis., C. Bis); porak (IIk.); Borneo (Sabah): via (Dusun); limpaga (Malay); melipaga (Dusun, Kwijau); surian; Malaya: serai wangi; Java: soeren (Sempol). Uses — In addition to its timber value, bark decoctions are considered to have astrin- gent and antiseptic properties, and are used to clean wounds or for gangrenous ulcers. Flower decoctions are also used as an antispasmodic [L.M. Perry, Medic. Pl. E & SE Asia (1980) 263]. TRIBUS XYLOCARPEAE Xylocarpeae Blume, Bijdr. (1825) 178; Noamesi, Rev. Xylocarpeae, unpubl. thesis, Univ. Wisconsin (1958) 103; T.D. Penn. & Styles, Blumea 22 (1975) 524. Carapeae Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) 267, 276. Trees. Indumentum of simple hairs. Flowers without a gynophore. Petals free. Sta- minal tube urceolate or cupular, margin lobed; anthers 8—10. Capsule + woody or leath- ery with a rudimentary columella. Seeds unwinged, with a corky or woody outer layer; cotyledons large, united; endosperm absent. Distribution — Two genera, tropics: Carapa (see Note | under Xylocarpus) and the following. XYLOCARPUS Xylocarpus Koen., Naturf. 20 (1784) 2; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 81; Noamesi, Rev. Xylocarpeae, unpubl. thesis, Univ. Wisconsin (1958) 103; T.D. Penn. & Styles, Blumea 22 (1975) 525; Mabb., Malays. For. 45 (1982) 448; Toml., Bot. Mangroves (1986) 274. — Granatum [Rumph., Herb. Amb. 3 (1743) 92, t. 61, 62, ex] Kuntze, Rev. Gen. Pl. 1 (1891) 110, p.p., nom. superfl., non St. Lag. (1880). Carapa auctt., p.p., non Aubl. Monosoma Griff., Notul. 4 (1854) 502. Semi-evergreen maritime trees. Twigs lenticellate, cicatrose. Leaves paripinnate with (1-)2—4(—5) pairs of leaflets, entire, glabrous. Flowers unisexual in short axillary thyr- ses. Calyx 4-lobed to about the middle, valvate. Petals 4, contorted and much longer than the calyx in bud. Staminal tube margin with 8 suborbicular, retuse or shallowly and irregularly divided lobes; anthers 8, included. Disk cushion-shaped, beneath or surrounding and united with ovary, red. Ovary 4- (or 5-)locular, each locule with 3 or 372 Flora Malesiana ser. I, Vol. 12 (1) (1995) 4(—6) ovules; style short, stylehead discoid, its margin crenellate and its upper surface with four radiating stigmatic grooves. Fruit a large pendulous subspherical capsule, tardily dehiscing by 4 (5) leathery valves from apex. Seeds 5—20, large, irregularly tetrahedral or pyramidal, outermost surface convex, attached to central columella, with aerenchymatous (?sarcotestal) coat; embryo with the radicle lying above the hilum; ger- mination cryptocotylar, the numerous subulate cataphylls verticillate or spirally arrang- ed; true leaves simple, entire, later ones trifoliolate (rarely first ones thus). 2n = 52. Distribution — Three species throughout the coastal regions of the Old World tropics from East Africa to the Western Pacific. Habitat & Ecology — Mangrove swamps and coastal woodlands on rock and other substrates. The seeds float just below the surface of the sea and are widely dispersed in currents. Xylocarpus was one of the first genera of trees to colonize Krakatoa after its eruption in 1883 (Harms, 1940: 27). Although not truly viviparous, the embryos, like those of Avicennia and Laguncularia, are well developed when they fall and emerge as seedlings soon afterwards [White, Veg. Afr. (1983) 261]. Organic solutes contributing to the osmotic potential of the leaves of the two mangrove species include proline [Popp et al., Vegetatio 61 (1985) 247]. In the Sundarbuns of India, the nectar produced by the flowers is an important source for bees (Apis dorsata), which move to the forests from March to July, a migration of some hundreds of kilometres [Chakrabarti & Chaudhuri, Sei: Cult: 38'C'972)/269): Uses — The wood of all three species is rich redbrown in colour and hard. When large enough pieces are available they are used for boat-building (still practised in the Maldives, where X. rumphii is largely used, Balasubramaniam) and construction as well as for good quality furniture, but the trees are often left in mangrove from which trees of other species have been removed for the production of charcoal. Details of the wood structure are provided by Panshin [Philipp. J. Sc. 48 (1932) 150] and T.D. Penn. & Styles (I.c.), where details of palynology are also presented. The bark is used in tanning. Of the commercial tan barks in the Philippines Filamor [Philipp. J. Anim. Ind. 26 (1965) 43] reports that Xylocarpus has the highest percent- age (13.13) tannin in the bark. With leather, it penetrates rapidly and gives a firm tex- ture and a tan colouration. It gives cloth an umber or red colour. Mixed with the tanbark of Ceriops, fixed with potassium dichromate, and coaltar fixed with petrol, the bark (‘nigi’) has been found to be the most efficacious preservative for fishing nets in the Philippines [Bersamin et al., Philipp. J. Sc. 96 (1967) 143]. A number of medical uses have been gathered together by L.M. Perry [Med. Pl. E & SE Asia (1980) 263]: all parts of the plant are astringent, especially the roots and bark, which has been widely used in the treatment of dysentery. Plants have been raised in Florida [Gill, Bull. Fairchild Trop. Gard. 24, 3 (1969) 10]. Notes — |. Xylocarpus is closely related to Carapa Aubl., a genus of the neotropics, west and central Africa, differing in its large inflorescences of flowers with imbricate sepals, its woody-coated seeds and leaves with 6—12(—18) pairs of leaflets and a dor- mant glandular (or rarely a fully developed) terminal leaflet, while 2n = 58. Neverthe- te Mabberley et al. — Meliaceae 37 less, the genera are closely allied, being geographical vicariants with differing ecology, for Carapa is a forest genus. 2. In view of the widespread nature of the three Xylocarpus species and because they have been widely confused (see Mabberley, |.c., for analysis) since the time of Rum- phius, who first made a published description of the genus, only literature pertinent to Malesia is cited below. 3. A guide to the voluminous literature on the ecology, silviculture and uses of the genus throughout its range may be found under the index entries ‘Carapa’ and ‘Xylocar- pus’ in B. Rollet, Bibliography on mangrove research 1600—1975, UNESCO (1981). 4. The species are most readily distinguished in the field and collectors have so far reported no intermediate specimens. In West Malesia, moreover, the local people have separate names for the two mangrove species which are frequently confused by herba- rium workers. Indeed, in the absence of ecological information, ripe fruit, details of the bark, buttresses and pneumatophores, all of which are distinguishing field characters, the museum worker may well be at a loss to pigeon-hole specimens, particularly sterile material, and those from young plants may be almost impossible to determine. Noamesi (1.c.) has shown that the features of the staminal tube formerly used to distinguish spe- cies are unreliable but the pedicel character is a useful one. He further suggests that X. moluccensis (*X. mekongensis’), which is intermediate in many of its characteristics between the other two, may have arisen as an hybrid between them. Furthermore, it would be tempting to explain away the intermediate herbarium specimens as being de- rived from hybridizations or even continuing introgression, but, without critical field studies, sampling within and between trees, this would be an unwarranted speculation [see White, Mitt. Bot. Staatssamml. Miinchen 10 (1971) 107]. Furthermore, as the most frequently confused species, the mangrove-living X. granatum and X. moluccen- sis grow together, there is also the possibility for mixed gatherings. KEY TO THE SPECIES la. Leaflets in 2—4 (5) pairs, ovate to cordate; pedicels not conspicuously swollen near calyx; fruit 6-8 cm diam.; tree of rocky coasts and sand, not mangrove, with neither conspicuous buttresses nor pneumatophores .......... 1. X. rumphii b. Leaflets in 1—3 (4) pairs, obovate to elliptic, apex rounded to obtuse; pedicels swol- len or not; fruit 6—25 cm diam.; trees of mangrove and other muddy substrates with either conspicuous buttresses or pneumatophores .................20-00-5 2 2a. Leaflets somewhat elliptic-ovate, not rounded at apex; thyrse 3—8(—13) cm long, usually with distinct main axis; pedicels not conspicuously swollen near calyx; fruit 6—11 cm diam.; buttresses not ribbon-like, pneumatophores erect, pointed ...... Di an Ns ase teeta Cah iva 1S jasc ret gat mara Allg Re ala Gr 2. X. moluccensis b. Leaflets usually rounded at apex, coriaceous; thyrse 1-6 cm long, often with dis- tinct main axis; pedicels conspicuously swollen near calyx; fruit 12—25 cm diam.; buttresses flattened, ribbon-like spreading across mud, pneumatophores absent 3. X. granatum 374 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 56. Xylocarpus rumphii (Kostel.) Mabb. a. Flowering shoot; b. half flower; c. bark. — X. moluc- censis (Lam.) M. Roem. d. Leaf; e. bark. Drawing R. Wise. Reproduced (rearranged) with permission from Tree Flora of Malaya 4. Nn Mabberley et al. — Meliaceae 37 1. Xylocarpus rumphii (Kostel.) Mabb. Xylocarpus rumphii (Kostel.) Mabb., Malays. For. 45 (1982) 450; in Fl. Nouv.-Caléd. et Dép. 15 (1988) 81, t. 15, f. 1, 2; in Tree Fl. Malaya 4 (1989) 260, f. 12A.— Carapa rumphii Kostel., Allg. Med. Pharm. Fl. 5 (1836) 1988. Carapa moluccensis auct. non Lam.: DC., Prodr. | (1824) 626, p.p.; Hiern in Hook. f., Fl. Brit. India 1 (1875) 567, p.p.; ?C.DC., Bot. Jahrb. 7 (1886) 461; in Engl., Bot. Ergebn. Gazelle (1890) 34; ?Warb., Bot. Jahrb. 13 (1891) 342; King, J. As. Soc. Beng. 64, 11 (1895) 87, p.p.; Backer, Schoolfl. Java (1911) 217, excl. var. elliptica Koord. & Valeton. Xylocarpus forstenii Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 62. Amoora naumannii sensu C.DC. in Engl., Bot. Ergebn. Gazelle (1890) t. 10, non C.DC., Bot. Jahrb. 7 (1886) 461; cf. Mabb., Blumea 31 (1985) 131. Aglaia zollingeri C.DC., Bull. Herb. Boiss. 2 (1894) 579; Backer, Schoolfl. Java (1911) 212; Koord., Exk. Fl. Java 2 (1912) 447; Backer & Bakh. f., Fl. Java 2 (1965) 127; cf. Mabb., Blumea 38 (1994) 309. — Amoora zollingeri (C.DC.) Koord., Exk. Fl. Java 2 (1912) 444. Xylocarpus granatum auct. non Koen.: Pierre, Fl. For. Cochinch. (1897) t. 359A; Harms in K.Schum. & Laut., Fl. Deutsch. Schutzgeb. Siidsee (1900) 379; ?Briq., Mém. Inst. Nat. Genev. 24 (1935) 38; Elmer, Leafl. Philipp. Bot. 9 (1937) 3395. °e a 4 co Fig. 57. Xylocarpus rumphii (Kostel.) Mabb., flowering shoot (Mabberley & Chan 1999). West Ma laysia, Port Dickson, July 1981. Photograph D.J. Mabberley. 376 Flora Malesiana ser. I, Vol. 12 (1) (1995) Xylocarpus moluccensis auct. non M. Roem.: Merr., Interpr. Rumph. (1917) 307, p.p.; Enum. Philipp. Flow. Pl. 2 (1923) 358; Ridley, Kew Bull. (1938) 291; Harms in Engl. & Prantl, Nat. Pflanzen- fam., ed. 2, 19b1 (1940) 83, t. 19, f. D, G—M; Bot. Jahrb. 72 (1942) 158; Noamesi, Rev. Xylocar- peae, unpubl. thesis, Univ. Wisconsin (1958) 137; Backer & Bakh. f., Fl. Java 2 (1965) 118, p.p.; Johnstone, Biotropica 13 (1981) 254; Toml., Bot. Mangroves (1986) 282. [Granatum litoreum latifolium Rumph., Herb. Amb. 3 (1743) 92, t. 62; Buc’Hoz, Hist. Univ. Veg. PI. IGLT7S) ta iS 215] Tree 4—12(—18) m with neither conspicuous buttresses nor pneumatophores; bole usually solitary, to 50 cm diam., frequently of poor form. Bark lenticellate to finely fissured, greyish; inner bark bright pink to red. Leaf rachis and petiole to 22 cm with terminal spike to | mm. Leaflets in 2—4 pairs, 5—10(—16) by (2—)3—5(—9.5) cm, ovate to cordate, sometimes + falcate, base broadly cuneate or rounded to truncate or cordate, asymmetric, apex acute to acuminate; venation prominent on both surfaces in sicco, conspicuous in vivo; petiolule (O—)1—3(—5) mm. Thyrses 10—18(—28) cm long, lax, pendent, main axis distinct; lateral branches to 8 cm; bracts and bracteoles c. 0.5 mm, narrowly triangular, persistent; pedicels 3—8 mm, not conspicuously swollen near calyx. Calyx lobes 1-1.5 mm long. Petals 3.5—6 by 2—2.5 mm, elliptic-oblong, creamy white. Staminal tube 2—2.5 mm diam., lobes apiculate or bifid to retuse. Fruit 6—8 cm diam., globose. Seeds 8—16, 3.6—7 cm long. First leaves compound (Turner). — Fig. 56a—c, 57. Distribution — Old World tropics from East Africa to Tonga; throughout Malesia but so far unrecorded from the Bornean (or New Caledonian) mainland and rare in Sumatra. Habitat — Cliffs and rocks near surf and sandy substrates above high watermark. Vernacular names — Niri, nyireh (Malay Peninsula, Sumatra, Java); piagau (Philip- pines: Tag., P.Bis., Mag.); New Guinea: kabahai (Pom), tawihi (Misima), wadawada (New Britain). 