FLORA MALESIANA SERIES I - SPERMATOPHYTA Flowering Plants Volume 11, part 3 Sapindaceae Index to revised families in Series I (Spermatophyta) Aceraceae Aue oO? Actinidiaceae s.s. ya i | Aizoaceae 4: 267 Alismataceae 5: 317; 6: 915 Alliaceae 1375 Alseuosmiaceae 10: 335 Amaranthaceae 4: 69,593; 6: 915; 8: 549 Amaryllidaceae £1353 Anacardiaceae 8: 395 Ancistrocladaceae rs Er Aponogetonaceae CALA ICT 7 ORNS Araliaceae—I 92a 5535 Araucariaceae 10: 419 Aristolochiaceae 10: 53 Balanophoraceae 12718329: 554 Basellaceae 5: 300 Bat(id)aceae 5: 414 Betulaceae 5: 207; 6: 917 Bignoniaceae 8: 114; i 554 Bixaceae s.s. "239 Burmanniaceae 4: 13, 592; 5435 ew yon Burseraceae 6: 917; 7: 820; 9: Butomaceae Ss Byblidaceae Te Callitrichaceae 4: Campanulaceae 6: 107, 928; 9: Cannab(in)aceae 4: Cappar(id)aceae 6: 61; 7: Caprifoliaceae 4: 175; 6: 928; 9: Cardiopteridaceae Ts Celastraceae 6: 227, 389, Centrolepidaceae SH Ceratophyllaceae 4: Chenopodiaceae 4: 99,594: 6: 932; 9: Chloranthaceae 10: Chrysobalanaceae 10: Clethraceae Ts Cochlospermaceae 4: Combretaceae 4: 533; 5: 564; 6: Coniferales 10: Connaraceae 521495;.16:'933; 9: 5: 208BO 555 118 135 251 556 222 822 556 93 930 421 4] DO 123 635 139 61 932 337 357 Convolvulaceae 4: 388, 599 Juglandaceae 6: 143 5: 558: 6: 936 Juncaceae 4: 210; 9: 566 7: 823; 9: 558 Juncaginaceae Ay 57 Coriariaceae 11: 385 Labiatae 8: 301; 9: 566 Cornaceae 8: 85 Legum.-Mimosoideae 11: 1 Corynocarpaceae Leeaceae i Pens, 4: 262; 5: 557 Lemnaceae Tp 219 Crassulaceae 4: 197; 9: 558 Lentibulariaceae 8: 275 Cruciferae 10: 541 Liliaceae s.s. 9: 189 Crypteroniaceae 8: 187 Linaceae 10: 607 Ctenolophonaceae 10: 629 Loganiaceae Cupressaceae 10: 442 6: 293, 953; S:a307 Cyperaceae 7: 435; 9: 107 Lophopyxidaceae ieee Datiscaceae 4: 382 Magnoliaceae 10: 561 Dichapetalaceae Malpighiaceae 5: 125 5: 305; 6: 941 Martyniaceae 4: 216 Dilleniaceae 4: 141; 7: 824 Menispermaceae 102 157 Dioscoreaceae 4: 293 Mimosaceae 1 Te Dipsacaceae 4: 290 Monimiaceae 10: 255 Dipterocarpaceae 9; 237 Moringaceae 4: 45 Droseraceae Myoporaceae 4: 265 RA 557; 9: 562 Myricaceae 4; 276 smu oe 105, ARID BRdaceae G2 157, AN Nyctaginaceae 6: 450 Se kk 10458, Nyssaceae 4: 29 aceai 6: 469, 943 Ochnaceae fe | 8: 549; 9: 562; 10: 716 Olacaceae 10: 1, 717 Erythroxylaceae Onagraceae 8: 98 5: 543; 8: 549 Opiliaceae 10: <231 Fagaceae 7: 265; 9: 563 Oxalidaceae Pulleys re52%9) Flacourtiaceae 5: 1, 565 Papaveraceae spe i 2 6: 943: 7: 827; 9: 563 Passifloraceae 7: 405 Flagellariaceae 4: 245; 9: 564 Pedaliaceae 4: 216; 7: 829 Geraniaceae 6: 445; 9: 565 Pentaphragmataceae 4: 517 Gnetaceae 4: 336; 6: 944 Pentaphylacaceae S27 128 Gonystylaceae 4; 349 Pentastemonaceae its, 393 Goodeniaceae 5: 335, 567 Philydraceae ye 6: 949; 7: 827; 9: 566 Phytolaccaceae 4: 228 Haemodoraceae Pinaceae 10: 447 5:111;10: 717 Pittosporaceae 5: 345; 6: 960 Haloragaceae 7: 239, 828 Plumbaginaceae 4: 107 Hamamelidaceae 5: 363 Podocarpaceae 10: 351 Hippocrateaceae 6: 389 Podostemaceae 4:65; 6: 963 Hydrocharitaceae 5: 381 Polemoniaceae 4: 195 6: 952; 7: 828 Polygalaceae 10: 455 9: 566; 10: 717 Pontederiaceae 4: 255 Hydrophyllaceae 4: 207 Portulacaceae gp fe i! Hypericaceae 8:1; 10: 717 Primulaceae 6: 173 Icacinaceae 7: 1; 9: 566 Proteaceae 5. 4y Iridaceae 8: 77; 10: 717 Ixonanthaceae 10: 621 (continued on inside back cover) FLORA MALESIANA SERIES I— SPERMATOPHYTA Volume 11 — part 3 — 1994 Sapindaceae by F. Adema, P. W. Leenhouts, P.C. van Welzen ERRATUM Flora Malesiana Series I, Volume 11, part 2 In the family Amaryllidaceae, under Curculigo, unfortunately a wrong legend of Fig. 5 has been printed on page 369. For a correct legend, see below. Fig. 5. Curculigo orchioides Gaertn. a. Flowering plant. — C. racemosa Ridley. b. Inflorescence. — C. erecta Laut. c. Inflorescence. — C. latifolia Dryand. var. latifolia. d. Inflorescence. — C. capitulata (Lour.) Kuntze. e. Inflorescence (a: Zippel 272; b: Jacobs 5610; c: Brass 6994; d: Geesink & Hattink 6401; e: de Wilde & de Wilde-Duyfjes 13539). Figure references given after the descriptions of these five Curculigo species on pages 367-370 should be changed as follows: C. capitulata = Fig. 5e; C. erecta = Fig. 5c; C. latifolia var. latifolia = Fig. 5d; C. orchioides = Fig. 5a; C. racemosa = Fig. 5b. CIP-GEGEVENS KONINKLIJKE BIBLIOTHEEK, DEN HAAG Flora Flora Malesiana. Series I, Spermatophyta : Flowering plants. - Leiden : Rijksherbarium / Hortus Botanicus, Leiden University. Vol. 11, pt. 3: Sapindaceae / by F. Adema, P.W. Leenhouts, P.C. van Welzen. - III. Comp. and publ. under the auspices of Foundation Flora Malesiana. - Met index, lit. opg. ISBN 90-71236-21-8 Trefw.: flora ; Zuidoost-Azié. All rights reserved © 1994 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 ser. I, Vol. 11 (3) (1994) 419-768 SAPINDACEAE (F. Adema, P.W. Leenhouts & P.C. van Welzen, Leiden, The Netherlands)! Sapindaceae Juss., Gen. P|. (1789) 246 (‘Sapindi’); Radlk. in Engl., Pflanzenr. 98 (1931— 1934) 1: Backer & Bakh. f., Fl. Java 2 (1965) 130; Abdulla in Fl. W. Pakistan 39 (1973) 1; S.T. Reynolds in Fl. Austral. 25 (1985) 4; Law Yuh-wu, Lo Hsin-shui, Wu Young-fen & Chen Te-chao in FI. Reip. Pop. Sinicae 47 (1) (1985) 1. — Type genus: Sapindus L. Trees, shrubs or lianas, rarely herbaceous climbers; monoecious, rarely dioecious or polygamous. /ndumentum usually of solitary, simple hairs, sometimes also of two-branched hairs, stellate bundles of hairs, or scale hairs (then young parts, buds, and inflorescences viscid). Leaves spirally arranged, rarely opposite or whorled, simple, biternate, digitate or (bi)pinnate; true stipules usually absent, pseudo-stipules sometimes present. Leaflets alternate to opposite, symmetric to distinctly asymmetric, entire or dentate to serrate or crenate. Inflorescences axillary, often together pseudoterminal, terminal or ramiflorous, thyrsoid, with or without branches; bracts and bracteoles present. Flowers usually uni- sexual, rarely bisexual, actinomorphic or zygomorphic. Sepals 4 or 5, rarely more, free to almost totally connate, equal to distinctly unequal, and then the outer | or 2 much smaller than the inner three, herbaceous to petaloid, in bud imbricate, valvate or apert. Petals absent or 2-6, free, usually clawed, often with | or 2 scales or auricles (= inrolled mar- gins), scales crested or not. Disc complete or interrupted, lobed or annular to semi-annu- lar, rarely with appendages or an erect (tubular) rim. Stamens 5—10(—74), usually 8, near- ly always inserted within the disc, often exserted in male flowers; filaments glabrous or hairy; anthers basifixed, opening introrsely or latero-introrsely lengthwise; in female flow- ers present as staminodes with non-opening anthers. Ovary superior, 1—3(—8)-celled, lobed or not; style usually apical, rarely inserted between the lobes, stigma entire with (1), 2 or 3 lines or grooves, or (1-), 2- or 3-lobed; in male flowers rudimentary. Ovules 1 or 2 per locule, ascending, anatropous, campylotropous or amphitropous. Fruits capsular or drupa- ceous, or consisting of 2 or 3 samaras, when capsular usually loculicidal, rarely septicid- al or septifragal. Seeds globose to obovoid, sometimes compressed, often with an arillode or a sarcotesta; endosperm absent; embryo usually thick, straight, sometimes sigmoid or convolute, cotyledons above each other (notorrhizal embryo) to laterally besides each other (lomatorrhizal embryo). Distribution — 140 genera with c. 1350 species, widespread in tropical and subtropi- cal regions of the world, especially well represented in South America. In Malesia 42 genera with ca. 235 species. 1) With contributions by: M. Davids (Paranephelium), J. van Dijk (Dictyoneura), B. Etman (Rhysotoechia), R.W.J.M. van der Ham (Mischocarpus, palynology), R. Hegnauer (phytochemistry), M. Jacobs (Pometia), R.K.W.M. Klaassen (wood anatomy), A.M. Schot (Lepiderema), H. Turner (Aryrera), M. Vente (Harpullia). (419) 420 Flora Malesiana ser. I, Vol. 11 (3) (1994) Habitat — Primary or secondary rain forest, forest edges, shrubland, savannahs, coastal vegetations, often along rivers or roads; in everwet or seasonal conditions; mainly low- land, but also montane up to 3600 m altitude; on all kinds of soil. F. Adema Ecological aspects — Floral biology. Sapindaceae have been recorded to be either monoecious or dioecious (see lists in Van der Ham 1990: 118, 119; and Van Welzen 1989: 41) except Dodonaea which can have bisexual flowers. Most Sapindaceae show dicho- gamy: three stages of floral development appear within the same inflorescence. The first flowers to appear are male (long filaments, dehiscing anthers, functional pollen, undevel- oped pistil). After they have dropped, the second flush of flowers, female, opens (short filaments, indehiscent anthers, non-functional pollen, well-developed pistil). After fruit- set the third phase of flowers opens, organs of both sexes present but flowers functionally male (intermediate to long filaments, dehiscing anthers, functional pollen, more or less well-developed ovary). In duodichogamy the second and third phase usually repeat as fourth and fifth phase. The consecutive phases may overlap. Dichogamous plants with synchronized flowering within the same plant are effective- ly dioecious too, which means that all Sapindaceae have to be cross-pollinated although self-pollination was reported for Xerospermum noronhianum (Appanah 1982; but see also Ha et al. 1988). There is apparently no self-incompatibility since interflower selfing occurs in several economically important plants (see references in Van der Ham 1990: 118). The Sapindaceae are mainly entomophilous; Dodonaea is the only genus within the family for which wind-pollination has been recorded (Keighery 1982; West 1982, 1984). The most important pollinators are bees, mainly of the genera Trigona (stingless bees) and Apis (honey bees). The former seem to be the main pollinators (Appanah 1982; Bawa 1977; Gondim 1984; Hawkeswood 1983; Subba Reddi et al. 1983; Uji 1987; Van Welzen et al. 1988). They may either be attracted by the nectar or by the pollen and many species are reported to have fragrant flowers; the attractant may be different for each species of Trigona (see discussion in Van der Ham 1990: 120). The trees seem to have three mechanisms to ensure cross-pollination by bees: a) The most important one is using the bees’ memory (see references in Van der Ham 1990: 121; and Van Welzen 1989: 43) by attracting them to the trees with the first male phase in the (duo)dichogamous stages, after which the bees will return to the less attractive female flowers and the attractive bisexual flowers (Van der Ham 1990: 121, 122). b) Fluctuations in nectar productivity alternating between trees with male/bisexual flowers and female ones during day-time (see references in Van Welzen 1989: 42). c) Differences in nectar composition between male and female flowers (Appanah 1982). However, female flowers are visited far less often than male or bisexual flowers, as the latter are more colourful due to the usually exserted and coloured stamens. P.C. van Welzen Fruits and dispersal. Young fruits of Sapindaceae are generally reported to be green, hard, and full of tannin. When mature, the colour changes, usually to red, and the fruit wall softens (Ha et al. 1988). Sapindaceae may fruit throughout the year, once or twice per year. They tend to have a good harvest one season, followed by a poor one the next (Appanah 1982; Bawa 1977; Van Welzen & Verheij 1991). Adema, Leenhouts, Van Welzen — Sapindaceae 42] Several fruit types occur in the Malesian Sapindaceae: a) Winged fruits, e.g. Atalaya and Dodonaea. b) Inflated fruits with a thin, papery wall, e.g. Cardiospermum and Koelreuteria. c) Drupaceous fruits, e.g. Allophylus and Lepisanthes. d) Berry-like fruits, e.g. Filicium and Sapindus. e) Dehiscent capsules, usually with (partial) arillodes around the seeds, e.g. in most Cupanieae. f) Indehiscent ‘capsules’, also usually with arillodes around the seeds, e.g. a few Cupanieae and most Nephelieae. The seeds or fruits are mainly dispersed by birds and mammals. The smaller fruits, like those of Allophylus (type c; Docters van Leeuwen 1932) and the seeds of the Cupanieae (type e), are mainly eaten by birds, which are attracted by the contrasts in colour between fruit (yellow to mainly red), arillode (mainly yellow to red), and seed (shiny dark brown to black). Several genera (Guioa, Mischocarpus, and to a lesser degree Sarcopteryx) add movement as an attractant, as an appendage of the arillode (pseudo-funicle) is attached to the basal corner of the fruit on which the seed remains dangling after dehiscence. The seeds of the larger fruits in this group are mainly eaten by parrots, parakeets, and presum- ably pigeons (Ridley 1930: 487; Van Welzen 1989: 44). The non-dehiscent and mainly large(r) fruits (type f) are eaten by mammals. In this category the commercially interesting species are found. These fruits, too, have the red colour as attractant. They are reported to be eaten and dispersed by man, monkey, fox- bat, sloth bear (Melursus ursinus), and pigs (Ridley 1930: 339-359). However, parakeets and parrots may eat these fruits too (Ridley 1930: 487). Fruits of the types a and b are reported by Ridley (1930: 195) to be dispersed by wind or (sea)water, just like the woody seeds of Pometia and the fruits of Sapindus (Ridley 1930: 268-270). P.C. van Welzen Germination and seedlings. Usually, the seeds of Sapindaceae germinate readily, within a week, and passage through animal intestines is not a prerequisite. The seeds are short- lived and do not show dormancy. During germination the radicle and petioles swell and become terete. The testa usually opens at the place where the embryonic radicle is present. The radicle is quite often enclosed in a testal pouch. The testa subsequently opens along the pleurograms. Then the petioles, hypocotyl, and radicle elongate. The cotyledons, turning green, remain loosely enclosed in the testa. This type of germination is classified as Hors- fieldia type and subtype (De Vogel 1980) and has been reported for several genera (Ade- ma 1991; De Vogel 1980; Van Welzen 1989). The seedlings may differ considerably in macromorphology from the mature plants. The first leaves are usually opposite (not alternate as in the adult). The rachis is usually slightly winged (wing usually absent). The margin of the leaflets is (crenate to) serrate (usually entire). There are no papillae and domatia on the lower surface of the leaflets. In size and number of leaflets the leaves of seedlings may differ dramatically from those in mature plants. Plants showing immature characters may flower and fruit (Van Wel- zen 1989: 45, 46), thus hampering species delimitation. P.C. van Welzen 422 Flora Malesiana ser. I, Vol. 11 (3) (1994) Ant associations. Several genera, Alectryon (Leenhouts 1988), Guioa (Van Welzen 1989: 218, 219), Harpullia (Leenhouts & Vente 1982), and Sarcopteryx (Van Welzen 1991) are reported to have associations with ants. The branchlets, which are hollow, are inhabited by ants. The branchlets are usually swollen below the nodes and in the swell- ings the nest openings can be found. Any profit to the plants in this possible symbiosis is sull unknown. P.C. van Welzen & F. Adema References: Adema, F., Leiden Bot. Series 15 (1991) 31. — Appanah, S., Biol. J. Linn. Soc. 18 (1982) 1-34. — Bawa, K.S., Evolution 31 (1977) 52-63. — Docters van Leeuwen, W.M., Trop. Natuur 21 (1932) 142. — Gondim, C.J.E., Acta Amazonica 14 (1984) 9-38. — Ha, C.O., V.E. Sands, E. Soepad- mo & K. Jong, Bot. J. Linn. Soc. 97 (1988) 295-316. — Ham, R.W.J.M. van der, Leiden Bot. Series 13 (1990) 118-122. — Hawkeswood, T.J., Victorian Nat. 100 (1983) 12-20. — Keighery, G.J., in W.R. Barker & P.J.M. Greenslade, Evolution flora fauna arid Australia (1982) 167-172. — Leenhouts, P.W., Blumea 33 (1988) 313-327. — Leenhouts, P.W. & M. Vente, Blumea 28 (1982) 1-51. — Ridley, H.N., Dispersal (1930) 195, 268-270, 339-359, 487. — Subba Reddi, C., E.U.B. Reddi, N.S. Reddi & P.S. Reddi, Proc. Indian Nat. Sci. Acad. 49b (1983) 57-72. — Uji, T., Suppl. Berita Biol. 3 (1987) 31-34 . — Vogel, E.F. de, Seedlings Dicotyl. (1980) 52-92. — Welzen, P.C. van, Leiden Bot. Series 12 (1989) 40- 46, 218, 219; Blumea 36 (1991) 87—103. — Welzen, P.C. van, A. Lamb & W.W.W. Wong, Nature Malay- siana 13 (1988) 10-25. — Welzen, P.C. van & E.W.M. Verheij, in E.W.M. Verheij & R.E. Coronel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991) 235-240. — West, J.G., in W.R. Barker & P.J.M. Greenslade, Evolution flora fauna arid Australia (1982) 329-333: Brunonia 7 (1984) 1-194. Morphology — /ndumentum. Usually consisting of solitary simple hairs, sometimes of two-branched hairs (T-hairs, Litchi), or stellate tufts of hairs (Dimocarpus, Glenniea, Harpullia); scale hairs (glandular scales, ‘Schilffern’) are found in several genera, then young axes and leaflets with a sticky (viscid) exudate. Various types of glandular hairs may be present (Adema 1991; Adema & Van der Ham 1993; Van Welzen 1989). (See also the section on leaf anatomy.) Leaves. Almost always spirally arranged, rarely opposite or whorled, usually pari- or imparipinnate, sometimes digitate, unifoliolate, or simple; if paripinnate often the rachis ending in an acumen. Stipules present in only a few genera (e.g. Cardiospermum); more often pseudo-stipules present: lowermost pair(s) of leaflets at the very base of the peti- ole, much smaller than and usually of a different shape as the other leaflets (e.g. Alectry- on repandodentata, Lepisanthes subg. Otophora, Pometia). See Weberling & Leenhouts 1966; Weberling 1976. Inflorescences. Axillary and often several together, pseudoterminal (terminal vegeta- tive bud present), or terminal, or rami- or cauliflorous; often more or less thyrsoid or paniculate; cymes (cymules) one- to many-flowered, either dichasia, cincinni, or bos- tryxes, various reductions may obscure their true form, however. Flowers. Usually 4- or 5-merous, actinomorphic to strongly zygomorphic. Sepals entirely free, outer one or two much smaller than the inner three, and in bud imbricate (e.g. Cupaniopsis, Guioa, Lepisanthes) to highly connate, all equal and in bud apert (Cu- bilia, Litchi). Petals free, with or without a claw, often the inside with | or 2 scales or auricles (inrolled margins), especially in the latter case the petals obliquely funnel-shaped (‘peltaten Kronblatter’, Leinfellner 1958); the scales may be provided with crests (e.g. Guioa, Sarcopteryx, Toechima) or not. Disc complete or interrupted, annular or saucer- Adema, Leenhouts, Van Welzen — Sapindaceae 423 like to semi-annular. Ovary 1-, 2-, or 3-celled, style shorter to longer than the ovary, stigma short or long, erect or recurved and lobed, or closed and with lines of stigmatic papillae on the outside. Fruits. 1—3-celled, lobed or not, outside smooth, or wrinkled, or with warts, knobs, or spines. Seeds. Usually covered by an arillode, either called a sarcotesta (when adnate to the exotesta) or an arillode (when free from the exotesta). See Van der Pijl (1955, 1957, 1966). In Guioa, Mischocarpus, and to a lesser extent Sarcopteryx the arillode is usually provided with an appendage (pseudo-funicle); endotesta often with a radicular pocket. Cotyledons either superposed above each other (notorrhizal embryo) or collateral beside each other (lomatorrhizal embryo), but there are many intermediates. PF. Adema References: Adema, F., Leiden Bot. Series 15 (1991) 1-190. — Adema, F. & R.W.J.M. van der Ham, Blumea 37 (1993) 173-215. — Leinfellner, W., Osterr. Bot. Zeitschr. 105 (1958) 443-514. — Pijl, L. van der, Proc. Kon. Ned. Akad. Wetensch., C, 58 (1955) 154-161. 162-312; Acta Bot. Neerl. 6 (1957) 618-641; Proc. Kon. Ned. Akad. Wetensch., C, 69 (1966) 615-640. — Weberling, F., Abhandl. Math.- Naturwiss. KI. Akad. Wissensch. und Lit. 2 (1976) 1-27. — Weberling, F. & P.W. Leenhouts, Abhandl. Math.-Naturwiss. Kl. Akad. Wissensch. und Lit. 10 (1966) 1-90. — Welzen, P.C. van, Leiden Bot. Series 12 (1989) 1-315. Palynology — A worldwide survey of Sapindaceae pollen has been made by Muller & Leenhouts (1976). Monographic pollen studies of Malesian taxa include those by Ade- ma (1991: Cupaniopsis), Adema & Van der Ham (1993: Cnesmocarpon, Jagera, Trigo- nachras), Van den Berg (1978: Cubilia, Litchi, Pometia), Van der Ham (1977: Mischocar- pus; 1990: Alectryon, Cubilia, Dimocarpus, Litchi, Nephelium, Pometia, Xerospermum), Van der Ham & Van Heuven (1989: Guioa) and Muller (1970: Lepisanthes; 1971: Dimo- carpus; 1973: Glenniea; 1985: Harpullia). Sapindaceae pollen grains are usually isopolar or subisopolar monads but occasional- ly the tetrad configuration persists in mature pollen. Permanent tetrads occur only in Magonia (S America). Several genera of Paullinieae (in Malesia only Cardiospermum) have distinctly heteropolar grains. Grain size is usually between 20 and 30 um. The pol- len of Cubilia is exceptionally small (av. 13 wm), while that of Cardiospermum is the largest in the family (up to 74 um). Grain shape is oblate to prolate. Colporate pollen is usually subprolate to prolate (P/E 0.75—2.00), whereas pollen with small apertures (po- rate, brevicolporate) or with connected apertures (syncolporate, parasyncolporate) is rel- atively broader, with a more oblate shape (P/E 0.50-1.00). The equatorial outline is bluntly triangular to almost circular; the meridional outline is more or less elliptic to almost circular. Generally Sapindaceae pollen is 3-aperturate, but often small percentages of grains with 2 and 4 apertures occur. The apertures are diverse, but lalongate, elliptic to subcircu- lar endoapertures are nearly always present. The pollen types recognized by Muller & Leenhouts (1976) are based largely on ectoaperture features. Colporate pollen is the com- monest type, being known from all 13 tribes; it is a relatively basic type like that of many other angiosperm families. Several probably derived types are restricted to one or a few tribes. The (para)syncolporate type is known only in the subfamily Sapindoideae (which is considered more derived than the Dodonaeoideae), being present in most Cupanieae 424 Flora Malesiana ser. I, Vol. 11 (3) (1994) (Van der Ham 1990), the Melicocceae (Castanospora, Tristira, Tristiropsis), the Nephe- lieae (Alectryon) and the Schleichereae (Schleichera). Parasyncolporate and syncolpo- rate (with and without apocolpial field, respectively) are not clear-cut character states. Moreover, several genera (e.g. Alectryon, Arytera, Cupaniopsis, Elattostachys) possess both colporate and (para)syncolporate pollen, and often intermediates as well. Reverse evolution, from (para)syncolporate to colporate, might have occurred in these groups. A trend towards (very) small ectoapertures includes the brevicolporate and porate types found in the Melicocceae, Lepisantheae (e.g. Lepisanthes), Schleichereae, Nephelieae (Pometia), Thouinieae (e.g. Allophylus) and Paullinieae. Pollen of Cardiospermum (Paul- linieae) is heteropolar and has short demicolpi on the proximal side of the grain. Pollen of several related American genera has a demisyncolporate aperture system, and that of a few Serjania and Urvillea species a syncolporate aperture on one side and short demi- colpi on the other. Therefore, (brevicol)porate Paullinieae pollen may have evolved from a syncolporate ancestral form while other (brevicol)porate groups (such as Lepisanthes spp.) might be derived from colporate forms. Distichostemon (N Australia) and a few Harpullia species have indistinct ectoapertures (cryptoaperturate). The shape of an aperture determines much of its harmomegathic motion, caused by dehydration and rehydration of the protoplasm. Colporate apertures fold their equatorial parts inwards (P/E increases), whereas (parasyncolporate apertures fold their polar parts inwards (P/E decreases). Grains with small apertures may fold nonapertural parts of the exine, or show a peristatic mechanism (Van der Ham 1990). The exine is usually clearly stratified (although this is not always visible with light microscopy), showing a tectum, a columellate infratectal layer and a nexine. In the Do- donaeeae |Diplopeltis, Distichostemon (both Australia) and Dodonaea] the infratectal layer is granular/columellate, which might relate to wind-pollination. Mostly the nexine consists of a distinctly delimited foot layer and endexine. The endexine is thin in non- apertural parts, thickens towards the apertures, and is maximal along/under the aper- tures. The ornamentation of the exine shows much variation. Striate, rugulate, psilate, and intermediate types are most common; (micro)echinate, scabrate and reticulate types are less frequent. In the subfamily Dodonaeoideae, rugulate pollen is rare and psilate pollen entirely absent. In the Sapindoideae, (micro)echinate and scabrate ornamentation 1s rare (only in the Malesian genera Cubilia, Dimocarpus, Jagera and Trigonachras), but it is common in the Dodonaeoideae in Malesia, e.g. Dodonaea (scabrate), Filicium and Ganophyllum (both microechinate). The tectum is usually perforated but less densely and less distinctly so towards the apertures. Perforate types are linked with more or less reticulate types. For example a rugulate/reticulate exine is found in Cupaniopsis spp., a (micro)echinate/reticulate exine in Harpullia spp., and a scabrate/reticulate exine in Jagera. Simple reticulate ornamenta- tion is rare (e.g. Cardiospermum, Pometia). Striate ornamentation is often associated with a colporate aperture system, and rugulate with a (para)syncolporate aperture sys- tem, which probably reflects a functional, harmomegathic relation. Ornamentation often varies within genera while Allophyllus cobbe (Muller 1979) and Dimocarpus longan (Van der Ham 1990, 1993) show remarkable infraspecific variation. Adema, Leenhouts, Van Welzen — Sapindaceae 425 Muller & Leenhouts (1976) stated that the Aceraceae and Hippocastanaceae are not distinct from the Sapindaceae. Pollen of both families was described as colporate with- out any derived character. However, pollen of the Hippocastanaceae has characteris- tic verrucae on its colpus membranes. Pollen of Handeliodendron (Harpullieae, China) shows the same feature, which supports inclusion of the Hippocastanaceae in the Har- pullieae. Fossil pollen. Several Sapindaceae pollen types are sufficiently characteristic to be recognized in dispersed (sub)fossil state. The (para)syncolporate type (Cupanietdites, Cupaniopsis type) is widespread, and known from the Turonian (Africa) onwards (Van der Ham 1990). The oldest Malesian data are Miocene records (New Guinea). Other fairly recognizable types are the pollen of Allophylus (not yet found in Malesia), Car- diospermum (not indigenous in Malesia; to be found only in recent and subrecent depos- its) and Pometia (Miocene Borneo). R.W.J.M. van der Ham References: Adema, F., Leiden Bot. Series 15 (1991) 25-30. — Adema, F. & R.W.J.M. van der Ham, Blumea 37 (1993) 173-215. — Berg, R.G. van den, Blumea 24 (1978) 369-394. — Ham, R.W.J.M. van der, Blumea 23 (1977) 301-335; Leiden Bot. Series 13 (1990); Palynosciences 2 (1993) 239-254. — Ham, R.W.J.M. van der & B.J. van Heuven, Blumea 34 (1989) 21-60. — Muller, J., Blumea 18 (1970) 507-561: Blumea 19 (1971) 133-145; Blumea 21 (1973) 105-117; Ann. Missouri Bot. Gard. 66 (1979) 593-632: Blumea 31 (1985) 161-218. — Muller, J. & P.W. Leenhouts, Linn. Soc. Symp. Ser. 1 (1976) 407-445. Wood anatomy — The description presented below is based on the preliminary re- sults of a detailed worldwide wood anatomical survey of the Sapindaceae (Klaassen in prep.). For general accounts see Metcalfe & Chalk (1950), and Solereder (1899). Infor- mation on individual genera is given by Anonymous (1986: Pometia); Balan Menon (1971: Guioa, Harpullia, Lepisanthes, Nephelium, Pometia, Tristira, Xerospermum); Burgess (1966: Pometia); Chowdhury & Ghosh (1963: Allophylus, Arytera, Dodonaea, Erioglos- sum [= Lepisanthes], Filicium, Harpullia, Lepisanthes, Mischocarpus, Nephelium, Paranephelium, Pometia, Sapindus, Schleichera, Xerospermum);, Desch (1954: Gutioa, Harpullia, Lepisanthes, Nephelium, Pometia, Tristira, Xerospermum); De Freitas (1958: Ganophyllum, Schleichera); Fundter & Wisse (1977: Pometia); Furuno (1977: Gano- phyllum, Pometia); Furuno (1979: Tristiropsis); Hayashi (1991: Sapindus); Tlic (1991: Alectryon, Allophylus, Arytera, Cupaniopsis, Dictyoneura, Dimocarpus, Diploglottis, Dodonaea, Elattostachys, Ganophyllum, Guioa, Harpullia, Jagera, Lepidopetalum, Le- pisanthes, Mischocarpus, Nephelium, Pometia, Sapindus, Schleichera, Tristiropsis, Xero- spermum); ITTO report (1991: Allophylus, Dodonaea, Ganophyllum, Harpullia, Koel- reuteria, Nephelium, Pometia, Sapindus, Schleichera); Kanehira (1924: Litchi, Pome- tia); Keating & Bolza (1982: Pometia); Koning-Vrolijk et al. (1962: Pometia); Martawi- jaya et al. (1986: Pometia); Meniado et al. (1981: Dimocarpus, Ganophyllum, Pometia); Moll & Janssonius (1911: Dodonaea, Ganophyllum, Guioa, Harpullia, Lepisanthes, Mischocarpus, Nephelium, Pometia, Sapindus, Schleichera, Xerospermum), Ogata (1983: Ganophyllum, Pometia, Tristiropsis); Pearson & Brown (1932: Filicium, Schleichera); Purkayastha et al. (1976: Pometia); Sudo (1970: Dimocarpus, Ganophyllum, Litchi, Nephe- lium, Pometia); Tang (1973: Amesiodendron, Litchi, Mischocarpus, Nephelium, Pome- tia). 