2. Xylocarpus moluccensis (Lam.) M. Roem. Xylocarpus moluccensis (Lam.) M.Roem., Synops. Monogr. | (1846) 124; Merr., Interpr. Rumph. (1917) 307, p.p.; J. Str. Br. Roy. As. Soc., spec. no. (1921) 3:18; Beck., Tectona 15 (1922) t. 34, f. 7; Craib, Fl. Siam. Enum. | (1926) 265, quoad var. gangeticus (Prain) Craib; Harms in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 19b1 (1940) 84, incl. var. ellipticus (Koord. & Valeton) Harms; Merr., Pl. Life Pac. World (1946) 31; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 888, ?p.p.; Backer & Bakh. f., Fl. Java 2 (1965) 118, p.p.; Mabb., Malays. For. 45 (1982) 450; Sant., Nat. Hist. Bull. Siam Soc. 31 (1983) 78; Mabb. in Tree Fl. Malaya 4 (1989) 260, f. 12C. — Carapa moluc- censis Lam., Encycl. Méth. | (1785) 621; Pers., Syn. 1 (1805) 416; Wood in Rees, Cyclop. 6, 2 (1806) Carapa n. 2; G. Don, Gen. Syst. | (1831) 686; Span., Linnaea 15 (1841) 183, p.p.; Schim- per, Indomal. Strandfl. (1891) 99; Karst., Bibl. Bot. 5, 22 (1891) 21, 23, 51, t. 6, f. 58-63, t. 8, f. 119-123, t. 11, f. 142; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 193, incl. var. elliptica; Koord., Minah. (1898) 384; Prain, Rec. Bot. Surv. India 2 (1903) 292, incl. var. gangetica Prain & var. ovalifolia Koord. ex Prain; Backer, Fl. Batavia (1907) 282, incl. var. elliptica; ?C.DC., Nova Guinea, Bot. 8 (1910) 426, incl. var. elliptica; Backer, Schoolfl. Java (1911) 217, quoad var. elliptica; Brown & Fisher, Bull. Philipp. Dep. Agr. Bur. For. 17 (1918) 32, t. 12 & 22 (1920) t. 12; Ridley, Fl. Malay Penins. 1 (1922) 414, p.p.; Groom & Wils., Ann. Bot. 39 (1925) 9, t. 8; Watson, Mal. For. Rec. 6 (1928) 70, 124, 184, 187, t. 34, 35; ?Burkill, Dict. Econ. Prod. Malay Penins. (1935) 455; Corner, Wayside Trees | (1940) 459; Blake, Austral. J. Bot. 2 (1954) 121; Corner, Wayside Trees, ed. 3 (1988) 498. — Granatum moluccensis (Lam.) Kuntze, Rev. Gen. Pl. 1 (1891) 110. Mabberley et al. — Meliaceae 297 Xylocarpus granatum auct. non Koen.: Sm. in Rees, Cyclop. 39, 1 (1818) Xylocarpus n. 1, p.p.; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 244, t. 20, f. 22, p.p.; Miq., Fl. Ind. Bat. 1, 2 (1859) 546, p.p.; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 62, p.p.; Schmidt, Bot. Tidsskr. 26 (1904) 68; Veg.-Bild. 3 (1906) t. 40; Whitford, For. Philipp. 2 (1911) 48; Koord., Exk. Fl. Java 2 (1912) 437, p.p.; Ridley, Kew Bull. (1938) 288. Carapa obovata auct. non Blume: King, J. As. Soc. Beng. 64, ii (1895) 87, p.p.; Pellegr. in Fl. Indo- Chine, Suppl. (1946) 719, quoad var. microphylla (Pierre) Pellegr.; Ridley, Fl. Malay Penins. 1 (1922) 414, p.p. Xylocarpus mekongensis Pierre, Fl. For. Cochinch. 5 (1897) t. 359B; Noamesi, Rev. Xylocarpeae, un- publ. thesis, Univ. Wisconsin (1958) 124; Johnstone, Biotropica 13 (1981) 253, t. 1, 254; Toml., Bot. Mangroves (1986) 281, f. 5.7, B40, 41. — Carapa mekongensis (Pierre) Pellegr., Fl. Indo- Chine | (1911) 777. Xylocarpus obovatus auct. non A. Juss.: Pierre, Fl. For. Cochinch. 5 (1897) sub t. 358, quoad var. mi- crophyllus Pierre. Carapa borneensis Becc., For. Born. (1902) 574. Xylocarpus gangeticus (Prain) C.E. Parkinson, Ind. For. 60 (1934) 140, t. 17. Xylocarpus australasicus Ridley, Kew Bull. (1938) 291; Johnstone, Biotropica 13 (1981) 254. Xylocarpus parvifolius Ridley, Kew Bull. (1938) 291. [Granatum litoreum parvifolium Rumph., Herb. Amb. 3 (1743), quoad t. 61; Buc’Hoz, Hist. Univ. Veg. Pl. 6 (1775) t. [24].] Tree 6—18(—30, Watson) m, with small buttresses and many pointed pneumato- phores; bole usually solitary, to 70(—210, Watson) cm diam. Bark rough with longi- tudinal fissures, falling as oblong flakes. Leaf rachis and petiole to 10 cm, sometimes Fig. 58. Xylocarpus moluccensis (Lam.) M. Roem. Base of sapling tree with pneumatophores. Silt bank near Pasuruan, E Java. Photograph J. Jeswiet, 1916. 378 Flora Malesiana ser. I, Vol. 12 (1) (1995) with persistent apical spike to 1 mm. Leaflets in (1) 2 or 3 (4) pairs, (S—)7—10(-17) by (2.5—)4—6.5(—7) cm, elliptic-oblong or lanceolate to oblanceolate, base cuneate, + asym- metric, apex acute to obtuse, venation prominent on both surfaces in sicco; petiolule (O—) 2—5 mm, sometimes swollen. Thyrses 3—8(—13) cm long, often produced with the new leaves, + lax, main axis clear; lateral branches to 4 cm; bracts and bracteoles c. 0.5 mm, + persistent; pedicels 3—8 mm, not conspicuously swollen near calyx. Calyx lobes 1—1.7 mm long. Petals 3.5—4 by 2—3 mm, oblong to obovate, creamy white. Staminal tube 2-3 mm diam., lobes acute to apiculate or bifid to retuse. Fruit 6—1 1 cm diam., flattened- globose. Seeds 5-10, 4—6.5 cm long. First leaves simple. — Fig. 56d, e, 58. Distribution — Tropical Asia from India (Sundarbans), Indochina and Thailand throughout Malesia to tropical Australia. The underlying ecological explanation for the restriction of X. moluccensis within Indomalesia (cf. the other species) is unknown, but it may be that it is less tolerant of seasonality than are the other two. Habitat — Mangrove swamps, most commonly in the upper reaches, often in only slightly brackish water (Corner, I.c.). Vernacular names — Nyireh batu (Malay Peninsula, Sumatra, Java); New Guinea: bubaia (Kinomeri), jasoguago (Wanigela), kabaju (Hattam), pihsih (Roro), teteliat (Tehid), uburi (Goaribari). Notes — |. This species has been widely confused with the next, the first clear dis- tinction being made by Beccari (1.c.). 2. The names of Xylocarpus species hinge on an interpretation of Rumphius’s plates in Herbarium Amboinense. His tab. 62 is clearly X. rumphii and the material figured in tab. 61 is referable to mangrove plants (Mabberley, |.c.). I have interpreted the plate as X. moluccensis, but, should the two branches depicted there ever be shown to represent both this species and X. granatum, it would be wise, in the interests of nomenclatural stability, to designate the smaller of them, the one which most closely resembles both Lamarck’s brief description and the plant here, as lectotype. 3. Xylocarpus granatum Koen. Xylocarpus granatum Koen., Naturf. 20 (1784) 2; Willd., Sp. Pl. 2 (1799) 328; Poir. in Lam., Encycl. Méth. 8 (1808) 806; Sm. in Rees, Cyclop. 39, 1 (1818) Xylocarpus n. 1, p.p.; A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 244, t. 20 f. 22, p.p.; Blanco, Fl. Filip. (1837) 298; ed. 2 (1845) 207 (‘Xilocarpus’); M.Roem., Synops. Monogr. | (1846) 124; Miq., Fl. Ind. Bat. 1, 2 (1859) 546, p.p.; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 62, p.p.; Blanco, Fl. Filip., ed. 3, 2 (1878) 24; Merr., Interpr. Rumph. (1917) 306; Sp. Blanc. (1918) 208; Brown & Fisher, Bull. Philipp. Dept. Agr. Bur. For. 17 (1918) 30, t. 11; 22, 920) 30; t. 11; Mem; J. Str. Bro RoywAsasocs spec. no. (1921) 318; Enum. Philipp. Flow. Pl. 2 (1923) 358; Beck., Tectona 15 (1922) 580, t. 34 f. 6; Lane-Poole, For. Res. Papua New Guinea (1925) 103; Merr., Univ. Calif. Publ. Bot. 15 (1929) 120; Ogura, Bot. Mag. Tokyo 54 (1940) 395, cum tab.; Harms in Engl. & Prantl, Nat. Pflanzen- fam., ed. 2, 19b1 (1940) t. 19 A—C (?E, F); Bot. Jahrb. 72 (1942) 158; Merr., Pl. Life Pacif. World (1946) 52, t. 56; K. Heyne, Nutt. Pl. Indon., ed. 3 (1950) 887; Quisumb., Med. Pl. Philipp. (1951) 489; Noamesi, Rev. Xylocarpeae, unpubl. thesis, Univ. Wisconsin (1958) 107; Backer & Bakh. f., FI. Java 2 (1965) 118; T.D. Penn. & Styles, Blumea 22 (1975) 514, fig. g, h; Mabb., Malays. For. 45 (1982) 450; Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 346, t. 265; Toml., Bot. Man- groves (1986) 278, figs. B.39, 42, 43; Mabb. in Fl. Nouv.-Caléd. et Dép. 15 (1988) 82, t. 15 f. 3, 4; Mabberley et al. — Meliaceae 379 in Tree Fl. Malaya 4 (1989) 260, f. 12B. — Carapa granatum (Koen.) Alston in Trim., Handb. FI. Ceylon 6 (1931) 45 (‘granata’); Burkill, Dict. Econ. Prod. Malay Penins. (1935) 452; Corner, Wayside Trees | (1940) 458; Life Pl. (1964) t. 36C; Wayside Trees, ed. 3 (1988) 497. — Carapa indica A. Juss. in Cuv., Dict. Sci. Nat. 7 (1817) 32, nom. superfl. Carapa moluccensis auct. non Lam.: DC., Prodr. 1 (1824) 626, p.p.; Hiern in Hook. f., Fl. Brit. India 1 (1875) 567, p.p.; Fern.-Vill., Nov. App. (1880) 44; Vidal, Sin. Atl. (1883) xx, t. xxix, f. F; King, J. As. Soc. Beng. 64, ii (1895) 87, p.p.; Kamerl., Teysmannia 22 (1911) 118, 129, cum tab.; Backer, Schoolfl. Java (1911) 217, quoad var. typica Backer; Ridley, Fl. Malay Penins. | (1922) 414, p.p. Carapa obovata Blume, Bijdr. (1825) 179; C.DC. in DC., Monogr. Phan. | (1878) 718; Schimper, Indomal. Strandfl. (1891) 99, t. 7 f. 1; King, J. As. Soc. Beng. 64, ii (1895) 87, p.p.; Koord. & Valeton, Bijdr. Booms. Java 3 (1896) 189; Koord., Minah. (1898) 384; Becc., For. Born. (1902) 574; Backer, Fl. Batavia 1 (1907) 280; Schoolfl. Java (1911) 217; Ridley, Fl. Malay Penins. 1 (1922) 414, p.p.; Watson, Mal. For. Rec. 6 (1928) 70, 124, 184, 187, t. 36, 37. — Xylocarpus obovatus (Blume) A. Juss., Mém. Mus. Nat. Hist. Nat. Paris 19 (1832) 244; M. Roem., Synops. Monogr. | (1846) 124; Miq., Fl. Ind. Bat. 1, 2 (1859) 546; Suppl. (1861) 197; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 62; Harms in Engl. & Prantl, Nat. Pflanzenfam. 3, 4 (1896) t. 157, f. A—C; Pierre, Fl. For. Cochinch. 5 (1897) sub t. 358, incl. var. macrophyllus Pierre; Harms in K. Schum. & Laut., Fl. Deutsch. Schutzgeb. Siidsee (1900) 379; Merr., Bull. Philipp. Bur. For. 1 (1903) 28; Perkins, Fragm. Fl. Philipp. (1904) 30; Whitford, For. Philipp. 2 (1911) 47; Koord., Exk. Fl. Java 2 (1912) 437. — Granatum obovatum (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 110. ?Xylocarpus carnulosus Zoll. & Mor., Nat. Gen. Arch. Neerl. Ind. 2 (1845) 582; Migq., Fl. Ind. Bat. 1, 2 (1859) 546. — ?Carapa carnulosa (Zoll. & Mor.) Kurz, J. As. Soc. Beng. 39, ii (1870) 72 (‘car- nosa’); Prelim. Rep. Veg. Pegu App. A (1875) xxxiil. Monosoma littorata Griff., Notul. 4 (1854) 502; Ic. 4 (1854) t. 588, f. 3. ?Guarea oblongifolia Griff., Notul. 4 (1854) 503. 2Amoora salomoniensis C.DC., Bot. Jahrb. 7 (1886) 461; in Engl., Bot. Ergebn. Gazelle (1890) 33. Xylocarpus benadirensis Mattei, Boll. Ort. Bot. Palermo 7 (1908) 99. Xylocarpus moluccensis auct. non M. Roem.: Schimper & Faber, Pflanzengeogr., ed. 3 (1935) 140, t.34: Xylocarpus minor Ridley, Kew Bull. (1938) 289. [Granatum litoreum parvifolium Rumph., Herb. Amb. 3 (1743) 93, non t. 61 (= X. moluccensis).] Tree or shrub (1—)6—15(—20) m with thin, branched, ribbon-like buttresses spread- ing out from base; bole solitary or occasionally more, to 90(—180, Watson) cm diam., often of poor form. Bark thin, smooth, scaling as irregular flakes, whitish to yellow- brown; inner bark reddish or pink. Leaf rachis and petiole to 9(—12) cm, usually much less. Leaflets in 1 or 2 (3) pairs, (3.5—)5—12 by (2—)3—6 cm, coriaceous, obovate or elliptic, base cuneate, apex rounded or obtuse, venation prominent on both surfaces in sicco, petiolules (2—)5—6(—11) mm, swollen. Thyrses (1—-)3—6 cm, + squarrose, borne on new and older twigs, frequently + forked with indistinct main axis; bracts and brac- teoles c. 0.5 mm, caducous; pedicels 3—9 mm, conspicuously swollen towards the calyx. Calyx lobes 1-3 mm long. Petals 3.5—5.5(—6.5) by 2—3 mm, oblong, creamy white or pinkish. Staminal tube 2—3.5 mm diam., lobes apiculate or bifid to retuse. Fruit 12—25 cm diam., flattened-globose. Seeds 8—16(—20), 4—6 cm long. First leaves simple. — Fig. 59, 60. Distribution — Old World tropics from East Africa and continental Asia throughout Malesia to Tonga. Habitat — Mangroves, usually estuarine and often associated with Nypa and Son- neratia. 380 Flora Malesiana ser. I, Vol. 12 (1) (1995) Vernacular names — Nyireh bunga (Malay Peninsula, Sumatra, Java); Philippines: bigi (Tagk.), tabigi (Tag., Bik., Tagb., P.Bis., S.L.Bis., C.Bis., Lan., Sul., Mag., official common name); New Guinea; awol (New Britain), ecahi (Labu), kaav (Bian), mokkemoffe (Arguni). Fig. 59. Xylocarpus granatum Koen. a. Flowering shoot; b. half flower; c. unripe fruit and seed; d. bark. Drawing R. Wise. Reproduced (rearranged) with permission from Tree Flora of Malaya 4. Meliaceae Mabberley et al. wet laa Sr Holttum standing next to R.E. ake roots, Dr. Fig. 60. Xylocarpus granatum Koen. Base of tree with sn the tree. Mangrove near Cilacap, Central Java. Photograph J. Jeswiet, 1929. 382 Flora Malesiana ser. I, Vol. 12 (1) (1995) Fig. 61. Aglaia odorata Lour. Male inflorescence. Campus Universiti Malaya, Kuala Lumpur. Photo- graph C.M. Pannell, 1978. Mabberley et al. — Meliaceae 383 CULTIVATED SPECIES Besides Aglaia korthalsii, Lansium domesticum, Melia azedarach, Azadirachta indica, and Sandoricum koetjape, considered above under the respective genera, cultivated Meliaceae are largely neotropical and mostly grown for timber or as street-trees. See also the paragraph ‘Uses’ at the beginning of the family account. See Pennington & Styles in Fl. Neotropica 28 (1981) for full descriptions and synonymy of the neotropi- cal species. Aglaia odorata Lour.: Burkill, Dict. Econ. Prod. Malay Penins. (1935) 74; Pannell, Kew Bull., Add. Ser. 16 (1992) 298. Bush or sometimes tree to 10 m. /ndumentum of yellowish-brown stellate scales, especially dense on shoot apex and fruit surface. Leaves imparipinnate, 1- or 2- (or 3-) jugate, rachis sometimes narrowly winged. /nflorescence with widely spaced, yellow, fragrant flowers. Fruits obovoid, c. 1 by 0.7 cm, red, brown or orange. — Fig. 61. Continental SE Asia. Commonly cultivated for its ornamental value, male trees more often than females. Strongly perfumed flowers used for scenting tea or clothes. Carapa guianensis Aubl.: Corner, Wayside Trees | (1940) 458 (‘guyanensis’) & ed. 3, 2 (1988) 497. Deciduous or semi-evergreen tree to 55 m in wild, flushing claret. Leaves paripin- nate, usually 4—8-jugate. Flowers creamy white in thyrses to 60 cm long. Fruit 5—10 cm diam. Seeds with woody sarcotesta. Tropical America. Source of valuable timber (crabwood) and oil (andiroba) and insec- ticide. Grown in Singapore. Fruits of C. guianensis from America have been washed up in the Carolines [Gunn & Dennis, Castanea 37 (1972) 199]. Carapa procera DC. from tropical America and Africa, with more rounded leaflets than in the previous species, has been cultivated in Java. Cedrela odorata L. (including C. mexicana M.Roem., C. glaziovii C.DC.): Backer, Schoolfl. Java (1911) 218; Corner, Wayside Trees | (1940) 495; 2 (1940) t. 134, 135; ed. 3 (1988) 498, t. 146, 147. Deciduous tree to 35(—60) m in wild. Leaves usually paripinnate, usually (5) 6—12 (—15)-jugate. Flowers creamy white, disagreeably scented, in thyrses to 40(—50) cm long. Stamens 5, free. Capsule with broadly 5-winged columella. Seeds 2—3 cm long, winged at one end. Tropical America. Source of a valuable timber (cedar); planted throughout the tropics. Malay Peninsula, Java, Philippines, New Guinea. Swietenia macrophylla King: Corner, Wayside Trees | (1940) 469; 2 (1940) t. 142; ed. 3 (1988) 507, t. 154; Backer & Bakh. f., Fl. Java 2 (1965) 118; Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 343, t. 262; Prawirohatmadjo et al. in Soeriane- 384 Flora Malesiana ser. I, Vol. 12 (1) (1995) gara & Lemmens (eds.), PI]. Res. SE Asia (PROSEA Handb.) 5 (1), Major commer- cial timbers (1993) 447, cum tab. — Swietenia mahagoni (L.) Jacq. var. macrophyl- la (King) Veitch, Travel Notes (1896) 53. Deciduous tree to 40(—60) m. Leaves usually paripinnate, (2) 3—6(—8)-jugate. Leaf- lets c. 15 cm long. Flowers pinkish or white in thyrses to 18(—20) cm long. Staminal tube 10-lobed; anthers 10. Capsule to 15 cm long. Seeds to 10 cm long, winged at one end. Tropical America. Introduced to Java in 1872 [see H.A. Fattah, Mahogany Forestry in Indonesia, in G. Hartshorn, Mahogany Workshop (1992)] and to Singapore in 1876 and by the 1930s widely planted there [Gard. Bull. Sing. 37 (1978) 114]; Malay Penin- sula, Borneo, Java, Philippines. The trees bear a rich epiphytic flora [Johnson & Barkat Awan, Malays. For. 35 (1972) 5] and are more resistant to shoot-borers than is S. ma- hagoni. Plants have been raised through tissue culture [Lee & Rao, Gard. Bull. Sing. 41 (1988) 11]. Swietenia mahagoni (L.) Jacq.: Corner, Wayside Trees 1 (1940) 463 (‘mahogani’ ); ed. 3 (1988) 508; Backer & Bakh. f., Fl. Java 2 (1965) 118; Prawirohatmadjo et al. in Soerianegara & Lemmens (eds.), Pl. Res. SE Asia (PROSEA Handb.) 5 (1), Major commercial timbers (1993) 447, cum tab. Tropical America. Similar to S. macrophylla but leaves 2—4-jugate, leaflets c. 3.5 cm long; thyrse to 5 cm; fruit smaller. Singapore, Java, Philippines, Lesser Sunda Islands (Flores, Sumbawa). DUBIOUS SPECIES Aglaia allocotantha Harms, Bot. Jahrb. 72 (1942) 175; Pannell, Kew Bull., Add. Ser. 16 (1992) 350. — Type destroyed (New Guinea). Aglaia araeantha Harms, op.cit. 162; Pannell, l.c. — Type destroyed (New Guinea). Aglaia bergmannii Warb., Bot. Jahrb. 8 (1891) 346; Pannell, ].c. — Type destroyed (New Guinea). Aglaia caroli Harms, Bot. Jahrb. 72 (1942) 176; Pannell, l.c. — Type destroyed (New Guinea). Aglaia everettii Merr., Philipp. J. Sc., Bot. 4 (1909) 271; Pannell, ].c. — Types destroy- ed (Philippines). Aglaia gracillima Harms, Bot. Jahrb. 72 (1942) 171; Pannell, l.c. — Type destroyed (New Guinea). Aglaia hiernii Koord., Minah. (1898) 381, nom. nud.; Pannell, op. cit. 351.— Type not seen. Mabberley et al. — Meliaceae 385 Aglaia huberti Harms, Notizbl. Bot. Gart. Berlin 15 (1941) 472; Pannell, l.c. — Type destroyed (Borneo). Aglaia koordersii Jain & Bennet, Ind. J. For. 9 (1987) 271, nom. nud.; Pannell, l.c. — Type not seen. Aglaia ledermannii Harms, Bot. Jahrb. 72 (1942) 165; Pannell, l.c. — Type destroyed (New Guinea). Aglaia leptoclada Harms, op. cit. 166; Pannell, l.c. — Type destroyed (New Guinea). Aglaia maboroana Harms, op. cit. 173; Pannell, l.c. — Type destroyed (New Guinea). Aglaia phaeogyna Harms, op. cit. 167; Pannell, l.c. — Types destroyed (New Guinea). Aglaia pycnoneura Harms, op. cit. 179; Pannell, l.c. — Type destroyed (New Guinea). Aglaia rodatzii Harms in K.Schum. & Laut., Fl. Deutsch. Schutzgeb. Siidsee (1900) 386; Pannell, |.c. — Types destroyed (New Guinea). Aglaia schraderiana Harms, Bot. Jahrb. 72 (1942) 168; Pannell, l.c. — Type destroyed (New Guinea). Aglaia schultzei Harms, op. cit. 177; Pannell, 1.c. — Type destroyed (New Guinea). Aglaia spaniantha Harms, op. cit. 168; Pannell, op. cit. 352. — Types destroyed (New Guinea). Aglaia steinii Harms, op. cit. 178; Pannell, l.c. — Type destroyed (New Guinea). Aglaia trichostoma Harms, op. cit. 169; Pannell, l.c. — Type destroyed (New Guinea). Aglaia urophylla Harms, op.cit. 174; Pannell, l.c. — Types destroyed (New Guinea). Aglaia vilamilii Merr., Philipp. J. Sc. 15 (1915) 536; Pannell, l.c. — Type destroyed (Philippines). Aglaia vulpina Harms, Bot. Jahrb. 72 (1942) 175; Pannell, l.c. — Type destroyed (New Guinea). Amoora naumannii C.DC., Bot. Jahrb. 7 (1886) 461 = ?Dysoxylum sp. See Mabb., Blumea 31 (1985) 131. 386 Flora Malesiana ser. I, Vol. 12 (1) (1995) Boswellia ? integra Blanco, FI. Filip., ed. 2 (1845) 242; ed. 3, 2 (1878) 86 = ?Dysoxy- lum. See Merr., Sp. Blanc. (1918) 210; Enum. Philipp. Flow. Pl. 2 (1923) 366. Boswellia ? obliqua Blanco, op. cit. 243, 87 = ?Dysoxylum. See Merr., Sp. Blanc. (1918) 210; Enum. Philipp. Flow. Pl. 2 (1923) 366. Dysoxylum microcricum Harms, Notizbl. Bot. Gart. Berlin 15 (1941) 477 = Dysoxy- lum sp. (sect. Cyrtochiton) — Type destroyed (New Guinea). The description suggests a relationship with the D. cyrtobotryum group. Lachanodendron album Reinw. ex Blume, Cat. Gew. Buitenzorg (1823) 70, nom. nud. = ?. See Mabb., Blumea 31 (1985) 140. Lansium montanum Jack ex Spreng., Syst. 3 (1826) 66 = ?Aglaia sp. See Mabb., Blumea 31 (1985) 143. Milnea montana (Jack, Trans. Linn. Soc. 14 (1823) 118, nom. provis.] (Spreng.) Steud., Nomencl., ed. 2, 1 (1841) 142; Miq., Fl. Ind. Bat. 1, 2 (1859) 544 = ?Aglaia sp. See Mabb., Blumea 31 (1985) 145. Selbya montana (Spreng.) M. Roem., Synops. Monogr. | (1846) 126 = ?Aglaia sp. See Mabb., Blumea 31 (1985) 143. EXCLUDED SPECIES Aglaia chartacea Kosterm., Reinwardtia 7 (1966) 261 = Lepisanthes tetraphylla (Vahl) Radlk. (Sapindaceae). See Mabb., Blumea 31 (1985) 143. Aglaia iloilo (Blanco) Merr., Philipp. J. Sc., Bot. 9 (1914) 533; Sp. Blanc. (1918) 212; Enum. Philipp. Flow. Pl. 2 (1923) 375; all excl. specim. cit. (= Aglaia argentea Blume). See Pannell, Kew Bull., Add. Ser. 16 (1992) 354, and also under Melia iloilo below. Aglaia intricatoreticulata Kosterm., Reinwardtia 7 (1966) 259, t. 12 = ?Lansium do- mesticum Corr. (q.v.) or = ?Lepisanthes (Sapindaceae). See Mabb., Blumea 31 (1985) 140, 142. Aglaia nivea Elmer ex Merr., Enum. Philipp. Flow. Pl. 2 (1923) 375, in syn. Aglaia iloilo. See Pannell, Kew Bull., Add. Ser. 16 (1992) 354, and under Melia iloila below. Aglaia pinnata (L.) Druce, Rep. Bot. Exch. Club Brit. Is. 1913, 3 (1914) 413 = Vitex pinnata L. (Labiatae—Viticoideae). Mabberley et al. — Meliaceae 387 Alliaria dasyphylla (Miq.) Kuntze, Rev. Gen. Pl. 1 (1891) 109 = Ailanthus integri- folia Lam. (Simaroubaceae). Alliaria euneuron (Miq.) Kuntze, l.c., see below under Dysoxylum euneuron. Amoora aphanamixis auct. non J.A. & J.H. Schultes: Miq., Fl. Ind. Bat., Suppl. (1861) 196 = Santiria griffithii (Hook. f.) Engl. (Burseraceae). See Leenh. & Kalkm. in Fl. Males. I, 6 (1972) 921. Amoora decandra (Wall.) Hiern (i.e. Sphaerosacme decandra (Wall.) T.D. Penn. from Himalaya), recorded in error from Malay Peninsula by Ridley, Fl. Malay Penins. 1 (1922) 400. Amoora forbesii S.Moore, J. Bot. Lond. 64, Suppl. (1926) 4 = Santiria apiculata Benn. (Burseraceae). See Mabb., Blumea 38 (1994) 309. Amoora roxburghiana Korth. ex Blume, Mus. Bot. Lugd.-Bat. 1 (1850) 211, nom. in syn. = Santiria laevigata Blume (Burseraceae). Amoora tomentosa Korth. ex Blume, |.c. = Santiria tomentosa Blume (Burseraceae). Antelaea canescens Cels ex Heynh., Nom. Bot. Hort. 1 (1840) 315; Nomencl. Alph. (1846/7) 38, i.e. sphalm. pro Anthelia (‘Anthelea’) canescens Cels; see Cels, Cat. Cult. Cels Prix Cour. 1842 (1842) 8 = ?Epipremnum sp. (Araceae). Azadirachta digyna Anon., Rév. Bot. Appl., Actes 8 (1928) 211, nom. nud. = ? Cedrela taratara Blanco, FI. Filip., ed. 2 (1845) 131; ed. 3, 1 (1877) 234 = mixtum. Teste Merr., Sp. Blanc. (1918) 213; Enum. Philipp. Flow. Pl. 2 (1923) 358, per- haps Pterocarpus (Leguminosae) wood with Dysoxylum or Aglaia herbarium material. Dysoxylum dasyphyllum Migq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 19; C.DC. in DC., Monogr. Phan. | (1878) 526 = Ailanthus integrifolia Lam. (Simaroubaceae). See Koord., Minah. (1891) 374; Noot. in Fl. Males. I, 6 (1962) 218. Dysoxylum euneuron Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 22; C.DC. in DC., Monogr. Phan. | (1878) 516. — Alliaria euneuron (Miq.) Kuntze = Euphorian- thus euneurus (Miq.) Leenh. (Sapindaceae). See Leenh. in Fl. Males. I, 11 (1994) 536. Flindersia R. Br. Now removed to Rutaceae (Flindersiaceae). 388 Flora Malesiana ser. I, Vol. 12 (1) (1995) Harpagonia Noronha, Verh. Batav. Gen., ed. 5, 1 (1790/1) art. 4, 2, nom. nud. = Psy- chotria L. (Rubiaceae). See Steenis, Regn. Veget. 7 (1970) 375. Melia iloilo Blanco, FI. Filip., ed. 2 (1845) 241 = ?Rutaceae. The plant described has opposite winged leaves which does not fit Aglaia to which Merrill referred it as Aglaia iloilo (Blanco) Merr. (see above). Although the vernacu- lar name iloilo is found on herbarium material of Aphanamixis polystachya (Wall.) R. Parker, the description does not match Aphanamixis either and the combination of features strongly suggests Rutaceae, perhaps (Van Balgooy, pers. comm.) Evodia. Owenia cerasifera F. Muell., Hook. J. Bot. 9 (1857) 305; Barr & Sug., Descr. Price List 1868 (1868) 6; Koch, Wochenschr. 11 (1868) 157; Robinson, Glean. Fr. Gard., ed. 2 (1869) 30; Forbes, Nat. Wand. (1885) 354; Hemsl., Bot. Chall. 