426 Flora Malesiana ser. I, Vol. 11 (3) (1994) The wood anatomy of the Sapindaceae of Malesia shows little variation, especially within the Cupanieae. This tribe contains two thirds of all the Sapindaceae genera and shows very little variation in vessel and pit dimensions, parenchyma distribution, ray composition and fibre type. On the other hand a wide variation is seen in the Nephelieae, the second largest tribe; all seven Malesian genera can be distinguished wood anatomi- cally, using vessel and pit diameter and parenchyma distribution. All genera in the Dodo- naeeae, Doratoxyleae, Harpullieae, Melicocceae and the Sapindeae have abundant pa- renchyma; some genera have apotracheal and vasicentric to aliform parenchyma while others have mainly aliform to aliform-confluent parenchyma. General wood anatomical description. Growth rings distinct to indistinct or absent, when present marked by differences in fibre wall thickness, or by marginal parenchyma. Ring porosity occurs probably only in temperate regions in Sapindus and Koelreuteria. Vessels diffuse, round to oval with a radial diameter 60—120 um, or wider than 120 um in Cubilia, Nephelium and Pometia, usually 10—-20/mm/, less than 10/mm? in Cubilia, Nephelium, Pometia and Tristira, and more than 20/mm2 in Dodonaea and Ganophyllum. Less than 70% of the vessels are solitary, the remaining ones in radial multiples of 2-4. Vessel member length 250-600 um. Perforation plates simple in horizontal or oblique end walls. Intervessel pits alternate, round to polygonal, usually 4—6 wm in horizontal diameter, pits larger in Allophylus, Cubilia, Koelreuteria, Tristiropsis and Zollingeria, and pits minute (smaller than 4 um) in Dimocarpus and Litchi; pit apertures slit-like to round. Fine helical wall sculpturing present in half of the genera; distinct helical wall sculpturing only when ring-porous (Sapindus and Koelreuteria). Vessel-ray and vessel- parenchyma pits similar to intervessel pits but half-bordered, in some genera unilaterally compound. Vessel dimorphism not related to ring porosity in Cardiospermum. Libriform fibres 600-1100 «um long, with simple or minutely bordered pits mainly con- fined to the radial walls, thin- to medium thick-walled; very thick-walled in Atalaya, Dodo- naea, Lepisanthes, Nephelium and Schleichera. Fibres septate in all genera except Dodo- naea, Harpullia and Tristira. Fibre dimorphism — irregular wavy tangential bands of alter- nating thin- and more thick-walled fibres — in Allophylus, Diploglottis and Paranephelium. Parenchyma. Half the genera have scanty paratracheal parenchyma. The others may have scarce, scarce to abundant or fully abundant apotracheal parenchyma (Atalaya, Dodo- naea, Filicium, Ganophyllum, Harpullia, Nephelium, Tristiropsis); marginal parenchy- ma (Filicium, Harpullia, Pometia, Tristira, Xerospermum); vasicentric to aliform paren- chyma (Dodonaea, Filicium, Ganophyllum, Harpullia, Pometia, Tristira, Tristiropsis, Xero- spermum); or aliform-confluent to banded parenchyma (Atalaya, Dodonaea, Lepisan- thes, Nephelium, Sapindus, Zollingeria). Rays typically 1 or 2 cells wide, predominantly biseriate or wider in some species of Atalaya, Dodonaea, Filicium, Harpullia, Sapindus, Tristiropsis and Zollingeria. Rays of two distinct sizes occur in Cardiospermum: short uniseriate rays and 2—4-seriate rays exceeding 1 mm in length. Rays range from homocellular with exclusively procumbent cells to heterocellular with procumbent body cells and indistinct to distinct marginal rows of mainly square but sometimes upright cells. Crystals typically in chambered crystalliferous strands, also in ray cells in Atalaya, Cardiospermum, Elattostachys, Koelreuteria, Nephelium, Paranephelium and Tristira. Adema, Leenhouts, Van Welzen — Sapindaceae 427 The wood anatomy of Gongrospermum and Sarcotoechia is not known. Useful timber species: Dimocarpus longan: The sapwood is lighter in colour than, but not sharply defined from. the reddish brown heartwood. The wood is strong, tough, very hard, difficult to split, highly durable, easily but slowly drying with little or no degradation. Dimocarpus is used for shuttle pipes, bearings, textile weaving stands and rifle butts (Keating & Bol- za 1982). Ganophyllum falcatum: The sapwood is similar in colour to the yellow-brown hard- wood. The wood is medium strong, heavy and durable, shrinks much in drying and is moderately difficult to work. Ganophyllum is used in house building, ship manufac- turing, sporting goods and agricultural implements (ITTO 1991, as Ganophyllum obliquum). Harpuilia arborea: The sapwood is similar in colour as the yellow-brown heartwood. The wood is medium weak, medium heavy and shrinks little on drying. Harpullia is used for house building, furniture, plywood, packing boxes (ITTO 1991; Chowdhury & Ghosh 1963). Pometia pinnata: The sapwood is lighter in colour than, but not sharply defined from the pink, red or red-brown hardwood. The wood is medium strong and medium hard, splits easily, is not durable, difficult to dry, and shrinks much on drying; in dry condition it is still susceptible to dimensional changes. Pometia is used for house and bridge con- struction, furniture, flooring, moulding, shipbuilding, tool handles, sporting goods, agri- cultural implements, packing boxes, bent wood, pulpwood (Anonymous 1986; Burgess 1966; Chowdhury & Ghosh 1963; Desch 1954; ITTO 1991; Keating & Bolza 1982; Mar- tawijaya et al. 1986; Purkayastha et al. 1976). Schleichera oleosa: The sapwood is distinct from the reddish heartwood. The wood ts very heavy, very strong, not durable and difficult to work. Schleichera is used for house building, ship manufacturing, tool handles, musical instruments, agricultural implements, oil presses, fuelwood (ITTO 1991; Chowdhury & Ghosh 1963). Tristiropsis acutangula and T. ferruginea: The pale pink or cream to pinkish brown sapwood is distinct from the grey-brown to red-brown heartwood. The wood is not strong, easy to dry and not durable. Tristiropsis is used for veneer and boards (Keating & Bolza 1982). Lesser used timbers: Arytera litoralis: The sapwood is similar in colour to the pinkish heartwood. The wood is hard, tough and heavy. Aryrera is used for agricultural implement wood and tool handles (Chowdhury & Ghosh 1963). Dodonaea viscosa: The sapwood is similar in colour to the yellowish brown to deep reddish brown heartwood, or sometimes lighter. The wood is hard, tough and strong, difficult to season because of splitting and cracking. Dodonaea is used for tool handles, fuelwood, walking sticks, turnery and engraving (Chowdhury & Ghosh 1963). Filicium decipiens: The reddish brown heartwood is distinct to indistinct from the greyish white sapwood. The wood is very hard, very heavy, strong, tough, durable and not easy to season. Filicium is used for furniture and as construction wood (Chowdhury & Ghosh 1963). 428 Flora Malesiana ser. I, Vol. 11 (3) (1994) Litchi chinensis: The reddish brown heartwood is distinct from the light pinkish to greyish brown sapwood. The wood is hard to very hard, heavy to very heavy, strong and tough and very durable. Litchi is used as general construction wood, for salt water piles, posts, beams, joists, rafters, flooring, keels and keelsons of ships (Chowdhury & Ghosh 1963). R.K.W.M. Klaassen References: Anonymous, 100 Malaysian Timbers (1986) 52. — Balan Menon, P.K., Mal. For. Rec. 27 (1971) 94-95. — Burgess, P.F., Sabah For. Rec. n. 6 (1966) 443-446. — Chowdhury, K.A. & S.S. Ghosh, Indian Woods 2 (1963) 207-239. — Desch, H.E., Mal. For. Rec. 15 (1954) 524-537. — Freitas, M. de, Estudo Madeiras Timor 5 (1958) 13—16. — Fundter, J.M. & J.H. Wisse, Meded. Landb. Hoge- school Wageningen 77 (1977) 157-161. — Furuno, T., Research Rep. Foreign Wood 6 (1977); ibid. 8 (1979). — Hayashi, S., Micrographic atlas of Japanese woods (1991). — Ilic, J., CSIRO Atlas Hard- woods (1991) 60-61, 421-430. — ITTO report, Identification, properties and uses of some SE Asian woods (1991) 165-168. — Kanehira, R., Ident. Philipp. woods by anatomical characters (1924) 73-81. — Keating, W.G. & E. Bolza, SE Asia, N Australia & Pacific. Charact., prop., uses of timbers. | (1982) 280. — Klaassen, R.K.W.M. (in prep.). — Koning-Vrolijk, G.M.C., S.M. Jutte & T. Hof, Nova Guinea, n.s. 10 (1962) 170-174. — Martawijaya, A., I. Kartasujana, K. Kadir & S.A Prawira, Indonesian Wood Atlas 1 (1986) 64-68. — Meniado, J.A., W.M. America, B.C. de Vela, F.N. Tamolang & F.R. Lopez, Wood ident. Handb. Philipp. Timbers 2 (1981) 94—96. — Metcalfe, C.R. & L. Chalk, Anat. Dicot. (1950) 419-431. — Moll, J.W. & H.H. Janssonius, Mikrogr. Holzes Java Baumarten 2 (1911) 316-406. — Ogata, P.K., Wood Industry 30-4 (38-10) (1983) 33. — Pearson, R.S. & H.P. Brown, Commercial Tim- bers India | (1932) 288-286. — Purkayastha, S.K., K.B.S. Juneja & S.M. Husain Kazmi, Indian For. Rec. 2 (1) (1976) 34. — Solereder, H., Syst. Anat. Dicot. (1899) 257-268. — Sudo, S., Trop. Asian Timbers (1970) 288-292. — Tang, Y., Trop. Subtrop. Woods Yunnan 7 (1973) 199-204. Leaf anatomy — A general survey of the leaf anatomy of Sapindaceae can be found in Solereder (1899, 1908), and Metcalfe & Chalk (1950). Remarks on genera and species can be found throughout the works of Radlkofer (see e.g. Radlkofer 1879, 1883, 1931— 1934). The tribe Cupanieae is best known up to now because of studies on Cupaniopsis (Adema 1991), Cnesmocarpon, Jagera and Trigonachras (Adema & Van der Ham 1993), and Elattostachys (unpublished data), and on Guioa (Van Welzen 1989). In surface view. Non-glandular hairs none to abundant, abaxially usually more dense than adaxially, usually unicellular, with thick, or more rarely thin, sclerified walls, rather variable in length, hairs two-armed in Litchi, tufted (‘stellate’) in Dimocarpus, Glenniea and Harpullia. Glandular hairs none to abundant, at least on abaxial surface, consisting of either a single large ellipsoidal cell (Cupaniopsis, Guioa), or of 1-3 uniseriate stalk cells and a large glandular top cell, or of 4-16 uniseriate stalk cells and a small glandular top cell (Cupaniopsis species, Diploglottis, Euphorianthus, Guioa hirsuta), or of | or 2 stalk cells and a large glandular head consisting of several cells (species of Jagera, Lep- isanthes, Sapindus). Scale hairs consisting of 1 stalk cell and a flat circular head of 6—10 thin-walled cells present in Dictyoneura, Dodonaea, Filicium, Ganophyllum, Schleichera. Papillae often present, sometimes connected by cuticular ridges (Cnesmocarpon, Gui- oa). Cuticle smooth to striate, if anticlinal walls undulate than often thinned in the loops of the undulations. Unspecialized epidermal cells polygonal, with straight to undulate anticlinal walls, often radiating around hairs and stomata; above midrib and veins square to rectangular, in rows parallel to the veins. Stomata predominantly cyclocytic, rarely anomo- or anisocytic, or in Harpullia paracytic; abaxially usually abundant, adaxially absent or rare, but rather common in Cupaniopsis. Adema, Leenhouts, Van Welzen — Sapindaceae 429 In transverse section. Lamina dorsiventral. Unspecialized epidermal cells square to flat, rectangular or erect, especially above midrib and along margin; delicate vertical secondary division walls present in Alectryon, Arytera, Cupaniopsis (but not observed by Adema 1991), Xerospermum. Hypodermis usually present only above midrib and veins, a continuous layer in Alectryon, Arytera, Cupaniopsis, Elattostachys and Harpullia. Mes- ophyll: palisade tissue composed of | or 2, rarely up to 4 layers of long erect cells, trans- versely septate in Alectryon, Nephelium, Pometia; spongy tissue compact to rather loose, sclerenchymatous fibres or sclerosed cells present in Cupaniopsis (but not observed by Adema 1991), Harpullia and Xerospermum. Midrib raised abaxially, flat or raised adax- ially, vascular bundles collateral, with a flat to arch-shaped adaxial strand and an abaxial arch, surrounded by a sclerenchyma sheath, in Cupaniopsis, Diploglottis and Euphorian- thus with extra vascular strands in the pith, pith consisting of large, round to transverse- ellipsoidal cells, often filled with starch grains. Minor veins usually embedded in the mesophyll; /arger veins may be vertically transcurrent. Margin with or without marginal vein, usually with normal mesophyll. Crystals none to abundant, usually rhomboidal, in ground tissue of midrib and veins, sometimes also in pith and/or phloem, rarely in pali- sade tissue or abaxial subepidermis. Druses are recorded for Guioa, Lepisanthes, Rhyso- toechia and Sapindus. Secretory idioblasts absent to usually abundant, small to very large, round to flat rectangular, or erect in palisade tissue, occurring in palisade tissue, spongy tissue, abaxial subepidermis, ground tissue of midrib; contents unknown, probably in most cases saponin (see also paragraph on Phytochemistry). F. Adema References: Adema, F., Leiden Bot. Series 15 (1991) 13-25. — Adema, F. & R.W.J.M. van der Ham, Blumea 37 (1993) 173-215. — Metcalfe, C.R. & L. Chalk, Anat. Dicot. 1 (1950) 419-422. — Radlkofer, L., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinch. 9 (1879) 457-678; Ueber die Methoden in der botanischen Systematik, insbesondere die anatomische Methode. Festrede (1883); in A. Engler, Pflanzenr. 98 (1931-1934) 1. — Solereder, H., Syst. Anat. Dicot. (1899) 257-268; ibid.. Erginzungsband (1908) 104-105. — Welzen, P.C. van, Leiden Bot.Series 12 (1989) 19-37. Chromosome numbers — Some tens of chromosome numbers of mainly South Amer- ican Sapindaceae are known and are listed below. Apparently polyploidy is scarce in the family. So far, only one certain case of tetra- ploidy has been found for Urvillea uniloba var. uniloba (Sapindoideae-Paullinieae, Fer- ruci 1981a). Ferruci (1985) also made two counts for Allophylus edulis (Sapindoideae- Thouinieae), once 2n = 14 was found and once 2n = 28. The first count, 2n = 14, is doubtful as it is exceptional among all other Sapindaceae; quite likely, a haploid set of chromosomes was counted erroneously as a diploid set. The species complex Allophylus cobbe (of which A. edulis is a synonym) is probably not a polyploid complex as suggest- ed by Leenhouts [Blumea 15 (1967) 301-358]. The basic chromosome numbers vary between x = 10 and x = 18. The tribes which are considered to show more derived characters, like Paullinieae and Thouinieae, tend to have the lower basic numbers. The following chromosome base-numbers are a compila- tion from literature. P.C. van Welzen & P.W. Leenhouts 430) Flora Malesiana ser. I, Vol. 11 (3) (1994) Dodonaeoideae x = 10,11, 12,14,15,16 Sapindoideae (ctd) Cossinieae ee dO) Llagunoa ING) Group B eens IES). 1106) Dodonaeeae ee AE lle (Ale) Cupanieae X= 14) 6 Dodonaea ea 1G enlSy Gls) Aporrhiza x = 14 Doratoxyleae NG Blighia Sle Filicium yore IG) Cupania x= 116 Harpullieae eS UIP Bs I) Nephelieae INAS NS NG Harpullia xe — tals) Alectryon X= ull6 Magonia ies) Dimocarpus es NS) Majidea se IY Litchi sales) Ungnadia x=) 16 Nephelium ees I Xanthoceras eg ellis) Xerospermum x = 16 Koelreuterieae Re LS AG: Schleichereae Xe aul eallG Koelreuteria yes Wil 5) ile Schleichera Ke oS slG Sapindoideae XH) ORES Group C xe 72) MLO ae: 14, 15, 16, 18 Paullinieae xi) LOE Group A x = 13,14, 15, 16, 18 Cardiospermum x = 10, 11 Lepisantheae Xo 13 4S 16 Houssayanthus x = 12 Chytranthus xr al Paullinia oe 1 Lepisanthes Koen SS aes Serjania Kee Pancovia x= 6 Urvillea xe — ell Melicocceae ee. lic) Thouinieae G2) ste Melicocca Ks wl Allophylus x= O24 Sapindeae 6 ellllwlseaks} Deinbollia Kee ls) Sapindus eS Mi ISS Sources: Ahuja & Natarajan, Curr. Sci. 26 (1957) 117. — Bhaduri & Bose, Proc. 36th Ind. Sci. Congr. pt. 3 (1949) 139-140. — Bowden, Amer. J. Bot. 32 (1945) 191-200. — Carr, Amer. J. Bot. 65 (1978) 238. — Chaudhuri, Curr. Sci. 9 (1940) 416. — Chen et al., J. Wuhan Bot. Res. 3 (1985) 423-428. — Diers, Z. Bot. 49 (1961) 437-488. — Eichhorn & Franquet, Comp. Rend. Acad. Sci. Paris 202 (1936) 1609. — Ferrucci, Bonplandia 5 (198 1a) 73-81; 5 (1981b) 164-174; Bolet. Soc. Argentina Bot. 24 (1985) 200-202. — Fritsch, Kulturpfl. 18 (1970) 194. — Gill, Bir, Sidhu & Singhal in Live, Taxon 33 (1984) 538. — Guervin, Rev. Cyt. Biol. Veg. 23 (1961) 4-87; Bull. Mus. Nat. Hist. Nat. Paris II, 33 (1961) 616— 619; 36 (1965) 858-868. — Hair & Beuzenberg, New Zeal. J. Sci. 2 (1959) 148-156. — Janaki-Ammal in Darlington & Wylie, Chrom. Atlas Flow. Pl. (1955) 197. — Kadry, Svensk Bot. Tidskr. 45 (1951) 414— 416. — Li Mao-xue, Act. Bot. Bor.-Occ. Sinica 7 (1987) 246-251, pl. II. — Maglio, Forni-Martins & da Cruz in Love, Taxon 33 (1984) 536. — Mangenot & Mangenot, Bull. Jard. Bot. Etat Brux. 27 (1957) 639-654; 28 (1958) 315-329; Rev. Cyt. Biol. Veg. 25 (1962) 411-447. — Mehra, Khosla & Sareen, Silvae Gen. 21 (1972) 96-102. — Miége, Ann. Fac. Sci. Univ. Dakar 5 (1960) 75-86. — Nanda, J. Ind. Bot. Soc. 41 (1962) 271-277. — Ono, Mem. Nat. Sci. Mus. Tokyo 10 (1977) 63-76, pl. 6-8. — Ramirez, Philipp. Agric. 45 (1961) 340-342. — Sarkar, Chakraborty, Saha & Das in Love, Taxon 25 (1976) 636. — Sarkar, Datta, Mallick & Chatterjee in Léve, Taxon 25 (1976) 649. — Sarkar, Datta, Chatterjee & Hazra in Love, Taxon 31 (1982) 578. — Semple, Ann. Missouri Bot. Gard. 61 (1974) 902. — Simmonds, Heredity 8 (1954) 139-146. — Singhal, Gill & Bir in Léve, Taxon 29 (1980) 356. — Sugiura, Bot. Mag. Tokyo 45 (1931) 353-355. — West, Brunonia 7 (1984) 1-194. Adema, Leenhouts, Van Welzen — Sapindaceae 43] AS iat al Phytochemistry and Chemotaxonomy — Chemical characters of the family have re- cently been discussed twice (Hegnauer 1973, 1990). Radlkofer (1890) fully discussed the taxonomic meaning of the chemical characters detected by him during his painstak- ing anatomical investigations. It should be stressed that he predicted the wide occurrence of saponins in the family, and showed their taxon-specific location in twigs, leaves, caly- ces, pericarp, testa, and embryo, and their deposition in ordinary parenchymatic cells or in idioblasts of different size and shape. These idioblasts (‘Secretzellen’) tend to occur in many, but by no means all taxa of the family and contain mainly resinoid compounds, mucilages, tannins, or (and) saponins; in fresh tissues the content of these idioblasts was assumed to be latex-like. Radlkofer also described the distribution, size and shapes of calcium oxalate deposits in the family. Leaf and twig epidermata of Sapindaceae are sometimes mucilaginous; Radlkofer gave an exact description of the mucilage cells of the epidermis and their largely erratic occurrence in the family. The chemistry of sapin- daceous mucilages is unexplored so far. Many taxa of Dodonaea, Filicium, Ganophyllum and Llagunoa have sticky, viscid exudates on leaves and twigs. Recent phytochemical results with sapindaceous plants demonstrate clearly that their chemical characters have much to offer to botanists looking for infrafamiliar and inter- familiar affinities. As already mentioned saponins occur widely in the family. The ichthyotoxic and de- tergent properties known from a large number of species are mainly derived from their saponins. Today the chemistry of these saponins is rather well known. Mono- and bidesmo- sidic saponins occur and their sapogenins are oleanene-type pentacyclic triterpenic acids such as oleanolic, medicagenic, and zanhic acids, and hederagenin, or polyhydroxylated derivates of beta-amyrin such as barringtogenol, R1-barrigenol, and camelliagenin-A. Sapindaceous saponins are often acylated in the sapogenin- and/or sugar-part; acetic, angelic, and other acids are acylating agents. Steroidal sapogenins have not so far been found in the family. The resinous exudates of Dodonaea taxa contain labdanoid and clero- danoid diterpenes, and triterpenes of the lupane-series. Lupeol, betulin, and betulinic acid were isolated from the bark of Schleichera oleosa. Accumulation of large amounts of quebrachitol, a monomethy! ether of l-inositol (= [-]-chiro-inositol) in leaves, barks, flowers, and fruits is highly characteristic of the family. This character is shared with Aceraceae and some Hippocastanaceae. Polyphenolic compounds seem to be accumulated in the family mainly in the form of coumarins (e.g. scopoletin), coumarinolignans (e.g. cleomiscosin-A), flavonoids, proan- thocyanidins (formerly leucoanthocyanidins), and tannins. Leaves of several taxa yield- ed glycosides of the flavonols kaempferol, quercetin, isorhamnetin, and sometimes myri- cetin, and of the flavone luteolin. Cuticular waxes and resinous exudates of Dodonaea species contain lipophilic flavonoids such as the prenylated and O-methylated kaempfer- ol derivatives viscosol and aliarin, and di- and trimethyl ethers of kaempferol (e.g. san- tin), and of 6-hydroxykaempferol (e.g. penduletin). A flavanone, pinocembrin, was also isolated from twigs of Dodonaea viscosa. Oligomeric proanthocyanidins and condensed tannins seem to be more or less ubiquitous; they are mainly based on procyanidins, but may also contain prodelphinidins. Hydrolysable gallotannins do also occur in Sapindaceae. Fruits of Harpullia pendula yielded gallic acid, m-digallic acid, and galloyl glucoses, 432 Flora Malesiana ser. I, Vol. 11 (3) (1994) and very recently the C-glucosidic gallic acid derivatives bergenin and | 1-O-galloylber- genin were isolated from leaves of Allophylus edulis var. edulis [Planta Medica 56 (1990) 679]. The tannin content of dry bark may reach 15—20% and tannin-rich sapindaceous barks are used locally as tanning agents. As previously mentioned, tannins are sometimes deposited in idioblasts or so-called tannin sacs. Sapindaceae store fatty oils, proteins, starch, and/or amyloid in taxon-specific com- binations and amounts in their embryos (there is no endosperm as was early noted by Radlkofer). Amyloid is known only from Cardiospermum. Large amounts of starch or fatty oils seem to occur vicariously. From a taxonomic point of view the fatty oils seem to be the most interesting storage products of seeds. Three main types of seed oils can be distinguished within the family: a) Oils which consist of the usual triglycerides only; b) oils which also contain non-cyanogenic cyanolipids; c) oils which consist of triglycer- ides and cyanogenic cyanolipids; b- and c-type oils are taxonomic markers of Sapinda- ceae. In cyanolipids the triol glycerin is replaced by a leucine-derived mono- or diol with a branched C,-skeleton, one double bond, and a terminal cyano group, e.g. HO-CH,- C(CH,)=CH-CN (= x). The difference between the non-cyanogenic (e.g. with alcohol x) and cyanogenic cyanolipids consists in the presence of a so-called cyanohydrin group in the alcohol part of the latter, e.g. CH,-C(CH,)-CH(OR)-CN (= y; R = H = alcohol, in casu a mono-ol). Cyanohydrins spontaneously release HCN; therefore an oil with the cyanolipid y with R = CO-(CH,),.-CH, yields on saponification the cyanogenic alcohol y with R = H. Obviously oils with cyanogenic cyanolipids are more or less toxic. Moreover, seed oils of Sapindaceae often contain unusual main fatty acids; arachidic, 11-eicosenoic, and dihydrosterculic acids are reported in literature. Arachidic and eicosenoic acids are pref- erentially combined with C.-alcohols, i.e. incorporated in cyanolipids, but may also be main fatty acids in oils without cyanolipids, such as the seed oil of Blighia sapida. A thorough analysis of the composition of the seed oils of a large number of taxa could be expected to disclose valuable characters for infrafamiliar classification. Cyanolipids are structurally and biosynthetically linked with a third group of toxic constituents, the cyanogenic glucosides. If in y R is a glucosyl residue, the hydrophilic cyanogenic glucoside heterodendrin of Heterodendrum oleaefolium and other taxa re- sults. Many Sapindaceae contain cyanogenic glucosides in dangerous amounts. Leaves of Heterodendrum oleaefolium can yield as much as 0.38% HCN (dry wt). The cyanoglu- cosides cardiospermin, its sulphate (ester with sulphuric acid), and heterodendrin (= di- hydroacacipetalin) have been isolated from Cardiospermum grandiflorum and Hetero- dendrum oleaefolium. In the former species such glucosides occur throughout the plant. Probably all parts (including defatted seeds) of sapindaceous plants, which are cyano- genic, contain such glucosides; these have aglyca (i.e. cyanohydrins) identical or nearly so with the alcohols esterified with fatty acids in cyanogenic cyanolipids. In ripe seeds of many Sapindaceae cyanoglucosides are replaced by cyanolipids; others may have cya- noglucosides in place of cyanolipids. Cyanoglucosides such as heterodendrin form a biochemical link between Sapindaceae and Leguminosae (acacipetalin and heteroden- drin in a number of Acacia species) and Rosaceae-Spiraeoideae (cardiospermin-type glucosides in Sorbaria). Adema, Leenhouts, Van Welzen — Sapindaceae 433 A fourth type of toxic principles in the family is represented by hypoglycin-A and biosynthetically related nonproteinogenic amino acids, such as alpha-(methylenecyclo- propyl)-glycine. They have a branched carbon chain of 6 or 7 C-atoms, and occur free or as glutamyl peptides in seeds and other parts of certain Aceraceae, Hippocastanaceae, and Sapindaceae. Hypoglycin-A was isolated for the first time from the ‘edible’ aril of Blighia sapida (Ackee tree) and subsequently traced as the cause of an intoxication known as ‘Jamaica vomiting sickness’. The biogenetic origin of these amino acids has not yet been established with certainty. They were thought to be derived from leucine or isoleu- cine, but results of recent biogenetic investigations (Kean & Lewis 1981) make another pathway more probable. Threonine, a C, hydroxyamino acid, and two C,-units supplied by methionine, would give the skeleton of the C, amino acid alpha-(methylenecyclopro- pyl)-glycine which is accumulated, for instance, in seeds of Litchi chinensis. Elongation of the chain of this amino acid by a mechanism known from glucosinolates should then result in hypoglycin-type C, amino acids. Finally some poorly known or erratically occurring classes of compounds should be mentioned. Essential oils were isolated from leaves of Serjania piscatoria, a fish poison plant, and S. serrata, from the wood of Exothea copalillo and trom Allophylus edulis (= A. cobbe), but their composition is not yet known. Radlkofer gave lengthy descriptions of glandular hairs and excretory cells (idioblasts) of sapindaceous plants. These anatomical characters fully agree with the production and accumulation of essential oils by some members of the family. Positive alkaloid reactions are reported in literature for a rather large number of Sa- pindaceae, but isolation and identification of alkaloid-like compounds are rare. Phenyl- acetamide was isolated from leaves of Allophylus cobbe and purine alkaloids (caffeine, theobromine, theophylline) accumulate in large amounts in seeds and other parts of some species of Paullinia (P. cupana, P. sorbilis, P. triantennata, P. yoco). Fresh pericarps of Blighia sapida yielded a quinonoid substance named blighione (C, H,,Og); its structure has not yet been elucidated. In summary, production and accumulation of quebrachitol, cyanolipids, cardiosper- min-type cyanoglycosides and hypoglycin-type amino acids are biochemical markers of the family which indicate affinities with Aceraceae and Hippocastanaceae on one side and with Leguminosae and the rosalean stock on the other. R. Hegnauer References: Hegnauer, R., Chemotaxonomie der Pflanzen 6 (1973) 271-287, 742, 786; 9 (1990) 486-496. — Kean, E.A. & C.E. Lewis, Phytochemistry 20 (1981) 2161. — Radlkofer, L., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinch. 20 (1890) 105-379 (especially p. 296-331, Anato- mische Charactere). Uses — The wood of several species is used for timber. (See also paragraph on wood anatomy, p. 425.) Various species are used in medicine, as vegetable, as soap (Sapindus saponaria) or fish poison. The family includes ornamentals such as species of Cardiosper- mum, Dictyoneura, Filicium, Koelreuteria, Lepisanthes. Some species are planted as gar- den fence (Dodonaea angustifolia), or as shade trees (Dodonaea angustifolia, Filicium). However, Sapindaceae are more important as a source of edible fruits and seeds. The juicy arillode or sarcotesta is particularly appreciated. Of economic importance, and hence 434 Flora Malesiana ser. I, Vol. 11 (3) (1994) widely cultivated, are Dimocarpus longan (Longan), Litchi chinensis (Litchi, Lychee) and Nephelium lappaceum (Rambutan). F. Adema Sources: Anonymous, The wealth of India. Raw materials. — Brown, W.H., Useful Pl. Philipp. 2 (1950) 356-369. — Burkill, I.H., Dict. Econ. Prod. Malay Penins. (1935); ed. 2 (1966). — Desch, H.E., Mal. For. Rec. 15 (1954) 528. — Heyne, K., Nutt. Pl. Indon. ed. 3 (1950) 988-1002. — Verheij, E.W.M. & R.E., Coronel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991). — Welzen, P.C. van, A. Lamb & W.W.W. Wong, Edible Sapindaceae in Sabah, Nature Malaysiana 13 (1988) 10—25. Taxonomy — The present system of the Sapindaceae follows that developed by Radlkofer (1890, 1931-1934) with only a few changes made by Muller & Leenhouts (1976). Radlkofer (1.c.) divided the family in two subfamilies (Eusapindaceae, Dyssapindaceae) and fourteen tribes. The subfamily Eusapindaceae was divided by him in two groups: Eusapindaceae nomophyllae and Eusapindaceae anomophyllae. In the view of Radl- kofer (1.c.) the Dyssapindaceae are derived from the Eusapindaceae. Muller & Leenhouts (1976) in their survey of the pollen types accepted most of the system of Radlkofer. The main differences are that Muller & Leenhouts combined the tribes Aphanieae and Lepisantheae and so have only thirteen tribes, and did away with the division of the Eusapindaceae, proposing a more informal grouping in three groups A, B, and C. The most important change is that the Dyssapindaceae (as Dodonaeoideae) are considered to be an assemblage of relicts and that the Eusapindaceae (as Sapindoi- deae) are more homogeneous and derived. The families Hippocastanaceae and Aceraceae are closely related to the Sapindaceae and may be included in that family, the former as a member of the tribe Harpullieae, the latter as a tribe of their own in the Dodonaeoideae. An enumeration of the subfamilies and tribus, and of the genera occurring in Malesia is given below. F. Adema A. Dodonaeoideae (= Dyssapindaceae) Cossinieae: — Dodonaeeae: Dodonaea Doratoxyleae: Filicium, Ganophyllum Harpullieae: Harpullia Sat Se ae oa 5. Koelreuterieae: Koelreuteria B. Sapindoideae (= Eusapindaceae) Group A 6. Lepisantheae: Glenniea, Lepisanthes, Zollingeria 7. Melicocceae: Tristira, Tristiropsis 8. Sapindeae: Atalaya, Sapindus Group B 9. Cupanieae: Amesiodendron, Arytera, Cnesmocarpon, Cupaniopsis, Dictyoneu- ra, Diploglottis, Elattostachys, Euphorianthus, Gloeocarpus, Gongrospermum, Guioa, Jagera, Lepiderema, Lepidopetalum, Mischocarpus, Paranephelium, Rhysotoechia, Sarcopteryx, Sarcotoechia, Synima, Toechima, Trigonachras Adema, Leenhouts, Van Welzen — Sapindaceae 435 10. Nephelieae: Alectryon, Cubilia, Dimocarpus, Litchi, Nepheltum, Pometia, Xero- spermum 11. Schleichereae: Schleichera Group C 12. Paullinieae: Cardiospermum 13. Thouinieae: Allophyllus References: Muller, J. & P.W. Leenhouts, A general survey of pollen types in Sapindaceae in relation to taxonomy, in I.K. Ferguson & J. Muller (eds.), The evolutionary significance of the exine (1976) 407— 445. — Radlkofer, L., Sitzungsber. Math-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinch. 20 (1890) 105— 379; in A. Engler, Pflanzenr. 98 (1932-1934) 3-18. KEY 1 TO THE MALESIAN GENERA (based on vegetative and flower characters) (F. Adema) la. Trees or shrubs, exceptionally lianas. Leaves simple, unifoliolate, (bi)pinnate or digitate. Inflorescences without basal tendrils ................--22++++-05- 2 b. Herbaceous or woody climbers. Leaves biternate. Inflorescences with basal tendrils Cardiospermum (p. 483) 2a. Leaves simple, unifoliolate, (im)paripinnate or digitate .................--. 3 PREY SOI INIMALC 20, se nra 5 ys Sais a) soe 4 ple Mints © w sis) ating es = Tristiropsis (p. 742) eee rT Gees TITIAN IPCI QUS ceo ae cg nwo cess alo aot nw wit Ie mm in ae chp oe rayne as siete - b. Leaves simple, unifoliolate, paripinnate or digitate ....................---. 5 4a. Pseudo-stipules present. Petals shorter than the sepals Lepisanthes subg. Otophora (p. 627) b. Pseudo-stipules absent. Petals longer than the sepals .. Paranephelium (p. 693) 5a. Scale hairs present (rarely only visible in inflorescences), often also with solitary Simple BAITS. YOUMS PaltS VISCIG osc ec -ce ne hain 2 Be ieee hans Fiche tule ae meee 6 b. Indumentum consisting of solitary, simple hairs, sometimes mixed with stellate hair tufts, or two-branched hairs (Litchi chinensis). Young parts not viscid ....... 11 N.B.: Lepisanthes fruticosa may have sticky inflorescences (race ‘glandulosa’ from Borneo). 6a. Leaves paripinnate: Ovules 1 or 2 per locule ...............-.--.--+-+s00: i) Pe eedA VES SIMpPle, OWIMES 2) Pek LOCC. scx a ath a ei atagt “hae sien Dodonaea (p. 522) BURP CANS DECSCHUI. cP as, a6 a cre gd wieione w nies Sse At wins mata ie ge oreo aa 8 Re PES AUSEME foe. cs cn Nk oe hom 2M epee me GOS eters EN eee = a gee 9 8a. Leaf-rachis winged. Disc hairy. Filaments glabrous. Ovary 2-celled Filicium decipiens (p. 754) b. Leaf-rachis not winged. Disc glabrous. Filaments hairy. Ovary 3-celled Lepiderema (p. 6/8) 9a. Leaflets entire. Sepals connate up to halfway, not petaloid. Stigma lobed... .. 10 b. Leaflets subentire to crenate. Sepals free, petaloid. Stigma grooved Dictyoneura (p. 507) Flora Malesiana ser. I, Vol. 11 (3) (1994) . Leaves (4—)5—8(—10)-jugate. Filaments of stamens glabrous. Ovules 2 per locule Ganophyllum falcatum (p. 538) Leaves (2—)3(—4)-jugate. Filaments of stamens sparsely hairy. Ovules | per locule Schleichera oleosa (p. 728) a. Leaves paripinnate, if unifoliolate than flowers regular, usually with 5 petals (Lepisanthes senegalensis) or without petals (Glenniea thorellii, Sarcotoechia plani- LECLYN cates RSs AP ROR eR IT AIS SR OA oe te SRI aS 12 . Leaves digitate, 1—5-foliolate. Flowers zygomorphic, 4-merous Allophylus cobbe (p. 459) | Lower side of leaflets’ with naked glands) 26 2 G21)..520 eee eee 13 > Lower side:of leaflets without naked: glands. 5 37..3..%- 4. 