1, 3 (1885) 132; Dyer, J. Linn. Soc., Bot. 21 (1885) 373; 374 (cerasifolia)sBoeuk Handl. | (1890) 671 = Pleiogynium timoriense (DC.) Leenh. (Anacardiaceae). See Ding Hou in FI. Males. I, 8 (1978) 474. Pistaciovitex L. ex Kuntze in T. Post & Kuntze, Lex. Gen. Phan. (1903) 442, excl. syn. Aglaia = Vitex L. (Labiatae—Viticoideae). The confusion derives from Burman’s giving a plate of Ag/aia odorata Lour. the name Vitex pinnata. Pistaciovitex pinnata (L.) Kuntze, |.c., excl. syn. Aglaia odorata Lour. = Vitex pin- nata L. (Labiatae—Viticoideae). Trichilia volubilis Blanco, Fl. Filip., ed. 2 (1845) 249; ed. 3, 2 (1878) 98 = mixtum (largely Leguminosae). See C.DC. in DC., Monogr. Phan. 1 (1878) 751; Merr., Sp. Blanc. (1918) 194; Enum. Philipp. Flow. Pl. 2 (1923) 380. Turraea trichostylis Migq., Fl. Ind. Bat., Suppl. (1861) 195, 502 = Leptonychia cau- data (G. Don) Burret (Sterculiaceae). See Veldk. & Flipphi, Blumea 32 (1987) 451. Walsura villosa auct. non Wall. ex Kurz: Ridley, Fl. Malay Penins. | (1922) 412 = Harpullia cupanioides Roxb. (Sapindaceae). See Leenh. & Vente, Blumea 28 C982) 27 Xylocarpus echinatus Calc. Cat. Exhib. 1862 ex Balf., Timb. trees E & S As., ed. 2 (1862) 268 & Cycl. India, ed. 2, 5 (1873) 221 = Artocarpus sp. (Moraceae). Watson, Descr. Cat. Ind. Dept. Int. Exhib. 1862 (1862) 161, has X. granatum followed by Artocarpus echinatus Roxb. of which X. echinatus is clearly a “‘portman- teau’ or even a new combination! Compare Lachanodendron domesticum under Lan- sium domesticum. 389 INDEX TO SCIENTIFIC PLANT NAMES Suprageneric names have been entered under the family name to which they belong preceded by the indication of their rank (subfamily, tribe, etc.). Infrageneric names have been entered immedia- tely under the generic name to which they belong, preceded by the indication of their rank (sub- genus, section, etc.). Infraspecific epithets have been entered under the specific name to which they belong preceded by the indication of their rank (subspecies, variety, forma, etc.). Synonyms have been printed in italics. Page numbers in bold type denote main treatment: an asterisk behind a page number denotes the presence of a figure of the concerned taxon. (Aglaia) Aengelenia Hassk. 30 (Aglaia) Aglaia Lour. 2, 4—6, 8, 18,21, 194, 197*, 386 sect. Aglaia 232 sect. Amoora (Roxb.) Pan- nell 212 sect. Beddomea (Hook. f.) Harms 232 sect. Euaglaia Harms 232 sect. Hearnia (F. Muell.) Harms 232 sect. Lansium (Correa) Kos- term. 314, 322 sect. Macroaglaia Harms 232 sect. Neoaglaia Harms 232 acariaeantha Harms 269 acida Koord. & Valeton 272 acuminata Merr. 259 acuminatissima Teijsm. & Binn. 268 var. kambangana Miq. 301 affinis Merr. 276 agglomerata Merr. & L.M. Perry 205, 249 agusanensis Elmer ex Mert. 271 aherniana Perkins 210, 291 allocotantha Harms 384 alternifoliola Merr. 228 ancolana Miq. 236 andamanica Hiern 228 angustifolia (Mig.) Miq. 202, 309 var. horsfieldiana C. DC. 309 anonoides Elmer ex B.D. Jackson 297 antonti Elmer 288 aphanamixis Pellegr. 189 var. frutescens (C. DC.) Pellegr. 189 apoana Merr. 288 aquea (Jack) Kosterm. 316 araeantha Harms 384 archboldiana A.C. Smith 300 argentea Blume 3, 203, 237, 386 var. angustata Miq. 237 var. borneensis Miq. 237 var. cordulata C. DC. 237 var. curtisii King 237 var. eximia (Miq.) Miq. var. hypoleuca (Miq.) Miq. 237 var. latifolia Miq. 236 var. microphylla Miq. 237 var. multijuga Koord. & Valeton 237 var. splendens Koord. & Valeton 237 var. stellatipilosa Adelb. 236 var. superba Miq. 237 argentea auct. 236 aspera Teijsm. & Binn. 268 var. sumatrana Baker f. 268 australiensis Pannell 224 badia Merr. 245 bamleri Harms 306 banahaensis Elmer ex Merr. 288 baramensis Merr. 288 barbanthera C. DC. 211, 292 barbatula Koord. & Valeton 234 batjanica Miq. 246 bauerlenii C. DC. 237 beccariana (C.DC.) Harms 309 beccarii C. DC. 228 beddomei(Kosterm.) Jain & Gaur 190 bergmannii Warb. 384 bernardoi Merr. 233 bicolor Merr. 245 boanana Harms 241 bordenti Merr. 297 borneensis Merr. 302 brachybotrys Merr. 228 brassii Merr. & L.M. Perry 207, 280 brevipeduncula C. DC. 284 brevipetiolata Merr. 277 breviracemosa (Kosterm.) Kosterm. 321 brownii Pannell 209, 305 bulusanensis Elmer ex Merr. 246 cagayanensis Merr. 228 calelanensis Elmer 268 canariifolia Koord. 243 caroli Harms 384 carrii Harms 286 caudatifoliolata Merr. 310 cauliflora Koord. 251 caulobotrys Quisumb. & Merr. 288 cedreloides Harms 259 celebica Koord. 251 ceramica (Miq.) Pannell 209, 285 chalmersii C. DC. 286 chartacea Kosterm. 386 cinerea King 267 cinnamomea Baker f. 211, 290 clarkii Merr. 235 390 (Aglaia) clemensiae Merr. & L.M. Perry 284 clementis Merr. 288 cochinchinensis (Pierre) Pellegr. 190 conduplifolia Elmer 213 conferta Merr. & L.M. Perry 290 confertiflora Merr. 251 copelandii Elmer 259 cordata Hiern 306 cordata form I Hiern 310 coriacea Korth. ex Miq. 210, 295, 296* costata Merr. 205, 248 crassinervia Kurz ex Hiern 204, 210, 267 cremea Merr. & L.M. Perry 205, 264 cucullata (Roxb.) Pellegr. 3, 204, 206, 213 cucullata auct. 214 cumingiana Turcz. 202,211, 293 cuprea Elmer 239 cupreolepidota Merr. 243 curranii Merr. 272 curtisii King 310 cuspidata C. DC. 211, 311 cuspidella Ridley 276 davaoensis Elmer 288 densisquama Pannell 203, 240 densitricha Pannell 208, 224 denticulata Turcz. 246 diepenhorstii Miq. 276 diffusa Merr. 272 diffusiflora Merr. 246 discolor Merr. 237 doctersiana Harms 249 domestica (Correa) Pellegr. 316 dookoo Griff. 316 dubia (Merr.) Kosterm. 326 dyeri Koord. 306 dysoxylifolia Koord. 251 dysoxylonoides Koord. 245, 2a edelfeldtii C. DC. 286 edulis (Roxb.) Wall. 11, 205, 207, 272 Flora Malesiana ser. I, Vol. 12 (1) (1995) (Aglaia) elaeagnoidea (A. Juss.) Benth. 4, 11, 204, 205, 243 var. glabrescens Valeton 243 var. pallens Merr. 243 elaphina Merr. & L.M. Perry 306 elliptica Blume 197*, 198%, 202, 211, 288 var. ceramica Miq. 285 elliptifolia Merr. 11, 246 elmeri Merr. 271 ermischit Warb. 283 erythrosperma Pannell 197*, AOS 219 euryanthera Harms 206, 207, 282 eusideroxylon Koord. & Valeton 228 everettii Merr. 384 exigua Merr. & L.M. Perry 286 eximia Mig. 203, 236 exstipulata (Griff.) Theob. 209, 303 ferruginea 10 flavescens C. DC. 206, 255 flavida Merr. & L.M. Perry 204, 207, 214 forbesiana C. DC. 269 forbesii King 208, 210,211, 265 forstenti Mig. 259 foveolata Pannell 211, 266 fragilis A.C. Smith 300 fraseri Ridley 276 fusca King 297 gamopetala Merr. 261 ganggo Miq. 259 gibbsiae C. DC. 283 gjellerupii C. DC. 284 glabrata Teijsm. & Binn. 206, 254 glabriflora Hiern 261 glabrifolia Merr. 261 glomerata Merr. 306 goebeliana Warb. 246 gracillima Harms 384 grandifolia 9 grandifoliola Merr. 228 grandis Korth. ex Mig. 203, 232 (Aglaia) griffithii (Hiern) Kurz 303 halmaheirae Mig. 103 hapalantha Harms 284 haplophylla Harms 298 harmsiana Perkins 288 hartmannii C. DC. 283 havilandii Ridley 288 hemsleyi Koord. 233 heptandra Koord. & Valeton 231 heterobotrys Merr. 298 heteroclita King 271 heterophylla Metr. 276 hexandra Turcz. 246 hiernii King 203, 208, 310, 384 huberti Harms 385 humilis King 265 hypoleuca Migq. 237 ignea Valeton ex K. Heyne 225 iloilo (Blanco) Merr. 386, 388 inaequalis Teijsm. & Binn. 288 insignis Schwartz 271 integrifolia Pannell 201, 308 intricatoreticulata Kosterm. 317, 386 irosinensis Elmer ex Merr. 291 Janowskyi Harms 190 kabaensis Baker f. 306 kinabaluensis Kosterm. 325 kingiana Ridley 260 koordersii Jain & Bennet 385 korthalsii Miq. 19,205, 251, 25222595 korthalsti (Miq.) Pellegr. 228 Kostermansii(Prijanto) Kos- term. 325 kunstleri King 271 laevigata Merr. 261 lagunensis Merr. 288 lanceolata Merr. 222, 246 lancifolia (Hook. f.) Harms 288 lancilimba Merr. 206, 242 langlassei C. DC. 288 lanuginosa King 232 Index to scientific plant names 39] (Aglaia) (Aglaia) (Aglaia) latifolia Miq. 272 var. feysmannii Koord. & Valeton 272 lauterbachiana Harms 255 lawii (Wight) Saldanha ex Ramamoorthy 13, 205-— 208, 210, 211, 228 laxiflora Mig. 202, 210,294 ledermannii Harms 385 leeuwenii Harms 249 lepidopetala Harms 206, 210, 223 lepidota Miq. 243 var. paupercula Miq. 243 lepiorrhachis Harms 205,242 leptantha Miq. 206, 210,211, 261, 262* var. borneensis C.DC. 261 leptoclada Harms 385 leucoclada C. DC. 202, 257 leucophylla King 207, 209, 210, 211, 271 littoralis Zippelius ex Miq. 228 llanosiana C. DC. 246 loheri Merr. 246 longifolia Teijsm. & Binn. 303 longipetiolata Elmer 288 longipetiolulata Baker f. 251 luzoniensis (Vidal) Merr. & Rolfe 202, 277 var. trifoliata Merr. & Rolfe 276 maboroana Harms 385 macrobotrys Turcz. 246 macrocarpa (Miq.) Pannell 206, 215 macrostigma King 208, 274*, 275 magnifoliola C. DC. 272 maingayi (Hiern) King 228 malaccensis (Ridley) Pan- nell 212*, 216 marginata Craib 288 matthewsti Merr. 298 megistocarpa Merr. 235 meliosmoides Craib 298 membranacea Kosterm. 315 membranifolia King 209, 301 menadonensis Koord. 288 meridionalis Pannell 224, 29] merrillii Elmer 317 micrantha Merr. 288 micropora Merr. 259 minahassae Koord. 272 mindanaensis Merr. ex Elmer 288 minutiflora Bedd. var. griffithii Hiern 303 miquelii Merr. 284 mirabilis Harms 298 mirandae Merr. 271 monophylia Perkins 277 monozyga Harms 212, 299 montana C. DC. 302 motleyana Stapf ex Ridley 272 moultonii Merr. 288 mucronulata C. DC. 272 multiflora Merr. 261 multifoliola Merr. 237 multinervis Pannell 202, 206, 222 myriantha Merr. 291 myristicifolia C. DC. 268 negrosensis Merr. ex Elmer 288 neotenica Kosterm. 298 nivea Elmer ex Merr. 386 novaguineensis (C. DC.) Cy DEZ283 nudibacca C. DC. 283 obliqua C.T. White & Fran- cis 259 ochneocarpa Merr. 310 odoardoi Merr. 298 odorata Lour. 11, 12,19, 201, 382*, 383 odoratissima Blume 200, 206, 208, 276 var. forbesii Baker f. 276 var. parvifolia Koord. & Valeton 276 var. pauciflora Koord. & Valeton 276 odoratissima auct. 243 oligantha C. DC. 297 oligocarpa Migq. 228 oligophylla Mig. 202, 210, 297 oligosperma (Pierre) Pellegr. 107 ovata Teijsm. & Binn. 288 oxypetala Valeton 288 pachyphylla Mig. 203, 234 palawanensis Merr. 288 palembanica Miq. 209, 304 var. borneensis Miq. ex Koord. 306 pallens (Merr.) Merr. 243 pallida Merr. 271 pamattonis Mig. 304 papuana (Merr. & L.M. Perry) Harms 95 parksii A.C. Smith 29] parviflora C. DC. 207, 269 parvifolia Merr. 243 parvifoliola C. DC. 286 pauciflora Merr. 288 pedicellata (Hiern) Kosterm. 228 penningtoniana Pannell 203, 212, 227 perfulva Elmer 233 perviridis Hiern 205, 260 phaeogyna Harms 385 pinnata (L.) Druce 386 pinnata (Blanco) Merr. 306 pirifera |] polyantha Ridley 297 polyneura C. DC. 207, 283 polyphylla Miq. 268 polystachya Wall. 188 porulifera C. DC. 284 procera C. DC. 269 pseudolansium Kosterm.324 puberulanthera C. DC. 202, 281 puncticulata Merr. 294 pycnocarpa Mig. 215 pycnoneura Harms 385 pyramidata | | pyricarpa Baker f. 267 pyriformis Merr. 210, 294 pyrrholepis Miq. 259 querciflorescens Elmer 288 racemosa Ridley 228 ramosii Quisumb. 245 ramotricha Pannell 208, 234 ramuensis Harms 306 rechingerae C. DC. 284 reinwardtiana Kosterm. 328 reinwardtii Miq. 288 reticulata Elmer ex Merr. 247 Flora Malesiana ser. I, Vol. 12 (1) (1995) 392 (Aglaia) (Aglaia) ridleyi P.T. Li & X.M.Chen 297 ridleyi (King) Pannell 221 rimosa (Blanco) Merr. 11, 205, 222, 246 rivularis Merr. 201, 279 rizalensis Merr. 277 robinsonii Merr. 288 rodatzii Harms 385 roemeri C. DC. 283 roxburghiana (Wight & Arn.) Mig. 11, 243 var. angustata Miq. 243 var. balica Miq. 243 var. paupercula (Miq.) Mig. 243 rubescens (Hiern) Dai 215 rubiginosa (Hiern) Pannell 203, 225, 226* rubra Ridley 281 rudolfi Harms 284 rufa Mig. 288, 306 var. celebica Mig. 288 rufa auct. 312 rufibarbis Ridley 208, 312*, 313% rufinervis (Blume) Bentv. 209, 302 rugulosa Pannell 212, 217 salicifolia Ridley 278 saltatorum A.C. Smith 270 samarensis Merr. 272 samoensis A. Gray 206, 256 sapindina (F. Muell.) Harms 207, 283 sapindina auct. 96 schlechteri Merr. & L.M. Perry 284 schraderiana Harms 385 schultzei Harms 385 schumanniana Harms 281 sclerocarpa C. DC. 228 scortechinii King 206, 254 sepalina (Kosterm.) Kos- term. 317 sexipetala Griff. 208, 210, 268 shawiana Merr. 298 sibuyanensis Elmer ex Merr. 228 silvestris (M. Roem.) Merr. 11, 205, 258*, 259 simplex Merr. 271 simplicifolia (Bedd.) Harms 202, 281, 298 sipannas Mig. 304 smithii Koord. 204, 245 sorsogonensis Elmer 288 sp. 1 Pannell 217 sp. 2 Pannell 219 sp. 3 Pannell 222 sp. 4 Pannell 251 sp. 5 Pannell 268 sp. 6 Pannell 267 sp. 7 Pannell 266 spaniantha Harms 385 spanoghei Blume ex Miq. 243 speciosa Blume 205, 250 var. macrophylla C. DC. 250 speciosa auct. 237 speciosa group 250 spectabilis (Miq.) Jain & Bennet 212, 220*, 221 splendens (Koord. & Val.) Koord. & Valeton 237 squamulosa King 203, 204, 239 squamulosa group 238 stapfii Koord. 288 steenisti Kosterm. 317 steinii Harms 385 stellatotomentosa Merr. 233 stellipila C. DC. 286 stenophylla Merr. 309 sterculioides Kosterm. 298 stipitata Li & Chen 228 subcuprea Merr. & L.M. Perry 204, 241 subgrisea Miq. 231 subminutiflora C. DC. 211, 286 submonophylla Mig. 228 subsessilis Pannell 210, 211, 286 subviridis Elmer ex Merr. 247 sulingi Blume 272 tarangisi Elmer 293 tayabensis Merr. 288 tembelingensis M.R. Hend. 288 tenuicaulis Hiern 209, 301 teysmanniana (Miq.) Miq. 197%; 208, 231 (Aglaia) tomentosa Teijsm. & Binn. 10, 209, 306, 307* tomentosa group 300 triandra Ridley 298 trichostemon C. DC. 302 trichostoma Harms 385 trimera Merr. 228 trimera Ridley 215 tripetala Merr. 213 triplex Ridley 215 trunciflora Merr. 288 turczaninowtl C. DC. 228 umbrina Elmer ex Merr. 248 unifoliolata Koord. 277 unifoliolata Ridley 298 urdanetensis Elmer ex Merr. 288 urophylla Harms 385 variisquama Pannell 204, 245 versteeghii Merr. & L.M. Perry 241 vilamilii Merr. 385 villosa (C. DC.) Merr. 288 vulpina Harms 385 winckelii Adelb. 302 yzermanni Boerl. & Koord. 113, 202, 278 zippelii Miq. 306 zollingeri C. DC. 375 Aglaiopsis Mig. 195 glaucescens Mig. 283 lancifolia (Hook. f.) Miq. 288 Aglaioxylon 2 Aglaja 246 Ailanthus integrifolia Lam. 387 flavescens Carriere 361 Ailantopsis Gagnep. 41 poilanei Gagnep. 43 Ailantus flavescens 361 Aitoniaceae (Harv.) Harv. 1,24 Aitonieae Harv. 1, 24 Alliaria |[Rumph.ex] Kuntze 61 sect. Didymocheton(Blume) Kuntze 66 sect. Typalliaria Kuntze 66 Alliaria Rumph. 107 acuminatissima (Blume) Kuntze 106 acutangula (Miq.) Kuntze 129, 130 Index to scientific plant names (Alliaria) alliacea (Forst. f.) Kuntze 90, 93 amooroides (Miq.) Kuntze 69 arborescens (Blume) Kuntze 103 beccariana (C.DC.) Kuntze 87 cauliflora (Turcez.) Kuntze 85 caulostachya (Miq.) Kuntze Te costulata (Miq.) Kuntze 106 cuneata (Hiern) Kuntze 87, 189 cyrtobotrya (Miq.) Kuntze 123 dasyphylla (Miq.) Kuntze 387 densiflora (Blume) Kuntze 81 euneuron (Miq.) Kuntze 387 fraterna (Miq.) Kuntze 106 gaudichaudiana (A. Juss.) Kuntze 68 glabra (C.DC.) Kuntze 107 grandis (Hiern) Kuntze 117 halmaheirae (Miq.) Kuntze 103 hiernii Kuntze 86 kunthiana (A. Juss.) Kuntze 103 lamponga (Miq.) Kuntze 106 lasiocarpa (Miq.) Kuntze 152, 155 latifolia (Benth.)Kuntze 124 lobbii (C. DC.) Kuntze 123 maingayi (Hiern) Kuntze 103 miqueliana (C.DC.) Kuntze 107 mollis Kuntze 90, 93 mollissima (Blume) Kuntze 90, 93 muelleri (Benth.) Kuntze 91, 93 multijuga (Blume) Kuntze 123 nageliana (C. DC.) Kuntze 107 nernstii (F. Muell.) Kuntze 103 nutans (Blume) Kuntze 71 (Alliaria) oppositifolia (F. Muell.) Kuntze 122 otophora (Miq.) Kuntze 69 procera (Hiern) Kuntze 109 ramiflora (Miq.) Kuntze 77 richii (A.Gray) Kuntze 90, 93 samoensis (A.Gray) Kuntze 90, 93 schizochitoides (Turcz.) Kuntze 91, 93 schultzii (C. DC.) Kuntze 129, 130 sessilis (Mig.) Kuntze 66 setosa (Span.) Kuntze 74 similis (Blume) Kuntze 106 spanoghei (Miq.) Kuntze 69 speciosa (A. Juss.) Kuntze ri teysmannii (C.DC.) Kuntze 91,98 thyrsoidea (Hiern) Kuntze 107 trichostyla (Miq.) Kuntze 81 turczaninowti (C. DC.) Kuntze 122 vrieseana (C. DC.) Kuntze 107 zippeliana Kuntze 91, 93 Amerina Noronha 194 Amoora Roxb. 18, 194, 212 sect. Amoora 212 sect. Aphanamixis (Blume) C.DGLS7 sect. Euamoora Miq. 187 sect. Pseudoguarea C. DC. 133 sect. Triamoora Kuntze 194 aherniana Merr. 213 amboinensis Miq. 189 aphana var. pubescens Miq. 189 aphanamixis Schultes & Schultes 189 aphanamixis auct. 387 auriculata Mig. 213 balansaeana C. DC. 135 beddomei Kosterm. 190 borneensis Mig. 193 caesifolia Elmer 179 canarana (Turez.) Hiern 13 cucullata Roxb. 213 cucullata auct. 189 393 (Amoora) cumingiana C. DC. 189 cupulifera Merr. 179 curtispica L.S. Gibbs 228 decandra (Wall.) Hiern 387 dysoxyloides Kurz 228 elmeri Merr. 190 forbesti S. Moore 387 fulva Merr. 174 ganggo (Miq.) Kurz 259 gigantea Pierre 222 grandifolia (Blume) Walp. 189 Janowskyi (Harms) Kosterm. 190 korthalsii Mig. 228 lactescens Kurz 228, 231 lanceolata Hiern 222 lauterbachii (Harms)C.DC. 190 lepidota Merr. 228 macrocalyx (Harms) C.DC. 190 macrocarpa Merr. 123 macrophylla Nimmo 189 maingayi Hiern 228 malaccensis Ridley 216 megalophylla C. DC. 190 var. frutescens C.DC. 190 mindorensis Merr. 179 myrmecophila Warb. 190 naumannii C.DC. 375, 385 nitidula Benth. 135 oligosperma Pierre 107 perrotetiana (A. Juss.) Steud. 189 polillensis Robins. 190 polystachya (Wall.) Wight & Arn. ex Steud. 188 racemosa Ridley 316 ridleyi King 221 rohituka (Roxb.) Wight & Arn. 9, 188 roxburghiana Korth. ex Blume 387 rubescens Hiern 215 rubiginosa Hiern 225 salomoniensis C. DC. 379 sogerensis Baker f. 190 spectabilis Miq. 221 sumatrana Miq. 194 teysmanniana Mig. 231 timorensis (A. Juss.) Wight & Arn. ex Steud. 189 394 (Amoora) tomentosa Korth. ex Blume 387 trichanthera Koord. & Vale- ton 215 vieillardii C. DC. 135 wallichii King 221 zollingeri (C. DC.) Koord. 375 Amura 189 Andersonia Roxb. 187, 194 cucullata Roxb. 213 rohituka Roxb. 188 Antelaea Gaertn. 329 azadirachta (L.) Adelb. 341 canescens Cels ex Heynh. 387 javanica Gaertn. 330, 334 Antelaea auct. 337 Anthocarapa Pierre 22, 61, 63, 133 balansaeana (C. DC.) Pierre 135 nitidula (Benth.) T.D. Penn. ex Mabb. 18, 134*, 135 sp) Ds Pennw!35 vieillardti (C. DC.) Pierre 135 Aphanamixis Blume 4—6, 8, 14,19, 22, 74, 187 agusanensis Elmer 190 amboinensis (Miq.) Harms 189 apoensis Elmer 190 blumei Span. 189 borneensis (Miq.) Merr. 188, 192*, 193 borneensis Harms 193 cochinchinensis Pierre 190 coriacea Merr. 190 cumingiana(C.DC.) Harms 189 davaoensis Elmer 190 elmeri (Merr.) Merr. 190 grandifolia Blume 189 humilis (Hassk.) Kosterm. 326 lauterbachii Harms 190 macrocalyx Harms 190 myrmecophila (Warb.) Harms 190 obliquifolia Elmer 190 pedicellata Ridley 193 perrotetiana A. Juss. 189 Flora Malesiana ser. I, Vol. 12 (1) (1995) (Aphanamixis) pinatubensis Elmer 190 polillensis (Robins.) Merr. 190 polystachya (Wall.) R.N. Parker 9) 10MI2 013 316. 21, 74, 87, 188, 388 pulgarensis Elmer 193 reticulosa Kosterm. 297 rohituka (Roxb.) Pierre 188 schlechteri Harms 190 sinensis How & Chen 190 sumatrana Harms 194 sumatrana (Miq.) Ridley 188, 194 sumatrana auct. 193 timorensis A. Juss. 189 trichanthera (Koord. & Valeton) Koord. 215 tripetala (Blanco) Merr. 189 velutina Elmer 190 Aphanamyxis 190 Aphanomyxis 187 Arbor Azadirachta Indica ... Breyne 341 Azadirachta Syriaca ... Breyne 331 fraxinifolio ... Bauhin 331 indica ... Bauhin 341 nussalavica Rumph. 71, 78 sebi Rumph. 69, 71 Argophilum Blanco 194 pinnatum Blanco 306 Aria Bepou Rheede 341 Artocarpus sp. 388 Azadirachta A. Juss. 5, 6, 18, 22,937 digyna Anon. 387 excelsa (Jack) Jacobs 18, 19, 337;33879340% foliis ramosis majoribus ... Breyne 332 foliis ramosis minoribus ... Breyne 332 Indica foliis Fraxini ... Breyne 341 indica A. Juss. 4, 9, 10, 12, [34195336 integerrima M. Roem. 43 Azedara Rat. 329 speciosa Raf. 330, 335 Azedarac Adans. 329 Azedaraca Raf. 329 amena Raf. 330, 335 Azedarach Dod. 331 Azedarach Mill. 329 commelini Moench ex Steud. 330, 335 florens Tourn. 332 floribus albis sempervirens Herm. 332 fructu polypyreno Herm. 332 Indica foliisnon deciduis ... Herm. 332 nigra Noronha 109 ramiflora Noronha 77 sambucina (Blume) Kuntze 331 sempervirens (L.) Kuntze 330, 332 var. australasica(A. Juss.) Kuntze 330, 334 var. dubia (Cav.) Kuntze 330, 334 var. glabrior (C. DC.) Kuntze 330, 335 tomentosa (Roxb.) Kuntze 143 vulgaris Gomez de la Maza 3305835 Azedarachaceae Schultes | Azedaracharum | Azederach deleteria Medik. 330, 335 odorata Noronha 331, 335 Baccaurea sylvestris Lour. 316 Beddomea Hook. f. 195 luzoniensis Vidal 277 racemosa Ridley 298 simplicifolia Bedd. 298 Beilschmiedia brassii Allen 36 Boboa seu Lanzones Kamel 317 Boswellia integra Blanco 386 obliqua Blanco 386 Brucea quercifolia Seem. 91, 93 sumatrana Roxb. 60 Buchanania paniculata Roxb. 189 spicata Roxb. ex Wall. 189 Cabralea A. Juss. 16, 61 canjerana ( Vell.) Mart. 12 subsp. polytricha(A. Juss.) TDs Renns2 Index to scientific plant names (Cabralea) eichleriana 12 microcarpus 12 pentandrus 12 polytricha 12 richardiana C. DC. 189 Cambania Comm. ex M. Roem. 61 Camunium [Rumph.] Roxb. 194 Carapa Aubl. 2,7, 14,16, 135 borneensis Becc. 377 carnulosa (Zoll. & Mor.) Kurz 379 guianensis Aubl. 383 indica A. Juss. 379 mekongensis (Pierre) Pel- legr. 377 moluccensis Lam. 376 var. elliptica Koord. & Valeton 376 var. gangetica Prain 376 var. ovalifolia Koord. ex Prain 376 var. typica Backer 379 moluccensis auct. 135, 375, 379 obovata Blume 379 var. microphylla (Pierre) Pellegr. 377 obovata auct. 377 procera DC. 13, 383 rumphii Kostel. 375 Carapa auctt. 371 Carapeae Harms 371 Castanospora longistipitata F.M. Bailey 185 Cedrela L. 2, 14, 358 sect. Joona Endl. 358 australis Mudie 367 australis F. Muell. 367 brevipetiolulata Haines 368 febrifuga Blume 364 f. ternatensis Miq. 367 f. timorensis Miq. 367 var. assamensis C. DC. 364 var. cochinchinensis (Pierre) C. DC. 368 var. glabrior C. DC. 364 var. inodora (Hassk.) C. DC. 370 var. pealii C. DC. 364 var. verrucosa C. DC. 364 (Cedrela) glabra C. DC. 361 glaziovii C. DC. 383 hainesii C. DC. 368 hexandra Wall. 367 inodora Hassk. 370 kingti C. DC. 368 var. birmanica C. DC. 368 longifolia Wall. ex C. DC. 361 var. kumaona C. DC. 361 mannii C. DC. 368 mexicana M. Roem. 383 microcarpa C. DC. 367 var. grandifoliola C. DC. 364 mollis Hand.-Mazz. 368 odorata L. 11, 13, 18, 383 odorata Blanco 370 serrata Royle 361 var. puberula C. DC. 361 serrulata Miq. 361 sinensis A. Juss. 360 var. hupehana C. DC. 361 var. lanceolata Li 361 var. schensiana C.DC. 361 sp. Wall. 355 sureni (Blume) Burkill 364 taratara Blanco 387 teysmannii Hassk. 367 toona Roxb. ex Rottl. & Willd. 9, 10, 12, 367 var. australis C. DC. 368 var. cuspidata C. DC. 368 var. gamblei C. DC. 368 var. hainesii (C. DC.) Haines 368 var. hasletii Haines 368 var. henryi C. DC. 364 var. latifolia Miq. ex C. DC. 368 var. multijuga Haines 368 var. nepalensis C.DC. 368 var. parviflora Benth. 367 var. pilipetala C. DC. 368 var. pilistaminea C. DC. 364, 368 var. pilistila C. DC. 364 var. puberula C. DC. 368 var. pubescens Franch. 368 var. pubinervis C.DC. 368 var. stracheyi C. DC. 368 var. sublaxiflora C. DC. 368 (Cedrela toona) var. talbotii C. DC. 368 var. vestita C.T. White 368 var. warburgii C.DC. 364 var. yunnanensis C. DC. 364 velutina DC. 367 Cedrelaceae R. Br. 1, 358 Cedrelopsis 7 Celastrus micrantha Roxb. 283 Chickrassia Wight & Arn. 354 nimmonit R. Graham ex Wight 355 tabularis (A. Juss.) Wight & Arn. 355 var. genuina Theob. 355 var. velutina (M. Roem.) Theob. 355 trilocularis (G. Don) M. Roem. 355 velutina M. Roem. 355 Chisocheton Blume 2, 4, 5, 7, 8, 14, 19, 21, 61, 63, 136 § Dasycolei Harms 175 § Graciles Harms 141 § Grandiflori Harms 171 § Pauciflori auct. 175 § Sandoricarpi Harms 175 § Schumanniani Harms 148 § Tetrapetali (Miq.) Harms 147, 160 sect. Chisocheton 147 series Paniculati Harms 160 series Pauciflori, Mabb. 171 series Schumanniani Harms ex Mabb. 148, 186 sect. Clemensia (Merr.) Airy Shaw 62, 141 sect. Cyrtochiton 62 sect. Dasycoleum (Turcz.) Harms 171, 175 sect. Euchisocheton Harms 141, 147 § Paniculati Harms 160 § Pauciflori Harms 171 sect. Hexapetalum Miq. 147 sect. Holopentas Mig. 62,171 series Pauciflori 171 series Sandoricocarpi Harms ex Mabb. 175, 186 396 Flora Malesiana ser. I, Vol. 12 (1) (1995) (Chisocheton) sect. Rhetinosperma (Radlk.) Mabb. 74, 184 sect. Tetrapetalum Miq. 147 acariianthus Harms 157 aenigmaticus Mabb. 139, 158 amabilis (Miq.) C.DC. 3, 63, 140, 163 var. sumatranum Migq. 163 amboinensis Valeton 164, 166 annulatus King 171 apoensis Elmer ex Merr. 167 archboldianus Merr. & L.M. Perry 157 balansae C. DC. 164, 166 barbatus C. DC. 167 beccarianus (Baill.) Harms 181 beccarianus auct. 182 benguetensis Elmer 164, 166 biroi Harms 153, 155 biroi Lane-Poole 179 biroi C.T. White 155 boridianus Harms 153, 155 brachyanthus Merr. 161 brachyanthus auct. 163 caroli Harms 153, 156 cauliflorus Merr. 139, 150 celebicus Koord. 