2.022 522 18 » eaves without pseudo-stipules). «2s bo scp teiaeblotens aie se oe 9 14 PE LCAVESMWALHAPSCUCO-StlDll CSiescaerscr-fow key) oacueah shone Sacken ksi ot: Pometia (p. 698) . Sepals free or basally connate, usually narrowly imbricate. Stamens usually 8 (6— LO) filamentssmostly bairy: to. ots erseusi seeds & a0 ss cee ele e ere 15 . Sepals almost totally connate, apert. Stamens 5, filaments glabrous or with few DEST Srey eecnan ss n ge tee ena oe Ae be hone Cans Sener Meme Cubilia cubili (p. 49/7) : Ovary 2-; rately 3-locullar, Walt <.2 = 34 n2k.. o sie snoters ols s oo) ays, 6 aaa ee 16 p Ovary, -5-lOcUlan SMO OU 7. 2's, 3.5 249 Se sppalayey ar avs Oey eon Sore copes see ican een 17 . Leaves 1- or 2-jugate, exceptionally unifoliolate or 3-jugate. Hairs if present main- ly in stellate tufts. Petals absent or present. Disc hairy .... Dimocarpus (p. 5//) . Leaves 1—7-jugate. Hairs if present solitary, simple. Petals present. Disc glabrous OG LARC DY NANI se tveret a, aera ees Ke, apes aes a monies Pete hs asia Xerospermum (p. 746) . Leaflets finely dentate to serrate. Ovary with irritant hairs...... Jagera (p. 6/4) . Leaflets entire. Ovary without irritant hairs ............ Trigonachras (p. 734) . Hairs often mainly in stellate tufts, mixed with solitary simple hairs (N.B.: In some cases these may be hard to find, especially when the leaflets are almost gla- [Sree eee ee eee ee ae ee ene en ee ty RN eee aos ado 6 sc 19 . Hairs solitary, simple, exceptionally two-branched (Litchi chinensis), if few small tufts of hairs present then leaves with pseudostipules (Pometia) ............ PL | . sepals connate-at the base-Petals absent or 1—5(—6)) oc cer. 3 ce oe ay eee 20 BSc pal suinee: shetal ScalwaVyiSws) si a-hepwer bey NS os ciara eae cna as Harpullia (p. 598) . Sepals valvate to narrowly imbricate in bud. Petals absent. Stamens 6 or 7, fila- ments clabrous, Ovary smooths «2 4. a2. oho eek eens: Glenniea (p. 540) . Sepals imbricate in bud. Petals absent or present. Stamens (6—)8(—10), filaments usually hamry.Ovary. tuberculate <5. 2.0) .see 52 oo oe Dimocarpus (p. 5//) 5 eavesswithipseudo-stipulles. 3 caeeeiitsscodhtseninm: 6 0%c's 3 oe apes, eee 22 p Leavesswathoutspseudo-Stipulesy: cojs2 6.5 6 .9ee = wey spies “p5y50 2bo,ctiskes yal ene 24 . Leaflets entire or dentate (Pometia pinnata). Sepals slightly to distinctly unequal, Mee OnUp to haliway connate. Petals spresemts 5 2i5 sy. ao ho oho cps creeyc eee 23 . Leaflets dentate. Sepals equal, connate, somewhat less than halfway. Petals absent Alectryon repandodentatus (p. 457) . Sepals free, imbricate, outer | or 2 distinctly smaller than inner 3. Stamens usually Si (ASO) eee. 38 ee nett sist 7 pease Lepisanthes subg. Otophora (p. 627) Adema, Leenhouts, Van Welzen — Sapindaceae 437 ee ee eee o Sepals up to halfway connate, not imbricate, slightly unequal. Stamens 5 (or 6) Pometia (p. 698) . Sepals free, imbricate, outer (1 or) 2 distinctly smaller than the innenshree/t) . 925 Sepals free or up to almost completely connate, apert, valvate or (slightly) imbri- cate, usually all equal, rarely slightly unequal ..........---- 0.0 eee eee eee 36 . Leaflets (apically) crenate, serrate or dentate ........-. 26... eee eee e eee 26 TST G oa a ae a an we a a Sa ae ahs fs cy 5 DO 28 . Branchlets straight. Petals with 2 scales ............2-. eee eee eee eee eee 27 Branchlets sinuate. Petal margin slightly folded, hairy Gloeocarpus patentivalvis (p. 546) _ Leaf rachis often slightly winged. Leaflets shallowly crenate, or with few teeth near the apex, rarely (sub)serrate. Sepals petaloid, (sub)glabrous outside. Petal scales HISHARIIGEOSECG on oo ws nym ah AES IS eee Oe Guioa (p. 548) Leaf rachis not winged. Leaflets dentate or serrate to crenate. Sepals not petaloid, at most the innermost one with a wide scarious rim, hairy outside. Petal scales not AERTS RT EO Me EE re LEST La Eger ct ae Cupaniopsis (p. 493) a. Ovary and pistillode 2-celled...... 1.2... 202s eee e eee cette renee 29 Ovary and pistillode 3- (or 4-)celled ... 1.2.2.5... eee eee ee eee eee eee 30 . Red glandular hairs present. Petals shorter than the sepals Cupaniopsis bilocularis (p. 497), celebica (p. 499), platycarpa (p. 503) _ No red glandular hairs. Petals as long as or longer than the sepals..........--. Lepisanthes tetraphylla (p. 630), subg. Aphania (p. 627) _ Petals with 2 scales, or with auricles, but then sepals glabrous outside ....... 3] Petals with | scale, rarely with a hairy ridge, if auricles present then sepals hairy pone A PTY Sa eR reer ea ee cence yite men ee ream re >| . Sepals glabrous or slightly hairy outside ............ 62+ sees eee este eee 32 Sepals hainy outsides minaqencteise) » ~~. 24 pees se eee Cupaniopsis (p. 493) _ Leaves bicoloured, usually punctate, domatia normally present. Petal scales crested or not, auricles rarely present. Disc complete or interrupted... . . Guioa (p. 548) Leaves concoloured, not punctate, domatia absent. Petal scales or auricles not crested. Pise: COMPpPIClE:< +5 5 sa ~ 405555 4445 S45 ee ee eee Rhysotoechia (p. 704) _ Petiole and rachis marginate to winged or not. Inflorescences terminal, sometimes also in axils of the upper leaves. Stamens not exserted. Stigma conical or elongate, grooved. Ovules sessile on a thickened placenta ...........+++++++++5505> 34 Petiole and rachis mostly not winged. Inflorescences terminal, axillary, rami- or cauliflorous. Stamens exserted or not. Stigma usually + capitate, globular or dome- shaped, slightly lobed, rarely elongate, grooved. Placenta with an obturator Lepisanthes (p. 627) . Leaves 1—5S-jugate. Petals 5. Disc complete, annular .......--------+++++5: BS Leaves 7—13-jugate. Petals 4. Disc interrupted, semi-lunate Sapindus rarak (p. 7/4) W Petalsauithvicsesles.cnctk. BGs Sapindus (p. 7/3) Flora Malesiana ser. I, Vol. 11 (3) (1994) a.0 Petalsisuallyabsentitarely up tor Sit) ye eee ee a, 37 » Petals*present, usually 5,sometimes reduced . : 7 .:eiepes see eee 43 . Ovary smooth or warty, |- or 2-, exceptionally 3-celled, usually lobed; stigma lobed, exceptionally grooved (Alectryon cardiocarpus: ovary 2-celled) ............ 38 . Ovary smooth, 3-celled, not lobed; stigma lobed or grooved ............... 40 . All hairs simple. Sepals connate or free, valvate or sligthly imbricate. Ovary LODE wiy.sdis nis Few SD ESS Bcd FOR a ios Win So weal WU oe aS Ge 39 . Two-branched hairs often present. Sepals connate, apert. Ovary not lobed Litchi chinensis (p. 654) . Ovary smooth. Sepals connate. Leaflets opposite or more rarely alternate, smooth belowsnoticlaucoustnme Aen Hee eee Alectryon subg. Alectryon (p. 450) . Ovary warty. Sepals free or connate. Leaflets alternate or more rarely to opposite, fmelyspapillate below;mostly glaucous .........55..55-4-. Nephelium (p. 669) . Twigs puberulous, pubescent or patently hairy. Leaves 1—5(—7)-jugate. Sepals free and imbricate or almost free to connate and apert or valvate. Stamens exserted. Stigma eroovedor short to longdobed sc 2)... 1. aseeecls hon sls | ene eee 4] . Twigs strigose, at least when young. Leaves unifoliolate or 1—2-jugate. Sepals free, slightly imbricate. Stamens hardly exserted. Stigma slightly lobed Sarcotoechia (p. 723) . Twigs puberulous or pubescent. Leaves 1—5(—6)-jugate. Sepals almost free to con- nate. apertionvValwatel) 2 /N0.2R RS Ae LO RUS 030 Se Oe 42 . Twigs patently hairy. Leaves 3—7-jugate. Sepals free, imbricate Tristira triptera (p. 740) . Leaves 1—5(—6)-jugate. Anthers glabrous or hairy. Ovary stipitate Mischocarpus (p. 658) . Leaves 2- or 3-jugate. Anthers glabrous. Ovary sessile Gongrospermum philippinense (p. 545) Betalsswathout.scales or auricles i392) .12 i. Sp LARI, eo erene 14 ‘. Petals*wathiscalestorauricles(. (sass (es eee ee 47 . Sepals free or up to halfway connate, calyx flat or cup-shaped with a wide mouth andidistinctilobes.(Stamens'4=10: sacieds bo Jou bo saan ae ee eee 45 . Sepals almost totally connate, calyx urceolate with a narrow mouth and minute lObeSMStainte nisi rail rt eI LN: APE SRE Cubilia cubili (p. 497) . Inflorescences axillary, together often pseudoterminal, rarely truly terminal. Ovary WAL Dy Ree iaine SW ONL een kha e See hee. Gee ie ee uD: ei nee 46 . Inflorescences ramiflorous. Ovary smooth ... Mischocarpus paradoxus (p. 662) . Leaflets alternate to more rarely opposite, lower surface mostly glaucous, without glands, with domatia. Stamens 4—10. Stigma lobed........ Nephelium (p. 669) . Leaflets opposite, lower surface not glaucous, with flat orbicular glands, without domatia. Stamens 8, exceptionally 7 or 9. Stigma grooved Xerospermum (p. 746) . Twigs smooth or striate to sligthly grooved. All leaves spirally arranged ..... 48 . Twigs with either 10-12 deep grooves or 6 slight grooves, rarely striate. At least upper leaves opposite or whorled, rarely spirally arranged. Ovary with irritant hairs Jagera (p. 6/4) Adema, Leenhouts, Van Welzen — Sapindaceae 439 mPetalsowith, lscale, this sometimes deeply cleft 2:3): 1.2m wie. s dak. fh eas 49 Beiicumit no RealesiOmainiGless s.:is% or te Le vs ola. wii ose ee 252 . Leaflets entire, rarely slightly sinuous (Nephelium compressum). Disc swollen, mostly free of the torus, without a rim or a collar. Anthers hairy. Ovary 2-, sometimes 3- ae eeamemranriniies StS 0 LRT Se OO Sa das eae ae ARH OREO 50 Leaflets hardly to coarsely crenate- to serrate-dentate, rarely entire. Disc composed of a flat ring, adnate to the torus except for the margin and with an erect rim or tubular collar. Anthers glabrous. Ovary 3-celled Amesiodendron chinense (p. 467) N.B.: Paranephelium with paripinnate leaves may key out here. From Amesioden- dron it differs in the swollen disc without rim or collar, from Nephelium in the smooth, non-papillate lower surface of the leaflets, from Lepidopetalum and Trigo- nachras in the warty ovaries with lobed stigmas. . Lower surface of leaflets smooth, not glaucous. Petal scale entire or bifid. Ovary Supelausiomaeropved ssls5edsntt vised shoving? want aes shetinge ss a Lower surface of leaflets finely papillate, glaucous. Petal scale bilobed. Ovary warty, stigma lobed, lobes spreading or recoiled................ Nephelium (p. 669) . Lower surface of leaflets with axillary hairy domatia, without glands. Ovary 2-, rarely 3-celled, stigma folded outwards like an overhanging roof Lepidopetalum (p. 620) Lower surface of leaflets without domatia, with glands. Ovary 3-celled, stigma cy- itadweall st) atten et. Hh ocrees iam x el Sea Sey Trigonachras celebensis (p. 737) aamenis@abrous. SttomaOTOOVEG!: isin) sider ASelSetss). es eee 53 Filaments hairy, exceptionally glabrous (Mischocarpus pentapetalus, triqueter: ovary stipitate, stigma distinctly lobed). Stigma grooved or lobed ................ 54 a. Sepals much connate. Petals with two scales Alectryon subg. Synalectryon (p. 450) Sepalsalmostitree. Petals with:aunicles, 2... © eee iene - 3 a. Fruit not winged. Unbranched tree. Leaflets finely dentate, with glabrous glands regularly along midrib at lower surface resembling insect damage Jagera (p. 6/4) Fruit basally winged. Branching tree. Leaflets entire or roughly dentate, without slands at lOWEr SUMACE.....maceee nase ni or Gc: «= Cnesmocarpon (p. 486) a. Leaves simple, unifoliolate, digitate or pinnate ...........-..--05 +e seers 5 . Leaves bipinnate (observe absence of axillary buds on secondary rachises) Tristiropsis (p. 742) a. Leaves all simple, unifoliolate, digitate, or imparipinnate..............----- 6 Leaves paripinnate or pseudo-imparipinnate (occasionally some leaves unifolio- EUS il ee De ee er Drak. 3 LES 10 Fruit not winged. Leaves digitate or imparipinnate ..........-..--.05++5++55 7 Fruit winged (Fig. 2c). Leaves simple............-..----- Dodonaea (p. 522) Leaves unifoliolate or imparipinnate. Fruit either (slightly) lobed and coriaceous, or more or less globose and very woody, larger than | by 1.5 cm; if about 1 by 1.5 oni then: rachis winged 340%. b 6. Tak Vachss 22 LR SOT. WF 8 Leaves (1—)3(—5)-digitate; rachis not winged. Fruit globose to obovoid, coriaceous, at most 0:4=1:3' by 013-018 emitiet). Cesare ee Allophylus cobbe (p. 459) 442 Flora Malesiana ser. I, Vol. 11 (3) (1994) ¢ a L/S Mees LY HP. (ee ie ay male iy ype ite ONG Fig. 1. Fruits with stinging hairs, spines, knobs, etc. — a. Fruits with stinging hairs; Jagera javanica (Blume) Blume ex Kalkman subsp. javanica. — b-j. Fruits with spines, knobs, etc. — b. Dimocarpus longan Lour. var. longan. — c. Ibid., var. malesianus Leenh. — d. Ibid., var. echinatus Leenh. — e. Litchi chinensis Sonn. —f. Nephelium uncinatum Radlk. — g. N. meduseum Leenh. —h. N. juglandifolium Blume. — i, j. Xerospermum noronhianum (Blume) Blume. Adema, Leenhouts, Van Welzen — Sapindaceae 443 Fig. 2 . Fruits with wings or crests. — a. Fruits crested; Alectryon repandodentatus Rad\k. — b-f. Fruits winged. — b. Atalaya salicifolia (DC.) Blume. — c. Dodonaea viscosa Jacq. — d. Sarcopteryx brachy- phylla Radlk. — e. Tristira triptera (Blanco) Radlk.; f. cross section. Sa. 9a. 10a. Haves never tr steilate MIS. Sone ee totes o teclete ct eae tem nae mines ence a 2 Hidiestmaliby tn stellate Uris": Onn se site 2 eee cee ones Glenniea thorelii (p. 544) Pseudo-stipules present or not; rachis winged or not. Fruit (slightly) lobed, smooth, coriaceous; wall glabrous inside ........ Lepisanthes subg. Otophora (p. 627) Pseudo-stipules absent; rachis not winged. Fruit subglobose, spiny to warty, woody: Wall PILOSE ISTE. oe ee mice ne ree ncaa ed ten eae crear sors Paranephelium (p. 693) Fruit winged (Fig. 2b, c, e), or with a crest on top of locules (Fig. 2a), or with a narrow sharp margin along the locules (Fig. 2d) ...............-.----045- 1] Fruit without wings, crests, or sharp margins, margins blunt .............-. 20 +44 Flora Malesiana ser. I, Vol. 11 (3) (1994) . Fruit winged (Fig. 2b-f) or with a sharp margin along the locules. Seeds naked, or with an arillode (seed cover free from testa), or with a smooth sarcotesta (seed Cover adnate. to testa) we tee oo od Pod oa a ees 3. oo a ee 12 Fruit with crest on top of locules (Fig. 2a). Seeds with a longitudinal strip of usually highly papillate sarcotesta (Fig. 3b) ....... Alectryon subg. Alectryon (p. 450) . Wings of fruit taller than broad (Fig. 2¢=e)) 3: .. «2... 2 ae eee 13 > Wings of fruit broader than tall (Fie: 2b) 3. 20... 3.) eee Atalaya (p. 479) a. Wings 4-10 mm broad. Seed naked .....2 2... 02.07%...» eee 14 . Wings less than 2 mm broad. Seed (partly) covered by an arillode or a sarco- EX] 3 |p ea ene Ree a aE tg, 0 ONE ee ey SEE PEER coe oo oc oo ooo 0er 15 . Fruit outside short pilose, inside glabrous, 1-locular, 3-carpellate Zollingeria borneensis (p. 754) . Fruit outside glabrescent, inside pilose, 3-locular, 3-carpellate (Fig. 2e, f) Tristira triptera (p. 740) . Fruit inside glabrous except sometimes some hairs on the fruit axis. Seed mainly to completely covered by an arillode’@ree trom testa)... .2. 222.74. serene 16 . Fruit inside pilose. Seed partly covered by a sarcotesta (adnate to testa)...... 17 . Indumentum absent to sericeous. Leaflets crenate to entire. Sepals dimorphic. Fruit coriaceous (G. pteropoda) to woody (G. contracta); pseudo-funicle absent (G. con- tracta) to present (G. pteropoda), long, curly, fruit axis beneath pseudo-funicle gla- DEOUSHHL eae seo accor n Guioa contracta (p. 569), G. pteropoda (p. 590) . Indumentum sericeous or hirsute. Leaflets entire. Sepals equal. Fruit coriaceous except for the woody base; pseudo-funicle usually short, straight (in S. caudata long and curly), fruit axis below pseudo-funicle hairy (in S. caudata glabrous) Sarcopteryx (p. 7/7) . Leaflets coriaceous, without domatia or with small pockets. Fruit 3-locular... 18 . Leaflets thin, with hair tufts as domatia. Fruit 2- (or very seldom 3-)locular Lepidopetalum (p. 620) . Fruits glabrous or shortly puberulous at most. Margin of leaflets entire to dentate/ Set ee eke: oh ae Teen nt AA wae eee Ee ne Bias. Ayo OloGne.D 6.0 0 6 19 . Fruits with appressed and patent hairs. Margin of leaflets dentate Cnesmocarpon dentata (p. 457) . Inflorescence a spike (cymules reduced to | or 2 flowers) Elattostachys (p. 527) . Inflorescence a thyrse (cymules di- or monochasial).......... Synima (p. 730) . Fruit warty to densely spiny or densely scaled (Fig. 1b-j) ................. P23 >; Enuitesmooth to wankled whem diye. : so. = <-citues cates ys to eee Di . Fruit inside glabrous. Seeds with sarcotesta (adnate to testa) or arillode (free from TES NN: SEENON APNE EMSC, SPM. 2 S83 As AOR on Oe COREY Renata co, c 22 PL Proitinside hatny. Seed MaKe ds. eee eres pen ec ee Paranephelium (p. 693) . Fruit completely covered with scales to simple spines ...................- 23 . Fruit with very few simple or branched spines...... Schleichera oleosa (p. 728) BAS CEG Wit SANCOLESTA, oi. cc. 5. 5 apse te pra ais aa a ia rc ona Eons ee 24 F eS Ce Guwithattl OG Gra atc3 5 ces ri hteates adedc dss Sadak Piped dabei sae 25 Adema, Leenhouts, Van Welzen — Sapindaceae 445 ee ——_ ae 24a. Leaves 1—5(—18)-jugate, usually papillate below (dull), often minutely sericeous; without glands, often with domatia. Sepals free to more than halfway up connate. Spines and scales on fruit usually higher than broad (Fig. 1f-h) Nephelium (p. 669) . Leaves 1—2(-3)-jugate, smooth below (more or less shiny), glabrous or hairy on midrib and nerves, not sericeous; often with scattered glands, without domatia. Se- pals free or only basally connate. Spines on fruit lower than broad (Fig. li, j) Xerospermum (p. 746) . Arillode covering seed (nearly) completely. Fruit indehiscent. Indumentum of tuft- Eee nranchned Or simplemairs so". 29. LSS ees oo. eh. Se 26 . Arillode covering lower half of seed only. Fruit dehiscent. Indumentum of simple ES TNR, eee ne. deere he St SE ee oe Cubilia cubili (p. 49/) . Indumentum often partly or mainly consisting of dense tufts of hairs. Glands present on lower side of leaflets near axils of veins (seldom absent in all leaflets). Seed MOGs Hiemeas broad =. 5. Els fe ee we ee oe Dimocarpus (p. 5//) . Indumentum consisting of solitary, simple or 2-branched hairs. Glands absent. Seed imetivman DGad. 8.2 Oe eet bey es GE Lee Litchi chinensis (p. 654) _ Seed naked. Placenta not thickened and cup-shaped below seed ............ 28 _ Seed with sarcotesta (can be a narrow basal ring around hilum) and/or arillode, or with thickened, cup-shaped placenta below seed .............--+.+-2--55- 36 . Leaves and twigs hairy and also covered with glandular scales (microscope!) . 29 Leaves and twigs without glandular scales, at most hairs only ..........---. 30 MEPehinel leaves WiNGed. coisa. niae sae oe heels Filicium decipiens (p. 754) Rachis of leaves without wing.............. Ganophyllum faleatum (p. 538) DAPSE MOICIOL Less Haihy ISIde S285) PP SIS aT, Re 31 Peer ptADLOUS IDSICE See Se ets ce ee eee cles rh er Sule wat atete heuer ene ate 34 PELEUMECOMMPIELely Hairy MSIAG 0 5).2 3) searre)-elelere' ate ete ere" «Sain a tabarenatm ner 32 b. Fruit inside hairy only around placenta.................... Sapindus (p. 7/3) _ Fruit sessile, more or less ellipsoid (to shortly obovoid) to subglobular, indehiscent to dehiscent with 3 or 4 usually unequal valves, or tearing apart at random; 2- or 3- locular:; either wall thick and hilum covering up to lower 3/4 of seed or wall thin and hilum covering less than lower 1/3 of seed ...................--+-.+-- 33 Fruit on broad stipe, obovoid, dehiscing loculicidally into 3 equal valves; 3-locular; wall thick, fleshy. Hilum covering less than lower 1/3 of seed Trigonachras (p. 734) a. Hairs short or none. Leaflets entire to serrate. Fruit capsular; wall 2.5—12 mm thick. Hilum covering up to lower 3/4 of seed ............. Paranephelium (p. 693) _ Hairs often more than 5 mm long. Leaflets entire. Fruit drupaceous; wall less than 2 mm thick. Hilum covering less than lower 1/5 of seed Lepisanthes tetraphylla (p. 630) . Fruit indehiscent. Endotesta not ruminate....................20200200025 35 . Fruit dehiscent. Endotesta ruminate .. Gongrospermum philippinense (p. 545) . Pseudo-stipules present or not; rachis winged or not; jugae | to more than 40. Outer (1 or) 2 sepals smaller. Fruit 2-, 3- (or 4-)locular, glabrous or pilose, less than 5 cm high; wall usually thin, sometimes thick and fleshy....... Lepisanthes (p. 627) 446 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 3. Fruits and seeds, special features. — a—c. Alectryon Gaertn. — a. A. glaber (Blume) Radlk., fruit dehiscing with an irregular calyptra. — b. A. ferrugineus (Blume) Radlk., fruit dehiscing with an irreg- ular calyptra, seed with a longitudinal strip of papillate sarcotesta. — c. A. connatus Radlk., fruit dehiscing septifragally. — d. Dictyoneura acuminata Blume subsp. acuminata, fruit valve inside with an extra, densely hairy, fleshy layer. b. Pseudo-stipules absent; rachis not winged; jugae 1—6. Sepals all equal. Fruit 2- locular, glabrous, either less than 2 cm high and wall rather thin or more than 6 cm hightand wall'very thick ‘and tleshty..-...-5.--+.+-c-- otto. Glenniea (p. 540) 56a) matrsin stellate tuits; also solitary”... secre ey ee eee Si), be Only solitary hairs’or plant @labrous.:.- 28-22... -02 oot sas. | eee 38 37a. Fruit lobed, loculicidally dehiscent with valves. Hilum small, covering less than WGIOlSCCUr ert tie ME emo rte oo core meter oe Harpullia (p. 598) b. Fruit not lobed, breaking up irregularly. Hilum very large, covering up to 1/2 of SCOURS eh ner Mes ACE ccc aecre ook Mera caters ones Dimocarpus (p. 5//) 38a. Pseudo-stipules absent. Fruit lobed or not; carpels usually thin to thick, coriaceous to woody, rarely mesocarp fleshy when fresh (Toechima).................- 3) b. Pseudo-stipules present (sometimes reduced or early caducous: look for scar). Fruit lobed, but sometimes simple by abortion of one cell; exocarp thin, hard, mesocarp ChICKeyWICY-WiCUEEKESIA: 6c ane ome ce ene ote chee mek mme a: Pometia (p. 698) 9a, Eruit completely hairy inside or hairs only alone suture .--..--+ 22+ 2-4 ace 40 b. Fruit glabrous inside but hairs sometimes present on the lower part of the axis 52 Adema, Leenhouts, Van Welzen — Sapindaceae 447 40a. Fruit inside completely hairy, or sometimes hairs only along suture, but then stigma Igpevanmernit notte haraly lobed ,GbOvale: 29.228. 2 6. eee gee ee 4] Fruit inside with hairs only along suture; fruit lobed, obcordate; stigma not lobed Arytera (p. 467) marmesnstde witout a fleshyilayeres £5 55010. iis chee te PIU D IS oe 42 . Fruit inside with a fleshy layer, thickest in middle of the valves, decreasing in thick- ness towards margin of valves and towards dissepiments (Fig. 3d) Dictyoneura acuminata (p. 508) . Fruit dehiscing loculicidally. Seed either (partly) covered by a smooth sarcotesta eidiatesto testa) or an anllode (free fromtesta) i -2.'. bs. -cn ee cence 43 Fruit either forming an irregular cap during dehiscence (Fig. 3a, b), then seed cov- ered with a longitudinal strip of usually highly papillate sarcotesta (subg. Alectry- on; Fig. 3b), or fruit dehiscing septifragally (Fig. 3c), then seed partly covered by a basal arillode, attached to a sarcotestal annulus (subg. Synalectryon) Alectryon (p. 450) a. Seeds (partly) covered by a sarcotesta (N.B.: upper margin can be free from seed), ARIMA SEE 2 3 Rei vs SS Bac Sheet yatta etna omereretie, sear 44 Seed (partly) covered by an arillode, sarcotesta restricted to the attachment around PAC UMMI Thad ots 502) es als ble Ye A, I eee epee ees se er eee 48 a. Fruit 2- or 3-locular, if 2-locular then fruit less than 1 cm high or outside gla- JUOUIS tosh; Set oe See an ens 3 Sn 2) a 45 Fruit 2-locular, more than 3 cm high, velutinous outside Cupaniopsis platycarpa (p. 503) . Fruit 2- or 3-locular, glabrous or very sparsely sericeous outside, not to strongly stipitate, if not stipitate fruit either less than 1.5 cm high or sarcotesta almost com- Fp EC OVELIN DSCC! Seti rs Cian ites eens Ses ers ae ee oa a ee ee 46 Fruit 3-locular, glabrous or tomentose/tomentellous outside, not stipitate, more than isssem high; ‘sarcotesta'a 2= (or 3=)lobed basal cupule.. ...4... 3.0 225: me 47 . Fruit glabrous outside. Leaflets with hair tufts forming domatia. Disc glabrous Lepidopetalum (p. 620) Fruit very sparsely sericeous outside. Leaflets without domatia. Disc pilose Sarcotoechia (p. 723) a. Leaflets with usually less than 10 major, spaced, strongly curved nerves. Fruit gla- brous or tomentose outside; wall more than 2 mm thick Toechima erythrocarpum subsp. papuanum (p. 732) Leaflets with more than 14 major, dense, rather straight nerves. Fruit tomentellous outside; wall usually c. | mm thick......... Euphorianthus euneurus (p. 536) a. Fruit usually not stipitate. Stigma erect, not lobed, only grooved (Fig. 4d). Arillode without basal Extension 44. 6 24)- leaks ie oes peneket eek Md Nee eee 49 Fruit usually with a narrow stipe. Stigma lobed, lobes spreading (Fig. 4a—c). Arillode usually with a pseudo-funicle (Fig. 4g)............... Mischocarpus (p. 658) a. Fruit wall thinly coriaceous to coriaceous, glabrous to pilose............... 50 Fruit wall woody; Slabrous).0g< 3/8 is eee boc ae be aes Elattostachys (p. 527) 448 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 4. Fruits stipitate or not, or lobed or not. — a-c. Fruits stipitate or not, not lobed. — a. Fruits sessile; Mischocarpus paradoxus Radlk. — b. Stipe short and broad; M. largifolius Radlk. — c. Stipe long and slender; M. pyriformis (F. Muell.) Radlk. subsp. papuanus (Radlk.) R.W. Ham. — d-f. Fruits lobed. — d. Fruits dehiscing loculicidally; Guioa diplopetala (Hassk.) Radlk. — e. Lepisanthes multijuga (Hook. f.) Leenh. — f. Just one lobe developed; Pometia pinnata Forst. & Forst. — g. Seed with a pseudo-funicle; Mischocarpus triqueter Radlk., ar = arillode, hi = hilum, pf = pseudo-funicle, t = testa. Adema, Leenhouts, Van Welzen — Sapindaceae 449 Ey euit>-lecular;elabrous to hairy, lobedjor not: !.).¢0!) 6.0 He eee eS a 5] . Fruit 2- (or 3-)locular, glabrous, lobed, lobes spreading Arytera brachyphylla (p. 47/) . Sepals unequal. Disc uninterrupted. Fruit wall coriaceous. Seed ellipsoid to obovoid; (lobed) arillode partly to completely covering seed....... Cupaniopsis (p. 493) . Sepals equal. Disc interrupted. Fruit wall thinly coriaceous. Seed lenticular; bilobed THlledescOvenne Seeds id niin ieee Lite. Diploglottis australis (p. 520) . Placenta seldom thickened below seed. Seed (partly) covered by sarcotesta and/or EM emebes ia. 34 ford. omen iri erie ashi dtiittes steht sae IRE ae 3 53 . Placenta thickened below seed, more or less cup-shaped. Seed naked Rhysotoechia (p. 704) mbrut msidewithoura fleshy lavertiraucwss 5 te beanie wort SeklOe... 54 . Fruit inside with a fleshy layer, thickest in middle of the valves, decreasing in thick- ness towards margin of vaives and towards dissepiments (Fig. 3d) Dictyoneura obtusa (p. 5/0) . Seeds covered with either an arillode (free from testa) or a glabrous sarcotesta (ad- nate to testa). Fruit indehiscent or irregularly or loculicidally dehiscing ...... 55 Seeds either covered with a longitudinal strip of usually highly papillate sarcotesta, locules of fruit forming an irregular cap during dehiscence (Fig. 3b, subg. Alectry- on); or seed partly covered with a smooth sarcotesta attached to an arillode, locules dehiscing septifragally from dissepiments (Fig. 3c, subg. Synalectryon) Alectryon (p. 450) PRICE Paltly) covered witha satcolestat f)i2 io. SMa eee cee eee 56 Meced (parity) covered withranarillode:. > 4 Rae ys Skee bree o etenee su) . Domatia absent; leaflets entire to crenate to dentate. Fruit 1-, 2-, or 3-locular . 57 . Domatia consisting of tufts of hairs; leaflets entire. Fruit 2-locular Lepidopetalum fructoglabrum (p. 622) Wbeatlets.entire Pratt 1=3-10Cilar Ale 22 PIAS a Be oe 58 . Leaflets crenate to dentate. Fruit 3-locular ... Amesiodendron chinense (p. 467) HeMTBITIS ES OCULAR &. ig iets 3) Alita & shtee tienes Nephelium maingayi (p. 685) Sea TEL EY 1 OT 9 =O GUNAR Ys 282 he Ses PRL baie veas, we aca ei Rhysotoechia applanata (p. 706) PGUIETHGSHISGEDD OF GENISCING ITSP UL AGI. arta as wteneper- raga ce 1a eto ca tate cua haan baker as 60 PALME OCMC Cally: GEHISCENE) 6 3).00 02 efe< iS Balers eases lige. eens afar bkieta nya Cree 6] a. Fruit a 1- or 2-celled dry berry, indehiscent, not to hardly stipitate, ellipsoid; endo- carp in dried state not detached from mesocarp ..... Schleichera oleosa (p. 728) . Fruit a 3-locular capsule (usually only 1 loculus developed), dehiscing irregularly, long-stipitate, globose; endocarp in dried state detached from mesocarp, forming cupule around arillode ..... Arytera macrobotrys (p. 474), A. bullata (p. 47/) a. Stigma a pyramidal remnant without recurved lobes on top of the fruit (check sev- SEAMUL LES cis, AWM ois V.083,abe raked tack te ak Penn ete RES EYE Go ct eM be th iE RGSS NS Leon hea 62 b Oromia WIth MECURVECLODES tic bite ote ts Biri ste leak Mischocarpus (p. 658) . Arillode with a basal pseudo-funicle (short and straight or long and sinuous). Fruit JODEG@OMINOL At: 4h Rete. SUES Bae SLRS tS AOE R MI RE See, GRC ES as 2 .ine: 63 450 Flora Malesiana ser. I, Vol. 11 (3) (1994) 63a. Fruit lobed (Fig. 4d), often clearly stipitate, wall coriaceous but woody in G. con- tracta. Pseudo-funicle long and sinuous. Sepals dimorphic ..... Guioa (p. 548) b. Fruit hardly lobed (Fig. 2d), stipe absent or inconspicuous, wall woody in lower half, more coriaceous in upper half. Pseudo-funicle usually short and straight, sel- dom long and sinuous (S. caudata). Sepals all equal...... Sarcopteryx (p. 7/7) 64a. Branchlets more or less straight. Leaves 2—10-jugate. Scale hairs absent or present. Inflorescence ramiflorous to axillary to terminal. Fruit sessile to stipitate..... 65 b. Branchlets usually appearing sinuous because of distichous phyllotaxis. Leaves 5— 14-jugate. Scale hairs absent. Inflorescence ramiflorous. Fruit hardly stipitate Gloeocarpus (p. 544) 65a. Scale hairs present in inflorescence. Pith in axis thin. Leaflets entire; apex not mucro- nulate. Disc glabrous. Fruit glabrous, wall thin, stipe very slender Lepiderema (p. 6/8) b. Scale hairs absent in inflorescence. Pith in axis thin to thick (up to 2/3 of diameter of twig). Leaflets entire to serrate to crenate; apex usually mucronulate. Disc gla- brous, or with 5 bundles of hairs, or pilose. Fruit glabrous to pilose, wall thin to thick stipe-absent.toibroadly cumeaten i. 2 als It Cupaniopsis (p. 493) ALECTRYON (P.W. Leenhouts) Alectryon Gaertn., Fruct. Sem. Pl. 1 (1788) 216, pl. 46; Radlk. in Engl., Pflanzenr. 98 (1933) 983; S.T. Reynolds, Austrobaileya | (1982) 472; in Fl. Austral. 25 (1985) 24; Austrobaileya 2 (1987) 332; Leenh., Blumea 33 (1988) 313. — Type species: Alectry- on excelsus Gaertn. Spanoghea Blume, Rumphia 3 (1847) 172. — Lectotype species (S.T. Reynolds, 1982): Spanoghea ferruginea Blume [= Alectryon ferrugineus (Blume) Radlk. ]. Medium-sized to small trees or shrubs, sometimes monoecious. Jndumentum of soli- tary simple hairs only. Branchlets terete. Leaves paripinnate, 1—8-jugate, unwinged; true stipules absent, lowermost pair of leaflets sometimes stipule-like. Leaflets opposite (or alternate), variably hairy (to glabrous), lower side smooth; base often oblique, then usu- ally in the lower leaflets the acroscopic side and in the upper ones the basiscopic side stronger developed; margin entire or serrate, dentate, or crenate; nerves ending free or the upper ones looped and joined. /nflorescences axillary (or together pseudo-terminal, rarely ramiflorous), a thyrse or panicle (or raceme). Flowers unisexual (or bisexual), actinomorphic. Sepals 4, 5 (or 6), from somewhat less than halfway to nearly completely connate, all equal, hardly or not petaloid, hairy on both sides (or inside glabrous). Petals absent or 4 or 5, about as long as to shorter than the sepals, short-clawed, with 2 scales without a crest, outside glabrous, margin ciliolate, inside hairy or glabrous. Disc uninter- rupted, annular or + lobed, without appendages, glabrous. Stamens (5—)8, exserted in male flowers; filaments hairy to glabrous; anthers basifixed, the base usually deeply cleft, usually glabrous, dehiscence latrorse or latro-introrse. Pistil sessile or short-stalked, densely hairy; ovary (1—)2—4(—5)-locular; style apical, columnar, mostly shorter than the ovary; Adema, Leenhouts, Van Welzen — Sapindaceae 45] stigma grooved or with recoiled lobes. Ovules | per locule. Fruits sessile or short-stipi- tate, 1- or more-lobed, capsular, dehiscing either with a loculicidal calyptra or septifra- gally (along septum), smooth or slightly warty, hairy or finally glabrous, inside glabrous (or hairy). Seeds black, partly covered by a red sarcotesta (see under the subgenera). — Fig. 5. Distribution — About 25 species in E Malesia, Australia, New Zealand, New Caledo- nia, New Hebrides, Solomon Islands, Fiji, Samoa, and Sandwich Islands. Habitat & Ecology — Often in monsoon forest, in forest edges, river banks, or coastal vegetations, often on limestone, in the lowland but sometimes also montane. The seeds, with the contrasting light yellowish greenish capsules, the red sarcotesta, and the shining black testa, are probably mainly dispersed by birds. Uses — Some species are good timber trees. Note — Alectryon seems to be closest to Heterodendrum Desf. from E Australia and Stadmania Lam. (incl. Smelophyllum Radlk.) from the Mascarene Islands, Madagascar, and SE and S Africa. KEY TO THE SPECIES la. Ovary 3—5-locular; fruits broadly ovoid, slightly lobed, septifragally dehiscent; seeds with a small sarcotesta around the hilum and the basal half enclosed by a cupular BEM CHC SH DOS VNGICCE YON). +0. crix cin oo apse Sie tgs ase Sik ny fee 6 sine ie URS een 2 b. Ovary 1- or 2-, exceptionally 3-locular; fruits with + spreading lobes, dehiscent by an irregular calyptra; seeds largely covered by a sarcotesta with only a narrow, lobed, Treo maatem (SUE, ATCCELYON) «. ~ ..<.tatee ie n> sis je 25 spose Fino. « © hcl etiRintng a ol