139, 158 ceramicus (Miq.) C. DC. 140, 179 ceramicus auct. 181, 182 chinensis Merr. 164, 166 clementis Merr. 179 cochinchinensis Pierre 164, 166 coriaceus Pierre 164, 166 costatus Hiern 117 crustularii Mabb. 138, 146 cumingianus (C. DC.) Harms 8, 140,164,165*, 166 subsp. balansae (C. DC.) Mabb. 164, 166 subsp. cumingianus 166 subsp. kinabaluensis (Merr.) Mabb. 138, 164, 166 curranii Merr. 140, 180 curranii auct. 181, 182 dempoense Baker f. 107 divergens Blume 167 (Chisocheton) diversifolius Miq. 139, 174 doctersii Harms 179 dysoxylifolius (Kurz) Hiern 171 erythranthus Merr. & L.M. Perry 157 erythrocarpus Hiern 140, 183 erythrocarpus auct. 157 eurycalyx Harms 153, 155 forbestti CoDC152; 155 formicarum Harms 153, 155 fragrans Hiern 167 frutescens C. DC. 153, 155 fulvus Merr. 167 gjellerupti Harms 153, 156 glirioides P.F. Stevens 139, 158 globosus Pierre 179 glomeratus Hiern 167 glomeratus auct. 158, 161 graciliflorus Harms 157 granatum Mabb. 139, 170 grandiflorus (Kurz) Hiern 171 grandiflorus auct. 117 hackenbergii Harms 163 harmandianus Pierre 164, 166 holocalyx Hiern 167 illustris Ridley 163 jJunghuhnii (Miq.) C. DC. 79 kinabaluensis Merr. 164, 166 kingii Harms 178, 179 kingti Koord. 184 koordersii Mabb. 140, 184 kunstleri King 145 lamekotensis Harms 153,156 lamii Diels ex H.J. Lam 153, 155 lansiifolius Mabb. 140, 168%, 169 laosensis Pellegr. 140, 160 lasio caroli Mabb. 155, 156 lasio entity formicarum Mabb. 154, 155 entity lasiocarpus Mabb. 154, 155 entity novoguineensis Mabb. 154, 155, 155 entity pachyrhachisMabb. 154, 156 (Chisocheton lasio) entity schlechteri Mabb. 154, 156 entity schumannii Mabb. 155, 156 entity trichocladus Mabb. 154, 156 entity versteegi1 Mabb. 154, 156 entity weinlandii Mabb. 155 lasiocarpus (Miq.) Valeton 4, 16, 140, 152, 155 lasiogynus Boerl. & Koord. 138, 162 lauterbachii Harms 152, 156 laxiflorus King 161, 167 ledermannii Harms 153, 156 leptopetalus Harms 157 longistipitatus (F.M. Bailey) L.S. Smith 140, 185, 186* macranthus (Merr.) Airy Shaw 138, 141 macrophyllus King 140, 175, TAS subsp. fulvescens Mabb. 178 subsp. macrophyllus 178, 179 macrophyllus (K. Schum.) Harms 152, 156 macrothyrsus King 179 medusae Airy Shaw 138, 142, 143* f. hiascens Jacobs 142 medusae auct. 141 mendozae Hildebr. 140, 171, 174 microcarpus Koord. & Vale- ton 181, 182 var. moluccanus Valeton 181 montanus P.F. Stevens 139, 151 morobeanus Harms 164,166 multijugis C. DC. 153 var. glabrior C. DC. 153, [55 myrmecophilus Merr. & Le M.Perry 15398 novobrittannicus P.F. Stevens 139, 151 novoguineensis C.DC. 152, is) Index to scientific plant names (Chisocheton) oreophilus Harms 153 pachycalyx Harms 179 pachyrhachis Harms 152, 156 pachyrhachis Hartley et al. 164, 166 paniculatus Hiern 164, 166 paohaniana Li & Chen 184 parvifoliolus Merr. 181,182 patens Blume 138, 140, 167 var. genuinus Valeton 167 var. minor Valeton 167 var. patens (Blume) Rid- ley 167 var. robustus Valeton 167 pauciflorus King 139, 173 paucijugus (Miq.) B.D. Jackson 181, 183 peekelianus Harms 179 penduliflorus Planch. ex Hiern 138, 140, 145 pentandrus (Blanco) Merr. 140, 180, 182 subsp. medius Mabb. 182 subsp. paucijugus (Miq.) Mabb. 182, 183 subsp. pentandrus 182 perakensis (Hemsl.) Mabb. 138, 171 philippinus (Turez.) Harms 181, 182 pilosus C. DC. 140, 157 pohlianus Harms 138, 152 pohlianus auct. 149, 157 polyandrus Merr. 138, 139, 144 polyanthus Harms 185 princeps Hemsl. 143 rex Mabb. 184 rhytidocalyx Airy Shaw 179 rigidus Ridley 105 ruber Ridley 3, 138, 160 rubiginosus King 143 rugosus Pierre 143 sandoricocarpus Koord. & Valeton 179 sapindinus P.F. Stevens 139, 159 sarasinorum Harms 139, 172 sarawakanus (C.DC.) Harms 139, 140, 161 sayeri (C. DC.) P. F. Stevens 139, 157 (Chisocheton sayeri) var. pilosus (C. DC.) P. F. Stevens 157 schlechteri Harms 153, 156 schoddei P. F. Stevens 139, 148, 149* schumannii C.DC. 153, 155 schumannii Harms 156 schumannii auct. 155, 157 setosus Ridley 138, 146 siamensis Craib 164, 166 sogerensis Baker f. 94 sorsogonensis Elmer ex Merr. 181, 182 species A, Mabb. 186 species B, Mabb. 186 spectabilis (Miq.) C. DC. 179 spicatus Hiern 181, 183 stellatus P.F. Stevens 140, 184 sumatranus Baker f. 267 tenuis P. F. Stevens 138, 149 tetrapetalus (Turez.) C.DC. 167 thorelii Pierre 164, 166 tomentosus (Roxb.) Mabb. 3, 138, 139, 143 torricelliensis Harms 153, 156 trichocladus Harms 153, 156 urdanetensis Elmer ex Merr. 167 versteegii C. DC. 153, 156 vindictae Mabb. 140, 175, 176* vrieseanus C. DC. 167 vulcanicus Elmer ex Merr. 179 warburgii Harms 139, 148 weinlandii Harms 152, 155 Chisochiton sp. Merr. 181 Chisogeton 117 Chukrasia A. Juss. 4, 5,18,21, 354 chickrassa Schultze-Motel 359 nimmonii (R. Graham ex Wight) Merr. & Chun tabularis A. Juss. 3, 10, 18, 355; 356* var. attopeuensis Pierre 355 397 (Chukrasia tabularis) var. dongnaiensis (Pierre) Pellegr. 355, 358 var. malaccana C.DC. 355 var. microcarpa (Pierre) Pellegr. 355 var. guadrivalvis Pellegr. 355 var. velutina (M. Roem.) Pellegr. 355 velutina (M. Roem.) C.DC. 355 var. dongnaiensis Pierre 359 var. macrocarpa C. DC. 355 Chuniodendron Hu 187 spicatum Hu 190 yunnanense Hu 190 Cipadessa Blume 7, 22, 57 baccifera (Roth) Miq. 58*, 59 var. cinerascens (Pellegr.) Pellegr. 59 var. integerrima Theob. 59 var. rothii Theob. 59 var. sinensis Rehd. & Wils. 59 borneensis Mig. 55 cinerascens (Pellegr.) Hand.-Mazz. 59 fruticosa Blume 59 var. cinerascens Pellegr. 59 sinensis (Rehd. & Wils.) Hand.-Mazz. 59 subscandens (Teijsm. & Binn.) Mig. 59 warburgii Perkins 59 Clausena chrysogyne Miq. 55 indica Oliv. 10 Clemensia Merr. 136, 141 macrantha Merr. 141 Cupania sp. Wall. 164, 167 Cuveraca Jones 358 Cyppadessa 60 Dasycoleum Turez. 136, 171 beccarianum Baill. 181, 183 cumingianum C. DC. 164, 166 forbesii Baker f. & Norm. $5153 philippinum Turez. 181, 182 sarawakanum C. DC. 161 sayeri C. DC. 157 398 Flora Malesiana ser. I, Vol. 12 (1) (1995) Didymocheton Blume 61, 62, 66 albiflorum (C. DC.) Harms 69 amooroides (Mig.) Harms 69 betchei (C. DC.) Harms 69 carrti Harms 73 decandrum (Blanco) Harms 68 funkii (C.DC.) Harms 91,93 gaudichaudianum A.Juss.68 hainanense (Merr.) Harms 91593 hirtum (Ridley) Harms 73 leschenaultianum A. Juss. 71 littorale Hassk. 31 maota (Reinecke) Harms 69 nutans Blume 71 pruriens Zipp. ex Span. 74 pumilum Blume ex Miq. 31 richii A. Gray 90, 93 sessile (Miq.) Kosterm. 66 spanoghei (Miq.) Harms 69 stellato-puberulum (C. DC.) Harms 73 suaveolens Blume ex Miq. 71 Didymochiton 90 Diplotaxis Wall. ex Kurz 136 Disoxylum 69 Disyphonia Griff. 61 Dracontomelon pilosum Seem. 193 Dysoxylon sp. 131 Dysoxylum Blume 2—5, 8, 14, 18, 22, 61, 63, 386 sect. Cleistocalyx F. Muell. 66 sect. Cyrtochiton Miq. 62, 63, 116, 133 sect. Dicrypta Migq. 116 sect. Didymocheton(Blume) Miq. 66, 75 sect. Dysoxylum 62, 63, 66 sect. Epicharis (Blume) Migq. 66, 75 sect. Goniocheton (Blume) Miq. 66 sect. Macrocheton Blume 66 sect. Prasoxylon(M. Roem.) Miq. 66 sect. Siphodysoxylum Pierre 116 (Dysoxylum) sect. Siphonodiscus F. Muell. 66 abo Hosakawa 91, 93 aculeatissinum Steud. 106 acuminatissimum Blume 106 acutangulum Miq. 16, 18,63, 129 subsp. acutangulum 130 subsp. foveolatum (Radlk.) Mabb. 130 acutangulum auct. 128 agusanense Elmer 85 alatum Harms 63, 76 albiflorum C. DC. 69 alliaceum (Blume) Blume 63, 65, 90, 106 var. angustifolium Valeton 106 var. genuinum Valeton 106 var. lanceolatum Koord. & Valeton 106 var. laxiflorum (Blume) CeDES106 var. laxiflorum Ridley 112 var. pauciflorum Koord. & Valeton 106 alliaceum (Forst. f.) Seem. 90593 alliaria (Buch.-Ham. ) Balakr. 90, 93 alternatum Ridley 123 alternifolium Elmer 131 altissimum Merr. 110 amooroides auct. 74 amooroides Miq. 69, 76 var. otophorum (Miq.) Koord. & Valeton 69 var. pubescens 69 var. typicum Koord. & Valeton 69 andamanicum King 123 angustifoliolum Merr. 87, 89 angustifolium King 3,4,62, 63, 65, 112 annae Mabb. 65, 94 apoense Elmer 107 arborescens (Blume) Miq. 16, 63, 64, 103, 104* f. ceramicum Mig. 103 f. timorense Mig. 103 var. timorense (Miq.) CaADGES103 (Dysoxylum) archboldianum Merr. & L.M. Perry 107 arnoldianum K. Schum. 110 aurantiacum Elmer 123 bakerarum Guillaum. 69, 71 bakeri Elmer 123 bamleri Harms 108 beccarianum C. DC. 87 benguetense Elmer 123 betchei C. DC. 69, 91, 93 biflorum Merr. 101 bijugum (Lab.) Seem. 93 biloculare Koord. & Valeton 123 blancoi Vidal 69 blumei Migq. 123 var. grandiflorum Koord. 123 var. typicum Koord. & Valeton 123 boridianum Mabb. 63, 65, 100 brachybotrys Merr. 62, 65, 128 brachycalycinum Harms 107 brachypodum Baker f.78, 80 brachystachys Ridley 128 brassil Merr. & L.M. Perry 62, 65, 114 brevipaniculum C. DC. 64, 84, 89 brevipes Hiern 106 callianthum Merr. & L.M. Perry 78, 80 capizense Merr. 122 carolinae Mabb. 63, 131,132* cauliflorum Hiern 16, 62,64, 86, 88* var. elongatum Ridley 86, 190 var. tomentellum Stapf 86 cauliflorum auct. 84,85, 190 cauliflorum group 62 caulostachyum Miq. 70, 77, 80 cerebriforme F. M. Bailey CO LATS championt Hook. f. & Thomson ex Thw. 6 cochinchinense Pierre 123 var. harmandianum (Pierre) Pellegr. 123 var. quocense Pierre 123 Index to scientific plant names (Dysoxylum) confertiflorum Merr. & L.M. Perry 124 corneri Hend. 117 costulatum (Miq.) Miq. 106 crassum Mabb. 62, 64, 98 cumingianum C.DC. 13, 16, 62, 64, 85 cuneatum Hiern 87, 189 cyrtobotryum Miq. 62, 66, 123 var. B forma borneensis 123 var. borneense Mig. 123 dasyphyllum Migq. 387 decandrum (Blanco) Merr. 68 dehiscens Elmer 141 dempoense (Baker f.) Harms 107 densevestitum C.T. White 78, 80 densiflorum (Blume) Miq. 64, 81, 82*, 83* var. minus Koord. & Vale- ton 81 distantinerve F. Muell. 124 dumosum King 64, 99 elmeri Merr. 81 enantiophyllum Harms 62, 63, 65, 126 esquirolii Lévl. 355 euneuron Miq. 387 euphlebium Merr. 107 excelsum Blume 63, 65, 109 var. genuinum Koord. & Valeton 109 var. glabriflorum Migq. 106 var. hasseltii Mig. 109,110 var. parvifolium Koord. & Valeton 109 var. pedicellatum Koord. & Valeton 109 filicifolium Li 91, 93 fissum C.T. White & Francis ex Lane-Poole 78, 80 flavescens Hiern 65, 128 floribundum Merr. 91, 93 forsteri C. DC. 90, 93 forsythianum Warb. 103 foveolatum Radlk. 129, 130 foxworthyi Elmer 87 fragrans Elmer ex Merr. 78 fraternum Mig. 106 (Dysoxylum) fulvum Airy Shaw 127 funkii C. DC. 69, 91, 93 gaudichaudianum (A. Juss.) Mig. 63, 64, 71, 76, 68 var. pubescens 68 gjellerupti C. DC. 103 glabrum C. DC. 107 glochidioides Elmer ex Merr. 78 gobara (Buch.-Ham.) Merr. 109 grande Hiern 62, 65, 117, 118* grandifoltum Merr. 123 grandifolium auct. 179 griffithii Hiern 81, 128 hainanense Merr.77, 91, 93 var. glaberrimum How & Chen 91, 93 halmaheirae (Miq.) C. DC. 103 var. subobovatum C. DC. 103 hamiltonii Hiern 90, 93 hapalanthum Harms 108 harmandianum Pierre 123 hasseltii (Mig.) Koord. & Valeton 110 havilandii Ridley 110 hexandrum Merr. 123 heyneanum Valeton ex K. Heyne 123 hirsutum C. DC. 71 hirtum Ridley 73 hongkongense (Tutch.) Merr. 63, 109 huberti Harms 110 ilocanum Merr. 122 inopinatum (Harms) Mabb. 62, 64, 97 intermedium Merr. & L.M. Perry 69, 71 interruptum King 117 kaniense Harms 62, 66, 125 kinabaluense Merr. 123 klanderi F. Muell. 63, 109 klemmei Merr. 107 kunthianum (A. Juss.) Miq. 103 lactiflorum Baker f. 74, 75 lampongum Mig. 106 lamproanthum Merr. & L.M. Perry 74 399 (Dysoxylum) lanceolatum Elmer 107 lasiocarpum Mig. 152, 155 lasiophyllum Baker f. 117 latifolium Benth. 62, 64, 124 latifolium auct. 122 laxiflorum Blume 106 laxum Merr. 101 ledermannii Harms 114 lenticellare Gillespie 10, 12, 13 leptorrhachis Harms 91 leschenaultianum (A. Juss.) Koord. & Valeton 71 leytense Merr. 77, 80 lobbii C. DC. 123 var. sumatranum Baker f. 117 loheri Merr. 78 longicalicinum C.DC.77, 80 longiflorum Merr. 77 longifolium Blume 106 longipetalum C. DC. 77 lukii Merr. 117 var. paucinervium How & Chen 117 macgregorii C. DC. 110 macrocarpum Blume 66, 116 f. sumatranum 116 var. sumatranum (Miq.) CYDC. Ti6 macrophyllum Teijsm. & Binn. 69 macrothyrsum Beehler 84 macrothyrsum auct. 109 var. microbotrys (King) Ridley 109 magnificum Mabb. 62, 65, 113 magnifolium C. DC. 74 maingayi Hiern 103 maota Reinecke 69, 71 megalanthum Hemsl. 77, 80 micranthum Merr. & L.M. Perry 96 microbotrys King 110 microcricum Harms 386 microstachyum Harms 89 miquelianum C, DC. 107 molle Mig. 91, 93 mollissimum Blume 3, 16, 18, 63, 64, 90 subsp. molle (Miq.) Mabb. 9, 12, 93 400 Flora Malesiana ser. I, Vol. 12 (1) (1995) (Dysoxylum mollissimum) (Dysoxylum) (Dysoxylum) subsp. mollissimum 93 var. halmaheirae Miq. 90, 93 var. sumatranum Mig. 90, 93 var. teysmannit (C. DC.) Koord. & Valeton 90, 93 monticola Harms 107 motleyanum (C. DC.) Ridley 110 muelleri Benth. 91, 93 multifoliolatum Li & Chen 123 multijugum (Blume) Adelb. 123 multijugum Arn. 164, 166 nagelianum C. DC. 107 nernstii F. Muell. 103 novoguineense Warb. 77, 80 novohebrideanum C. DC. 103 nutans (Blume) Miq. 64, 71 f. tomentosum Mig. 71 var. sumatranum Miq. 71 var. tomentosum (Miq.) CsD@r7A nymanianum Harms 94 oblongifoliolum Quisumb. & Merr. 123 octandrum (Blanco) Merr. 90795 oppositifolium F. Muell. 62, 63, 122 otophorum Migq. 69 oubatchense Harms 135 pachyrhache Merr. 62, 65, MSO 121 pachystachyum Harms 125 palawanense Merr. 122 pallidum Merr. 110 panayense Merr. 123 paniculatum Arn. ex Wight 164 papillosum King 65, 116 papuanum (Merr. & L.M. Perry) Mabb. 65, 95 parasiticum (Osb.) Kosterm. 63, 64, 76, 79* pauciflorum Merr. 65, 101, 102* paucijugum C. DC. 71 peerisiae Kosterm. 110 pettigrewianum F. M. Bailey 16, 62, 64, 84 phaeotrichum Harms 65,111 pilosum A.C. Smith 91, 93 platyphyllum Merr. 107 procerum Hiern 109 var. integrum C. DC. 109 var. macranthum C. DC. 109 var. motleyanum C. DC. 109, 110 pubescens Teijsm. & Binn. 68 pulchrum Ridley 107 pyriforme Merr. 85 quaifei C. DC. 69 quercifolium (Seem.) A.C. Smith 91, 93 racemosum King 123 var. arboreum King 123 ramiflorum Miq. 77 ramostt Merr. 122, 133 randianum Merr. & L.M. Perry 65, 115 revolutum Elmer 131 richardsonianum Merr. & L.M. Perry 78 richii (A. Gray) C.DC. 9, 12, 90, 93 rigidum (Ridley) Mabb. 65, 105 robinsonii Merr. 78, 80 roemeri C. DC. 77, 80 roseum | 1 rostratum Merr. 107 rubrum Merr. 103 rufum (Rich.) Benth. 58, 68 var. glabrescens Benth. 68 rufum auct. 71 rugulosum King 62, 65, 127 rumphii Merr. 78, 80 salutare Fern.-Vill. 69 samoense A. Gray 90, 93 sattelbergense Merr. & L.M. Perry 107 schiffneri F. Muell. 77, 80 schizochitoides (Turcz.) CDEs91F 93 schultzii C. DC. 126, 129, 130 sericeum (Blume) Adelb. 77 sericiflorum C.T. White 74, 76 sericopetalum Harms 115 sessile Miq. 62, 64, 66, 67* setosum (Span.) Miq. 64,71, 74 siargaoense Merr. 85 sibuyanense Elmer 103 simile Blume 106 sorsogonense Elmer 164, 166 sp. | 131 sp 2433 sp2ouls3 sp. 4 133 sp. 385 sp. 1 131 spanoghei Miq. 69 sparsiflorum Mabb. 65, 99 speciosum (A. Juss.) Mig. 77 spectabile (A. Juss.) Hook. f. 910 spiciflorum Zipp. ex Span. 189 steinitt Harms 89 stellatopuberulum C. DC. 21, 64, 73 sulphureum Elmer ex Merr. 85 tafaense Merr. & L.M. Perry 74, 76 testaceum Elmer 85 teysmannii C. DC. 91, 93 thaumasianthum Harms 74 thyrsoideum Hiern 107 var. andamanicum King 107 tpongense Pierre 123 triangulare Merr. 85 trichostylum Mig. 81 f. glabrum Mig. 81 f. pubescens Migq. 81 trukense Kaneh. 91, 93 turbinatum King 110 turczaninowti C. DC. 122 turczaninowil auct. 123 undulatum Hend. 127 urens [Teijsm. & Binn.] Valeton 66 variabile Harms 64, 93, 94 velutinum Koord. 91, 93 venosum Merr. 122 venulosum King 123 verruculosum Merr. 119 vestitum Warb. 69 Index to scientific plant names (Dysoxylum) vrieseanum C. DC. 107 wenzelii Merr. 122 whiteanum Merr. & L.M. Perry 84 yunzaingense Merr. & L.M. Perry 65, 96 Ebermeyera 31 Ebermaiera pulchella Zipp. ex Span. 31 Ekebergia 7 capensis Sparrm. 9, 10 indica Roxb. 59 integerrima Wall. 59 senegalensis 9, 10 Entandrophragma C. DC. 2, 13, 14, 18 caudatum Sprague | 1 cylindricum (Sprague) Sprague 11, 12 sapele 18 utile 18 Epicharis Blume 61, 66 alata (Harms) Harms 76 altissima Blume 81 brachybotrys Harms 89 brachypoda (Baker f.) Harms 78 calliantha (Merr. & L.M. Perry) Harms 78 cauliflora Blume 77 caulostachya (Miq.) Harms 77 cumingiana (C.DC.) Harms 85 cuneata (Hiern) Harms 87, 189 densevestita (C.T. White) Harms 78 densiflora Blume 81 dubiosa Span. ex Mig. 109 exarillata Nimmo 228 glabra Blume ex Mig. 81 hierniana Harms 86 (Epicharis) megalantha(Hemsl.) Harms 77 mollis Wall. ex Voigt 90, 93 pachyrhachis (Merr.) Harms 119 procera (Hiern) Pierre 109 pyriformis (Merr.) Harms 85 ramiflora Pierre 77 robinsonii (Merr.) Harms 78 rumphii (Merr.) Harms 78 schiffneri (F. Muell.) Harms i sericea Blume 77 setosa Span. 74 siargaoensis (Merr.) Harms 85 sp. Kurz 117 speciosa A. Juss. 77, 80 triangularis (Merr.) Harms 85 triangularis auct. 87 trichostyla Zipp. ex Miq. 77 Epipremnum sp. 387 Eumacrochiton 66 Euphoria exstipulata Griff. 303 Euphorianthus euneurus (Miq.) Leenh. 387 Flindersia R. Br. 387 Goniocheton Blume 61, 66 arborescens Blume 103 aurantiacum Zipp.ex Miq.69 Gonioscheton 103 Granatum [Rumph. ex] Kuntze 371 litoreum latifolium Rumph. 376 litoreum parvifolium Rumph. 377, 379 moluccensis (Lam.) Kuntze 376 obovatum (Blume) Kuntze 379 hyacinthodora Harms 78, 80 Guarea L. 2, 7, 14, 16, 61, 63 kunthiana A. Juss. 103 leytensis (Merr.) Harms 77 loheri (Merr.) Harms 78 longicalicina (C. DC.) Harms 77 longiflora (Merr.) Harms 77 longipetala (C.DC.) Harms 77 macrocarpa Migq. 215 acuminata Wall. 109 alliacea Blume apud Nees 106 alliaria [Aiken] Buch.-Ham. 90, 93 amaris Buch.-Ham. 189 axillaris Blume 106 cauliflora Reinw. ex Blume FUT | 401 (Guarea) densiflora (Blume) Spreng. 81 disyphonia Griff. 109 foetida Blume 93, 106 gobara | Aiken] Buch.-Ham. 109, 164 grandis Wall. 117 guidonia (L.) Sleumer 113 macrocarpa Blume apud Nees 116 mollis Wall.ex Hiern 90, 93 oblonga Wall. 109 oblongifolia Griff. 379 paniculata Buch.-Ham. 93 paniculata Roxb. 164, 166 paniculata auct. 90 procera Wall. 109 sericea (Blume) Spreng. 77 Hantol Kamel 347 Harpagonia Noronha 61, 388 Harpullia cupanioides Roxb. 388 Hartighesia 90 Hartighsea A. Juss. 61 acuminata Migq. 103 alliaria (Buch.-Ham.) Arn. 90,93 angustifolia Mig. 309 cauliflora Turez. 85 costulata Miq. 106 excelsa (Blume) A. Juss. 109 forsteri A. Juss. 90, 93 gobara Wight & Arn. ex Voigt 109 mollissima (Blume) A. Juss. 90, 93 ramiflora Griff. 81, 128 schizochitoides Turcz. 91,93 sp. Turcz. 122 sumatrana Miq. 103 trichosiphone F. Muell. ex CysDG. 94; 93 Hearnia F. Muell. 195 beccariana C. DC. 309 cumingiana (Turez.) C.DC. 293 elliptica (Blume) C. DC. 288 glaucescens (Miq.) C. DC. 283 var. novaguineensis C. DC. 283 lancifolia (Hook. f.) C. DC. 288 402 Flora Malesiana ser. I, Vol. 12 (1) (1995) (Hearnia) macrophylla C. DC. 283 sapindina F. Muell. 283 sarawakana C. DC. 251 villosa C. DC. 288 Heckeldora Pierre 61 Heynea Roxb. ex Sims 2, 8, 22,41 affinis A. Juss. 43 cochinchinensis Baillon 48 connaroides (Wight & Arn.) Voigt 43 fruticosa Teijsm. & Binn. 43 multijuga Blume 123 quinquejuga Roxb. 43 quinquejuga Spreng. 302 sumatrana Mig. 43 trijuga Roxb. ex Sims 12,41, 42*, 44% f. pubescens (Kurz) Craib 4] var. bijuga C. DC. 41 var. microcarpa Pierre 41 var. multijuga C. DC. 41 var. pilosula C. DC. 41 var. velutina Gagnep. 41 velutina How & Chen 41 Heyneoxylon 2 Heynia Roxb. 41 Khaya 6, 13, 18, 19 grandifolia C. DC. 11 senegalensis A. Juss. 13 Lachanodendron album Reinw. ex Blume 386 domesticum Nees 316 Lamiofrutex Laut. 34 papuanus Laut. 36 Lance Bontekoe 317 Lansium Correa 6, 7, 18, 22, 314 sect. Neolansium Harms B22. aqueum (Jack) M.Roem. 316 breviracemosum Kosterm. 315, 321 cinereum Hiern 324 domesticum Correa 9, 12, NS OH22eAS1S S17 *, 318*, 386 var. aqueum Jack 315 var. langsat Jack 316 var. pubescens Koord. & Valeton 316 var. typicum Backer 316 (Lansium) dubium Merr. 326 humile Hassk. 326 javanicum Koord. & Valeton ex Moll & Janss. 316 javanicum M. Roem. 316 kostermansii Prijanto 325 membranaceum (Kosterm.) Mabb. 315 monophyllum Merr. ex Perkins 328 montanum Jack ex Spreng. 386 montanum Rumph. 190 parasiticum (Osb.) Sahni & Bennet 77, 317 var. aqueum (Jack) Sahni & Bennet 317 pedicellatum Hiern 228 pedicellatum Kosterm. 317 sepalinum Kosterm. 317 silvestre [Rumph.] M. Roem. 259 sp. Merr. 326 Lansium Rumph. 317 Lansones Blanco 317 Lepisanthes 386 forbesii Baker f. 87 tetraphylla (Vahl) Radlk. 386 Leptonychia caudata (G. Don) Burret 388 Lovoa 18 Macrocheton (Blume) M. Roem. 61 sect. Macrocheton M.Roem. 66 sect. Schoutensia M.Roem. 66 excelsum (Blume) M.Roem. 109 forsteri M. Roem. 90, 93 Macrochiton 61, 66, 90, 109 Mallea A. Juss. 57 baccata Heyne ex Wall. 59 integerrima Wall. ex Voigt 59 montana Herb. Madras ex Wall. 59 parviflora Moon 59 rothii A. Juss. 59 subscandens Teijsm. & Binn. 59 Megaphyllaea Hemsl. 2, 62, 136 (Megaphyllaea) annulata (King) Ridley 171 perakensis Hemsl. 171 sp. Merr. 142 Melia L. 5, 6, 14, 20, 329 angustifolia Schum, & Thonn. 331, 335 argentea Hiern 331, 334 arguta DC. 331, 335 australasica A. Juss. 331, 334 australis Sweet 331, 334 azadirachta L. 341 azedarach L. 4, 9-13, 18, 19, 330, 333*, 336 var. acuminatissima Miq. 3305335 var. australasica (A. Juss.) C. DE, 3305334 var. biglandulosa Pierre 330 var. cochinchinensis (Pierre) Pellegr. 330, 334 var. floribunda (Carr.) Morren 330, 335 var. glabrior C. DC. 330, 335 var. glandulosa Pierre 330, 334 var. incisa Mig. 330, 335 var. intermedia (Mak.) Mak. 330, 335 var. japonica (G. Don) Mak. 330, 335 subvar. intermedia Mak. f. albiflora Mak. 330, 335 subvar. semperflorens (Mak.) Mak. 330, 335 subvar. toosendan (Sieb. & Zucc.) Mak. 330, 335 var. javanica Koord. & Valeton 330, 334 var. sambucina (Blume) Mig. 330, 335 var. sempervirens L. 330, 335 var. squamulosa C. DC. 330, 334 var. subtripinnata Miq. 3804335 var. toosendan (Sieb. & Zucc.) Mak. 330, 335 Index to scientific plant names (Melia azedarach) var. umbraculifera Knox ex Sarg. 330, 335 var. umbraculiformis Hort. ex Berck. & Bail. 330, 335 f. umbraculifera (Sarg.) Rehd. 330, 335 azedarach sensu Blanco 334 baccifera Roth 59 birmanica Kurz 331, 334 bogoriensis Koord. & Vale- ton 331, 334 bukayun Royle 331, 335 candollei A. Juss. 331, 334 chinensis Sieb. ex Mig. 331, 335 cochinchinensis M. Roem. commelini Medik. 330, 335 composita Willd. 331, 334 var. biglandulosa 330 var. cochinchinensis Pierre 331 dubia Cav. 330, 334 excelsa Jack 337 flaccida Zipp. ex Span. 331, 334 floribunda 335 florida Salisb. 330, 335 foliis decompositis L. 332 foliis duplicato-pinnatis L. 332 foliis pinnatis L. 341 fraxinifolia Salisb. 341 guineensis G.Don 331, 335 hasskarlii K. Koch 331, 335 iloilo Blanco 388 indica (A. Juss.) Brandis 341 integerrima Buch.-Ham. 43 Japonica G. Don 331, 335 var. albicans C. DC. 331 var. semperflorens Mak. 331 japonica Hassk. 331, 335 javanica M.Roem. 331, 335 koetjape Burm. f. 346 orientalis M.Roem. 331, 335 parasitica Osb. 76 pendula Reinw. ex Miq. 167 penduliflora Wall. 145 pinnata Stokes 341 pubescens Reinw. ex Miq. 71 (Melia) pumila Moon 30 robusta Roxb. 331, 334 sambucina Blume 331, 335 sempervirens (L.) Sw. 330, 335 superba Roxb. 331, 334 tomentosa Miq. 334 tomentosa Roxb. 143 tomentosa auct. 331, 355 toosendan Sieb. & Zucc. 11, 331/335 Meliacea rugosa Wall. 143 singapureana Wall. 297 wightiana Wall. 189 Meliaceae Juss. 1 subfam. Melioideae 5, 16, 18, 20, 24 tribus Aglaieae Blume 6, 16, 20, 62, 187 tribus Guareeae T. D. Penn. 7, 16, 20, 60, 62 tribus Melieae DC. 6, 16, 329 tribus Sandoriceae T. D. Penn. 16, 344 tribus Trichilieae DC. 6, 16, 20, 40 tribus Turraeeae Harms 6, 7, 16, 24 tribus Vavaeeae Harms 6, 16, 34 subfam. Neomangenotioi- deae Leroy 16 subfam. Swietenioideae Harms 4, 5,7, 16, 18, 20, 354 tribus Cedrelacearum 354 tribus Cedreleae (R. Br.) DC. 1, 16, 358 tribus Swietenieae A. Juss. 16, 354 tribus Xylocarpeae Blume 16, 20, 371 Meliadelpha Radlk. 61 Meliae | Meliapollis 7 Melioschinzia K. Schum. 136 macrophylla K.Schum. 152, 156 Melospermum Scort. ex King 55 rubro-stamineum Scort. ex King 55 403 Milnea Roxb. 194 argentea Reinw. 237 blumei Teijsm. & Binn. 276 dulcis Teijsm. & Binn. 288 edulis Roxb. 272 lancifolia Hook. f. 288 montana Teijsm. & Binn. 231 montana (Spreng.) Steud. 386 roxburghiana Wight & Arn. 243 sp. Kurz 228 sulingi (Blume) Teijsm. & Binn. 272 Mioptrila Raf. 358 odorata Raf. 361 Monocyclis Wall. 45 robusta Wall. 47 Monosoma Griff. 371 littorata Gritf. 379 Munronia Wight 6, 7, 30 sect. Philastrea (Pierre) Harms 34 sect. Pseudoturraea Harms 34 humilis (Blanco) Harms 21, 30;,32;335 javanica Benn. 31 neilgherrica Wight 31 palawanensis (Merr.) Harms 32 pinnata (Wall.) Theob. 19, 21, 30 pseudoturraea Harms 32 pumila Wight 30 timoriensis Baill. 31 wallichii Wight 30 Napeodendron Ridley 45 altissimum Ridley 49 Naregamia Wight & Arn. 2 Nemedra A. Juss. 194 elaeagnoidea A. Juss. 243 Nimmonia lawii Wight 228 Nymania 7 Olea malabarica Nimbo ... Ray 341 malabarica Fraxineo ... Pluk. 341 Owenia F. Muell. 3 cerasifera F. Muell. 