  • 3. A. kangeanensis 3a. Leaflets + acuminate; nerves not distinctly connected; fruit wall hard, inside gla- “DON DLT ip Ab ge ergo an eet nee ene O PRES. AUN mRNA CRIS PENS 5 1. A. affinis b. Leaflets broadly rounded to slightly emarginate; nerves looped and joined; fruit wall pergamentaceous, inside hairy. .................0.5--- 2. A. connatus Aq. Mowermiost pair of leaflets not stipule-like..- we ee ee 2 b. Lowermost pair of leaflets stipule-like, attached near the base and much smaller Sate OLHCTSW: otoris deta eT. - > els a std LARMVEY- 9. A. repandodentatus Bal Egil? ono AOCUlans. acme. WA... 14). eae a es 6 ‘Siggel x SiG yy BBV S/EcT1 ae nan dau ae genres See 2 ets) ae 10. A. reticulatus 6a. Leaflets entire; fruits usually with only 1 lobe developed, c. 15 by 12.5—15 mm diam., the wall corky, c. 1.5—2 mm thick; seeds with a smooth sarcotesta...... 7 b. Leaflets nearly always incised; often 2 fruit lobes developed, these c. 8-10 mm diam., the wall woody, 0.5—1 mm thick; seeds with a papillose sarcotesta ..... 8 7a. Leaflets acuminate at apex; midrib above prominulous; branchlets hollow, inhabit- eg Byants .acciaire th node Salle. vated aid 8. A. myrmecophilus b. Leaflets rounded at apex; midrib above slightly sunken; branchlets solid, without Rens ee ee en eet Eran oi Morera eye ae ote OMe wep tend es acne 6. A. fuscus 452 Flora Malesiana ser. I, Vol. 11 (3) (1994) 8a. Twigs terete to slightly grooved, up to 5 mm thick; petiole halfway along its length up-to, 2:5.mm thickleaflets. up toWl6:subysO.eme se eee -1e ee 9 b. Twigs strongly 5-grooved, 4-10 mm thick; petiole halfway along its length 2.5—8 mimithickwWeatletsmpito4s by Miscmyede a Seeee Bae eee aoe 5. A. ferrugineus 9a. Margin of leaflets with some distant teeth mostly only in the upper half; fruits not ononlysshchtlycordateshssiea haere eee lie ee oe ee 7. A. glaber b. Margin of leaflets rather densely serrate-dentate nearly from the base; fruits deeply Condatenn saiutratiy ae see is Cee nis a ee 4. A. cardiocarpus Subgenus Synalectryon Alectryon Gaertn. subg. Synalectryon (Radlk.) Leenh., Blumea 33 (1988) 315. — Alec- tryon Gaertn. sect. Synalectryon Radlk., Sapind. Holl.-Ind. (1879) 93. — Lectotype species (Leenhouts 1988): Alectryon connatus (F. Muell.) RadIk. Twigs tomentellous, glabrescent; branchlets black or grey, glabrous. Petiole semi- terete; rachis above flat, + carinate; leaf axes tomentellous, glabrescent or not. Leaflets pergamentaceous, entire. Inflorescences axillary, a thyrse, hairy, sparsely branched. Se- pals 5, connate for halfway or more, hairy on both sides. Petals 5 (or absent in male flowers), the blade rounded, ciliolate, scales woolly. Disc especially in male flowers some- times hardly developed. Stamens in male flowers 8 (always?) (to fewer in female flow- ers), filaments and anthers glabrous. Pistil 3—S-merous; stigma grooved. Fruits short- stipitate, broadly ovoid, slightly lobed, the base slightly concave, the apex apiculate, dehiscence septifragal; wall either pergamentaceous or hard, inside glabrous or hairy. Seeds with a sarcotesta around the hilum and the basal half enveloped by the cup-shaped arillode. Distribution — Apart from the 3 Malesian species two more species belong to this subgenus, viz. A. coriaceus (Benth.) Radlk. and A. subcinereus (A. Gray) Radlk. from Queensland and New South Wales. 1. Alectryon affinis Radlk. in Engl. & Prantl, Nat. Pflanzenfam., Nachtr. 3 (1907) 205; Bot. Jahrb. 56 (1920) 275; in Engl., Pflanzenr. 98 (1933) 1001; Leenh., Blumea 33 (1988) 315. — Lec- totype (Leenhouts 1988): W. Fitzgerald 30 (M holo), New Guinea. pressed short-hairy to glabrous; base symmetric or in upper leaflets the basiscopic side slightly more developed, acute to obtuse, slightly attenuate; apex hardly to slightly acuminate, very apex rounded; midrib above flat to prominulous, nerves 0.5—1 cm apart, curved, ending free or the upper ones + looped and joined near the margin, prominulous Erect shrub with spreading, often pendulous — on both sides, intersecondary nerves few, veins and branches, c. 3 m high, or tree up to 14 m high, dbh c. 19 cm; outer bark grey, shallowly fissured; in- ner bark red. Twigs grooved, c. 2.5 mm thick. Leaves |—3-jugate; petiole 3—7.5 cm long, c. 1.5 mm thick; petiolules 2—4 mm long, above broadly grooved or flat. Leaflets opposite (to alternate), ovate to elliptic, 7-16 by 2.5—8 cm, index 2-3.5, above glabrous, beneath fairly densely short-hairy on midrib and nerves, in between sparsely ap- veinlets minutely reticulate, prominulous. /nflores- cences up to 16 cm long, sparsely hairy, peduncle 1—3.5 cm long; branches few, up to 4 cm long but usually much shorter; cymules short-stalked, sev- eral-flowered; pedicels 2—5 mm long. Sepals c. | mm high. Petals c. 0.8 mm long, on both sides gla- brous. Stamens 7 or 8; filaments c. 0.8 mm long; anthers c. 1.5 mm long, yellow to pink. Fruits c. | by 1.25 cm, yellowish brown, slightly warty, out- Adema, Leenhouts, Van Welzen — Sapindaceae 4 Nn is) icm i imm Fig. 5. Alectryon Gaertn. Habit and fruits. — A. cardiocarpus Leenh. a. Fruit. — A. connatus Radlk. b. Habit of fruit bearing twig; c. unopened fruit; d. partly dehisced fruit. — A. ferrugineus (Blume) Radlk. e. Fruit. — A. glaber (Blume) Radlk. f. Fruit. — A. myrmecophilus Leenh. g. Fruit. — A. repan- dodentatus Radlk. h. Fruit. — A. reticulatus Radlk. i. Fruit (a: BW 4804; b, c: Pullen 6895; d: Byrnes & Clarkson 3622; e. Fallen, Wiakaba & Lelean 339, f: Alston 16075; g: NGF 45253; h: Pullen 6871; i: Hartmann s.n.). 454 side very sparsely hairy to glabrous, inside gla- brous, wall hard, c. 0.5 mm thick. Distribution — Malesia: Papua New Guinea. (Central and Northern Provinces). Habitat & Ecology — Rather dry forest on rocky slope, patches of secondary forest in /mperata grassland. Altitude 150-750 m. Fl. July—Sept.; fr. Sept. 2. Alectryon connatus Radlk., Sitzungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 8 (1878) 340; in Engl., Pflanzenr. 98 (1933) 1000; S.T. Reynolds, Austrobaileya | (1982) 473, f. 37c; in Fl. Austral. 25 (1985) 25, f. 4f-h, map 25; Leenh., Blumea 33 (1988) 317, f. 1. — Spanoghea connata F. Muell., Trans. Philos. Inst. Victoria 3 (1859) 26. — Type: Hill & Mueller s.n. (MEL holo), Australia, Queens- land. Tree up to 20 m high, dbh c. 20 cm (ora shrub). Twigs grooved, 1.5—4 mm thick. Leaves 1—3-jugate; petiole 1—4.5 cm long, strongly flattened like the rachis; petiolules absent to up to 3 mm long, above broad and flat. Leaflets opposite to alternate, el- liptic (to slightly obovate), 4.5—12.5 by 1.75—5 cm, index |.5—3, above glabrous or sparsely hairy on midrib, nerves, and veins, beneath at first hairy all over, glabrescent with the exception of midrib and nerves; base symmetric to oblique, then the basi- scopic side broadest, acute to obtuse, attenuate or not; apex rounded to slightly emarginate; midrib above angular to rounded, nerves 0.75—1 cm apart, nearly straight, lower ones not joined, upper ones looped and joined near the margin, prominulous on both sides, intersecondary nerves often strong- ly developed, veins rather laxly reticulate, veinlets densely reticulate beneath. /nflorescences 6-12 cm long, fairly densely hairy; peduncle 1.5—3 cm long; pedicels c. 1 mm long. Sepals c. 1 mm high. Petals 0.5—1 mm long, inside with some woolly hairs; in male flowers sometimes absent. Stamens in male Flora Malesiana ser. I, Vol. 11 (3) (1994) flowers 8, in female flowers 5; filaments up to 1.5 mm long; anthers c. 1.5 mm long. Fruits 0.5—0.8 by 1—1.2 cm, outside sparsely tomentellous, inside sparsely woolly; wall thin-pergamentaceous. — Fig. 5b-d. Distribution — Australia (Queensland) and Malesia: Papua New Guinea (Central Prov.). Habitat & Ecology — Low monsoon scrub in the lowlands. Fr. mainly Apr.—July. 3. Alectryon kangeanensis Leenh., Blumea 32 (1987) 222; 33 (1988) 317. — Type: Backer 29552 (L holo; BRI, K, SING, U, UC), Kangean Archipelago. Twigs grooved, 3.5—4.5 mm thick. Leaves 1—3- jugate; petiole 1-5 cm long, strongly flattened like the rachis; petiolules absent to up to 2 mm long, above broad and flat. Leaflets opposite to alternate, ovate to elliptic (to obovate), 8-15 by 2.5—5 cm, index 2.54, glaucous below, usually with scattered hairs on the midrib; base symmetrical to slightly oblique, acute; apex narrowly rounded; midrib above angular, nerves 0.5—1.5 cm apart, slightly curved, all (or except the lower ones) looped and joined near the margin, above hardly, beneath dis- tinctly prominent, intersecondary nerves often well developed, veinlets densely reticulate, indistinct. Inflorescences 10-15 cm long, fairly densely hairy; branch 2-6 cm long; pedicels c. 3 mm long. Se- pals c. | mm high. Petals c. 0.5 mm long, scale densely hairy. Stamens: filaments in male flowers c. 2 mm long; anthers c. 1.5 mm long, dehiscence latrorse. Fruits c. 0.75 by 1 cm, outside sparsely tomentellous, inside glabrous; wall pergamenta- ceous. Distribution — Malesia: Kangean Archipelago (Paliat I., one collection only). Habitat & Ecology — Altitude 50 m. Fl. and young fr. in May. Note — The present species fairly strongly re- sembles A. coriaceus Radlk. from NE Australia. Subgenus Alectryon Alectryon Gaertn. subg. Alectryon: Leenh., Blumea 33 (1988) 318. — Alectryon Gaertn. sect. Eualectryon Radlk., Sapind. Holl.-Ind. (1879) 93. — Alectryon Gaertn. sect. Spanoghea (Blume) Radlk., Sapind. Holl.-Ind. (1879) 93. — Type species as the ge- nus. Twigs variably hairy, + glabrescent; branchlets grey to black (to brown), glabrous (or hairy). Petiole terete to semiterete (to triangular in cross section); rachis terete (to angu- lar above, flat and strongly ribbed, or triangular in cross section); leaf axes hairy (or Adema, Leenhouts, Van Welzen — Sapindaceae 455 glabrous). Leaflets mostly variably pergamentaceous (to herbaceous or papyraceous or thin-coriaceous); margin serrate, dentate, or crenate at least in the upper half (or entire). Inflorescences axillary (or ramiflorous), a sparsely branched panicle or thyrse (or ra- ceme), densely hairy (to nearly glabrous). Sepals 4, 5, (6), somewhat less than halfway to nearly completely connate, hairy on both sides or on the outside only. Petals usually absent, if present blade kidney-shaped, inside woolly. Stamens 5—8; filaments variably hairy (or glabrous); anthers glabrous (or sparsely ciliate). Pistil (1- or) 2- (or 3-)locular; stigma lobed (to grooved). Fruits sessile, lobed, the + spreading lobes often slightly car- inate and shouldered, the wall woody or corky, dehiscence by an irregular calyptra. Seeds from around the hilum to near the apex with a narrow to usually very broad strip of strongly papillose sarcotesta with a lobed free margin. Distribution — Like the genus; about 20 species. 4. Alectryon cardiocarpus Leenh., Blumea 32 (1987) 221; 33 (1988) 320. — Type: BW (Ver- steegh) 4804 (L holo; BO), New Guinea. Alectryon reticulatus auct. non Radlk.: Rehder, J. Arnold Arbor. 14 (1933) 63. Tree up to 11 mhigh, dbh c. 20 cm; bark smooth, brown with conspicuous grey or white patches. Twigs terete, 4-5 mm thick, glabrescent. Leaves 3—5-jugate; petiole slightly flattened above, 4—7.5 cm long, 1.5—2.5 mm thick; petiolules 4-10 mm long, above broad and flat with the midrib raised; axes hairy, glabrescent. Leaflets (sub)opposite, ovate to elliptic, 6.5—16.5 by 2.5—6 cm, index 2— 3.25, stiff-pergamentaceous; glabrous, or sparsely hairy on the midrib (also with a few hairs on the nerves beneath); base hardly to very oblique, the acroscopic side broader than the basiscopic side, rounded to acute, slightly attenuate ot not; margin serrate-dentate; apex acute, mucronate; midrib above prominulous; nerves 1—1.5 cm apart, oblique- ly patent, slightly curved, ending in marginal teeth, prominulous on both sides; intersecondary nerves hardly developed, veins and veinlets laxly reticu- late. Inflorescences axillary, panicles, up to c. 6 cm long; branches few, obliquely patent to patent, up to 6 cm long, rather many-flowered, densely hairy; peduncle c. 5 mm long; pedicels very short. Sepals \-1.Z mm high, nearly completely connate, hairy at both sides to only inside sparsely hairy mainly basally. Petals absent. Stamens: filaments 0.8—1.75 mm long; anthers c. 1.25 mm long. Pistil 2-locular, stigma apparently grooved. Fruits 2- lobed, deeply cordate especially when young, smooth, lobes globular, c. 8 mm in diam., densely fulvous-tomentellous; wall c. 0.5 mm thick. — Fig. Sa. Distribution — Malesia: New Guinea. Habitat & Ecology — In monsoon scrub and sec- ondary forest, on coastal limestone rocks, river banks, and slopes, from sea level to 40 m altitude. Fl. May—June; fr. May—Aug. Note — The present species is close to A. ferru- gineus; it differs primarily by the small, deeply cordate fruits. Furthermore, A. ferrugineus is gen- erally far more coarse and more hairy, and the hairs are not so reddish brown as in A. cardiocarpus. 5. Alectryon ferrugineus (Blume) Radlk., Sapind. Holl.-Ind. (1879) 14; in Engl., Pflanzenr. 98 (1933) 995; P. Royen, Man. For. Trees Papua & New Guinea 2 (1964) f. 2; Leenh., Blumea 33 (1988) 321. — Spanoghea ferruginea Blume, Rumphia 3 (1847) 173. — Nephelium ferrugi- neum F. Muell., Descr. Notes Papuan Pl. 1 (1876) 21. — Lectotype (Leenhouts 1988): Zip- pelius 158a (L holo), New Guinea. Alectryon strigosus Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 255; in Engl., Pflanzenr. 98 (1933) 991.— Type: J. Chalmers s.n. (MEL, n.v.), New Guinea. Jagera latifolia Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 264. — Type: W. Sayer s.n. (M holo; MEL), New Guinea. Alectryon mollis Radlk. in K. Schum. & Laut., Nachtr. Fl. Schutzgeb. Siidsee (1905) 308; in Engl., Pflanzenr. 98 (1933) 992. —Type: Hell- wig 3 (B¥ holo; BO, K), New Guinea. Alectryon macrophyllus Kanehira & Hatusima, Bot. Mag. Tokyo 52 (1938) 413, f. 3. — Type: Kane- hira & Hatusima 4034 (FU, n.v.), New Guinea. Tree up to 20 m high, dbh c. 25 cm, but mostly much smaller, or shrub; (with small buttresses); bark smooth, brown or grey to black, (patchy). Twigs strongly 5-grooved, 0.4—2 cm thick, fairly densely hairy, usually glabrescent. Leaves 2—8-ju- gate; petiole terete (to semiterete or triangular in 456 cross section), 2-29 cm long, 2.5—8 mm thick; pet- iolules 1-10 mm long, above flattened (or broadly grooved with or without a median rib, or terete). Leaflets opposite (to alternate), ovate to elliptic (to slightly falcate), 5.5—45 by 3-17 cm, index 1.75— 3.5, pergamentaceous (or herbaceous or thin-co- riaceous); mostly above densely hairy on midrib and nerves, beneath sparsely hairy all over to gla- brous; base slightly to strongly oblique, in lower pairs of leaflets broader at the acroscopic side to basiscopic side broader in upper pairs, subcordate or rounded to acute, (slightly attenuate); margin dentate to serrate at least in the upper half (to sub- entire); apex rounded to acute-acuminate; midrib above prominulous or sometimes flat; nerves 0.75— 2.5 cm apart, oblique-patent to widely spreading, curved to nearly straight, ending in the marginal teeth or not, above prominulous (to grooved), be- neath prominent; intersecondary nerves variably developed; veins and veinlets laxly to densely re- ticulate, either prominulous on both sides or only beneath. Inflorescences axillary (or ramiflorous), panicles or thyrses, up to 30 cm long; branches few, patent, up to 12 cm long, often densely flowered, densely hairy; peduncle up to 2 cm long; pedicels 0-1.5 mm long. Sepals 0.75-1.5 mm high, con- nate, inside hairy. Petals absent (but see note). Sta- mens: filaments 0.5—2.25 mm long; anthers 1.25— 2 mm long. Pistil 2-locular with a lobed stigma. Fruits 2-lobed(to only | lobe developed), carinate and angled to shouldered, fairly densely hairy but finally glabrescent, the lobes about globular, 8—10 mm diam.; wall 0.5—0.75 mm thick. — Fig. Se. Distribution — Malesia: Halmahera, Ternate, and New Guinea (mainly the eastern half). Habitat & Ecology — In primary and secondary forests, often along edges, on stream banks, some- times periodically flooded or in swampy places, often on limestone; sea level up to 300(—1950) m altitude. Fl. and fr. throughout the year. Note — Clemens 8489 from Papua New Guinea, Morobe Prov., Boana, is the only specimen that has flowers with petals, albeit strongly reduced; they are c. 0.8 mm long and the scale is represent- ed by a rim of hairs only. 6. Alectryon fuscus Radlk., Philipp. J. Sc., Bot. 8 (1914) 461; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 999; Leenh., Blumea 33 (1988) 318. — Type: Ahern 747 (M holo; BO, NY), Philippines. Tree? Twigs terete, 2.5—4 mm thick, blackish. Leaves 2—4-jugate; petiole semiterete, 2.5—5 cm long, c. 2 mm thick; petiolules 3—5 mm long, grooved above; axes glabrous or petiole and Flora Malesiana ser. I, Vol. 11 (3) (1994) rachis puberulous, especially above. Leaflets (sub)opposite, elliptic to somewhat ovate, 6.5—12 by 3-4 cm, index 2—3.5, pergamentaceous; glabrous or beneath on base of midrib with some hairs; base symmetrical, obtuse to acute; margin entire; apex rounded; midrib slightly grooved above; nerves 0.75—1 cm apart, patent, curved, mainly ending free, above inconspicuous, beneath prominulous; inter- secondary nerves variably developed, veins and veinlets finely reticulate, prominulous on both sides. Inflorescences axillary, together pseudoter- minal, widely branched thyrses, up to 10 cm long, puberulous. Flowers unknown. Stigma with 2 spreading lobes. Fruits 2-locular but usually only 1 lobe developed; lobes nearly globular, c. 15 by 12.5 mm, _ fulvous-tomentellous, glabrescent, smooth; wall corky, c. 1.5 mm thick. Seeds: sar- cotesta smooth. Distribution — Malesia: Philippines (Luzon). Habitat & Ecology — “In thickets and forests at low altitude” (Merrill 1923). Fr. Jan. (ripe?). 7. Alectryon glaber (Blume) Radlk., Sapind. Holl.-Ind. (1879) 14; in Engl., Pflanzenr. 98 (1933) 993; Leenh., Blumea 33 (1988) 323, f. 2. — Spanoghea glabra Blume, Rumphia 3 (1847) 174. — Type: Spanoghe 52 (L holo), Lesser Sunda Islands. Alectryon serratus Radlk., Sapind. Holl.-Ind. (1879) 15,48; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 203; Atlas 1 (1913) pl. 126; Radlk. in Engl., Pflanzenr. 98 (1933) 994; Backer & Bakh. f., Fl. Java 2 (1965) 139. — Type: Zollinger 2726 (FI holo; BO, P), Java. Alectryon sphaerococcus Radlk., Sapind. Holl.-Ind. (1879) 15, 49; in Engl., Pflanzenr. 98 (1933) 992. — Type: Beccari it. sec. 2 (FI, n.v.), SE Celebes. Alectryon excisus Radlk., Philipp. J. Sc., Bot. 8 (1914) 460; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 994. — Type Ahern 470 (M holo; BO), Philippines. Alectryon inaequilaterus Radlk., Philipp. J. Sc., Bot. 8 (1914) 460; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 990. — Type: Merrill 5393 (M holo), Philippines. Alectryon ochraceus Radlk., Philipp. J. Sc., Bot. 8 (1914) 460; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 994. — Type: FB (Curran) 17455 (M holo), Philippines. Alectryon celebicus Radlk. [in Engl. & Prantl, Nat. Pflanzenfam., Nachtr. 3 (1907) 205, nom. nud. |] Adema, Leenhouts, Van Welzen — Sapindaceae 457 in Fedde, Rep. 18 (1922) 341; in Engl., Pflan- zenr. 98 (1933) 989. — Type: Koorders 18817 (M holo; BO, L), Celebes. Alectryon ferrugineus auct. non Radlk.: Koord., Minah. (1898) 401. (Shrub or) tree up to 3 m high, dbh c. 30 cm; stem slightly fluted, with rather inconspicuous thick buttresses; bark smooth, pale brown to ash grey. Twigs slightly grooved to terete, up to 5 mm thick, usually early glabrescent. Leaves 2—5(—7)-jugate; petiole semiterete to terete, 1—-8.5 cm long, (0.5— )l1-2 mm thick; petiolules 1-8 mm long, terete to above flattened to hollowed with | or 3 ribs. Leaf- lets opposite (to alternate), ovate to elliptic, 4-14 by 1—-S.5 cm, index 24.5, pergamentaceous to pa- pyraceous; glabrous (to slightly hairy on the mid- rib and beneath on the nerves to, exceptionally, (sub)tomentose); base symmetrical to oblique, in lower leaflets the acroscopic, in upper ones the basiscopic side larger, acute to (broadest side) rounded, somewhat attenuate; margin mainly in the upper half only slightly serrate, dentate, or crenate (or entire); apex rounded to acute; midrib above prominulous; nerves 0.5—2 cm apart, spreading to fairly steep, curved to nearly straight, ending free or in marginal teeth, or looped and joined near the margin, prominulous on both sides (or flat above); intersecondary nerves variably developed, veins and veinlets laxly to minutely reticulate, prominulous on both sides (or beneath indistinct). /nflorescenc- es axillary, racemes, panicles, (or thyrses), up to 9 cm long, simple or with some up to 4 cm long branches, few-flowered, hairy; peduncle 0.75—2.5 cm long; pedicels 1—1.5 mm long. Sepals c. 1 mm high, subconnate, inside hairy. Petals absent. Sta- mens: filaments short; anthers c. 1.2 mm long. Pistil 2- (or 3-)locular with a lobed stigma. Fruits 2-lobed and then often cordate (or only | lobe developed), carinate and shouldered (to slightly grooved), densely to sparsely fulvous (or ferrugineous tomen- tose or tomentellous), the lobes almost globular, 0.9-1 by 0.75—0.9 cm; wall 0.5—1 mm thick. — Fig. oi Distribution — Malesia: E Java, Lesser Sunda Islands (Sumbawa, Flores, Roti), Philippines (Palawan, Mindoro, Luzon, Negros, Sulu Islands, Mindanao), Celebes, Ceram, Kai Islands. Habitat & Ecology — Preferably on coral lime- stone cliffs along the sea shore, also more inland on limestone ridges; from sea level to 1000 m alti- tude. Fl. Feb., Apr., May; fr. Feb.—June, Aug., Sept.— Nov. Uses — Once reported as a good timber. Note — The ‘A. celebicus’ form differs only in being far more hairy. 8. Alectryon myrmecophilus Leenh., Blumea 32 (1987) 223; 33 (1988) 319. — Type: Brass 29489 (L holo), New Guinea. Tree up to 18 m high, dbh up to 30 cm; bark dark brown or grey green, smooth. Twigs deeply grooved, up to c. 8 mm thick, fairly densely to sparsely hairy, glabrescent, branchlets redbrown to black. Leaves 2—4-jugate; petiole semiterete to terete, 6-11 cm long, 2—2.5 mm thick; petiolules 3-10 mm long, above hollowed. Leaflets subop- posite to alternate, elliptic, 7.5—20 by 3.5—7 cm, index 2—3, papyraceous to pergamentaceous; gla- brous or beneath slightly puberulous on midrib and nerves; base symmetrical, acute to rounded, slightly attenuate; margin entire; apex tapering into a rath- er long and slender acute acumen; midrib above slightly raised; nerves 1—1.5 cm apart, spreading, curved, ending free, above slightly sunken, beneath prominent, intersecondary nerves exceptional; veins mainly transverse, veinlets laxly reticulate, rather inconspicuous. /nflorescences axillary, a thyrse, up to 25 cm long, widely branched from the base, branches up to 10 cm long, obliquely patent; flowers in stalked, many-flowered cymules, densely minutely hairy. Sepals c. 0.8 mm high, somewhat less than halfway connate, inside gla- brous or sericeous. Petals 5 (or absent), unguicu- late, the blade kidney-shaped, c. 0.5 by 0.7 mm, outside glabrous, inside woolly. Stamens: filaments short; anthers c. | mm long. Pistil 2- (or 3-)locu- lar. Fruits with mostly only | lobe developed, obovoid-globular, c. 16 by 14 mm, slightly cari- nate close to the style remnant, otherwise smooth, densely ferrugineous-tomentellous; wall corky, c. 2 mm thick. Seeds: sarcotesta smooth. — Fig. 5g. Distribution — Malesia: Papua New Guinea (Morobe Prov.). Habitat & Ecology — On river banks in forest at 650—1000 m altitude, in understorey of dense for- est and in roadside scrub. Fl. June, Dec.; fr. Sept. The hollow branches are inhabited by ants. Notes — |. The present species is close to A. fuscus. 2. The collection Jaheri 445 from the Kai Is- lands (BO) probably belongs to this species. It dif- fers in the following characters: leaflets widest below the middle, the base slightly oblique, mid- rib above slightly sunken, nerves c. 0.75 cm apart, the fruit lobes c. 20 by 15 mm, fulvous tomentel- lous, the wall c. 4 mm thick. 3. The collection NGF 5279 lacks petals. 9. Alectryon repandodentatus Radlk., Bot. Jahrb. 56 (1920) 274; in Engl., Pflanzenr. 98 (1933) 992: S.T. Reynolds, Austrobaileya | (1982) 479, 458 f. 37e; in Fl. Austral. 25 (1985) 31, f. 41-j, map 34; Leenh., Blumea 33 (1988) 325, f. 3. — Type: Loher s.n. (M holo), New Guinea. Treelet, up to 10 m high, dbh up to 20 cm, but often much smaller; bark rather smooth, light grey, often patchy. Twigs terete, 3—5 mm thick, fairly densely short fulvous-hairy; branchlets glabrous. Leaves |—3- (or 4-)jugate; petiole terete, 0.75—4 cm long; petiolules 0-5 mm long, terete. Leaflets: basal pair often strongly falcate, upper pairs ellip- tic to obovate, 1.5—15 by 1-10 cm, basal pair stip- ule-like and much smaller than the others, middle and upper pair either of about the same size or the upper one distinctly bigger, index 1.25—2, fairly thin (to stiff pergamentaceous or + bullate); mid- rib and nerves on both sides densely hairy, in be- tween very sparsely above, beneath rather sparse- ly hairy; base symmetrical to variably oblique, in the basal pair the acroscopic side more strongly developed to in extreme cases the basiscopic side not developed at all, in the upper pair the basiscopic side stronger developed, the base rounded to slight- ly cordate (to obtuse to acute); margin dentate; apex rounded (or obtuse), mucronulate; midrib promi- nent and rounded on both sides; nerves 0.75—1.5 cm apart, + obliquely patent, slightly curved to nearly straight, ending in marginal teeth, above slightly, beneath a bit more prominent; veins and veinlets above rather densely reticulate, beneath laxer. Inflorescences axillary, widely branched panicles, 3-10 cm long: branches up to 7 cm long, spreading, racemoid, densely hairy. Sepals 4, c. 1.5 mm high, somewhat less than half connate, inside sparsely hairy mainly on the lobes. Petals absent. Stamens 6 or 7; filaments c. 2.5 mm long; anthers 1—1.25 mm long. Pistil 2-locular, stigma slightly 2-lobed. Fruits 2-lobed, the lobes globular, 5-7 mm diam., above with a triangular to falcate, up to 7 mm long wing, densely shortly fulvous hairy. — Fig. Sh. Distribution — Malesia: Papua New Guinea (Central Prov., around Port Moresby) and the Mur- ray Islands in Torres Strait. Habitat & Ecology — Mainly in coastal mon- Flora Malesiana ser. I, Vol. 11 (3) (1994) soon scrub, in and along the mangrove, also along the beach and in savannahs, on sandy as well as on rocky hills, from sea level to 150 m altitude. FI. Mar., May, Aug.; fr. Apr., May, Aug., Oct. Note — The present species most resembles A. subdentatus (Benth.) Radlk. f. pseudostipularis Radlk. from SE Queensland. 10. Alectryon reticulatus Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 255; in Engl., Pflanzenr. 98 (1933) 993; Leenh., Blumea 33 (1988) 327, f. 4. — Type: Anonymous s.n. (M holo), New Guinea. Tree to shrub? Twigs angular or terete, 1—2.5 mm thick, subglabrous, glabrescent. Leaves 2- or 3-jugate; petiole semiterete, 2—3.5 cm long; peti- olules (OQ—)3—5 mm long, above grooved with a fine rib. Leaflets alternate, elliptic, 5—9 by 2—3 cm, in- dex 2.5—3, pergamentaceous; the midrib beneath sparsely appressed short-hairy, otherwise glabrous; base symmetrical or in upper leaflets slightly ob- lique, acute, attenuate; margin entire; apex not or slightly acuminate, narrowly rounded to slightly emarginate; midrib above prominulous; nerves 0.5— 1 cm apart, spreading, nearly straight, indistinctly looped and joined towards the margin, prominu- lous on both sides; intersecondary nerves well developed, veins and veinlets laxly reticulate, prominulous on both sides. /nflorescences axillary, a Slender thyrse up to 11 cm long, in fruit still slight- ly hairy; peduncle c. | cm long; branches 1 or 2, patent; cymules (sub)sessile. Flowers unknown. Sepals probably 5, fairly highly connate, outside slightly hairy. Pistil 1-locular. Fruits subglobular, c. 7.5 mm diam., with a small stylar hook laterally halfway, stigma not lobed: wall rugose, subgla- brous, hard, c. 0.5 mm thick. — Fig. 5i. Distribution — Malesia: Islands in the Gulf of Papua and Torres Strait. Note — Closely allied to A. unilobatus S.T. Rey- nolds, which is known from a few localities in Queensland only. Adema, Leenhouts, Van Welzen — Sapindaceae 459 ALLOPHYLUS (P.W. Leenhouts) Allophylus L., Sp. Pl. (1753) 348; Gen. Pl. ed. 5 (1754) 164; Radlk., Sitzungsber. Math.- Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 201; in Engl., Pflanzenr. 98 (1932) 455; Leenh., Blumea 15 (1968) 301. — Type species: Allophylus zeylanicus L. [= Allophylus cobbe (L.) Raeuschel]. Gemella Lour., Fl. Coch. (1760) 648, nom. illeg., non Hill (1761). — Type species: Ge- mella trifolia Lour. [= Allophylus cobbe (L.) Raeuschel]. Schmidelia L., Mant. Pl. 1 (1767) 10, nom. illeg., non Boehm. in Ludwig (1760). — Type species: Schmidelia racemosa L. [= Allophylus cobbe (L.) Raeuschel]. Usubis Burm. f., Fl. Ind. (1768) 89. — Type species: Usubis triphylla Burm. f. [= Allo- phylus cobbe (L.) Raeuschel]. Aporetica Forst. & Forst., Char. Gen. Pl. (1775) 66, t. 66. — Type species: Aporetica ternata Forst. & Forst. [= Allophylus cobbe (L.) Raeuschel]. Ornitrophe Comm. ex Juss., Gen. (1789) 247. — Type species: Ornitrophe borbonica Gmel. [= Allophylus cobbe (L.) Raeuschel]. Shrubs, small trees, or sometimes woody climbers, monoecious or dioecious. Indu- mentum sometimes of stellate-fascicled hairs. Leaves digitate, (1—)3(—5)-foliolate. Inflo- rescences axillary, usually either simple or composed of a few long and slender, racemi- form thyrses, sometimes paniculate. Flowers unisexual, obliquely zygomorphic. Sepals 4, free, imbricate, outer 2 distinctly narrower than the inner ones. Petals 4, nail-shaped to spathulate, inside above the claw with a 2-lobed scale (much smaller than to nearly as big as the blade) which is usually bearded. Disc + interrupted abaxially, mostly lobed or restricted to glands in front of the petals, in female flowers sometimes saucer-shaped. Stamens 8, rarely fewer, all with about the same length, exserted in male flowers; fila- ments mostly sparsely woolly in the lower part. Ovary deeply 2- (or 3-)lobed, 2- (or 3-) celled, 1 ovule per cell; style 1, inserted between the lobes, more or less deeply 2- (or 3-) branched, the branches exceptionally also forked at the apex. Fruits drupaceous, mostly consisting of | mericarp only, globular to obovoid, thin-walled, mostly almost glabrous when ripe. Seeds without aril. — Fig. 6. Distribution — Probably monotypic, though up to 250 species may be accepted. Cir- cumtropical, in some regions penetrating into the Subtropics. See Herzog in Hannig & Winkler, Pfl. Areale 4 (1936) map 32b. Note — Closely related to the Central and South American genus Thouinia Sm., which differs mainly by its winged fruits. (1940) 584, f. 211, 213; Backer & Bakh. f., FI. Java 2 (1965) 133. — Rhus cobbe L., Sp. Pl. (1753) 267; Burm. f., Fl. Ind. (1768) 75. — Allophylus cobbe (L.) Raeuschel, Nomencl. ed. 3 (1779) 108; Hiern in Hook. f., Fl. Br. India 1 (1875) 673; Fern.