388 Philastrea Pierre 30 Picroderma Gagnep. 41 laotica Gagnep. 43 404 Flora Malesiana ser. I, Vol. 12 (1) (1995) Piper hyalinum Reinw. ex Mig. 189 Piptosaccos Turcz. 61 hypophyllantha Turez. 77 Pistaciovitex L. ex Kuntze 195, 388 pinnata (L.) Kuntze 388 Plagianthus humilis Blanco 32 Plagiotaxis Wall. 354 chickrassa Wall. 355 velutina Wall. 355 Pleiogynium timoriense (DC.) Leenh. 388 Plutea Noronha 314 Portesia Blanco 194 Prasoxylon M. Roem. 61, 66 alliaceum (Blume) M.Roem. 106 Pseudobersama Verdc. 2 Pseudocarapa Hemsl. 2, 61 inopinata Harms 97 nitidula (Benth.) Merr. & L.M. Perry 135 var. latifolia Merr. & L.M. Perry 135 papuana Merr. & L.M. Perry 95 Pseudocedrela 6 Pseudoclausena T.P. Clark 2, 22,55 chrysogyne (Migq.) T. P. Clark 50*,55 f. chrysogyne 57 f. velutina (Ridley) T. P. Clark 57 Pseudosycomorus Camer. 331 Psychotria L. 61, 388 Ptaeroxylon 7 Pterorhachis 7 Quivisia 29 Reinwardtiodendron Koord. 4—6, 22, 322 anaimalaiense (Bedd.) Mabb. 322 celebicum Koord. 21, 324, 328 cinereum (Hiern) Mabb. 323%, 324* dubium (Merr.) X.M. Chen 326 humile (Hassk.) Mabb. 324, 326, 327* kinabaluense (Kosterm.) Mabb. 324, 325 (Reinwardtiodendron) kostermansii (Prijanto) Mabb. 324, 325 merrillii Perkins 328 Rhetinosperma Radlk. 136, 184 longistipitata (F.M. Bailey) Radlk. 185 Rhus blinii Léy. 59 Ricinocarpodendron Boehm. 187 borneense (Miq.) Mabb. 193 cumingianum (C. DC.) Mabb. 189 polystachyum (Wall.) Mabb. 188 sumatranum (Miq.) Mabb. 194 Ruagea Karst. 16, 61 Sandoricum Cav. 5, 18, 21, 344 beccarianum Baill. 63, 344, 348*, 353 borneense Miq. 3, 4, 63, 344, 351 borneense auct. 347 caudatum Mabb. 344, 352 dasyneuron Baill. 345 domesticum Rumph. 347 emarginatum Hiern 353 glaberrimum Hassk. 347 harmandianum Pierre ex Laness. 347 harmandii Pierre 347 harmsianum Perkins 347 indicum Cav. 346 var. cochinchinense Pierre 346 var. velutinum Hiern 346 koetjape (Burm. f.) Merr. 8, 19, 345, 348* ledermannii Harms 347 maingayi Hiern 347 var. quadripetalumC.DC. 347 nervosum Blume 347 nervosum (Vahl) M. Roem. 346 radiatum King 347 serratum G. Don 347 ternatum Blanco 347 venosum (Spreng.)M. Roem. 347 vidalii Merr. 347 Santiria apiculata Benn. 387 egriffithii (Hook. f.) Engl. 387 laevigata Blume 387 tomentosa Blume 387 Sapindus multijugus Wall. 355 Schizochiton Spreng. 136 sect. Euschizochiton Kuntze 147 sect. Dasycoleum (Turcz.) Kuntze 17] diversifolium (Miq.) Miq. 174 amabile Mig. 163 ceramicum Migq. 179 divergens (Blume) Spreng. 167 grandiflorum Kurz 117 Junghuhnii Mig. 179 paniculatum (Hiern) Kurz 164, 166 patens (Blume) Spreng. 167 paucijugum Mig. 181, 183 spectabile Mig. 179 tetrapetalum Turcz. 167 Schizochiton Wall. 167 Schizogiton 167 Scutinanthe boerlagii Hochr. 47 engleri Elmer 43 Scyphostigma philippense M. Roem. 90, 93 Scytalia glabra Buch.-Ham. ex Wall. 47 Selbya M. Roem. 195 montana (Spreng.) M. Roem. 386 Soymida 14 febrifuga (Roxb.) A. Juss. 10; 19 Sphaerosacme Wall. ex Royle 187 decandra (Wall.) T.D. Penn. 187, 387 polystachia Wall. 188 spicata Wall. 189 Surenus Rumph. 358 australis Kuntze 367 febrifuga (Blume) Kuntze 364 glabra (C. DC.) Kuntze 361 inodora (Hassk.) Kuntze 370 microcarpa (C.DC.) Kuntze 367 rubra Rumph. 364 Index to scientific plant names (Surenus) serrata (Royle) Kuntze 361 serrulata (Miq.) Kuntze 361 sinensis (A. Juss.) Kuntze 360 teysmannii (Hassk.) Kuntze 367 toona (Rottl. & Willd.) Kuntze 367 velutina (DC.) Kuntze 367 Surwala M. Roem. 45, 46 robusta (Roxb.) M.Roem. 47 Swietenia 14, 18 chickrassa Roxb. 355 macrophylla King 18, 383 mahagoni (L.) Jacq. 13, 18, 384 var. macrophylla (King) Veitch 384 sotrophola Buch.-Ham. ex Wall. 355 sureni Blume 364 toona Stokes 367 trilocularis Roxb. ex Buch.- Ham. 355 velutina Wall. ex Kurz 355 villosa Wall. ex Kurz 355 Synoum A. Juss. 7, 61, 98 Taeniochlaena polyneura Schellenb. 317 Toona (Endl.) M. Roem. 2—4, 18, 21, 358 australis Harms 367 calantas Merr. & Rolfe 360, 370 ciliata M. Roem. 3,9, 10,12, 18, 360, 366 subsp. nepalensis (C.DC.) Panigrahi 368 subsp. velutina (DC.) Ba- hadur 367 var. australis (C. DC.) Ba- hadur 368 var. brevipetiolulata (Haines) Mishra & Pani- grahi 368 var. candollei Bahadur 364 var. grandifoliola(C.DC.) Bahadur 364 var. hainesti (C. DC.) Pa- nigrahi & Mishra 364, 368 var. hasletii (Haines) Ba- hadur 368 405 (Toona ciliata) var. hasletii (Haines) Pa- nigrahi & Mishra 368 var. henryi (C. DC.) C.Y. Wu 364 var. hexandra (Wall.) Ba- hadur 367 var. kingii (C. DC.) Baha- dur 368 var. latifolia (C. DC.) Chandr. 368 var. mollis (Hand.-Mazz.) Bahadur 368 var. parviflora (Benth.) Bahadur 367 var. pubescens (Franch.) Hand.-Mazz. 368 var. pubinervis (C. DC.) Bahadur 368 var. sublaxiflora (C. DC.) C.Y. Wu 368 var. vestita (C.T. White) Bahadur 368 var. yunnanensis (C. DC.) C.Y. Wu 364 febrifuga (Blume)M.Roem. 364 var. cochinchinensis Pierre 368 var. griffithiana Pierre 368 var. ternatensis Pierre 368 var. glabrior (C. DC.) Pierre 364 glabra (C. DC.) Harms 361 hainesit (C. DC.) Harms 368 hexandra (Wall.) M. Roem. 367 inodora (Hassk.) Hochr. 370 kingii (C. DC.) Harms 368 longifolia {Wall. ex M. Roem. ex] Harms 36] mannii (C.DC.) Harms 368 microcarpa (C.DC.) Harms 367 var. denticulata A. Chev. 361 var. grandifolia A. Chev. 361 var. pilipetala (C. DC.) Bahadur 364, 368 var. sahnii Bahadur 364 mollis (Hand.-Mazz.) A. Chev. 368 paucijuga Merr. 370 (Toona) philippinensis Elmer 370 serrata (Royle) M. Roem. 361 serrulata (Miq.) Harms 361 sinensis (A. Juss.) M. Roem. 3, 360, 362* var. denticulata (A.Chev.) Bahadur 361 var. glabra (C. DC.) Ba- hadur 361 var. grandifolia(A. Chev.) Bahadur 361 var. grandis Pamp. 361 var. hupehana (C. DC.) A. Chev. 361 var. incarvillei A. Chev. 361 var. puberula (C. DC.) Bahadur 361 var. schensiana (C. DC.) Wu 361 sureni (Blume) Merr. 360, 362* 73632365" var. cochinchinensis (Pierre) Bahadur 368 var. inodora (Hassk.) Ba- hadur 370 var. philippinensis Baha- dur 370 var. pubescens (Franch.) Chun 368 var. feysmannii (Hassk.) Bahadur 367 ternatensis (Miq.) Bahadur 367 velutina (DC.) M. Roem. 367 Trichilia L. 2, 5, 6, 7, 8, 14 alliacea Forst. f. 90, 93 alliacea auct. 106 arborescens (Blume) Spreng. 103 connaroides (Wight & Arn.) Bentv. 43 f. glabra Bentv. 43 var. microcarpa (Pierre) Bentv. 43 excelsa (Jack) Spreng. 109, 337 hexandra Blume ex Miq. 167 hirta L. 12 humilis Zipp. ex Mig. 31 longissima Wall. 167 406 Flora Malesiana ser. I, Vol. 12 (1) (1995) (Trichilia) macrocarpa (Blume) Spreng. 116 mollissima (Blume) Spreng. 90, 93 nervosa Vahl 346 pentandra Blanco 180, 182 rimosa (Blanco) Blanco 246 rufinervis Blume 302 similis (Blume) Spreng. 106 sinensis Bentv. 41 tripetala Blanco 189 venosa Spreng. 347 volubilis Blanco 388 Trichilia auct. 41 Turraea L. 2, 6, 7, 21, 24 billardierei (DC.) A. Juss. 25 breviflora Ridley 2, 28*, 29, 30 brownii C. DC. 25 concinna Benn. 25 decandra Blanco 68 humilis (Blanco) Merr. 32 indica C. DC. 25 membranacea Merr. 25 nilotica Kotschy & Peyr. 9 octandra Blanco 90, 93 palawanensis Merr. 32 pinnata Span. 30 pinnata Wall. 30 pubescens Hell. 25, 26* var. billardierei (DC.) Pellegr. 25 pumila Benn. 32 trichostylis Mig. 388 villosa Benn. 25 virens L. var. billardierei DES25 virens auct. 25, 68 zollingeri C. DC. 25, 32 Turraeanthus Baill. 61 Vavaea Benth. 4, 6, 21, 34 sect. Grandiflorae T. D. Penn. 39 sect. Vavaea 35 amicorum Benth. 16, 35 archboldiana Merr. & L.M. Perry 36 ardisiodes Elmer 36 australiana S.T. Blake 36 bantamensis (Koord. & Va- leton) Koord. & Merr. 36 bougainvillensis B.L. Burtt 36 (Vavaea) brassii (Allen) Kosterm. 36 brevipedunculata Kurata 36 chalmersii C. DC. 36 var. angustifolia Merr. & L.M. Perry 36 harveyi Seem. 35 heterophylla Merr. 36 kajewskii Merr. & L.M. Perry 36 lamii Steenis 36 ledermannii Harms 36 luzonensis Elmer 36 megaphylla Wright 35 oligantha B.L. Burtt 36 pachyphylla Merr. 36 papuana F. M. Bailey 35, 38%, 39 pauciflora Ridley 36 pauciflora Volk. 36 pilosa Merr. 36 retusa Merr. 36 scaevoloides Guillaumin 36 sorsogonensis Elmer ex Merr. 36 surigaoensis Elmer 36 tubiflora T.D. Penn. 35, 38, 39, 40* vitiensis Seem. 36 Vitex L. 388 bantamensis Koord. & Vale- ton 36 pinnata L. 386, 388 Walsura Roxb. 2, 4—6, 18, 22, 45 sect. Heynea (Roxb. ex Sims) Harms 41 sect. Neowalsura Harms 55 sect. Ruswala T. P. Clark 53 sect. Surwala (M. Roem.) Hook. f. 46 sect. Walsura 48 aherniana Perkins 48 angulata Craib 49 borneensis Merr. 55 brachybotrys Merr. 55 celebica C. DC. 55 chrysogyne (Miq.) Bakh. f. 35) cochinchinensis (Baillon) Harms 48 dehiscens T.P. Clark 46, 53 (Walsura) elata Pierre 48 glabra Merr. 55 glauca C. Fischer 49 grandifolia Ridley 49 hosei Ridley 55 hypoleuca Kurz 48 intermedia Craib 43 monophylla [Elmer ex] Merr. 3, 19, 21, 46, 52, 63 multijuga King 55 neurodes Hiern 48 pachycaulon Mabb. ex T. P. Clark 46, 51 palawanensis Elmer 55 pallida Craib 43 perrottetii C. DC. 43 pinnata Hassk. 46, 48, 50* pubescens Kurz 43 punctata Suess. 43 var. papillosa Siiss. & Heine 193 quadrilocularis Valeton 55 quinquejuga Kurz 43 robusta Roxb. 46, 47 sarawakensis T. P. Clark 46, 52 spec. A 54 sumatrana (Migq.) Harms ex Koord. 43 tenuifolia Ridley 43 trijuga (Sims) Kurz 41 var. genuina 41 var. pubescens (Kutz) Kurz 41 velutina Ridley 57 villamilii Merr. 49 villosa auct. 388 Xilocarpus 378 Xylocarpus Koen. 3—5, 7, 14, 16; 18, 21) 183msoRszil australasicus Ridley 377 benadirensis Mattei 379 carnulosus Zoll. & Moritzi 379 echinatus 388 forstenii Miq. 375 gangeticus (Prain) C.E. Par- kinson 377 granatum Koen. 10,12, 373, 378, 380*, 381* granatum auct. 375, 377 mekongensis Pierre 377 minor Ridley 379 Index to scientific plant names (Xylocarpus) moluccensis (Lam.) M. Roem. 12, 373, 374*, 376, 377* var. ellipticus (Koord. & Valeton) Harms 376 var. gangeticus (Prain) Craib 376 moluccensis auct. 376, 379 (Xylocarpus) obovatus A. Juss.379 var. macrophyllus Pierre 379 var. microphyllus Pierre 377 obovatus auct. 377 parvifolius Ridley 377 rumphii (Kostel.) Mabb. 12, 373. STAFSSi5* 407 Zanthoxylon 43 Zanthoxylum connaroides Wight & Arn. 43 Zederachia Heist. ex Fabr. 329 Zizipha candida Monspellien- sium Lobel. 331 Zygophyllaceae 24 Zygophyllum 43 PIV g> ee. (continued from inside front cover) WNL UU 3 5185 00206 6387 Podostemaceae 4:65; 6: 963 Scyphostegiaceae Thymelaeaceae Polemoniaceae 4: 195 5: 297; 6: 967 4: 349; 6: 1,976; 7: 830 Polygalaceae 10: 455 Simaroubaceae 6: 193, 968 — Trapaceae 4: 43 Pontederiaceae AS 255 Sonneratiaceae Trigoniaceae 4: 59 Portulacaceae we U2} 4; 280,513; 6: 973 Trimeniaceae 10: 327 Primulaceae 6: 173 Sparganiaceae 4: 233;10: 718 — Triuridaceae 10: 109 Proteaceae 5: 147 Sphenocleaceae 4: 27 ~~ Turneraceae 45 235 Punicaceae 4: 226 Sphenostemonaceae 10: 145 Typhaceae 4: 242; 6: 982 Restionaceae 5: 416 Stackhousiaceae 4: 35 Ulmaceae eS | Rhizophoraceae Staphyleaceae 6: 49 Umbelliferae 4: 113, 595 5: 429; 6: 965; 8: 550 Stemonaceae 11: 399 555995; '6e, 983 Rosaceae 11: 227 Stylidiaceae 4:529; 6: 976 7: 830; 9: 569 Sabiaceae £03) 551 Styracaceae 4: 49; 9: 568 — Valerianaceae 4: 253 Salicaceae 5: 107 Symplocaceae Violaceae 7: 179, 831; 10: 720 Salvadoraceae 4: 224 8: 205; 9:569;10: 718 Xyridaceae Sapindaceae 11: 419 Taccaceae 7: 806 4: 366, 598; 9: 571 Sarcosperma(ta)ceae 4: 32 Taxaceae 10: 347 Zygophyllaceae 4: 64 Saururaceae 4: 47 Index to revised families in Series II (Pteridophyta) Cyatheaceae 1: 65 Lindsaea group 1: 177 — Tectaria group Fd Gleicheniaceae 1: 1 Lomariopsis group 1: 255 Thelypteridaceae 22°33 Isoetaceae 1: 62 Schizaeaceae Ls 37 Editorial Committee: C. Kalkman P.F. Stevens D.W. Kirkup W. J. J.O. de Wilde H.P. Nooteboom Board of the Foundation: Aprilani Soegiarto, Jakarta, chairman P. Baas, Leiden, vice-chairman M.C. Roos, Leiden, secretary / treasurer B.A. Barlow, Canberra O. Gideon, Lae K. Iwatsuki, Tokyo K. Larsen, Aarhus A. Latiff Mohamad, Bangi ISBN 90 -71236 - 26-9 FLORA MALESIANA Compiled and published under the auspices of Foundation Flora Malesiana G.LI. Lucas, Kew D.A. Madulid, Manila Ph. Morat, Paris E.S.P. Ng, Rome D.H. Nicolson, Washington P.H. Raven, St. Louis M.A. Rifai, Bogor S.H. Sohmer, Fort Worth Requests for scientific information to be addressed to the Editors, at the address below. Subscription orders and requests for sample copies to be sent to: Publications Department Rijksherbarium / Hortus Botanicus P.O. Box 9514 2300 RA Leiden The Netherlands Fax (31) (71) 273511