-Vill., Nov. App. (1880) 51; Vidal, Phan. Cuming. (1885) 104; Kuntze, Rev. Gen. Pl. 1 (1891) 141; Ridley, J. Str. Br. Roy. As. Soc. 33 (1900) 66; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 146; Radlk. in Engl., Pflanzenr. 98 (1932) 594; Corner, Gard. Bull. Str. Settl. 10 (1939) 38; Wayside Trees [Ampacus litorea prima Rumph., Herb. Amb. 2 (1741) 188. — Ampacus litorea (angustifolia) minor Rumph., Herb. Amb. 2 (1741) 189]. See Merr., Int. Rumph. (1917) 336, sub Allophylus ternatus. — Schmidelia cobbe (L.) DC., Prod. 1 (1824) 610, nom. illeg.; Wight, Ic. (1845) t. 460 964/2. — Type: Herb. Hermann vol. 2, fol. 46 (‘Kobbae’) (BM), Sri Lanka. Allophylus zeylanicus L.,Sp.P1.(1753) 348; Hiern in Hook. f., Fl. Br. India | (1875) 673; Fern.- Vill., Nov. App. (1880) 51; Radlk. in Engl., Pflanzenr. 98 (1932) 553. — Type: Herb. Her- mann 140, vol 3, fol. 54 (BM holo), Sri Lanka. Usubis triphylla Burm. f., Fl. Ind. (1768) 89, t. 32, f. 1. — Allophylus triphyllus (Burm.) Merr., Philipp. J. Sc. 19 (1921) 363. — Schmidelia racemosa L., Mant. Pl. 1 (1767) 67, nom. illeg. — Allophylus racemosus (L.) Boerl., Handl. 1 (1890) 284, nom. illeg., non Swartz (1788); Radlk. in Engl., Pflanzenr. 98 (1932) 568; Merr., Comm. Lour. (1935) 246. — Allophylus cobbe (L.) Raeuschel var. racemosus (L.) Ridley, J. Str. Br. Roy. As. Soc. 33 (1900) 66. — Ornitro- phe schmiedelia Pers., Syn. Pl. 1 (1805) 412. — Type: Herb. Linnaeus (Kleinhoff?) (L), Java. Paullinia seriana auct. non L.: Burm. f., Fl. Ind. (1768) 90, p.p. Aporetica ternata Forst. & Forst., Char. Gen. Pl. (1775) 66, t. 66; Blanco, Fl. Filip. (1837) 290. — Schmidelia ternata (Forst. & Forst.) Cambess., Mém. Mus. Nat. Hist. Nat. Paris 18 (1829) 24, nom. illeg. — Allophylus blancoi Blume, Rumphia 3 (1847) 129. — Allophylus ternatus (Forst. & Forst.) Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 313, nom. illeg., non Lour. (1790); in Engl., Pflanzenr. 98 (1932) S72: Allophylus ternatus Lour., Fl. Coch. (1790) 232; Roxb., Hort. Beng. (1814) 88 (‘lanatus’); Rid- ley, Fl. Malay Penins. | (1922) 489. Gemella trifolia Lour., Fl. Coch. (1790) 649. — Aporetica gemella DC., Prod. 1 (1824) 610, nom. illeg.; Blanco, FI. Filip. ed. 2 (1845) 203. Ornitrophe integrifolia Willd., Sp. Pl. 2, 1 (1799) 322; Roxb., Fl. Ind. ed. Carey (1832) 268. — Allophylus integrifolius Blume, Rumphia 3 (1847) 129, nom. illeg. — Type: Commerson 407, Borbonia. Schmidelia timoriensis DC., Prod. 1 (1824) 611, nom. illeg. — Allophylus timoriensis (DC.) Blume, Rumphia 3 (1847) 130; Radlk. in Engl., Pflanzenr. 98 (1932) 587. — Type: Riedlé s.n., Timor. Schmidelia bantamensis Blume, Bijdr. (1825) 231, nom. illeg. — Allophylus fulvinervis (Blume) Blume var. bantamensis (Blume) Blume, Rumphia 3 (1847) 136. — Allophylus cobbe (L.) Raeuschel subf. bantamensis (Blume) Backer, FI. Batavia | (1907) 336. — Type: Anonymous g.n. (L), Java. Schmidelia fulvinervis Blume, Bijdr. (1825) 231, nom. illeg. — Allophylus fulvinervis (Blume) Flora Malesiana ser. I, Vol. 11 (3) (1994) Blume, Rumphia 3 (1847) 133, incl. var. bulla- tus, burmannianus, fuscus, hircinus, montanus, reclinatus, sericeus, velutinus;, Miq., Fl. Ind. Bat. 1, 2 (1859) 576; Ridley, Fl. Malay Penins. | (1922) 489. — Allophylus cobbe (L.) Raeuschel f. fulvinervis (Blume) Backer, Fl. Batavia | (1907). — Type of species: Blume 1550 (L, U), Java. Schmidelia javensis Blume, Bijdr. (1825) 232, nom. illeg. — Allophylus javensis (Blume) Blume, Rumphia 3 (1847) 126, incl. var. cuspidata, racemosa, rigida, robusta, striata; Radlk., Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 231; in Engl., Pflan- zenr. 98 (1932) 578. — Allophylus javensis (Blume) Blume var. genuinus Radlk., Sitzungs- ber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 38 (1909) 239. — Type of species: Blume 1803 (L), Java. Schmidelia littoralis Blume, Bijdr. (1825) 232, nom. illeg. — Allophylus littoralis (Blume) Blume, Rumphia 3 (1847) 124, incl. var. re- panda, serrulata. — Allophylus cobbe (L.) Raeuschel f. /ittoralis (Blume) Backer, FI. Batavia | (1907) 337. — Type of species: Blume 1683 (L), Java. Ornitrophe glabra Roxb. [Hort. Beng. (1814) 28, nom. nud.], Fl. Ind. ed.Carey (1832) 267, nom. illeg. — Schmidelia glabra (Roxb.) Steud., Nomencl. ed. 2, 2 (1841) 531, nom. illeg.; Benth.in Hook. Lond. J. Bot. 2 (1843) 213. — Allophylus glaber Boerl., Handl. 1 (1890) 284, nom. illeg.; Ridley, Fl. Malay Penins. | (1922) 489; Radlk. in Engl., Pflanzenr. 98 (1932) 566. — Allophylus cobbe (L.) Raeuschel var. glabra (Roxb.) Prain, Rec. Bot. Surv. India 1 (1898) 236, non Kuntze (1891). — Type: Roxburgh s.n., India. Ornitrophe repanda Roxb. [Hort. Beng. (1814) 88, nom. nud.], Fl. Ind. ed. Carey (1832) 269. — Type: Roxburgh 2639, Moluccas(?). Ornitrophe villosa Roxb. [Hort. Beng. (1814) 28, nom. nud.], Fl. Ind. ed. Carey (1832) 265. — Allophylus villosus (Roxb.) Blume, Rumphia 3 (1847) 132; Ridley, Fl. Malay Penins. 1 (1922) 490; Radlk. in Engl., Pflanzenr. 98 (1932) 560. — Allophylus cobbe (L.) Raeuschel f. villosus (Roxb.) Hiern in Hook. f., Fl. Br. India 1 (1875) 674. — Type: Roxburgh s.n., India. Aporetica penicellata Blanco, Fl. Filip. (1837) 291. — Neotypes: Merrill Sp. Blanc. 962, 1028, Luzon. Schmidelia macrophylla Zipp. ex Span., Linnaea 15 (1841) 180, nom. nud., non DC. (1824). — Type: Zippelius s.n. (L), Timor. Schmidelia parviflora Zipp. ex Span., Linnaea 15 Adema, Leenhouts, Van Welzen — Sapindaceae 461 Se ______$ (1841) 180, nom. nud. — Type: Spanoghe s.n. (L), Timor. Allophylus amboinensis Blume, Rumphia 3 (1847) 129. — Type: Zippelius s.n. (L), Ambon. Allophylus cambessedei Blume, Rumphia 3 (1847) 129, nom. illeg. Allophylus celebicus Blume, Rumphia 3 (1847) 128. — Type: Forsten s.n. (L, U), Celebes. Allophylus ligustrinus Blume, Rumphia 3 (1847) 126. — Schmidelia ligustrina Blume ex Teijsm. & Binn., Cat. Hort. Bog. (1866) 391, nom. il- leg. — Syntypes: Korthals s.n. (L), Sumatra; Kuhl & van Hasselt s.n. (L), Java. Allophylus rufescens Blume, Rumphia 3 (1847) 137. — Allophylus fulvinervis (Blume) Blume var. rufescens (Blume) Migq., Fl. Ind. Bat. 1, 2 (1859) 576. — Type: Anonymous s.n. (L), Bor- neo. Allophylus rufescens Blume var. cuspidatus Blume, Rumphia 3 (1847) 137. — Type: Korthals s.n. (L), Borneo. Allophylus rufescens Blume var. heterodon Blume, Rumphia 3 (1847) 137. — Type: Korthals s.n. (L), Borneo. Allophylus rugosus Blume, Rumphia 3 (1847) 138. — Type: Anonymous s.n. (L), Java. Allophylus sessilis Blume, Rumphia 3 (1847) 125. — Type: Junghuhn s.n. (L), Java. Allophylus sumatranus Blume, Rumphia 3 (1847) 132, incl. var. crenulatus, elongatus, glabrius- culus, politus; Radlk. in Engl., Pflanzenr. 98 (1932) 586. — Type of species: Anonymous S.n. (L), Sumatra. Schmidelia leptostachya Blume, Rumphia 3 (1847) 141, nom. illeg. — Allophylus leptostachyus (Blume) Boerl., Handl. 1 (1890) 284. — Type: Anonymous s.n. (L), Java. Schmidelia mutabilis Blume, Rumphia 3 (1847) 140, nom. illeg.; Miq., Sumatra (1861) 199, 511. — Allophylus mutabilis (Blume) Boerl.,Handl. 1 (1890) 284. — Syntypes: Anonymous s.n. (L), Blume 1205 (L), both Java. Schmidelia obovata A. Gray, U. S. Expl. Exp. Bot. 1 (1854) 249, nom. illeg.; cf. Merr., Philipp. J. Sc., Bot. 3 (1908) 79. — Type: Cuming 1502 (FI, L, M), Philippines. Schmidelia grossedentata Turcz., Bull. Soc. Nat. Moscou 31 (1858) 401, nom. illeg. — Allophy- lus grossedentatus (Turcz.) Fern.-Vill., Nov. App. (1880) 51; Lecomte in Fl. Indo-Chine 1 (1912) 1013; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 585. — Allophylus cobbe (L.) Raeuschel var. grossedentata (Turcz.) Vidal, Phan. Cu- ming. (1885) 105. — Type: Cuming 640 (FI, L, M), Luzon. Allophylus sundanus Migq., F1. Ind. Bat. 1, 2 (1859) 575, nom. illeg. Schmidelia tomentosa Hook. f., Trans. Linn. Soc. 23 (1860) 164, nom. illeg. — Allophylus tomen- tosus Boerl., Handl. 1 (1890) 284. — Type: Motley 245, Borneo. Schmidelia mutabilis Blume var. subglabra Miq.. Sumatra (1861) 199, 511.— Type: Diepenhorst HB 2330 (BO, L, U), Sumatra. Schmidelia fulvinervis Blume var. macrophylla Teijsm. & Binn., Cat. Hort. Bog. (1866) 391, nom. nud. Allophylus dimorphus Radlk., Sapind. Holl.-Ind. (1879) 17, 56; Merr., Fl. Manila (1912) 304; Enum. Philipp. Flow. Pl. 2 (1923) 494; Radlk. in Engl., Pflanzenr. 98 (1932) 602; Gagnep. in Fl. Indo-Chine, Suppl. 1 (1950) 929. — Type: Lobb 456 (FI, L), Malaya. Allophylus filiger Radlk., Sapind. Holl.-Ind. (1879) 17, 56; in Engl., Pflanzenr. 98 (1932) 583. — Type: Lobb 472 (FI, L), Malaya. Allophylus leptococcus Radlk., Sapind. Holl.-Ind. (1879) 12, 56; in Engl., Pflanzenr. 98 (1932) 581. — Type: Beccari 2828 (FI), Kei Islands. Allophylus cobbe (L.) Raeuschel f. blancoi Fern.- Vill., Nov. App. (1880) 51. — Allophylus cob- be (L.) Raeuschel var. blancoi (Fern.-Vill.) Vidal, Phan. Cuming. (1885) 104. — Type un- known. Allophylus macrostachys Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 56; in Engl., Pflanzenr. 98 (1932) 584. — Syntypes: Cuming 826 (FI, L, M): Warburg 13339, 13340, 13343, all Luzon. Allophylus quinatus Radlk. in Perkins, Fragm. FI. Philipp. 1 (1904) 57; in Engl., Pflanzenr. 98 (1932) 603. — Type: Cuming 1270 (FI, L, M), Luzon. Allophylus setulosus Radlk. in Perkins, Fragm. FI. Philipp. 1 (1904) 68; in Engl., Pflanzenr. 98 (1932) 581. — Type: Warburg 14919, Philip- pines, Jolo. Allophylus micrococcus Radlk. in K. Schum. & Laut., Nachtr. Fl. Schutzgeb. Siidsee (1905) 307; in Engl., Pflanzenr. 98 (1932) 599. — Syntypes: Hellwig 294 (BO); Lauterbach 1385 (SING), 2224 (WRSL), 2340 (WRSL), 2690; Rodatz & Klink 2; all Papua New Guinea. Allophylus unifoliolatus Radlk. in Elmer, Leafl. Philipp. Bot. 1 (1907) 208; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 498; Radlk. in Engl., Pflanzenr. 98 (1932) 555. — Type: Elmer 7329 (M), Leyte. Allophylus apiocarpus Radlk., Sitzungsber. Math.- Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 38 (1909) 227, 238; in Engl., Pflanzenr. 98 (1932) 556. — Type: FB 6120 (M, in protologue 462 Flora Malesiana ser. I, Vol. 11 (3) (1994) erroneously cited as 6420), Philippines, Cebu. Allophylus chlorocarpus Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 232, 239; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 494; Radlk. in Engl., Pflanzenr. 98 (1932) 586. — Type: FB 3473 (M), Palawan. Allophylus insignis Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 234, 240; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 604. — Type: Cuming 1447 (FI), Lu- zon. Allophylus dasythyrsus Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 231, 239; in Engl., Pflanzenr. 98 (1932) 582. — Type: Cuming 1332 (FI, L, M), Luzon. Allophylus hymenocalyx Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 229, 238; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 565. — Type: Copeland 1635 (M), Mindanao. Allophylus largifolius Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 226, 238; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 552. — Type: Merrill 5475 (BO, L, M), Mindanao. Allophylus malvaceus Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 38 (1909) 231, 239; in Engl., Pflanzenr. 98 (1932) 583. — Type: Cuming 102] (FI), Lu- zon. Allophylus leucocladus Radlk., Philipp. J. Sc., Bot. 6 (1911) 181, nom. illeg. — Allophylus leuco- chrous Radlk., Philipp. J. Sc., Bot. 8 (1914) 444; in Engl., Pflanzenr. 98 (1932) 582. — Type: BS 6880 (M), Philippines, Polillo. Allophylus leptocladus Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1602; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 593. — Type: Elmer 11957 (BO, FI, L, M, U), Mindanao. Allophylus repandodentatus Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1603; in Engl., Pflanzenr. 98 (1932) 600. — Type: Elmer 11827 (BO, L, M), Mindanao. Allophylus simplicifolius Radlk. in Elmer, Leaf]. Philipp. Bot. 5 (1913) 1601; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 496; Radlk. in Engl., Pflanzenr. 98 (1932) 558. — Syntypes: Elmer 11138, 13590 (both BO, FI,L,M,U), Mindanao. Allophylus subincisodentatus Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1603; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 497; Radlk. in Engl., Pflanzenr. 98 (1932) 599. — Type: Elmer 11728 (BO, FI, L, M,U), Mindanao. Allophylus brevipetiolatus Radlk., Philipp. J. Sc., Bot. 8 (1914) 449; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 494; Radlk. in Engl., Pflanzenr. 98 (1932) 557. — Type: Vidal 3693 (K), Panay. Allophylus granulatus Radlk., Philipp. J. Sc., Bot. 8 (1914) 451; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 577. — Type: BS 12560 (M), Luzon. Allophylus peduncularis Radlk., Philipp. J. Sc., Bot. 8 (1914) 450; in Engl., Pflanzenr. 98 (1932) 557. — Syntypes: FB 1/266] (PNH, in protologue erroneously cited as /266), Philippines, Mas- bate; Vidal 3716, 3740 (both K), Philippines, Ticao. Allophylus laetevirens Ridley, Trans. Linn. Soc. Bot. 9 (1916) 32; Radlk. in Engl., Pflanzenr. 98 (1932) 604. — Type: Kloss s.n. (SING), Irian Jaya. Allophylus samarensis Merr., Philipp. J. Sc., Bot. 11 (1916) 192; Enum. Philipp. Flow. Pl. 2 (1923) 496; Radlk. in Engl., Pflanzenr. 98 (1932) 553. — Type: FB 24402, Samar. Allophylus scandens Ridley, J. Str. Br. Roy. As. Soc. 75 (1917) 26; Radlk. in Engl., Pflanzenr. 98 (1934) 1489. — Syntypes: Haviland 9&7 (SING); Hose 123 (BM); Ridley s.n. (SING); all Singapore. Allophylus stenophyllus Merr., Philipp. J. Sc. 14 (1919) 417; Radlk. in Engl. Pflanzenr. 98 (1932) 603. — Type: BS 32839 (BM, L, SING), Lu- zon. Allophylus lopezii Merr., Philipp. J. Sc. 26 (1925) 469; Radlk. in Engl., Pflanzenr. 98 (1934) 1490. — Type: BS 41387, Philippines, Busuanga. Allophylus betongensis Craib, Kew Bull. (1926) 359; Fl. Siam. Enum. | (1926) 322; Radlk. in Engl., Pflanzenr. 98 (1934) 1489. — Type: Kerr 7665, Thailand. Allophylus obliquus Radlk. in Merr., Pl. Elm. Born. (1929) 173, nom. nud. — Type: Elmer 21263 (L), Borneo. Allophylus bartlettii Merr., Pap. Mich. Ac. Sc. 23 (1938) 183. — Type: Rahmat 6933, Sumatra. Allophylus cobbe (L.) Raeuschel var. glaber, limo- sus, marinus, velutinus, villosus Corner, Gard. Bull. Str. Settl. 10 (1939) 40-42. Allophylus simplex Quis., Philipp. J. Sc. 76, 3 (1944) 44, nom. illeg., non Baillon (1893). — Type: Merrill 7262, Palawan. This is only a partial list of synonyms, restricted to names used in Malesian context. Adema, Leenhouts, Van Welzen — Sapindaceae 463 Mostly a shrub (sometimes straggling to lian- oid) or treelet, more rarely a tree up to 25 m high, dbh 30 cm. Twigs terete, 3-8 mm in diameter, rather smooth to pustular-lenticellate, either glabrous with the exception of the sparsely appressedly hairy ter- minal bud, or + persistently thinly to densely cov- ered with often stellately tufted, appressed to pat- ent, short, fulvous hairs; the glabrous parts at first shining reddish to blackish brown, later on silvery- to yellowish grey to reddish brown. Leaves mostly 3-foliolate, sometimes part or all of them 1- or 5- foliolate (if 1-foliolate, then lateral leaflets often represented by minute, subulate appendages); pet- iole terete (rarely quadrangular), mostly flattened to grooved above especially at apex and base, 4.5— 20 cm long, 1—3.5 mm thick, indumentum usually like the twig at a slightly earlier stage; petiolules 0-25 mm, the terminal one either of the same length, or up to 4 times as long as the lateral ones, indumentum like that of the petiole. Leaflets ellip- tic or oblong, more rarely lanceolate, laterals of- ten ovate and oblique, terminal sometimes obovate, 2.5—35 by 1.5S—22 cm (terminal one slightly to dis- tinctly larger than the laterals), nearly membranous to fleshy or coriaceous, when dry above greenish to greyish or dark brown and dull to shining, be- neath light green to reddish brown and dull (some- times even glaucous) to shining, glabrous to (es- pecially beneath) densely velvety, moreover often bearded in the axils of the nerves (when glabrous often with a distinct gland) and sometimes also of the veins; base cuneate to rounded, often slightly decurrent, in the terminal leaflet always narrower than in the laterals, in the latter hardly to distinctly oblique; apex + tapering acuminate; margin entire or (mostly sparsely) serrate, crenate, or dentate, sometimes lobed-dentate (mainly in Philippine rac- es); midrib hardly prominent to keeled above, prom- inent beneath, nerves 6—15 per side, slanting, looped and joined near the margin or not, venation usually rather inconspicuous above, hardly conspic- uous to prominent beneath. /nflorescences axillary, solitary or rarely 2 in one axil, simple to thyrsoid, up to 40 cm long, glabrous to densely pubescent, hairs mostly not distinctly stellately fascicled; pe- duncle usually about 1/3—1/5 of the total length, laxly to densely branched, flowers in subsessile to short-stalked, few- to rather many-flowered, um- bel-like dichasia or sometimes in comb-like cincin- ni, in the upper part often solitary; bracts mostly minute, sometimes longer than the pedicels, subu- late, slender. Sepals 1—2.5 by 0.8—2 mm (the inner ones hardly longer, but distinctly broader than the outer), green to whitish, entire to denticulate, most- ly ciliolate, outside glabrous to sparsely appressed short-hairy mainly in the central part. Petals nail- shaped to spathulate, 1—2.2 mm, white, the claw about 2/3 of the entire length, blade entire to bilobed, scale very small to nearly equalling the blade, nearly glabrous to densely woolly along the whole margin of the petal, and especially of the scale, the latter moreover often bearded. Disc in female flowers sometimes saucer-shaped, usually lobed, 0.2—0.8 mm high, glabrous or sometimes puberulous, orange. Stamens c. 8, subequal, in male flowers exserted; filaments mostly sparsely woolly in the lower part; anthers about 0.5 mm. Pistil in female flowers: ovary deeply 2- (or 3-)lobed, lobes obovoid, smooth or rough, glabrous or minutely stellate-fascicled hairy, or sparsely to densely pi- lose by appressed, long, stiff hairs; style 1—-1.5 mm long, glabrous or pilose up to the stigmatic lobes: stigma with 2 (or 3) spreading, fairly long, rarely bifurcate lobes. Fruits mostly with only | meri- carp developed, globular (smaller ones) to obovoid and narrowed at the base (larger ones), 4.5—12.5 by 3.5-8 mm, smooth to slightly wrinkled, red (black to brown when dry), somewhat fleshy and almost glabrous when ripe. — Fig. 6. Distribution — Pantropical, in S America, S Af- rica, Madagascar, and SE Asia slightly penetrat- ing into the Subtropics; throughout Malesia. Habitat & Ecology — Under everwet as well as under seasonal conditions; on sandy beaches and coastal rocks, in and along the Barringtonia for- mation, in brackish as well as freshwater swamps, in open places, shrubberies, and along and in sec- ondary as well as primary forests of all kinds, as well on limestone outcrops as on granitic boulders; from sea level up to 1500(—2000) m altitude. FI., fr. Jan._Dec. The flowers are visited by bees, the fruits are eaten by birds (see, e.g., Docters van Leeuwen, Trop. Natuur 21, 1932, 142, who men- tions Aplonis panayensis strigatus Horsf., the Glossy Starling). For galls see Docters van Leeu- wen, Zoocecidia (1926) 333, f. 601-603. Uses — The wood is reported to be very hard but not very durable; it is mainly used as a timber for temporary structures and indoors, e.g. for raft- ers. Canes and hilts are made from the wood, as well as beaters for the cotton fruits (Mindoro). In the Bismarck Archipelago, poles and branches are used for floats of outrigger canoes and for mark- ing the location of fish traps. Also used as fire- wood. The pulped leaves, or an extraction or decoc- tion of them, as well as a decoction of the roots and the bark, are used in native medicine against stomach-ache and fever. In Perak, a disease inside the mouth of children is cured with it. In Minda- nao, the scraped bark is applied to rigid abdomen, the bark to burns. The berries, though a bit sour, 464 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 6. Allophylus cobbe (L.) Raeuschel. a. Habit; b. male flower; c. petal from inside; d. female flower; e. fruit (a, e: Forbes 2557; b—d: Maxwell 77-375). are eaten. In New Guinea fruits are used as a fish poison. See Burkill, Dict. Econ. Prod. Malay Penins. (1935) 104; Heyne, Nutt. Pl. Indon. ed. 3 (1950) 988. Chromosomes — 2n = 28: Ferrucci, Bolet. Soc. Argentina Bot. 24 (1985) 200-202. Notes — Like so many widely distributed spe- cies with a wide ecological amplitude, A. cobbe comprises a great number of local races. This is even so within Malesia, though the variability is here rather restricted compared with that shown by this species in Africa and South America. Within a restricted region each of these races usually char- acterizes a well-circumscribed habitat. The picture Adema, Leenhouts, Van Welzen — Sapindaceae 465 is simplest in W Malesia, especially in the Malay Peninsula, Sumatra, and Java. But even here most of the races turn out to be not sharply delimited against each other. For the Malay Peninsula, Corner (Gard. Bull. Str. Settl. 10, 1939, 40) distinguished between 5 varieties; he already cited, however, several inter- mediates that broke down their limits. Turning to Malesia as a whole, the picture be- comes far more complicated, the delimitation of infraspecific taxa of more than local importance completely impossible. Thus *A. javensis’ and ‘A. sumatranus’, both inland species from Java resp. Sumatra, represent a pair of well-circumscribed, closely related but in some points distinctly dif- ferent races. ‘Allophylus cobbe var. glaber Corner’ from the Malay Peninsula is intermediate between these two, overlapping both; the characters that consistently distinguish the first two races, are here variable. On the other hand, “A. swmatranus’ en- compasses nearly the whole range of variability of ‘A. cobbe var. glaber Corner’ and ‘A. cobbe var. villosus Corner’, thus breaking down the differ- ences between these two races from the Malay Peninsula. Even the differences between races which in one region seem not to be closely related, characteriz- ing very different habitats, may fade away in other regions. A fine example is given by “A. timorensis’ and ‘A. racemosus’. In Java, these two races are well distinguishable; the former is restricted to the sandy beach and coastal rocks, the latter is an in- land form. ‘A. timorensis’ is distributed from the Malay Peninsula to far in the Pacific, and is rather uniform; only in New Guinea the range of varia- bility is somewhat wider and here it intergrades completely with “A. micrococcus’ , a form of coastal as well as inland habitats. Typical “A. racemosus’ is nearly restricted to Java; towards the East it grades into ‘A. fernatus’ which, in turn, in New Guinea also grades into ‘A. micrococcus’. It is thus impossible to subdivide A. cobbe for the whole of Malesia. On a local scale it may be possible to distinguish between the more impor- tant races, but the distinctions will not hold in stud- ies that encompass a broader geographical range. AMESIODENDRON (P.W. Leenhouts) Amesiodendron Hu, Bull. Fan Mem. Inst. Biol. 7, Bot. (1936) 207; Yap in Tree Fl. Ma- laya 4 (1989) 437. — Type species: Paranephelium chinense Merr. |= Amesiodendron chinense (Merr.) Hu]. Tree, monoecious. Indumentum of solitary simple hairs only. Leaves paripinnate, 3— 6-jugate; leaf axes sparsely minutely puberulous (to glabrous). Leaflets opposite to alter- nate, smooth beneath; the margin hardly to coarsely crenate- to serrate-dentate, especial- ly in the upper half (to entire); domatia absent; nerves ending in the teeth or in the inci- sions, in the former case alternating with intersecondary nerves ending in the incisions or not, in entire parts nerves looped and joined near the margin. /nflorescences terminal and in the upper leaf axils. Flowers unisexual, regular. Sepals 5, + valvate in bud, all equal, not petaloid, hairy on both sides, margin entire. Petals 5, shorter than or about equal to the sepals, not or hardly clawed, glabrous or inside slightly hairy in the upper part, entire, just above the base with a scale varying from somewhat shorter to much longer than the petal itself, woolly on both sides. Disc a ring adnate to the torus, flat except for the margin, with an erect rim to tubular collar up to c. 0.5 mm high, thin-fleshy, glabrous. Stamens (6) 7, 8 (9), distinctly exserted in male flowers; filaments sparsely woolly in the lower half (or for the greater part); anthers glabrous, dehiscence latrorse. Pistil short- stalked or sessile, densely hairy, 3-locular; ovules 1 per locule; style apical, somewhat longer than the ovary, apically with two stigmatic lines (or in fruit, probably only after fertilization, with two minute, hardly spreading lobes). Fruits capsular, hardly stipitate, with up to 3 partly connate globular lobes, loculicidally dehiscent, often + crested along 466 Flora Malesiana ser. I, Vol. 11 (3) (1994) SWessencinan sas Fig. 7. Amesiodendron chinense (Merr.) Hu. a. Habit; b. fruit (a: KEP 97776; b: de Wilde & de Wilde- Duyfjes 19226). Adema, Leenhouts, Van Welzen — Sapindaceae 467 the suture outside, when young slightly coarsely warty, when mature a bit rough to scurfy; wall thick and woody, glabrous on both surfaces. Seeds with a large glabrous sarcotesta around the hilum. — Fig. 7. Distribution — China (Guizhou, Yunnan, Guandong, Guangxi, Hainan), Indo-China (Laos, Annam), and Malesia: Malay Peninsula, Sumatra. Probably monotypic. Habitat — Lowland rain forest. Note — As far as can be judged from the descriptions and figure the species A. in- tegrifoliolatum and A. tienlinensis, both published by Lo, Acta Phytotax. Sin. 17, 2 (1979) 36, f. 3, show no essential differences from A. chinense. Amesiodendron chinense (Merr.) Hu, Bull. Fan Mem. Inst. Biol. 7, Bot. (1936) 209; How & Ho, Acta Phytotax. Sin. 3 (1955) 399; Lo in FI. Hainan. 3 (1974) 88, f. 586; Ming in Fl. Yun- nan. | (1977) 280, t. 66 f. 1 & 2. — Paranephe- lium chinense Merr., Lingnan Sc. J. 14 (1935) 30, f. 10; Gagnep. in Fl. Indo-China, Suppl. | (1950) 971, f. 122: 14-18; Yap in Tree Fl. Ma- laya 4 (1989) 437. — Type: S.K. Lau 13] (P iso), Hainan. Tree up to 25 m high, dbh up to 60 cm, with buttresses up to 1.80 m high, crown dense, widely spreading, bark grey-brown, scaly. 7wigs terete, 3— 8 mm in diam., light greyish to yellowish (or red- dish) brown, glabrous or sometimes minutely pu- berulous, densely warty lenticellate, later becom- ing more greyish and more smooth. Leaves: peti- ole 2-5 cm long, + flattened above; rachis flattened above; petiolules 2-8 mm long, deeply grooved above. Leaflets: lower ones symmetrical, ovate, 4— 15 by 1.254 cm, index 34, middle and upper ones asymmetrical and often slightly falcate, acroscop- ic side wider, elliptic, 8—15 by 24 cm, index 3.5— 4, pergamentaceous to coriaceous (or chartaceous), glabrous (or above on the basal part of the midrib sparsely to fairly densely minutely puberulous); base acute, in the higher leaflets acute at the nar- rower side, rounded and attenuate at the broader side; apex often long-tapering, acute, mucronulate; midrib above a slender rib which may be slightly sunken towards the base, beneath prominent; nerves 5—7.5(—15) mm apart, straight or slightly curved, strongly curved towards the margin, above mostly prominulous, beneath prominent; veins and vein- lets rather reticulate, above often only the stronger veins visible, beneath prominulous and distinct. Inflorescence a widely and laxly branched thyrse, up to 30. cm long, densely puberulous; main branch- es few, erecto-patent, sparsely branching again. Flowers pink (outside Malesia apparently mostly white), in often + opposite, nearly sessile, about 5-flowered cymes, in the lower parts of the branches rather distant, clustered towards the apices; pedi- cels of lateral flowers less than | mm long, of cen- tral flower up to 2.5 mm long; bracts caducous, bracteoles deltoid, minute. Sepals deltoid, c. | mm high. Perals transversely elliptic to orbicular, 0.8- 1.2 by 0.6—1.4 mm; scale subquadrangular to tri- angular or heart-shaped, 0.5—1.2 by 1.2-1.4 mm, erect to bent. Stamens c. 4 mm long; filaments thread-like; anthers suborbicular, 0.33-0.5 mm long, red. Pistil: style c. 4 mm long, slender; loc- ules glabrous inside. Fruits dark brown, usually only | or 2 lobe(s) developed, these globular, 3-— 3.5 cm in diam., connate for c. 2.5 cm; wall 2-5 mm thick. Seeds globular, 2.5—2.75 cm in diam., testa hard, smooth, shining reddish brown, around the hilum with an orbicular to kidney-shaped sar- cotesta 1.5-2.5 by 1.5 cm. — Fig. 7. Distribution — China, Indo-China, Peninsular Thailand, and Malesia: Sumatra (Aceh) and the Malay Peninsula (Perlis, Perak, Pahang, Selangor). Habitat & Ecology — In and along primary rain forest on well-drained soils; up to 600 m altitude. Fl. (Feb.), Apr., May; fr. Jan., Nov. ARYTERA (H. Turner) Arytera Blume, Rumphia 3 (1849) 169; Radlk., Sapind. Holl.-Ind. (1879) 44; Sitzungs- ber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 551; 20 (1890) 267, 293; in Engl., Pflanzenr. 98 (1933) 1268; R.W. Ham, Blumea 23 (1977) 289; S.T. Reynolds, Austrobaileya 2 (1985) 158; in Fl. Austral. 25 (1985) 87, 198. — Lec- 468 Flora Malesiana ser. I, Vol. 11 (3) (1994) totype species (S.T. Reynolds 1985): Arytera xerocarpa (Blume) Adelb. (= Arytera litoralis Blume). Zygolepis Turcz., Bull. Soc. Nat. Moscou 21 (1848) 573. — Type species: Zygolepis rufescens Turcz. Trees or rarely shrubs. Indumentum of solitary, simple hairs; glandular scales always absent in the Malesian species. Twigs terete, smooth to slightly rough, hairy at least when young. Leaves spirally arranged, paripinnate, 1|—6-jugate, without pseudo-stipules, peti- ole (hemi)terete, pulvinate; rachis (hemi)terete; petiolules usually consisting of a pulvi- nus only, usually 1-grooved. Leaflets opposite to subopposite, rarely alternate, symmet- ric, usually not falcate, margin entire to slightly repand, not revolute, both surfaces smooth; domatia usually present in axils of major nerves, rarely completely absent. /nflorescenc- es axillary to pseudoterminal, rarely ramiflorous; bracts and bracteoles triangular, out- side hairy, inside usually glabrous; pedicels hairy. Flowers seemingly bisexual, probably functionally unisexual. Calyx 5-dentate to -partite, teeth triangular to ovate, apert to nar- rowly imbricate, all equal, rarely slightly unequal, margin entire. Petals (2—)5(—6), slightly longer to slightly shorter than the calyx; scales present, rarely completely absent, free or adnate to margin, or enation of margin, not crested. Disc complete, glabrous or hairy. Stamens 8 (6-10); filaments at least basally pilose; anthers pilose or glabrous. Ovary sessile, 2- or 3-locular, smooth, hairy; stigma apical, not lobed, with 2 or 3 stigmatic lines, or 2- or 3-lobed with lobes distinctly recurved in fruit. Ovules | per locule. Fruits with 1-3 well-developed lobes, loculicidal, central axis not thickened transversely, [in A. bullata and A. macrobotrys opening irregularly (loculicidal), central axis thickened trans- versely,| short- or long-stipitate, dissepiments complete, lobes laterally not or slightly flattened; exocarp thick, coriaceous, mesocarp thick, coriaceous to woody, endocarp thin, chartaceous, hairy at least on sutures of carpels (in A. bullata and A. macrobotrys thick, radially striate from the attachment of the seed up to 1/2—2/3 the height of the fruit, not covering the sutures or the fruit axis, sclerenchymatous, detaching from the fruit wall when mature, glabrous). Seeds (partly) covered by an apically open arillode, consisting of 1 or 2 layers, the outer or only one soft, yellow, the inner one firm, dark brown when dry; hilum basal; sclerotesta rather thin, coriaceous, endotesta even thinner, more mem- branous. — Figs. 8, 9. Distribution — One widespread species from NE India throughout SE Asia up to Hainan and eastward across Malesia up to the Solomon Islands; in New Guinea (includ- ing the Bismarck Archipelago) 12 additional species, of which 4 also in Australia, and | other also in the Solomon Islands. Seven additional species in Australia, eight more in the Pacific. Habitat — In various forest types. KEY TO THE SPECIES la. Fruit wall glabrous inside, endocarp thick, sclerenchymatous, radially striate from the attachment of the seed and separating from the pericarp in ripe fruit. Arillode very thick. Calyx and petals punctate, calyx margin slightly membranous. Leaves 3-6-jugate, leaf rachis always glabrous. Inflorescence at least 17 cm long ..... 2 Adema, Leenhouts, Van Welzen — Sapindaceae 469 I b. 3a. 4a. Sa. 6a. Ta. Sa. 9a. 10a. Fruit wall hairy inside, at least on the sutures of the carpels, endocarp thin, charta- ceous, not radially striate, not separating from the pericarp in ripe fruit. Arillode thin or thickened only towards the base. Calyx and petals usually not punctate, calyx margin not membranous. Leaves 1—4-jugate, leaf rachis usually hairy, at least when young. Inflorescence up to 18 cm long............--.--2 2s ese ee eeee 3 . Stipe of fruit S-6 mm long. Arillode not alate. Petioles up to 5.5 cm long; leaflets very coriaceous. Petal claw 0.4—0.6 mm long.................-- 3. A. bullata Stipe of fruit 2-3 mm long. Arillode alate. Petioles longer than 6 cm; leaflets subco- riaceous to chartaceous. Petal claw 0.2-0.3 mm long ....... 6. A. macrobotrys Indumentum of patent, crispate hairs ... 2... 0... eee cece eee eee tees - Indumentum of straight, usually rather appressed hairs..............--.---- 8 Leaves 1- or 2-jugate, rachis (excluding petiole) up to 5.5 cm long; nerves margin- ally indistinctly looped. Calyx teeth (sub)glabrous inside. Flowering twigs 1.5—5 RMI CRTO te AU Sis, AVINASH ee Pe ee 5 Leaves 4-jugate, rachis (excluding petiole) over 13 cm long; nerves marginally dis- tinctly looped. Calyx teeth densely puberulous inside. Flowering twigs more than 5 MI ANS IE 0 hss, CPST, RO SAL, DANONE, Et 10. A. multijuga Bracts triangular, 0.3—1 mm long; pedicels at anthesis longer than 1.5 mm. Calyx deeply incised. Claw of petal usually longer than 0.2mm ...............--- 6 Bracts narrowly triangular, 0.8—1.5 mm long; pedicels at anthesis up to 1.5 mm long. Calyx connate up to 1/3. Claw of petal up to 0.2 mm long . 4. A. densiflora Inflorescence rachis up to 15 cm long. Often several petals more or less reduced, at most hairy on outside and on margin, less than 1 mm long, about as long as the eM aa... SLC Shot toe Pou share. SalI) Dems HOMe te tite Jaa. Cua: Sergiomel <> 7 Inflorescence rachis not longer than 5 cm. Petals all developed, hairy on both sides, more than 1.5 mm long, slightly longer than the calyx ....... 8. A. morobeana Petal blade ovate to suborbicular to obovate; scales free, almost linear, often one reduced, apex usually forked. Apex of leaflets acuminate. 5. A. lineosquamulata Petal blade elliptic to semiorbicular; scales adnate to margin up to halfway, equal in size, apex broadened. Apex of leaflets rounded to slightly acuminate 13. A. pseudofoveolata Leaves (2-) 3- or 4-jugate, rachis (excluding petiole) up to 10.5 cm long; leat- let index up to 5, apex acuminate to caudate. Pedicels elongating up to 10 mm in Prererny oz Alas | ay ae std oe). whoa Ee et ee 9 Leaves all 1- or 2-jugate, rachis (excluding petiole) up to 5(—7.5) cm long; leaf- let index less than 3.6, apex retuse to acuminate. Pedicel elongating up to 7 mm in Beisel St nto canihwueeTiea east: «bis od yeeieteeaeieh ioe eA SeAREEIe pas 10 Domatia large sacs, opening on top. Calyx 0.8—1.1 mm high. Petals never longer than calyx, (sub)pilose inside. Disc pilose. Margin of hypocotyl! pilose 12. A. novaebrittanniae . Domatia often pustular sacs to pockets, rarely pits, opening in front, rarely on top. Calyx 1-1.5 mm high. Petals often slightly longer than calyx, glabrous inside. Disc glabrous, rarely pilose. Margin of hypocotyl glabrous ........... 6. A. litoralis Arillode thick towards base, 2-layered. Leaves 1- or 2-jugate; domatia usually present; nerves basally indistinctly looped, midrib below usually (sub)puberulous, tertiary 470 Flora Malesiana ser. I, Vol. 11 (3) (1994) lla. 13a. venation slightly to distinctly scalariform. Stigma not or very slightly lobe ... 11 Arillode thin, 1-layered. Leaves always 1-jugate; domatia always absent; nerves distinctly looped, midrib below glabrous, tertiary venation reticulate. Stigma dis- finctly dobedsm fruites Jas. wieehh}s.even. Leora. 1. A. bifoliolata Cotyledons (obliquely) above each other. Petals about as long as calyx, petal blades triangular to rhomboid to ovate, scales free. Anthers up to 1.3 mm long. Nerves indistinctly looped’ submareimallys..) s/s, 252-1... 20ae 2 ela. os) eee V2 Cotyledons beside each other. Petals shorter than calyx, scales adnate to margin. Anthers c. 0.6 mm long. Nerves with distinct submarginal loops towards the apex (001 ape ned Sirah a Mere SOMBRE Cen MAbs ri AIRS 3 iis Braet 3 8. A. miniata a. Petals inside (sub)glabrous. Disc usually pilose. Venation above about the same colourias laminanStyle upto 2) mm lone i fruit 22.25. Sasi. 2 6 Soe 13 . Petals inside rather hairy. Disc glabrous. Venation above usually yellowish to red- dish. Style up to 3 mm long in fruit. (Anther curved, connective slightly protruding) 11. A. musca Fruit wall inside completely pilose. Domatia many (sunken) sacs, opening on top. Leaflets broadly oblong-elliptic to -obovate, index up to 2.1; nerves on lower sur- face of leaflets almost flat, up to 14 mm apart; both surfaces of leaflets the same colour whenrdricd A Aye tt aie sec ee BER Beri eee IEE Oe 2. A. brachyphylla Fruit wall inside hairy on sutures only, rarely also on valves, but then always with a glabrous patch in the centre. Domatia many to few (rarely completely absent), often pustular sacs or pockets, opening in front. Leaflets ovate to elliptic, index up to 3.8; nerves on lower surface of leaflets raised, up to 35 mm apart; lower surface of leaflets usually more brownish than upper surface when dried..... 6. A. litoralis 1. Arytera bifoliolata S.T. Reynolds in Fl. Aus- tral. 25 (1985) 198; Austrobaileya 2 (1985) 161. — Type: Hyland 2533 (BRI holo), Australia. Tree or shrub 5—10 m high, dbh c. 15 cm; bark smooth (or flaky), dark brown. /ndumentum short, appressed, straight. Branchlets hairy when young; flowering twigs 1-4 mm thick. Leaves 1-jugate; petiole 0.6—5.5 cm long, hairy when young; peti- olules 2-10 mm long, not to slightly 1-grooved. Leaflets opposite, ovate to elliptic (obovate), 5.3— 18.6 by 1.8-7 cm, index 1.93.6, not to slightly falcate, coriaceous to chartaceous, (punctate), up- per surface glabrous, lower surface glabrous, con- colorous with upper surface; base symmetric, acute to attenuate; apex rounded to slightly acuminate, very apex retuse to rounded (acute), not mucronu- late; venation flat above and below, concolorous with lamina to midrib reddish-brown, midrib raised below, nerves distinctly looped submarginally, veins laxly reticulate, not distinct; domatia absent. Inflorescences axillary to pseudoterminal, branch- ing basally and along rachis; rachis flattened, 2— 9.5 cm long, hairy when young; first-order branches up to 3.5 cm long; cymules pleiochasial, 1—7-flow- ered; bracts 0.5—0.6 mm long, bracteoles 0.2—0.4 mm long; pedicels |—S5 mm long, to 7 mm in fruit, hairy. Flowers 2-3 mm in diameter. Calyx 0.9-2 mm high, deeply incised, outside hairy, inside gla- brous to subpuberulous, teeth equal, not punctate, margin not membranous, apex acute. Petals 5, ovate to rhombic, 1.6—2.5 by 0.5—1.3 mm, not punctate, claw 0.2—0.5 mm long, blade gradually decurrent into claw, margin entire, pilose, apex acute to acu- minate, outside pilose, inside (sub)glabrous; scales 0.5—1.1 mm long, free, basally not auriculate, (apex broadened), rather densely pilose. Disc glabrous. Stamens 7—9; filaments 2.5—4 mm long, basally pi- lose; anthers 0.5—1.1 mm long, straight, pilose, con- nective not protruding. Ovary 2-locular; style and stigma elongating to 0.8—1.5 mm in fruit, in fruit distinctly 2-lobed, upper 0.8—1.5 mm stigmatic. Fruits with | or 2 well-developed lobes, 0.6—1.3 by 0.5—1.7 cm, loculicidal, slightly to distinctly rugose to verrucose, subglabrous, stipe 0.5—2 mm long, broadly cuneate, lobes slightly flattened lat- erally, 6—11 by 4—7.5 mm, dorsally sharply angled to keeled; endocarp thin, chartaceous, pilose on sutures. Seeds ellipsoid to ovoid, flattened lateral- ly, c. 10 by 5 mm; arillode completely covering Adema, Leenhouts, Van Welzen — Sapindaceae 47] seed, lobed, not folded towards the base inside, thin, chartaceous, |-layered. Distribution — Malesia: SE Irian Jaya; Australia: Northern Territories, N Queensland. Habitat & Ecology — Vine forests on lateritic soils, also on dunes, adjacent to mangroves, and along creeks among rain forest trees; altitude from sea level up to 250 m. FI. Apr. (in Irian Jaya), Aug.— Dec.; fr. Nov., Dec. 2. Arytera brachyphylla Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 552; in Engl., Pflanzenr. 98 (1933) 1277. — Cupania brachyphylla F. Muell., Notes Pap. Pl. 6 (1885) 6. — Type: D’Albertis s.n. (FI holo; M), New Guinea. Tree. Indumentum short, appressed, straight. Branchlets hairy when young; fruiting twigs 4—4.5 mm thick. Leaves 2-jugate; petiole 4—5.8 cm long: rachis 2.5-4.1 cm long, terete, glabrous to hairy; petiolules 49.5 mm long, |-grooved. Leaflets op- posite, oblong-elliptic to -obovate, 8.3—15 by 4.5— 8.5 cm, index |.5—2.1, not falcate, thickly charta- ceous, not punctate, upper surface glabrous, lower surface (slightly) hairy on major nerves, colour slightly different from above; base symmetric, acute to almost rounded; apex rounded to shortly acumi- nate, very apex obtuse to rounded, not mucronu- late; venation flat above, midrib slightly raised, concolorous with lamina, venation almost flat be- low, midrib distinctly raised, nerves indistinctly looped submarginally, veins scalariform, lax, not very distinct; domatia (sunken) sacs opening on top. Inflorescences axillary to pseudoterminal, branching basally and along rachis; rachis terete, 6.5-14 cm long, hairy when young; first-order branches up to 7 cm long; bracts and bracteoles not observed; pedicels to 4-6 mm long in fruit, hairy. Flowers not observed. Calyx c. 1.2 mm high, deeply incised, outside hairy, inside glabrous; teeth equal, not punctate, margin not membranous, apex acute. Ovary 2- (or 3-)locular; style and stigma elongating to 2—2.5 mm in fruit, not lobed, in fruit upper 0.7—1.5 mm stigmatic. Fruits with | or 2 well- developed lobes, 0.8—1.3 by 0.9-2.1 cm, loculicidal, smooth to slightly verrucose, slightly hairy, stipe c. 2mm long, slender, lobes not flattened laterally, 7.5-11 by 5.5-9.5 mm, dorsally rounded; endo- carp thin, chartaceous, pilose. Seeds (sub)orbicular to slightly obovoid, not flattened laterally, 6—10 by 6—9 mm; arillode covering 1/2—2/3 of the seed, dentate to slightly lobed, not to slightly folded to- wards the base inside, thick towards base, coria- ceous, 2-layered. Distribution — Malesia: Papua New Guinea, along the Fly River. Habitat & Ecology — Unknown. Note — Known only from the type specimen. 3. Arytera bullata H. Turner, Blumea 38 (1993) 137. — Type: Hartley 12077 (A holo; CANB, K, LAE), New Guinea. Tree, c. 36 m high, dbh c. 90 cm; bark smooth, light grey. Indumentum short, appressed, straight. Branchlets hairy when young; flowering twigs 6 mm thick. Leaves 3—6-jugate; petioles 4—-5.5 cm long; rachis 4.5—9.5 cm long, hemiterete, glabrous; petiolules 6—9 mm long, 2-grooved. Leaflets sub- opposite to alternate, oblong-elliptic, 6.7—10.9 by 3-4 cm, index 2.2—2.9, slightly falcate and bullate, very coriaceous, punctate, upper surface glabrous, lower surface glabrous, colour slightly different from above; base symmetric, slightly attenuate to acute; apex obtuse to slightly acuminate, very apex retuse, not to minutely mucronulate; venation above slightly sunken, midrib raised, concolorous with lamina, venation below raised, nerves marginally distinctly looped, veins laxly reticulate, distinct; domatia large pits to sacs opening on top. /nflores- cences pseudoterminal, branching along rachis; rachis flattened, 17—22.5 cm long, hairy when young; first-order branches up to 10 cm long; cy- mules dichasial, 7—15-flowered; bracts and bracte- oles slightly punctate, bracts 0.3—0.8 mm long, bracteoles 0.2—-0.3 mm long; pedicels 0.6—1 mm long, elongating up to 3 mm in fruit, hairy. Flow- ers 2mm in diameter. Calyx 0.8—1 mm high, deeply incised, outside hairy, inside glabrous, teeth equal, slightly punctate, margin slightly membranaceous, apex acute to obtuse. Petals 5, oblong-elliptic, 0.9— | by 0.7—1 mm, slightly punctate, claw 0.4—0.6 mm long, blade abruptly decurrent into claw, margin entire, pilose at base of blade, apex rounded, out- side glabrous, inside subpuberulous; scales absent or present, 0—-0.2 mm long, free, basally auricu- late, apex not broadened, pilose. Disc slightly lobed, glabrous. Stamens 7 or 8; filaments 2—2.3 mm long, basally pilose; anthers 0.3 mm long, straight, gla- brous, connective not protruding. Ovary 3-locular; style and stigma elongating up to 0.7—1 mm in fruit, 3-lobed, in fruit upper 0.2—0.3 mm stigmatic. Fruits with | or 2 well-developed lobes, 2.7—3 by 2.6— 4.5 cm, opening irregularly, smooth to slightly ru- gose, glabrous, stipe 5—6 mm long, slender, lobes laterally not flattened, c. 24 by 17—18 mm, dorsal- ly rounded; endocarp sclerenchymatous, radially striate from the attachment of the seed, detaching from the fruit wall when mature, glabrous. Seeds orbicular, not flattened laterally, c. 17 by c. 19 mm; arillode not alate, completely covering seed, not 472 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1cm 1mm 1mm , Wesson dore9> 1cm Fig. 8. Arytera Blume. Flower and fruits. — A. bullata Turner. a. Fruit; b. fruit wall from inside. — A. litoralis Blume. c. Female flower; d. petal from inside; e. fruit (a, b: Hartley 12077, c, d: Gibbs 2664, e: van Balgooy 6099). lobed, not folded towards the base inside, very thick especially towards base, fleshy, coriaceous towards the base, 1-layered. — Fig. 8a, b. Distribution — Malesia: Papua New Guinea (E Highlands Prov.). Habitat & Ecology — In oak forest; altitude c. 1500 m. FI. July. 4. Arytera densiflora Radlk., Bot. Jahrb. 56 (1920) 301; in Engl., Pflanzenr. 98 (1933) 1278. — Type: Ledermann 9555 (B holo, probably lost; K, L, M), New Guinea. Tree 2-5 m high. Jndumentum patent, crispate. Branchlets hairy when young; flowering twigs 3-5 mm thick. Leaves 2-jugate; petioles 3—9 (—-18) cm long; rachis 2.5-4.5 cm long, (hemi)terete, densely hairy; petiolules 5-7 mm long, 1-grooved. Leaflets opposite, elliptic, 6.6-20.7 by 4-8.8 cm, index 1.5—3.1, not falcate, chartaceous to slightly coriaceous, (punctate), upper surface glabrous, lower surface hairy especially on major nerves, colour slightly different from above; base slightly asymmetric, basiscopic side broader (symmetric), slightly attenuate to acute; apex acuminate to cus- pidate, very apex retuse to obtuse, not mucronu- late; venation flat above, midrib slightly raised, con- colorous with lamina to slightly reddish brown, venation raised below, nerves marginally indistinct- ly looped, veins scalariform, lax, distinct; domatia pockets to sacs opening in front. /nflorescences axillary to pseudoterminal to ramiflorous, branch- ing basally and along rachis; rachis terete to slightly flattened, 4.5-16 cm long, densely hairy when Adema, Leenhouts, Van Welzen — Sapindaceae young; first-order branches up to 7.5 cm long; cy- mules dichasial to cincinnate, 1—6-flowered; bracts and bracteoles narrowly triangular, bracts 0.8—1.5 mm long, bracteoles 0.5—0.7 mm long; pedicels 0.8—1.5 mm long, hairy. Flowers 1.5—3 mm in di- ameter. Calyx 0.9-1.3 mm high, connate up to 1/3, outside hairy, inside glabrous; teeth equal, not punc- tate, margin not membranous, apex acute to ob- tuse. Petals 5, triangular to rhomboidal to almost orbicular, 0.7—1.4 by 0.5—0.9 mm, not punctate, claw 0.1—0.2 mm long, blade abruptly decurrent into claw, margin entire, pilose, apex rounded to acute, outside subpilose, inside (sub)glabrous: scales 0.2—0.5 mm long, free, basally not to slight- ly auriculate, apex broadened, densely pilose. Disc glabrous to subpilose on rim. Stamens 8; filaments 1—-1.6 mm long, densely pilose; anthers 1.1—1.6 mm long, incurved, densely pilose, connective slightly protruding. Ovary 2-locular; style and stigma not lobed. Fruits not observed. Distribution — Malesia: Papua New Guinea (central mountain range). Habitat & Ecology — Primary forest, on well- drained volcanic soils; 600-850 m altitude. Fl. Oct. Note — Schodde 2438 from Lake Kutubu has very large, sac-like domatia. 5. Arytera lineosquamulata H. Turner, Blumea 38 (1993) 138. — Type: Carr 14969 (L holo; A, BM, K, NY), New Guinea. Tree c. 12 mhigh. /ndumentum patent, crispate. Branchlets hairy when young; flowering twigs 2— 3 mm thick. Leaves 1- or 2-jugate; petioles 2-6 cm long; rachis 1.5—3.5 cm long, (hemi)terete, hairy; petiolules 5-8 mm long, |-grooved. Leaf- lets opposite to subopposite, ovate to elliptic, 6.7— 16.2 by 2.8-6 cm, index 2.2—2.8, not falcate, chart- aceous to coriaceous, not punctate, upper surface glabrous, lower surface glabrous to sparsely hairy, especially on major nerves, colour slightly differ- ent from above; base symmetric, slightly attenu- ate to acute; apex acuminate, very apex obtuse to rounded, not mucronulate; venation flat above, midrib slightly raised, colour reddish, venation raised below, nerves marginally indistinctly looped, veins scalariform, lax, distinct; domatia small pock- ets to sacs opening in front. /nflorescences axil- lary to pseudoterminal to ramiflorous on young branches, branching basally and along rachis; ra- chis terete, 5.5—15 cm long, densely hairy when young; first-order branches up to 8 cm long; cy- mules dichasial to monochasial, 1—4-flowered; bracts 0.3—1 mm long, bracteoles 0.1—0.3 mm long; pedicels 1-2 mm long, densely hairy. Flowers 1.5— 3 mm in diameter. Calyx 0.9-1.2 mm high, deeply 473 incised, outside hairy, inside (sub)glabrous, teeth equal, not punctate, margin not membranous, apex acute to somewhat obtuse. Petals 2-5, often more or less reduced, obovate to ovate to suborbicular, 0.3—1 by 0.2—0.7 mm, not punctate, claw 0.2—0.3 mm long, blade abruptly to gradually decurrent into claw, margin entire, (sub)pilose, apex obtuse to acute, outside (sub)glabrous, inside glabrous; scales almost linear, often one or both reduced, 0.3—0.8 mm long, almost free, (basally auriculate), apex often forked, not broadened, sparsely pilose. Disc pilose on rim. Stamens 7 or 8; filaments in female flower 0.7—1.5 mm long, pilose; anthers in female flower 0.5—0.7 mm long, straight, pilose, connec- tive not protruding. Ovary 2- (or 3-)locular; endo- carp pilose on sutures of lobes; style and stigma not lobed. Fruits not observed. Distribution — Malesia: Papua New Guinea (Central Prov.); Australia: N Queensland. Habitat & Ecology — Secondary and semi-de- ciduous forests; altitude c. 1000 m. Fl. Nov. 6. Arytera litoralis Blume, Rumphia 3 (1849) 170; H. Turner, Blumea 38 (1993) 144. — Eu- phoria xerocarpa Blume, Bijdr. (1825) 234, p.p. (excl. fruits, see note 1), comb. illeg. — Nephe- lium xerocarpum Cambess., Mém. Mus. Nat. Hist. Nat. Paris 18 (1829) 30. — Ratonia lito- ralis Teijsm. & Binn., Cat. Hort. Bogor (1866) 216. — Arytera ochracea Blume ex Koord., Exk. Fl. Java 2 (1912) 542, in syn. — Arytera litoralis f. genuina Radlk. in Gibbs, J. Linn. Soc. Bot. 42 (1914) 65, nom. illeg. — Arytera xero- carpa (Blume) Adeib., Blumea 6 (1948) 324. — Lectotype (Turner 1993): Blume 1314 (L holo), Nusa Kambangan, Indonesia. Zygolepis rufescens Turcz., Bull. Soc. Imp. Nat. Moscou 21 (1848) 709. — Ratonia zygolepis Turez., Bull. Soc. Imp. Nat. Mosc. 36 (1863) 586. — Arytera rufescens Radlk., Sapind. Holl.- Ind. (1879) 44. — Ratonia rufescens Fern.-Vill., Nov. App. (1880) 52. — Aryrera litoralis f. rufescens Radlk. in Gibbs, J. Linn. Soc. Bot. 42 (1914) 65. — Type: Cuming 176] (MW holo, n.v.; A, BM, K, MO, P), Cebu, Philippines. Arytera gigantosperma Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 674. — Type: Beccari s.n. (FI holo; M), Sumatra. Arytera angustifolia Radlk., Sapind. Holl.-Ind. (1879) 44. — Arytera litoralis ft. angustifolia Radlk. in Gibbs, J. Linn. Soc. Bot. 42 (1914) 65. — Type: Teijsmann s.n. (U holo, n.v.; L), Java. Guioa geminata Laut. & K. Schum. in K. Schum. 474 & Laut., Fl. Schutzgeb. Siidsee (1900) 420. — Arytera geminata Radlk. in K. Schum. & Laut., Nachtr. Fl. Schutzgeb. Stidsee (1905) 308. — Type: Lauterbach 2305 (B holo, probably lost; WRSL), New Guinea. Arytera litoralis var. major King, J. As. Soc. Beng. 65, II (1896) 446. — Arytera litoralis f. major Radlk. in Gibbs, J. Linn. Soc. Bot. 42 (1914) 66. — Syntypes: King’s coll. 695, S885, 4456; Ridley 1609, 5995; Scortechini 20, Wray 3163, Perak, Malaya. Nephelium mutabile auct. non Blume: Miq., Fl. Ind. Bat., Suppl. 1 (1860) 198, 508. More complete synonymy in H. Turner (1993). Trees, rarely shrubs, 2-40 m high, dbh 7-91 cm; bark smooth, greyish green to reddish to al- most black. /ndumentum short appressed, straight. Branchlets hairy when young; flowering twigs 1— 7 mm thick. Leaves 1—3(—4)-jugate; petioles 1.3— 9.5 cm long; rachis 0.8—11.5 cm long, (hemi)terete, glabrous to hairy when young; petiolules 2-14 mm long, slightly to distinctly l-grooved. Leaflets (sub)opposite, ovate to elliptic (to obovate), 4.2— 31.1 by 1.4-12 cm, index 1.6—4.5, not falcate, slightly coriaceous to chartaceous, not to densely punctate, upper surface glabrous, lower surface glabrous to hairy especially on major nerves, col- our the same as to (slightly) different from above; base symmetric, rarely oblique, then acroscopic side usually broader, (rounded to) acute to slightly attenuate; apex acuminate to cuspidate (retuse or rounded), very apex retuse to obtuse to rounded, not mucronulate; venation flat above, midrib slight- ly raised, concolorous with lamina to reddish brown or yellowish, venation raised below, nerves mar- ginally indistinctly looped, veins (slightly) scalar- iform to almost reticulate, lax, not distinct; doma- tia often pustular, large to small pockets to sacs (pits) (absent), opening in front (on top). /nflores- cences axillary to pseudoterminal (ramiflorous), branching along rachis (basally or unbranched); rachis terete to slightly flattened, 1.5—17 cm long, hairy when young; first-order branches up to 10 cm long; cymules dichasial (monochasial), 1—7- flowered; bracts 0.3—1.2 mm long, bracteoles 0.1— 0.6 mm long; pedicels 1-5 mm long, hairy. Flow- ers 1—3.5 mm in diameter. Calyx 0.8—2 mm high, deeply incised, outside hairy, inside glabrous, teeth equal, not punctate (punctate), margin not mem- branous, apex acute to acuminate. Petals (2—)5(— 6), triangular to rhomboidal to (ob)ovate, 0.5—2.2 by 0.3-1.9 mm, not punctate, claw 0.1—0.4 mm long, blade usually gradually decurrent into claw, margin entire, (sub)pilose, apex obtuse to acute to acuminate, outside glabrous to pilose, inside Flora Malesiana ser. I, Vol. 11 (3) (1994) (sub)glabrous (to subpilose); scales 0.2—1.2 mm long, free to basally adnate to margin, (slightly auriculate basally), apex broadened (irregular), slightly to densely pilose. Disc glabrous to pilose. Stamens 6—8(—10); filaments 2—4 mm long, pilose; anthers 0.7—1.1 mm long, straight, pilose, connec- tive not protruding. Ovary 2- (or 3-)locular; style and stigma elongating up to 3 mm in fruit, not to slightly 2 (or 3)-lobed, in fruit upper 0.5—2.5 mm stigmatic. Fruits with 1 or 2 (3) well-developed lobes, 0.7—3.6 by 0.5—2.3 cm, loculicidal, smooth to slightly rugose to verrucose, glabrous to slight- ly hairy, stipe 0-3 mm long, slender to broadly cuneate, lobes not to slightly flattened laterally, 8— 23 by 5-21 mm, dorsally rounded to slightly an- gled; endocarp thin, chartaceous, pilose on sutures. Seeds ellipsoidal to orbicular, not to slightly flat- tened laterally, 6-24 by 5-19 mm; arillode cover- ing the seed halfway to completely, dentate to lobed, not to slightly folded towards the base inside, thick towards base, coriaceous, 2-layered. — Fig. 8c—e. Distribution — From NE India (Bay of Bengal) all over SE Asia up to S China (Hainan), through- out Malesia, up to the Solomon Islands. Habitat & Ecology — In primary and secondary forests on all kinds of soils; altitude 0-1500 m. Fl., fr. throughout the year. Uses — For a description of the timber, see p. 427. Notes — 1. For a discussion of the nomenclatu- ral problems regarding A. litoralis, see H. Turner (1993). 2. A variable species, which cannot be divided into smaller entities, because intermediates between different forms can always be found. On the Less- er Sunda Islands east of Lombok and in New Gui- nea the leaves are generally 2-jugate with smaller domatia, and a less scalariform venation; also, the disc is usually rather densely pilose and the lower side of the leaflets often has a denser indumentum on the nerves. Here too, though, more ‘typical’ forms also occur. In Irian Jaya this species is re- ported to form buttresses. 3. Rarely (e.g. NGF 5238, 15418, 29771) the endocarp is almost completely pilose, with a gla- brous patch near the centre of the valves only. These specimens have sometimes been identified as A. brachyphylla, from which they can be distin- guished, however, by their less oblong leaflets with prominent lateral veins on the lower side. 7. Arytera macrobotrys (Merr. & L.M. Perry) R.W. Ham, Blumea 23 (1977) 291; S.T. Rey- nolds in Fl. Austral. 25 (1985) 90; Austrobaileya 2 (1985) 160. — Mischocarpus macrobotrys Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) Adema, Leenhouts, Van Welzen — Sapindaceae 524. — Type: Brass 7618 (A holo; BRI), New Guinea. Tree up to 20 m high, buttressed; bark slightly fissured, lenticellate, brown; sapwood corrugated. Indumentum short, appressed, straight. Branchlets hairy when young; flowering twigs 5—8 mm thick. Leaves 3—6-jugate; petioles 6—10.5 cm long; ra- chis 8.5—32.5 cm long, (hemi)terete, glabrous; pet- iolules 6-12 mm long. Leaflets subopposite to al- ternate, elliptic to slightly obovate, 7.7—18 by 3.3- 5.9 cm, index 2.1—3.4, not falcate, subcoriaceous to chartaceous, densely punctate, upper surface glabrous, lower surface subglabrous, approximately concolorous with upper surface; base symmetric, attenuate to acute; apex acuminate, very apex re- tuse, minutely mucronulate; venation flat above, midrib slightly raised, colour yellowish to the same as that of lamina, venation raised below, nerves marginally distinctly looped, veins laxly reticulate, distinct; domatia sacs opening on top. /nflorescenc- es axillary to pseudoterminal, branching along ra- chis, the latter flattened, 17-40 cm long, hairy when young; first-order branches up to 20 cm long; cy- mules dichasial, 3—7-flowered; bracts and bracte- oles punctate, bracts 0.3-1 mm long, bracteoles 0.1—0.4 mm long; pedicels 1.5—2 mm long, hairy. Flowers 1.5—1.7 mm in diameter. Calyx 0.5—0.9 mm high, connate up to 1/3, outside hairy, inside glabrous, teeth equal, punctate, margin slightly membranous, apex acute to obtuse. Petals 5, ovate, 0.8—1.1 by 0.6—1 mm, punctate, claw 0.2—0.3 mm long, blade gradually decurrent into claw, margin entire to slightly denticulate, pilose, apex rounded to acute, outside glabrous, inside pilose; scales minute to absent, up to 0.4 mm long, adnate to or enation of margin, (basally auriculate), apex not broadened (slightly broadened and forked), sparsely pilose. Disc glabrous. Stamens 7 or 8; filaments 1.8—2.5 mm long, pilose; anthers 0.3—0.4 mm long, straight, glabrous, connective not protruding. Ovary 3-locular; style and stigma elongating up to more than 2 mm in fruit, in fruit 3-lobed, upper c. 0.5 mm stigmatic, papillose. Fruits with | or 2 well- developed lobes, 1.8—3 by 1.8—2.8 cm, loculicidal or Opening irregularly, smooth, glabrous, stipe 2— 3 mm long, slender, lobes laterally not flattened, 1.8—2.8 by 1.8—3 cm, dorsally rounded; endocarp sclerenchymatous, radially striate from the attach- ment of the seed, detaching from the fruit wall when mature, glabrous. Seeds ovoid, slightly flattened laterally, c. 13 by 12 mm; arillode not alate, com- pletely covering seed, not lobed, not folded towards the base inside, very thick, especially towards the base, fleshy to spongy, |-layered. Distribution — Malesia: Papua New Guinea (Western Prov.); Australia: N Queensland. 475 Habitat & Ecology — Substage or canopy tree, common on ridges; altitude 75—80 m. FI. July, Aug.; fr. Oct., Nov. 8. Arytera miniata H. Turner, Blumea 38 (1993) 138. — Type: Carr 11554 (L holo; A, BM, CANB, K), New Guinea. Tree or shrub 2-10 m high, dbh 2.5—12.5 cm; bark slightly suberose, grey to pale brown. /ndu- mentum short, appressed, straight. Branchlets hairy when young; flowering twigs |.5—2 mm thick, fruit- ing twigs 24.5 mm thick. Leaves 1- or 2-jugate; petioles 1-5 cm long; rachis 1.5—3.5 cm long, terete to hemiterete, (with a slight to distinct longitudi- nal ridge), hairy; petiolules 3—9 mm long, not to slightly 1-grooved. Leaflets opposite to suboppo- site, ovate to elliptic, 4-11.6 by 1.9-6 cm, index 1.4—2.5, not falcate, (slightly) coriaceous to some- what chartaceous, not to slightly punctate, upper surface glabrous, lower surface slightly hairy es- pecially on major nerves, slightly browner than above; base symmetric (asymmetric, basiscopic side broader), slightly attenuate to acute; apex re- tuse to slightly acuminate, very apex retuse to rounded, not mucronulate; venation flat above, midrib slightly raised, concolorous with lamina to reddish-yellow, venation raised below, nerves mar- ginally indistinctly looped to more or less distinctly looped apically, veins more or less scalariform, lax, distinct; domatia somewhat pustulate (pockets to) sacs opening in front. /nflorescences axillary to pseudoterminal, branching along rachis; rachis terete, 3-10 cm long, hairy when young; first-or- der branches up to 9 cm long; bracts 0.4—0.8 mm long, bracteoles 0.2—0.3 mm long; pedicels 1.5—4 mm long in fruit, hairy. Flowers not observed. Calyx 0.6-1.5 mm high, deeply incised, outside hairy, inside glabrous, teeth equal, not punctate, margin not membranous, apex acute. Petals 5, obo- vate, c. | by 0.6 mm, not punctate, claw c. 0.3 mm long, blade gradually decurrent into claw, margin entire, pilose, apex obtuse, outside and inside gla- brous; scales c. 0.5 mm long, adnate to margin, basally not auriculate, apex broadened, pilose. Disc probably glabrous. Stamens: filaments in female flower c. 1 mm long, basally pilose; anthers in fe- male flower c. 0.6 mm long, straight, pilose, con- nective not protruding. Ovary 2-locular; style and stigma elongating up to 1-2 mm in fruit, apically minutely 2-lobed, in fruit upper 1—1.5 mm stig- matic. Fruits with | or 2 well-developed lobes, 0.7— 1.3 by 0.7-1.7 cm, loculicidal, slightly rugose to verrucose, slightly hairy, stipe 1-3 mm long, slen- der, lobes not to slightly flattened laterally, 7-10 by 4-7 mm, dorsally rounded; endocarp thin, chart- aceous, pilose along sutures. Seeds ellipsoid to 476 slightly ovoid, not flattened laterally, 8-9 by 5-6 mm; arillode completely covering seed, lobed, not folded towards the base inside, thin to slightly thick- ened towards base, coriaceous, 2-layered. Distribution — Malesia: Papua New Guinea (Central Prov.). Habitat & Ecology — Rain forest and semi-de- ciduous monsoonal forest, also on edge of man- grove, rare; altitude 0-30 m. FI. probably Sept.; fr. Jan.—Mar. 9. Arytera morobeana H. Turner, Blumea 38 (1993) 139. — Type: LAE (Katik & Taho) 74816 (L holo; A, BRI, CANB, LAE), New Guinea. Arytera litoralis auct. non Blume: Hartley et al., Lloydia 36 (1973) 269. Tree c. 6-8 m high, dbh c. 8 cm; bark light grey to brown. Indumentum patent, crispate. Branchlets hairy when young; flowering twigs 1.5—5 mm thick. Leaves 1- or 2-jugate; petioles 2.5—7 cm long; ra- chis 1.8—3.5 cm long, (hemi)terete, hairy; petiolules 4-9 mm long, |-grooved. Leaflets opposite, slightly ovate to elliptic to slightly obovate, 9.3-21.6 by 3.8—7.2 cm, index 2.1—3.2, not falcate, chartaceous, punctate, upper surface glabrous, lower surface subglabrous to hairy especially on major nerves, colour slightly to distinctly different from above; base symmetric to slightly asymmetric, then acro- scopic side broader, slightly attenuate to acute; apex acute to acuminate (slightly retuse), very apex re- tuse to rounded to obtuse, not mucronulate; vena- tion flat above, midrib slightly raised, concolor- ous with lamina, midrib reddish, venation raised below, nerves marginally indistinctly looped, veins scalariform, lax, distinct; domatia pockets to sacs opening in front. Inflorescences axillary to pseu- doterminal, branching basally and along rachis; rachis terete to slightly flattened, 3—S cm long, hairy when young; first-order branches up to 1.5—2 cm long; cymules monochasial, 1- or 2-flowered; bracts 0.3-0.9 mm long, bracteoles 0.2—0.4 mm long; pedicels 1.5—2 mm long, hairy. Flowers 2—2.5 mm in diameter. Calyx 1-1.5 mm high, deeply incised, outside hairy, inside (sub)glabrous, teeth equal, not punctate, margin not membranous, apex acute. Petals 5, elliptic, 1.5—1.8 by 0.8—1.2 mm, not punc- tate, claw 0.3—0.4 mm long, blade gradually de- current into claw, margin entire, pilose, apex ob- tuse to acute, outside rather densely pilose, inside subglabrous to pilose; scales 0.8—1.2 mm long, free, basally not auriculate, apex broadened, rather densely pilose. Disc subglabrous to pilose on rim. Stamens 8 or 9; filaments in female flower 0.8—1.4 mm long, densely pilose; anthers in female flower 1—1.5 mm long, incurved, densely pilose, connec- Flora Malesiana ser. I, Vol. 11 (3) (1994) tive not protruding. Ovary 2-locular, inside pilose on sutures; style and stigma elongating up to 3 mm in developing fruit, minutely 2-lobed, upper 2 mm stigmatic. Fruits not observed. Distribution — Malesia: Papua New Guinea (Morobe Prov.). Habitat & Ecology — Lowland rain forest, alti- tude c. 100 m. Fl. Mar., Apr. 10. Arytera multijuga H. Turner, Blumea 38 (1993) 140. — Type: ANU (Flenley) 2546 (L holo; A, BRI, CANB, K, LAE), New Guinea. Tree c. 8 m high, dbh c. 10 cm; bark brown. Indumentum patent, crispate. Branchlets hairy when young; flowering twigs 5-10 mm thick. Leaves 4- jugate; petioles 9.5-13 cm long; rachis 13.5—-18.5 cm long, terete, slightly 2-grooved, hairy; petiolules 3-10 mm long, not or indistinctly 2-grooved. Leaf- lets subopposite to alternate, elliptic to slightly obovate, 10.6—20.4 by 4.7—7.2 cm, index 2.3—3, not falcate, coriaceous, slightly punctate, upper surface slightly to densely hairy on major nerves, lower surface hairy, especially on major nerves, approx- imately concolorous with upper surface; base asym- metric, basiscopic side broader, acute; apex slightly cuspidate, very apex rounded, not mucronulate; venation flat above, midrib slightly raised, con- colorous with lamina, venation raised below, nerves marginally distinctly looped, veins scalariform, lax, distinct; domatia minute pockets opening in front. Inflorescences axillary, branching along rachis; rachis terete, 4-1 1 cm long, hairy when young; first- order branches up to 7.5 cm long; cymules dicha- sial, 1—3-flowered; bracts 0.7—1 mm long, bracte- oles 0.2—0.5 mm long; pedicels 1.5—3 mm long, hairy. Flowers 2.5—3 mm in diameter. Calyx deep- ly incised, outside hairy, inside densely puberu- lous, teeth slightly unequal, not punctate, margin not membranous, apex obtuse, 2 outer smaller teeth 1.1-1.4 mm high, 3 inner larger ones 1.7—2 mm high. Petals 5, elliptic to ovate, 1.1—1.9 by 0.8—1.2 mm, not punctate, claw c. 0.1 mm long, blade grad- ually decurrent into claw, margin entire (to slight- ly denticulate near apex), pilose, apex obtuse to acute, outside (sub)glabrous, inside subglabrous to subpuberulous; scales 0.4—0.9 mm long, adnate to margin, basally not auriculate, apex broadened, densely pilose. Disc glabrous. Stamens 7 or 8; fil- aments 2-3 mm long, pilose; anthers 1.1—1.4 mm long, straight, glabrous, connective slightly pro- truding. Ovary 3-locular; style and stigma not lobed. Fruits not observed. — Fig. 9. Distribution — Malesia: Papua New Guinea (W Highlands Prov.). Habitat & Ecology — In rain forest, on slope, in deep shade; altitude c. 2200 m. FI. June. Adema, Leenhouts, Van Welzen — Sapindaceae 477 gat! FN ci imm Imm icm Fig. 9. Arytera multijuga Turner. a. Habit; b. male flower; c. petal from inside (a—c: ANU (Flenley) 2846). 478 11. Arytera musca H. Turner, Blumea 38 (1993) 140. — Type: Brass 7620 (L holo; A, BM, BO), New Guinea. Arytera divaricata auct. non F. Muell.: Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 522. Tree 8-15 m high, dbh c. 12.5 cm; bark smooth or flaky, grey or brown. Indumentum short, ap- pressed, straight. Branchlets hairy when young; flowering twigs 3—7 mm thick. Leaves 2-jugate; petioles 2.5—10.5 cm long; rachis 1.5—5 cm long, (hemi)terete to flattened with 2 more or less dis- tinct longitudinal grooves, hairy to glabrescent; petiolules 4-9 mm long, 1-grooved. Leaflets op- posite to subopposite, elliptic to slightly obovate, 4.5-19 by 2.2-8.8 cm, index 1.8—2.9, not falcate, thinly coriaceous to chartaceous, usually not punc- tate, upper surface glabrous, lower surface slight- ly hairy on major nerves, colour slightly different from above; base symmetric to slightly asymmet- ric, then basiscopic side broader, acute to slightly attenuate; apex obtuse to slightly acuminate, very apex retuse to obtuse, not mucronulate; venation flat above, midrib slightly raised, colour yellow- ish to reddish, venation raised below, nerves mar- ginally indistinctly looped, veins weakly scalar- iform, lax, distinct; domatia pockets (to sacs) open- ing in front. Inflorescences axillary to pseudoter- minal, branching along rachis (basally); rachis terete to slightly flattened, 4-12.5 cm long, hairy when young; first-order branches up to 6 cm long; cymules dichasial, 1—7-flowered; bracts 0.3—0.7 mm long, bracteoles 0.2—0.3 mm long; pedicels 1— 4.5 mm long, hairy. Flowers 1.5—2 mm in diame- ter. Calyx 0.7—1.2 mm high, deeply incised, out- side hairy, inside glabrous, teeth equal, not punc- tate, margin not membranaceous, apex (sub)acute. Petals 2-5 (6), elliptic (to orbicular), 0.9-1.3 by 0.4-1 mm, not punctate, claw 0.1—0.3 mm long, blade gradually decurrent into claw, margin entire, pilose, apex obtuse to acute (to slightly acuminate), outside and inside subpilose; scales 0.5—0.8 mm long, free, basally not or slightly auriculate, apex (slightly) broadened, densely pilose. Disc glabrous. Stamens 8; filaments 1.5—2.5 mm long, pilose; an- thers 1.1-1.3 mm long, incurved, pilose, connec- tive slightly protruding. Ovary 2- (or 3-)locular; style and stigma elongating up to 1.5—3 mm in fruit, with a distinct thickening between style and stig- ma, not to slightly 2-lobed, in fruit upper 0.5—1 mm stigmatic. Fruits with | or 2 well-developed lobes, 0.7—1.3 by 0.7—2.6 cm, loculicidal, smooth to slightly rugose, slightly hairy, stipe 0.5—2 mm long, slender, lobes sometimes flattened laterally, 9-15 by 6-10 mm, dorsally rounded; endocarp thin, chartaceous, pilose on sutures. Seeds orbicular, flat- Flora Malesiana ser. I, Vol. 11 (3) (1994) tened laterally, c. 6 by 6 mm; arillode completely covering seed, lobed, slightly folded towards the base inside, thick towards base, coriaceous, 2-lay- ered. Distribution — Malesia: Papua New Guinea (Western Proy.). Habitat & Ecology — Rain forest and monsoon forest; altitude 15—30 m. FI. Sept.; fr. Dec. 12. Arytera novaebrittanniae H. Turner, Blumea 38 (1993) 141. — Type: LAE (Stevens et al.) 58188 (L holo; A, BRI, CANB, E, K, LAE, M, NSW), Papua New Guinea. Tree 7-21 m high, dbh 25-30 cm, not buttressed; bark rugose, somewhat scaly, not to slightly fis- sured, dark grey to brown. /ndumentum short, ap- pressed, straight. Branchlets hairy when young; flowering twigs 2—3 mm thick, fruiting twigs 3.5— 7 mm thick. Leaves 2—4-jugate; petioles 1-9 cm long; rachis 1.5-10.5 cm long, hemiterete, often with a ridge above, glabrous to slightly hairy when young; petiolules 1-7 mm long, not to 1-grooved. Leaflets opposite to subopposite, ovate, 4.4—-17.8 by 1.4-5.9 cm, index (2.3—)3-4.9, not to slightly falcate, coriaceous to chartaceous, (punctate), up- per surface glabrous, lower surface glabrous to slightly hairy on major nerves, colour the same as to slightly different from above; base symmetric, acute; apex acuminate to caudate, very apex round- ed, not mucronulate; venation flat above, colour the same as lamina, raised below, nerves margin- ally indistinctly looped, veins weakly scalariform, lax, distinct; domatia few to many large sacs open- ing on top. Inflorescences axillary to pseudoter- minal, branching along rachis; rachis flattened when young, terete when in fruit, 3-18 cm long, hairy when young; first-order branches up to 3.5— 4 cm long; cymules (irregularly) dichasial, 1—7- flowered; bracts 0.3-1 mm long, bracteoles 0.1— 0.3 mm long; pedicels 3—5 mm long, elongating up to 5-8 mm in fruit, hairy when young. Flowers 2 mm in diameter. Calyx 0.8—1.1 mm high, deeply incised, outside hairy, inside glabrous, teeth equal, not punctate, margin not membranaceous, apex acute. Petals 5, rhomboid to obovoid, 0.6—0.8 by 0.5—0.6 mm, not punctate, claw 0.1—0.3 mm long, blade gradually decurrent into claw, margin entire, pilose, apex obtuse, outside pilose, inside subgla- brous to pilose; scales 0.6—0.7 mm long, free, ba- sally not auriculate, apex broadened, densely pi- lose. Disc pilose. Stamens 7 or 8; filaments 2.2— 2.8 mm long, pilose; anthers 0.8—0.9 mm long, straight, pilose, connective not protruding. Ovary 2- (or 3-)locular; style and stigma elongating up to Adema, Leenhouts, Van Welzen — Sapindaceae 1 mm stigmatic. Fruits with | or 2 well-developed lobes, 1.5—2.2 by 1-2.9 cm, loculicidal, smooth, subglabrous to slightly hairy, stipe 1.5-2.5 mm long, slender, lobes laterally not flattened, 15-19 by 10-12 mm, dorsally angled; endocarp thin, chartaceous to coriaceous, (sub)puberulous on su- tures. Seeds ovoid, not flattened laterally, c. 14 by 9 mm; arillode covering 1/2—3/4 of the seed, lobed, not folded towards the base inside, thick towards base, coriaceous, 2-layered. Distribution — Malesia: Papua New Guinea (W New Britain); Solomon Islands. Habitat & Ecology — Forest, on coral limestone; altitude 125-1200 m. Fl. May; fr. Apr., May. 13. Arytera pseudofoveolata H. Turner, Blumea 38 (1993) 142. — Type: Brass 5560 (A holo; BM, BO, NY, US), New Guinea. Arytera sp.: S.T. Reynolds in Fl. Austral. 25 (1985) 93; Austrobaileya 2 (1985) 165. Arytera foveolata auct. non F. Muell.: Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 523. Small tree. Indumentum patent, crispate. Branchlets hairy when young; flowering twigs 2.5— 3 mm thick. Leaves 2-jugate; petioles 3.8—7.5 long; rachis 1.8—4.8 cm long, (hemi)terete, (with 2 lon- gitudinal grooves), hairy; petiolules 3-10 mm long, 1-grooved. Leaflets opposite, ovate to obovate, 5.4— 17.7 by 2—7.4 cm, index 2—3.2, not falcate, coria- ceous to chartaceous, punctate or not so, upper surface glabrous, lower surface hairy on major nerves, colour different from above; base symmetric to slightly asymmetric, basiscopic (sometimes acro- scopic) side broader, slightly attenuate to acute; 479 apex rounded to slightly acuminate, very apex re- tuse to rounded, not mucronulate; venation flat above, midrib usually slightly raised, concolorous with lamina to reddish or yellowish, venation raised below, nerves marginally indistinctly looped, veins scalariform, lax, distinct; domatia small, few pock- ets to (pustular) sacs opening in front. /nflores- cences axillary to pseudoterminal, branching along rachis or not branching; rachis terete to slightly flattened, 3.5—14 cm long, hairy when young; first- order branches up to 5.5 cm long; cymules dicha- sial, 1—5-flowered; bracts 0.3—0.7 mm long, bracte- oles 0.1—0.2 mm long; pedicels 1.5—3 mm long, hairy. Flowers 1.5—2 mm in diameter. Calyx 0.7— 0.9 mm high, deeply incised, outside hairy, inside glabrous, teeth equal, not punctate, margin not membranous, apex acute to obtuse. Petals 3-5, el- liptic to almost semiorbicular, 0.6—1 by 0.5—0.7 mm, not punctate, claw 0.1—0.4 mm long, blade gradually decurrent into claw, margin entire, sub- glabrous to subpilose, apex obtuse, outside and inside glabrous; scales 0.3-0.7 mm long, adnate to margin up to halfway, basally not auriculate, apex slightly broadened, sparsely pilose. Disc pilose on rim. Stamens 6-8; filaments 3—3.7 mm long, sparse- ly pilose; anthers 0.5—0.6 mm long, straight, sub- glabrous to subpilose, connective not protruding. Ovary 2-locular; style and stigma not observed. Fruits not observed. Distribution — Malesia: Papua New Guinea (Central Prov.); Australia: Cape York area. Habitat & Ecology — Vine forests, on soils de- rived from basic volcanic rocks and basalt; up to 100 m altitude. Fl. Nov. ATALAYA (P.W. Leenhouts) Atalaya |Span., Comp. Bot. Mag. | (1836) 345, nom. nud.] Blume, Rumphia 3 (1847) 186; Radlk. in Engl., Pflanzenr. 98 (1932) 605; S.T. Reynolds, Austrobaileya 1 (1981) 398. — Type species: Atalaya salicifolius (DC.) Blume. Small trees or shrubs, monoecious. Indumentum never stellate-fascicled. Leaves paripinnate (towards the inflorescence sometimes ternate to 1-foliolate), up to 5-jugate; stipules absent; petiole and rachis often partly marginate to sometimes winged. Leaflets entire; petiolules short to very short, swollen. /nflorescences terminal and sometimes in the upper leaf-axils, paniculate. Flowers regular or nearly so. Sepals 5, free, imbricate, margin in Malesian species petaloid, outer 2 smaller. Petals 5 (in one Australian species 4), imbricate, (sub)equal, slightly longer than the calyx, clawed, outside variably pilose, inside above the claw provided with a small, long-hairy scale. Disc annular, complete (in one Australian species interrupted), slightly undulate, hairy or glabrous. Stamens 8, free, 480 Flora Malesiana ser. I, Vol. 11 (3) (1994) all about equal, not exserted; filaments subulate, hairy at least in the lower half; anthers basally attached, oblong, emarginate at base, dehiscence latero-introse. Ovary triangular, 3-celled; style conical, slightly twisted, in Malesian species with 3 stigmatic lines; pistil- lode in male flowers minute or absent. Ovules | per cell, anatropous, apotropous, angu- lar, attached about halfway the cell, sessile on a protuberance of the placenta. Fruits like 3-winged Acer-fruits (often, however, | or 2 cells abortive), breaking up to samarae with a dorsal, straight to curved, oblique wing, glabrous or tomentose. Seeds without aril, rootlet in a pocket. — Figs. 10, 11. Distribution — 12 species: 9 in N and E Australia, of which one also in the Lesser Sunda Islands; 1 in SE New Guinea; 2 species in S Africa. Habitat & Ecology — Tropical and subtropical regions with a periodically dry cli- mate; light forests. Notes — 1. The uppermost parts of the inflorescences are double bostryxes. 2. The genus seems to be closest to the Central American genus Thouinidium Sm. 3. Both Malesian species belong to sect. Atalaya (= Euatalaya Radlk.). KEY TO THE SPECIES la. Fruits densely short-hairy. Leaflets mostly distinctly petiolulate, mostly oblique at the base. Sepals hairy outside. Petals distinctly clawed.......... 1. A. papuana b. Fruits glabrous. Leaflets subsessile, hardly or not oblique at the base. Sepals gla- brous outside. Petals hardly clawed. . . 1. Atalaya papuana (Radlk.) Leenh., Blumea 13 (1965) 126. — Sapindus papuana Radlk. in Engl., Pflanzenr. 98 (1932) 661. — Type: Lister Turner s.n. (BRI, L), 1925?, New Guinea. Guioa eriantha Merr. & L.M. Perry, J. Arnold Ar- bor. 21 (1940) 513; Welzen, Leiden Bot. Series 12 (1989) 304. — Type: Brass 8244 (A, L), New Guinea. Tree up to 23 m high, dbh up to 20 cm. Branch- lets greyish brown, terete, rather densely inconspic- uously lenticellate, young parts densely appressed fulvous short-hairy, early glabrescent. Leaves (1—) 2-3-jugate, glabrous; petiole slender, 2.5-6 cm, flattened, swollen at base; rachis up to 5 cm, to- wards the apex slightly broadened and sometimes marginate. Leaflets subsessile (or petiolules up to 1 cm long, swollen at base), opposite, elliptic, 5.5— 22 by 3-8.5 cm, stiff-chartaceous, brownish- or yellowish- to greyish green when dried, beneath lighter and more dull; base cuneate and attenuate, often distinctly oblique; apex obtuse; midrib above slightly carinate, prominent beneath, nerves c. 12— 14 per side, spreading, straight to slightly curved, only the upper ones looped and joined at the mar- gin, slender, hardly prominent on either surface or prominent beneath, between every two nerves at least 1 intermediate vein + parallel and often near- 2. A. salicifolia ly as strongly developed as these, veinlets prominu- lous on both surfaces. Inflorescences pyramidal thyrsoid, 8-30 cm long, densely short fulvous-hairy, the branches ascending, in bigger inflorescences repeatedly thyrsoid, for the greater part bearing several short-stalked few-flowered cymes; bracts lanceolate, 4 mm long. Flowers reported to be creamy-white. Sepals with a broad, membranous, short-ciliate (partly glandular) margin, outside rath- er densely appressed shorty-hairy, outer obovate (-orbicular), 2.5—3.2 by 2—2.8 mm, inner nearly or- bicular to obovate, 3—4 by 2.3-3 mm. Petals 0.8— 1.5 mm long clawed, ovate to lanceolate, 2.5—3.6 by 1.5—2.5 mm, inside at base incurved-auriculate, outside the claw and at least the basal part of the blade appressed-hairy, inside the claw, the ‘scales’ and at least the basal half of the blade rather densely to sparsely woolly. Disc annular, low and thick, with undulate margin, rather densely short-hairy. Stamens in male flowers 4 mm long, filaments 2— 3 mm long; in female flowers 2.5 mm long, fila- ments 1.5 mm long; anthers | mm long. Pistil 4.5 mm long; ovary 3.5 mm long, densely velvety; style 1 mm long, columnar, twisted, glabrous; stigma truncate, vaguely lobed; in male flowers rather re- duced. Fruits densely appressed short-hairy, sama- rae 4 cm long, wing 1.5 cm wide, slightly curved upwards. — Fig. 10. Adema, Leenhouts, Van Welzen — Sapindaceae 481 | | A m—---7- x Fig. 10. Atalaya papuana (Radlk.) Leenh. a. Habit; b. flower; c. petal from inside; d. petal from outside; e. stamen; f. disc; g. ovary; h. fruit (a: Schodde 2740; b-—g: Schodde 2742; h: Schodde 2807). 482 Flora Malesiana ser. I, Vol. 11 (3) (1994) Lem Fig. 11. Atalaya salicifolia (DC.) Blume. a. Petal from inside; b. fruit (a, b: Kooy 1388). Distribution — Malesia: Papua New Guinea (Western and Central Provinces). Habitat & Ecology — Coastal monsoon dune scrub, savannah forest, stunted pyric swamp for- est, poor primary rain forest; lowland. Fl. June— Sept.; fr. Aug.—Sept. Note — Apparently most closely related to A. hemiglauca (F. Muell.) Benth. from tropical and subtropical Australia. 2. Atalaya salicifolia (DC.) Blume, Rumphia 3 (1847) 186; Radlk. in Engl., Pflanzenr. 98 (1932) 609; Meijer Drees, Comm. For. Res. Inst. 33 (1951) 108; S.T. Reynolds, Austrobaileya 1 (1981) 400, f. 27A. — Sapindus salicifolius DC., Prod. 1 (1824) 608. — Cupania salicifo- lia (DC.) Descaisne, Nouv. Ann. Mus. Hist. Nat. Paris 3, 3 (1834) 443. — Type: Gaudichaud s.n. (?Riedlé) in Herb. DC., Timor. Atalaya bijuga Span., Comp. Bot. Mag. | (1836) 345, nom. nud. Treelet up to 15 m high, dbh up to 40 cm. Branchlets greyish brown, terete, rather densely minutely lenticellate, glabrous except for the sparsely appressed short-hairy tips, furthermore glabrous. Leaves |—3-jugate (according to Blume sometimes ternate or 1-foliolate); petiole slender, 2.5—6 cm, terete at base, flattened and carinate at apex; rachis up to 7.5 cm, in the upper part flat- tened and broadened (to marginate). Leaflets sub- sessile (or with petiolules up to 3 mm long, swol- len at base), opposite or subopposite, (elliptic or) oblong (rarely ovate-oblong) to lanceolate, some- times slightly falcate, 6-16.5 by 1.5—-4 cm, when dried greyish green to brown and shining above, lighter and dull beneath; base cuneate-attenuate; apex obtuse to rounded; midrib above slightly car- inate, beneath more or less prominent, nerves 12— 28 per side, nearly transverse, straight to slightly curved, looped and joined near the margin, slen- der, barely prominent on both sides, between eve- ry two nerves | or 2 intermediate veins nearly as strongly developed as these, veinlets almost as prominent. Inflorescences paniculate, 15—25 cm long; branches long, erect, branched again, bear- ing in their upper half stalked, few-flowered cymes; bracts lanceolate, 1-2 mm long. Sepals with a broad, membranous, short-ciliate margin, outer nearly semi-orbicular, c. 2 mm diam., inner nearly orbicular, c. 3 mm diam. Petals obovate-cuneate, concave, short-clawed, c. 3 by 2.5 mm, outside sparsely hairy near the base, rather long ciliate at the base; scales nearly orbicular to deeply bilobed, c. 0.5 mm long, densely long-hairy. Disc annular, c. 0.5 mm high, thin, glabrous. Stamens in male flowers 3 mm long, filaments 2.5 mm long; in fe- male flowers 2 mm long, filaments 1 mm long; anthers 1 mm long. Pistil 2 mm long; ovary hairy along the upper half of the angles; style slightly shorter, long-conical with thickened stigmatic lines in the upper part; in male flowers strongly reduced to absent. Fruits glabrous, samarae c. 2.5 cm long, wing | cm wide, straight. Seeds subovoid, c. 7.5 by 4 mm. — Fig. 11. Distribution — Malesia: Lesser Sunda Islands Adema, Leenhouts, Van Welzen — Sapindaceae 483 (Sumbawa, Sumba, Timor, and Leti); Australia. Note — The petals are unequal: the two upper Habitat & Ecology — Dryland forest; up to500 —_ ones are very concave and slightly oblique and their (—1000) m altitude. Monsoon climate. Possibly scales are deeply bilobed, the others are slightly preferably on limestone. Fl. Aug., Nov., Dec.; fr. | concave, not oblique, and have nearly orbicular Nov., Dec. scales. CARDIOSPERMUM (P.W. Leenhouts) Cardiospermum L., Sp. Pl. (1753) 366; Gen. Pl. ed. 5 (1754) 171; Radlk. in Engl., Pflan- zenr. 98 (1932) 370-413. — Type species: Cardiospermum halicacabum L. Suffruticose or herbaceous climbers (some American species undershrubs), monoe- cious. Indumentum never stellate-fascicled. Leaves biternate (Malesian species), with minute stipules at the base. Inflorescences axillary, thyrsoid, mostly (some American species excepted) provided with a pair of tendrils. Flowers unisexual, obliquely zygo- morphic. Sepals 5 or 4 (by coalescence of the abaxial two), free, imbricate, outer two smaller. Petals 4 (abaxial one missing), provided with a scale inside slightly above the base which is only slightly smaller and narrower than the petal itself; scales of adaxial petals almost simple, of abaxial ones hood-shaped, bearded, and crested. Disc a gland at the base of every petal, abaxial ones bigger and in some species at one side long-cornic- ulate (sect. Ceratadenia Radlk. to which C. grandiflora belongs). Stamens 8, slightly curved upwards, unequal; in female flowers only slightly reduced. Ovary 3-angled, 3- celled, with a short style and a 3-lobed stigma; ovules 1 per cell, attached basally. Fruits capsular, 3-lobed, inflated, 3-celled, septicidal, papyraceous. Seeds with heart-shaped to orbicular hilum (sometimes described as arillode). — Fig. 12. Distribution — About 12 species, mostly restricted to tropical and subtropical Amer- ica; one species (C. grandiflorum) extending to Africa; C. halicacabum is a worldwide tropical and subtropical weed. See Herzog in Hannig & Winkler, Pfl. Areale 4 (1936) 36, map 32a. Habitat — Apparently light-demanding plants of forest-edges, hedges, shrubberies, savannahs, etc., at low altitudes. Ecology — For pollination see A.G. Hamilton, Rep. 7th Meet. Austral. Assoc. Adv. Sc. (1899) 559-560. The mode of dispersal of the American/African species C. grandiflorum, which is sometimes cultivated as an ornamental in Malesia, is very peculiar, and hardly mentioned in botanical literature. The broadly spindle-shaped, inflated fruits are septicidal and sep- tifragal; the dissepiments split into three elliptic, membranous wings, of nearly the same length and width as the fruit. Each seed is well-affixed to the centre of one of these wings. This is in clear contrast to the dispersal of C. halicacabum, the fruits of which are more strongly inflated — globular or even broader — and which seem to be dispersed as a whole, like little balloons. Morphology — The inflorescence in its most complete form consists of a rather long peduncle with a pair of tendrils (subtended by bracts) slightly below its apex, and a rather short rachis with some pseudo-whorls of 3 lateral branches and a terminal partial inflo- rescence. The latter, as well as the partial inflorescences terminating the lateral branches, 484 Flora Malesiana ser. I, Vol. 11 (3) (1994) is a few-flowered bostryx. Branches and flowers all have small, lanceolate bracts. For the ontogeny of the flower see Payer, Organog. de la Fleur (1857) 149-153, t. 32. For ovule and seed (especially the hilum) see Van der Pijl, Acta Bot. Neerl. 6 (1957) 624-627; Nair & Joseph, J. Ind. Bot. Soc. 39 (1960) 176-194 (n.v.). For embryology see Kadry, Svensk Bot. Tidskr. 40 (1946) 111-126. Notes — 1. The synonym Physalis Norofia, cited by Radlkofer with doubt on author- ity of Hasskarl (Tijd. Nat. Gesch. Phys. 11, 1844, 226) is probably erroneous. Norona, Verh. Bat. Gen. K. W. 5 (1791) 23, cited: “Physalis Halicacabum, Daun capo, nov. cogn.” Apparently, he did not intend to describe a new species, but referred to Physalis hali- cacabum Crantz (Inst. 2, 1766, 370), a name often treated as synonymous with Physalis alkekengi L. (Solanaceae); the Sundanese vernacular daun capo also refers to Physalis. 2. The genus is most closely related to Urvillea Kunth (Central and South America). KEY TO THE SPECIES la. Flowers 2-4 mm long; abaxial disc-lobes not corniculate. Inflorescences mostly consisting of one pseudo-whorl of 3 lateral branches. Fruits 3-lobed, globular, 2-4 em an idiametersaly .{Alse Bi fac) ee C. halicacabum b. Flowers 8-10 mm long; abaxial disc-lobes long-corniculate. Inflorescences with several pseudo-whorls. Fruits 3-lobed, ellipsoid, up to 6.5 by 3.5 cm. Tropical and subtropical America and Africa. Ornamental, sometimes naturalized Cardiospermum halicacabum L., Sp. Pl. (1753) 366; Sims, Bot. Mag. 25 (1807) t. 1049; Wight, Ic. 2 (1841) t. 508; Blume, Rumphia 3 (1847) 185; Griff., Notul. 4 (1854) 546; King, J. As. Soc. Beng. 65, II (1896) 421; Ridley, J. Str. Br. Roy. As. Soc. 45 (1906) 186; Fl. Malay Penins. 1 (1922) 488; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 493; Craib, Fl. Siam. Enum. 1 (1926) 321; Backer, Onkruidfl. Suikerrietgr. (1930) 419; Radlk. in Engl., Pflanzenr. 98 (1932) 379, f. 8A—C; Adelb., Blumea 6 (1948) 322; Hend., Mal. Wild FI. Dicot. (1949) 59, f. 52; Steenis, FI. Sch. Indon. (1949) 251; Gagnep. in FI. Indo- Chine, Suppl. 1 (1950) 933; Hend., Comm. Mal. Wild Fl. (1961) 10, t. 2; Backer & Bakh. f., Fl. Java 2 (1965) 132. — [Halicacabus peregrinus Rumph., Herb. Amb. 6 (1750) 61, t. 24 f. 2.] — Type: Herb. Clifford 151, Cardiospermum I (BM). Cardiospermum corindum L., Sp. Pl. ed. 2 (1762) 526; Radlk. in Engl., Pflanzenr. 98 (1932) 397. — Neotype (Pennington 1993): Houstoun s.n. (BM), Jamaica. Cardiospermum microcarpum H.B.K., Nov. Gen. ed. 4, 5 (1841) 104. — Cardiospermum hali- cacabum L. var. microcarpum (H.B.K.) Blume, C. grandiflorum Swartz Rumphia 3 (1847). — Type: Humboldt & Bon- pland 1184, Venezuela. Cardiospermum luridum Blume, Rumphia 3 (1847) 184. — Cardiospermum halicacabum L. var. lu- ridum (Blume) Adelb., Blumea 6 (1948) 322. — Type: Anonymous s.n. (L), Java. Annual or perennial herbs or subscrubs, often much branched mainly near the base. Stems deep- ly 5-sulcate, slender, glabrous to sparsely hairy. Leaves c. 5-8 by 5-8 cm, sparsely appressed-short- hairy to subglabrous; petiole 1.5—3 cm, slender, grooved; lateral petiolules c. 0.5 cm, terminal one c. 1 cm long, both narrowly winged; stipules lan- ceolate, caducous. Leaflets herbaceous, mostly about 3-partite and pinnately lobed, lobes and apex aristulate. Inflorescences patent, sparsely short- hairy, 5-14 cm long; peduncle 7-10 cm, slender, slightly above the tendrils terminated by a pseu- do-whorl of 3 spreading, bracteate, long-stalked, few-flowered bostryxes (moreover rarely with an ebracteate, terminal bostryx); bracts lanceolate (lower) to elliptic (higher), 1-2 mm. Flowers 2— 3.5 mm long, slender pedicelled. Sepals 4, con- cave, thin, subglabrous, green, tinged red, outer pair broadly ovate to suborbicular, 1-1.5 by 1.2 mm, Adema, Leenhouts, Van Welzen — Sapindaceae 485 0 a ne ee Fig. 12. Cardiospermum halicacabum L. a. Habit; b. stipules; c. fruit; d, e. petals from inside with scales; f. disc; g. ovary; h. seed (a, b, d—g: de Wilde 2765, c, h: de Raadt 54). 486 inner pair suborbicular to broad-elliptic, 2—2.5 by 1.5—2 mm, with white margins. Petals obovate- cuneate to orbicular, rounded and slightly emar- ginate at apex, |.5—2.5 by 1-2 mm, at the base with some woolly hairs at the margin, otherwise gla- brous, white to creamy with yellowish margin; scales glabrous but for the bearded apical part, those of adaxial petals narrow-obovate, somewhat ob- lique, 1.2—2 by nearly 1 mm, rounded at apex, slightly thicker than the petal itself, those of abax- ial petals oblong, hood-shaped, about |.2 mm long with a transverse crest of about 0.2 mm. Disc gla- brous, abaxial lobes not corniculate. Stamens: fil- aments band-shaped, 0.8—2.5 mm, slightly hairy: anthers almost versatile, elliptic, 0.5 mm, yellow. Ovary obovoid, 2—3 mm, nearly glabrous to densely pubescent; style columnar, glabrous or with some hairs; stigma lobes short, thick; in male flowers pistil strongly reduced. Fruits 3-lobed, globular, 1.5—4 cm in diam., mostly sparsely short-hairy, green, reddish at base or with reddish veins. Seeds subglobular, c. 4 mm diam., dull-black, smooth, glabrous; hilum cordate, rather large, white. — Fig. 12. Distribution — Probably of American origin, but now a fairly common weed of the Tropics and Sub- tropics; throughout Malesia. Habitat & Ecology — Under everwet as well as under seasonal conditions, on acid as well as on basic soils, dry to marshy or even periodically flooded with fresh water, preferably in sunny plac- es: waste ground, road-sides, grassland, shrubland, hedges in cultivated areas, also along forest edges and sometimes along sandy beaches. Altitude up to c. 1500 m. The fruits are dispersed by sea, by rivers, over short distances by the wind, but main- ly by man. See Ridley, Dispersal (1930) 73, 268, Flora Malesiana ser. I, Vol. 11 (3) (1994) 491. Fl., fr. Jan—Dec. Uses — Used as a vegetable and in medicine; baskets are made from the stems, the seeds are used as beads. For more details see Burkill, Dict. Econ. Prod. Malay Penins. (1935) 457; Dalziel, Useful Pl. W. Trop. Afr. (1937) 332; Heyne, Nutt. Pl. In- don. ed. 3 (1950) 988. Chromosomes — 2n = 22: Kadry, Svensk Bot. Tidskr. 45 (1951) 414. Notes — 1. Usually the flower developing first in every bostryx is female, all others are male. 2. In Elbert 1985 the sepals are leafy and green, and the two lower ones are not completely con- nate. 3. The scales of the adaxial petals are probably also crested, but the crest is here lateral, turned to- wards the abaxial side of the flower. The suture is inconspicuous in typical C. halicacabum, often more clearly visible and even more or less con- cave (the scale then becoming laterally hood- shaped) in several forms of C. corindum. The scales of the abaxial petals have a central part which is more or less fleshy, and broad membranous wings; their crests are slightly concave. Probably, these scales are coherent in the flower. 4. For the reduction of C. corindum to C. hali- cacabum see Standley & Steyermark, Fieldiana Bot. 24, VI (1949) 240. Cariospermum corindum is a very variable species according to Radlkofer’s monograph where it is subdivided into 14 forms. All Malesian specimens represent typical hali- cacabum, a form which is surprisingly uniform all over the world. There is slight variation in the sizes of flowers and fruits: var. microcarpum re- presenting the relatively small-fruited, var. luridum the big-fruited specimens. In my opinion these varieties do not deserve any taxonomic status. CNESMOCARPON (F. Adema) Cnesmocarpon Adema, Blumea 37 (1993) 195. — Type species: Guioa dasyantha Radlk. [= Cnesmocarpon dasyantha (Radlk.) Adema]. Small or medium-sized trees, twigs, petioles and rachises lenticellate. Indumentum of solitary simple hairs. Twigs terete, striate to grooved. Leaves spirally arranged, paripin- nate, 1—-8-jugate, without pseudo-stipules, neither petiole nor rachis winged. Leaflets al- ternate to opposite, asymmetric, papillate below; petiolules pulvinate; margin entire or remotely dentate; midrib not or slightly prominent above; domatia absent or pocket-like. Inflorescences axillary or rarely ramiflorous, bracts and bracteoles subulate to triangular or ovate, both sides appressed-hairy. Flowers unisexual, regular. Sepals 5, free, imbricate, not petaloid, slightly unequal, both sides appressed-hairy. Petals 5, spathulate, shorter to Adema, Leenhouts, Van Welzen — Sapindaceae 487 longer than the sepals, clawed, with 2 scales or auricles (in C. dentata sometimes absent). Disc complete or interrupted, glabrous. Stamens 8; filaments patently hairy; anthers gla- brous or sparsely hairy. Ovary 3-celled, hairy; style apical with 3 stigmatic lines. Fruits 3- celled, basally 3-winged, loculicidal, outside velutinous and densely covered with irritat- ing hairs, inside tomentose to appressed-hairy. Seeds obovoid, testa shiny black, sarcotes- ta carunculoid; cotyledons unequal, parallel or obliquely superposed. — Figs. 13, 14. Distribution — Four species in Australia and Malesia: Papua New Guinea. Habitat — Primary forest, lowland to montane (up to 2000 m altitude). KEY TO THE SPECIES [ee ecatiets.entire, lower side whitish or-glaucous.....-... 2.2 ee ee eee eee ees 2 Bwreailcissemotely dentate; lower side Sreemiiminir. oe se 2. C. dentata 2a. Indumentum strigose. Upper side of leaflets totally glabrous ................ 3 b. Indumentum puberulous to tomentose. Upper side of leaflets usually with a densely ACD (6 | oo eee 2 ee a ae | eo 3. C. discoloroides 3a. Leaves (3—)4—5-jugate. Leaflets 6-18.5 by 2.5-5.5 cm, nerves 6—12 per side, 7— 20(—28) mm apart. Up to 1000 m altitude................... 1. C. dasyantha b. Leaves 2—3(-4)-jugate. Leaflets 8.5—19 by 3.5—-8 cm, nerves 5—10 per side, 9-30 mm apart. 1600-2000 m altitude..... 1. Cnesmocarpon dasyantha (Radlk.) Adema, Blumea 37 (1993) 197. — Guioa dasyantha Radlk., Bot. Jahrb. 56 (1920) 277; in Engl., Pflanzenr. 98 (1933) 1159; Welzen, Leiden Bot. Series 12 (1989) 304. — Jagera dasyantha (Radlk.) S.T. Reynolds, Austrobaileya 3 (1991) 500. — Type: Ledermann 10365 (L), Papua New Guinea, Sepik area. Jagera discolor L.S. Smith ex S.T. Reynolds, Aus- trobaileya | (1981) 407, f. 28A; in Fl. Austral. 25 (1985) 67. — Type: L.S. Smith 4977 (BRI, L), Australia, Queensland. Trees 5—28 m high, dbh 15—30(—92.5) cm; bark smooth, light grey to red brown, inner bark pink to orange or red brown; wood pale creamy pink to reddish. Twigs striate to grooved, 2-5 mm in diam., (thinly) strigose, soon glabrous. Leaves (3—)4—5- jugate; petiole 1.5—9.5 cm, strongly pulvinate; ra- chis 2.5—19 cm, both flattened above, rounded be- low, striate, strigose to glabrous; petiolules 3-16 mm, flattened above, rounded below, grooved above, strigose to glabrous. Leaflets alternate or opposite, elliptic to ovate, 6—-18.5 by 2.5—5.5 cm, index 1.9-3.4, mid or dark green above, greyish green or glaucous below, thickly chartaceous, above glabrous, below glabrous to thinly strigose, whit- ish when dry; base cuneate to rounded; apex acu- minate, rarely cuspidate; margin entire; midrib not prominent above, nerves 6—12 per side, 7—20(—28) mm apart, angle to midrib 40—60°; domatia absent Pe PO Eo) ee 4. C. montana or small, pocket-like. Inflorescences axillary, 6— 10 cm long, with | to many branches, in fruit 5.5— 15 cm long; bracts and bracteoles subulate to tri- angular or ovate, 0.2—2.5 by 0.1—1 mm; pedicels c. 3.7 mm long, articulated at midpoint, strigose. Flower buds + globular, 1.9-2.5 by 2.2—2.5 mm; flowers white or cream. Sepals (broadly) ovate to triangular, 1.7—3.7 by 1.6—2.5 mm. Petals 2.1—2.4 by 1.2-1.7 mm, claw 0.5—0.9 mm, outside ap- pressed-hairy at the claw, margin ciliate, inside appressed-hairy except apex; auricles woolly. Disc entire. Filaments of staminodes 1.5—2.1 mm; an- thers 0.6 mm, glabrous. Style 1.2—1.4 mm, thinly hairy, stigma 0.2 mm. Fruits subglobular, 3-angled in cross section, 15 by 16 mm, wall at base very thick, thinning upwards, tomentose inside, but ap- pressed-hairy by the seeds. Seeds 9 by 5 mm; cot- yledons obliquely superposed. — Fig. 13. Distribution — Malesia: Papua New Guinea (Sepik, W New Britain, Central and Western Prov.): Australia: N Queensland, between Mt Lewis and Mt Fox. Habitat & Ecology — Primary forest, altitude 400-1000 m. Fl. Mar., Sept.; fr. Mar. 2. Cnesmocarpon dentata Adema, Blumea 37 (1993) 197. — Type: Jacobs 9524 (BISH, BO, L), Papua New Guinea, Mt Bosavi. Trees 6-26 m high; bark moderately smooth, patchy light and dark grey and brown, blaze thin, 488 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1cm 1com Fig. 13. Cnesmocarpon dasyantha (Radlk.) Adema. a. Habit; ». detail of lower surface of leaflet (a, b: NGF 21918). Adema, Leenhouts, Van Welzen — Sapindaceae 489 Fig. 14. Cnesmocarpon Adema. Leaflets, fruits and seeds. — C. dentata Adema. a. Leaflet lower surface; b. fruit; c. seed. — C. discoloroides Adema. d. Leaflet lower surface; e. fruit; f. seed. — C. montana Adema. g. Leaflet lower surface (a—c: Jacobs 9524; d-f: LAE 58053; g: Carr 13384). red brown. 7wigs striate, 2—S mm in diam., short tomentose, glabrescent. Leaves 1—3-jugate; peti- ole 1.5—5 cm, + pulvinate, flattened above, round- ed below; rachis (O—)1.5—12 cm, about terete, both striate, short tomentose; petiolules 3-8 mm, flat- tened above, rounded below, grooved, short tomen- tose. Leaflets opposite to alternate, + elliptic, 4.5— 17.5 by 2.5-8 cm, index 1.8—2.8, green below, chartaceous, above and below almost glabrous, midrib densely, nerves thinly puberulous; margin dentate; midrib slightly prominent above, nerves 7-12 per side, mostly ending in a tooth, 5—20 mm apart, angle to midrib 45—75°; domatia absent. /n- florescences axillary, 3-8 cm long, with or with- out branches, in fruit S—13.5 cm long; bracts and bracteoles triangular, 0.3—1.9 by 0.2—1.2 mm; pedi- cels c. 3 mm long, articulated at about midpoint, short tomentose. Sepals triangular to deltoid, 1.7— 2.5 by 1-1.8 mm. Petals 1.2—3 by 0.6—1.9 mm, claw 0.4—0.6 mm, outside appressed-hairy up to half- way, margin ciliate, inside appressed-hairy in lower half; appendages absent, or either 2 scales or auri- cles, woolly; corolla white. Disc interrupted. Fila- ments of staminodes |.9—2.2 mm; anthers 0.6 mm, glabrous or thinly hairy. Style 2-3 mm, glabrous or thinly hairy, stigma 0.5—0.9 mm. Fruits about globular, basally 3-winged, glossy bright red to orange, succulent, very sour, 22 by 20 mm, wall c. 6 mm thick, succulent, + appressed hairy inside. Seeds black with yellow aril, 11 by 5 mm; cotyle- dons parallel. — Fig. 14a—c. Distribution — Malesia: Papua New Guinea (W & S Highlands Prov.). Habitat & Ecology — Primary forest, altitude 10-700 m. Volcanic soil. Fl., fr. Oct. 490 Flora Malesiana ser. I, Vol. 11 (3) (1994) 3. Cnesmocarpon discoloroides Adema, Blumea 37 (1993) 199. — Type: LAE 58053 (P.F- Ste- vens) (A, BISH, L), Papua New Guinea, Mt Shungol. Trees 5-16 m high, dbh 5—25 cm; bark (green- ish) grey to brown or black, slightly cracked, inner surface straw to dark (reddish) brown, under bark red or greenish, inner bark red or orange; wood white to orange brown. Twigs striate to grooved, 2—5(—10) mm in diam., shortly tomentose, glabres- cent. Leaves (2—)4—5(-8)-jugate; petiole 3-11 cm, strongly pulvinate, flattened above, rounded below, towards the apex terete; rachis (2.5—)9.5—30.5 cm, about terete, both striate, shortly tomentose, gla- brescent; petiolules 2-15 mm, flattened above, rounded below, grooved above, shortly tomentose, glabrescent. Leaflets alternate to opposite, elliptic to ovate, rarely obovate, mid or dark green above, pale green, greyish or glaucous below, 6—22 by 2.5— 8.5 cm, index |.7—3.2(—4.2), thickly chartaceous, above glabrous to thinly puberulous, midrib more densely so, below almost glabrous to thinly puber- ulous, midrib and nerves more densely so, + whit- ish when dry; base cuneate to rounded; apex short- or long-acuminate; margin entire, exceptionally un- dulate; midrib not prominent above, nerves 8-16 per side, 5-20 mm apart, angle to midrib 45-65°; domatia inconspicuous, pocket-like. /nflorescenc- es axillary or ramiflorous, c. 1 cm long, branched or not, in fruit 3-17 cm long; bracts and bracteoles subulate to deltoid, 0.2—1.2 by 0.1—1.1 mm; pedi- cels 1.5 mm long, articulated 1/4 up their length, shortly tomentose. Flower buds about globular, yellow green, 1.7 by 1.9 mm; flowers only known from buds and from remains below the young fruits, white. Sepals + elliptic, inner ones apically with scarious rims, 1.7—3.1 by 1.5—2.2 mm, ciliate. Pet- als 3 by 1.2 mm, claw 1.2 mm, outside appressed- hairy at the claw, ciliate at the base of the blade, inside appressed-hairy except apex; auricles woolly. Disc entire. Filaments of staminodes 2.2-2.5 mm; anthers 0.5—0.6 mm, glabrous. Style 1-1.9 mm, glabrous, stigma 0.4-0.7 mm. Young fruits red, 3- angled in cross section, tomentose inside. — Fig. 14d-f. Distribution — Malesia: Papua New Guinea (W New Britain, Morobe, and Milne Bay Prov.). Habitat & Ecology — Montane or hill forest, 0— 1300 m altitude. Stony ground. Fl. May, Oct.; fr. Jan.—May, Oct.—Dec. 4. Cnesmocarpon montana Adema, Blumea 37 (1993) 199. — Type: Carr 14154 (BM, G, L, SING), Papua New Guinea, Alola. Trees 8-10 m high. Twigs grooved, 2—3 mm in diam., strigose. Leaves 2—3(—4)-jugate; petiole 4.5— 7 cm, strongly pulvinate, flattened above, rounded below, rarely towards the apex terete; rachis 4.5— 12 mm, flattened above, rounded below, rarely terete, both striate, (thinly) strigose; petiolules 5— 10 mm, flattened above, rounded below, grooved above, strigose. Leaflets alternate to opposite, el- liptic to ovate, 8.5-19 by 3.5—8 cm, index 1.8—3, coriaceous, above glabrous, below rather densely strigose, whitish when dry; base cuneate to round- ed; apex shortly and obtusely acuminate; margin entire; midrib not prominent above, nerves 5—10 per side, 9-30 mm apart, angle to midrib 40-45°; domatia absent. Inflorescences axillary, 16-19 cm long; branches in fruit at least 7 cm long; bracts and bracteoles + triangular, 0.4 by 0.4 mm; pedi- cels 4 mm long, articulated at midpoint, strigose. Flower buds brownish, flattened globular, 2 by 2— 2.5 mm; flowers green or white. Young fruits red- dish, tomentose inside. — Fig. 14g. Distribution — Malesia: Papua New Guinea (Central Prov.). Habitat & Ecology — Forest, altitude 1600-2000 m. Fl. Sept.—Nov.; fr. Jan. CUBILIA (P.W. Leenhouts) Cubilia Blume, Rumphia 3 (1849) 100; Radlk. in Engl., Pflanzenr. 98 (1932) 921-924; Leenh., Blumea 24 (1978) 297, 298. — Type species: Cubilia blancoi Blume [= Cu- bilia cubili (Blanco) Adelb.]. Trees medium-sized, monoecious. Jndumentum of solitary, simple hairs; no glandular scales. Leaves spirally arranged, paripinnate, 3—7-jugate, without pseudo-stipules, gla- brous, neither petiole nor rachis winged. Leaflets opposite to alternate; base equal-sided to oblique, in the latter case broadest at the acroscopic side; margin entire; nervation Adema, Leenhouts, Van Welzen — Sapindaceae 49] mainly open but somewhat irregular. /nflorescences terminal or pseudo-terminal, cymules many-flowered, bracts inconspicuous. Flowers unisexual, regular. Calyx urceolate, the narrow mouth surrounded by 5 minute lobes, densely tomentellous on both surfaces, not petaloid. Petals 5, included in the calyx, hardly clawed, sericeous on both sides, the mar- gin sometimes slightly inflexed just above the base but without a clear scale. Disc annu- lar, broad, adnate to the receptacle, glabrous. Stamens 5 (or 6), hardly exserted; filaments flattened, glabrous or with a few hairs; anthers adnate, glabrous, dehiscing introrse. Pistil sessile, 2-carpellate, divided to near the base, warty and densely hairy; stigma sessile, inserted between the lobes, 2-lobed Ovules 1 per cell, attached at the base. Fruits 2- parted, the parts obovoid, loculicidal, glabrous, densely aculeate, pericarp coriaceous, glabrous inside. Seeds attached basally, up to about halfway enveloped by a thin-fleshy + entire arillode, hilum large, nearly orbicular. — Figs. 15, 16. Distribution — Monotypic. Cubilia cubili (Blanco) Adelb., Blumea 6 (1948) 325; Backer & Bakh. f., Fl. Java 2 (1965) 143; Leenh., Blumea 24 (1978) 397. — Cubilia blan- coi Blume, Rumphia 3 (1847) 101, nom. illeg.; Merr., Int. Rumph. (1917) 338; Sp. Blanc. (1918) 240; Enum. Philipp. Flow. Pl. 2 (1923) 505; Radlk. in Engl., Pflanzenr. 98 (1932) 923, f. 22; Madulid, Nat. Mus. Papers 2, 1 (1991) 56. — Euphoria cubili Blanco, Fl. Filip. (1837) 287, nom. illeg.; ed. 2 (1845) 200; ed. 3, 2 (1878) 10. — Neotype (Leenhouts 1978): Merrill Sp. Blanc. 705 (BO, L), Luzon. Cubilia rumphii Blume, Rumphia 3 (1847) 100; Koord., Minah. (1898) 402; Ic. Bogor. 1, 4 (1901) 51, t. 92, 93; Backer, Schoolfl. (1911) 268; Koord., Suppl. Cel. 2 (1922) t. 52, 53, 3 (1922) 26. — Boa Massy Rumph., Herb. Amb. Auct. (1755) 5, t. 3. — Type: Rumphius, Herb. Amb. Auct. (1755) 5, t. 3. Trees up to 25(—50) m high, dbh up to 75 cm, with up to 6 m high buttresses; bark usually smooth and reddish (sometimes greyish) brown. Branch- lets terete, 4-8 mm in diam., redbrown to cinna- mon or purplish black, smooth to slightly grooved, lenticels inconspicuous, the tip appressed short ful- vous hairy, otherwise glabrous. Leaves often with 2 buds, mostly only the lower one developing; pet- iole terete or sometimes slightly flattened above, smooth or (thicker ones) canaliculate, strongly swollen at base, 5-25 cm long, 1.5—3 mm thick; rachis slightly flattened above, otherwise terete, sometimes carinate above towards the apex; peti- Olules terete, above with a narrow groove, 3-7.5 mm long, mostly slender. Leaflets elliptic (to ovate), (S—)10-15(—40) by (2.5—)3-5(—10) cm, index 2-4, the upper often, sometimes all slightly falcate, chartaceous, often with a naked gland in or in front of some of the nerve axils beneath; base rounded and attenuate to acute; apex (obtuse or) tapering (rarely rather abruptly) short to rather long broad- ly (rarely slender) obtuse- to acute-acuminate, mucronate or not; midrib slender, above prominu- lous to sunken, beneath mostly sharp-triangular, sometimes + rounded, nerves 1—2(—4) cm apart, angle to midrib 55—75°, slightly, towards the mar- gin strongly curved, mostly free but for the upper- most ones, sometimes a few looped and joined to- wards the margin, prominulous above, more so beneath, intercalary veins variable, mostly feeble, veins and veinlets reticulate, above mostly faint, beneath prominulous. /nflorescences thyrsoid to co- rymbiform, up to 30 cm long, densely and minute- ly appressed brown hairy; branches sparsely branched; cymules with stalks to 1 cm long and up to c. 25-flowered; bracts triangular and hardly 0.5 mm to subulate and 3 mm; pedicels slender, 3-4 mm long, slightly thickening towards the apex. Flowers copper. Calyx in male flowers c. 3 mm high, 2 mm in diam., the opening c. | mm in diam., lobes minute; in female flowers 1.5—2 mm high, 3—4 mm in diam., the opening wider than in male, lobes broad-triangular, c. 0.8 mm high. Petals in male flowers elliptic, c. 1 by 0.6 mm, in female subovate, 1.5 by 1.5 mm. Stamens: filaments c. | by 0.2 mm; anthers c. 0.3 mm; stamens in female flowers hardly reduced. Pistil c. 1.5 mm high; stig- ma-lobes triangular, dorso-ventrally flattened, c. 1 mm long. Fruit parts 3-4(—5) by 2—2.5 cm, green to brown, the warts pyramidal to triangular, up to 2 mm long, pericarp c. 0.8 mm thick, endocarp smooth, white. Seeds ellipsoid-ovoid, 2.5 by 1.7 cm, testa smooth, shining dark brown, arillode to 6—10 mm high. — Figs. 15, 16. Distribution — Malesia: eastern half of Borneo (incl. P. Laut), Philippines (Mindoro, Luzon, Sa- mar, Biliran, Mindanao), Celebes, W Moluccas. Outside its area rarely cultivated in Java. Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 15 . Cubilia cubili (Blanco) Adelb. Habit (FB 1996). Adema, Leenhouts, Van Welzen — Sapindaceae 493 Fig. 16. Cubilia cubili (Blanco) Adelb. a. Male flower; b. ibid., longitudinal section; c. female flower; e. petal; f. fruit; g. open fruit with seed; h. longitudinal section of seed (a-e: PNH 22872; f: NIFS Cel. V/236 = Waturandang 221; g: after Radlkofer (1931-1933), fig. 22; h: Kostermans 7011). Habitat & Ecology — Primary and secondary for- ests, apparently mostly on poorly aerated, often basic soils, often along rivers; mostly at low alti- tudes, up to 600 m (Koorders 22616, Celebes, Mi- nahassa, according to the label from 1700 m). FI. Jan.—Dec.; fr. probably also Jan.—Dec. Uses — The whitish to light reddish brown tim- ber is used for indoor construction. The arillode is edible; the cooked or roasted seeds are eaten and are said to be comparable with but more delicious than Castanea seeds. See Brown, Useful PI. Philipp. 2 (1950) 357, f. 174; Heyne, Nutt. Pl. Indon. ed. 3 (1950) 1002; Kraemer, Trees West. Pacific Reg. (1951) 212, f. 75. Note — Though the species as a whole is rather uniform it shows clinal variation in a few charac- ters. In Borneo the leaves are 3—5-jugate, the glands on the lower side of the leaflets are uncommon, and the apex of the leaflets is obtuse or obtuse- acuminate; in the Philippines the leaves are some- times 6-jugate, glands are always present, the apex of the leaflets is mostly acute-acuminate; in Celebes the leaves are 5- or 6-, sometimes 7-jugate, glands are always present (though rare in some specimens), the apex of the leaflets is sometimes acute-acumi- nate. The few trees grown in Java represent the Celebes form. CUPANIOPSIS (F. Adema) Cupaniopsis Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 483, 498, 584; 20 (1890) 291, 357; Bot. Jahrb. 56 (1920) 283; in Engl., Pflanzenr. 98 (1933) 1177; S.T. Reynolds, Austrobaileya 2 (1984) 44; in Fl. Austral. 494 Flora Malesiana ser. I, Vol. 11 (3) (1994) 25 (1985) 55; A.C. Smith, Fl. Vitiensis Nova 3 (1985) 603. — Cupaniopsis sect. Elat- topetalum Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 584; in Engl., Pflanzenr. 98 (1933) 1183. — Lectotype species (S.T. Rey- nolds 1984): Cupania anacardioides A. Rich. [= Cupaniopsis anacardioides (A. Rich.) Radlk. ]. Cupaniopsis sect. Macropetalum Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 357; in Engl., Pflanzenr. 98 (1933) 1182. — Type species: Cupaniopsis macropetala Radlk. Shrubs or small to medium sized, often unbranched palmoid trees (“Schopfbaume’ ). Twigs terete, smooth to striate or grooved. /ndumentum of simple solitary hairs, some- times mixed with red clavate glands, in many Pacific species mainly consisting of scale hairs. Leaves spirally arranged, paripinnate, 1—28-jugate; pseudo-stipules absent; petiole pulvinate; petiolules pulvinate, usually distinct. Leaflets opposite to alternate, lower ones usually smaller than the upper ones, thinly papyraceous to coriaceous, often punctate; margin entire to serrate or dentate, rarely lobed; upper surface often darker and shinier than the lower, lower surface rarely papillate; domatia absent or present. /nflorescences thyrsoid, axillary, often together pseudoterminal, branched or not, laxly to densely flow- ered; cymules dichasial, 1- to several-flowered; bracts and bracteoles acicular to deltoid, usually not persisting. Flower buds (flattened) globose to obovoid; flowers regular, uni- sexual, rarely bisexual, male flowers with a minute pistillode, female ones with rather large staminodes, otherwise not much different. Sepals (4 or) 5(—7), outer 2 distinctly smaller than inner 3, free, imbricate, often concave, especially the inner sepals with a narrow to wide petaloid rim to almost totally petaloid, usually persisting, margin usualy ciliate and often also with small glandular hairs. Petals 5, elliptic to orbicular or spathu- late, rhomboid or obovate, rarely clawed, often dentate at apex; scales | or 2, rarely crested, free or up to 3/4 connate with the margin of the petals, rarely auricles instead of scales. Disc entire, lobed. Stamens (5—)8—14, usually exserted in male flowers; filaments patently hairy, usually up to halfway, rarely glabrous; anthers shorter to longer than the filaments, glabrous to rather densely hairy. Ovary 2- or 3-locular, smooth, glabrous to densely hirsute, sessile or with a short gynophore; style apical, shorter than the ovary, with 2 or 3 stigmatic lines, rarely stigma lobed. Ovules 1 per locule;. Fruit a loculicidal capsule, rarely distinctly lobed, (1-) 2- or 3-celled, sessile or with a short stipe, usually rounded in cross section, rarely keeled; pericarp coriaceous to woody, thin to rather thick, outside smooth to rugose, glabrous to hirsute, inside glabrous to tomentose or appressed- hairy. Seeds ellipsoid or globose to obovoid, often dorso-ventrally, rarely laterally flat- tened, arillode half to completely covering the seed, rarely (C. platycarpa) a sarcotesta, hilum oval, (sub)basal; sclerotesta woody, thin, usually black, endotesta membranous, brownish. — Figs. 17-19. Distribution — 60 species; in Malesia: Celebes, Moluccas and New Guinea; N and E Australia, Caroline Islands (Truk Tol), Pacific from the Solomon Islands to Samoa and New Caledonia. Habitat & Ecology — Secondary or primary forests, often in forest margins, road- and riversides, on floodplains and beaches. Rather indifferent to soil type. Altitude: sea level to lower montane zones. Usually rare. Adema, Leenhouts, Van Welzen — Sapindaceae 495 8a. 9a. 10a. lla. KEY TO THE SPECIES . At least young parts with red glands. Lower side of leaflets sericeous. Pistillode, auary dapairuits 2-celled; stisma with 2 Lines m.ceu wsteeind oes stets & Hi oteieileys « 2 . Red glands absent. Lower side of leaflets glabrous to puberulous, if appressed hairy then not sericeous. Pistillode, ovary and fruits 3-celled; stigma with 3 lines.... 4 . Axial parts villose to tomentose. Petals glabrous outside, with | scale. Stamens Pee Ne WoC Ie at <1) en. are ide 6 ite «cies ete eae, cate tl le noel ipa Pe 3 . Axial parts strigose. Petals hairy outside, with 2 scales. Stamens not exserted. Celebes 5. C. celebica a. Leaflets usually with small, pocket-like domatia. Petalar scales crested. Fruits el- lipsoid, lenticular or obcordate, 4-8 by 4-6 cm, stipe c. 10 mm long. Seeds 3.5—4 ee MEM rahe SAL COLES tay ss OS. Res oe eee ee ee Smee 11. C. platycarpa . Leaflets without domatia. Petalar scales not crested. Fruits transversely ellipsoid to obreniform, 2 by 3 cm, stipe | mm long. Seeds | by 0.5 cm, with an arillode 3. C. bilocularis PPC S COR SIANTINOUCS)KOl a. ake e: ; =; Shetek ah let ee ee. een) ee 5 motancAs (Or stamimlodes) 10-14: oe... SS PP Eee. 12. C. rhytidocarpa fees CRUTS srarely, OOSCULELY Chenate sy: cen a eee 19 > Weatlets' below puberulous‘omihe veins: 2 >... eee ele tegen 14. G. hospita . Leaflets below glabrous, sericeous or hirsute, without small red glandular hairs 20 . Leaflets below hirsute with small red glandular hairs besides simple hairs 13. G. hirsuta aikeatlets: below papillate:: 24.0505 ou. cht. ae sn. 2 21 Ugleeatlets below SmOOtl) 2298 * . o.isce sce thelr s ait cl tistd 2 0 esp. 2 eee oe rr 4] . Disc uninterrupted (if in fruit check basal ring around several fruits; Fig. 35d) 22 Discsmectupted (Fig. 35C)e oso ace oi. ee ease Siete oo ps] . Leaflets below glabrous to sericeous to hirsute, domatia absent to many; margin entire. Petal with folded auricles or scales (Fig. 36a—d); crest absent to present (Fig. 36h-k). Fruit 0.7—1.6 cm high, stipe slender to broadly obconical (Fig. 37) ... 23 . Leaflets below hirsute, domatia many; margin usually with a few subapical teeth. Petal with scales (Fig. 36a—c); crest absent (Fig. 36h). Fruit 0.6—0.8 cm high, stipe Slender (Rigs Sila)ree. aie tge ne hee come are ete ieee 25. G. oligotricha Petals with scales (Fig. 36a—c). Stipe of fruit (unknown in G. multijuga) slender to broadly obconical (Fig. 37). Cotyledons (unknown in G. multijuga) subcollateral (ESS Sb) Ne eae es ek Sea S| ee SIM pcttacee a fae eee keen 24 . Petals with folded auricles (Fig. 36d). Stipe of fruit broadly obconical (Fig. 37c). Cotyledonsisuperposed(Figy3Sa)ays &. wives Jota ae ake eho 39. G. subsericea Branchlets (not leaves or inflorescences) glabrous to sericeous. Leaflets symmetri- cal to basally asymmetrical, domatia absent to many, if many then apex of petal sealesi(very much broadenedi(Figs36a)* 2. APN lih ah. chs 1 Ee ee 25 . Branchlets hirsute (to sericeous). Leaflets asymmetrical, especially base and apex, domatia many. Apex of petals scales not to hardly broadened (Fig. 36b) 30. G. pleuropteris Leaflets 2.3-24.3 cm long, lower surface glabrous (to slightly sericeous), domatia absent to many; very apex mucronulate or not. Petals 0.5—4 mm high and scales with a very broadened apex (Fig. 36a) or petals 0.3—1.8 mm high and apex of scales not broadened (Fise36b)yistsde she he Ba eA ee tre Be ae 26 . Leaflets 3.8-6.3 cm long, lower surface very sparsely subsericeous, at least a few leaflets with a single domatium, very apex mucronulate. Petals 1.5—1.7 mm long, apex of scalesmot.broadened: (Figg 56b)1... 38 «ag ds teaeean-e ie 21. G. multijuga Adema, Leenhouts, Van Welzen — Sapindaceae 553 26a. Leaflets usually not mucronulate; domatia (absent to) many. Petals 0.5—4 mm long; apex of scale very broadened (Fig. 36a); crest seldom present, then part of pilose bifid scale apex (Fig. 361i, j). Fruit with a 2-5 mm high stipe . 10. G. diplopetala b. Leaflets usually mucronulate; domatia in at least some leaflets | (or 2). Petals 0.3- 1.8 mm long; apex of scales not broadened (Fig. 36b); crest seldom present, then clavate, glabrous (Fig. 36k). Fruit stipe 0-3 mm lon........... 2. G. acutifolia 27a. Leaflets without domatia or with small pockets or sacs (Fig. 34a, b)......... 28 b. Leaflets with one highly domed sac (Fig. 34d) ......... 24. G. novobritannica 28a. Leaflets symmetrical to asymmetrical/falcate, venation usually raised only on low- er surface, concolorous with lamina above, laxly to densely reticulate ....... 29 b. Leaflets falcate, venation conspicuously raised on both sides, discolorous with lamina mice censely TeHeCulate:, . weelly sexu ricas Pee AMSA Le 36. G. reticulata 29a. Leaflets ovate to elliptic to obovate, if obovate: branchlets at most sericeous, doma- tia absent to many. Petal blade obovate to circular, gradually or sharply decurrent into claw (Fig. 36e, f); crest absent to present, pilose part of bifid scale apex, or menmeus:andiclavate (Brg <3 6bsk)iveis). Anos cl. ect eas atu i. oper bei 30 b. Lower leaflets often ovate, upper ones (elliptic to) obovate, domatia many. Branch- lets hirsute (to sericeous). Petal blade obovate, gradually decurrent into claw (Fig. 36e); crest usually present, pilose part of bifid scale apex (Fig. 36i, j) 30. G. pleuropteris METS HAUIN UN CLALE «IE, 50 5y'eiocof ovorenord Bet4s) Wav wlar adndn Wacc-ay scl AEA Gy SE 31 PP omicepHnctatec: sv ish ser. Uotahiine. 20ea./assau see iG) Cueatisele. AAT oe 39 3la. Leaflets ovate (to elliptic), usually falcate, coriaceous to very coriaceous, margin flat (to revolute), apex (obtuse to) acuminate to caudate, usually mucronulate; domatia 2 DOE NOM 0a op RC A ae Ree Ee Res RE VO” eee GMP Ue eS b. Leaflets elliptic (to obovate), not falcate, very coriaceous, margin revolute, apex rounded to acuminate, not mucronulate; domatia absent or a single sac 26. G. palawanica Een aly SUDMESHIC. Pais PIADROUS, 21.525 Oona Olga deta ee coef tees ee OS b. Ovary densely hirsute. Fruits sericeous, glabrescent ......... 35. G. pubescens 33a. Claw of petal 0.4—-1.3 mm long; crest (absent to) pilose part of bifid scale apex, or subglabrous and clavate (Fig. 361-k). Fruit with 2-5.5 mm long, slender stipe (Fig. 37a; unknown of G. discolor: leaflets strongly bicoloured) ................ 34 b. Claw of petal c. 0.5 mm long; crest glabrous, clavate (Fig. 36k). Fruit with ac. 2 mm long, broadly obconical stipe (Fig. 37c). Leaflets bicoloured 31. G. plurinervis 34a. Lower surface of leaflets somewhat differently coloured from upper surface, gla- brous to subsericeous (to hirsute), domatia absent to many . 15. G. koelreuteria b. Lower surface of leaflets very differently coloured from upper surface, sericeous, domahaabsentensinelene: «2 ao). P9en o odd te 11. G. discolor 35a. Leaflets ovate to obovate. Inflorescence subglabrous to usually shortly sericeous (to hirsute), hairs brown. Blade of petals elliptic to obovate, gradually decurrent into claw (Fig. 36e); those of G. parvifoliola unknown, this species with a subseri- CEQUSARTOLESCENCE) cals annals stgestom treaty agus Selita tt stachide ts alertore uncieney alte travian 36 554 Flora Malesiana ser. I, Vol. 11 (3) (1994) WO 1 t L S/S Fig. 36. Guioa Cav. Petals and petal scales, crests on petal scales. — a. Scale free, apex very broad; b. scale free, without auricles, apex narrow; c. scale free, with auricles, apex narrow; d. scales as folded margins of petals; e. transition between claw (y) and blade (x) gradual; f. transition between the claw and the blade abrupt; g. claw almost absent, auricles very small; h. scale without crest; i. crest very small; j. crest linear; k. crest clavate. < < Fig. 37. Guioa Cay. Fruit shape (x: width of lobes; y: length of lobes; z: heigth of stipe). — a. Stipe high and slender, upper and lower margin of lobes parallel, straight; b. stipe almost absent, broad, lower and especially upper margin of lobes highly convex; c. stipe short, broadly cuneate, margin of lobes not parallel, straight. Adema, Leenhouts, Van Welzen — Sapindaceae Nn in nn b. 40a. 4la. Leaflets ovate (to elliptic). Inflorescence densely long (sericeous to) hirsute, hairs golden. Blade of petals orbicular, sharp transition with claw (Fig. 36f) 22. G. myriadenia Domatia absent to | (to many); if many: crest on petal scales absent or pilose when present; cotyledons subcollateral (Fig. 38b)s! sec. eoivce eoee ens ed . 57 Domatia 2 to many. Crest on petal scales glabrous. Cotyledons (obliquely) super- paneer 88a) | eos 3; olemialed ders Bune gel ans Bot 8. G. comesperma Sepals pilose on margin, outside (sub)glabrous. Leaflets 2-20 cm long, ovate to elliptic, apex (obtuse to) acuminate to caudate, mucronulate or not.......... 38 Sepals pilose on margin and outside. Leaflets 1.6—3.9 cm long, elliptic to obovate, Bpexuobtusenotimucronulate code to. ee eadd ote ek wales 27. G. parvifoliola Leaflets ovate (to elliptic), usually asymmetrical, very apex usually mucronulate. Peat sealesasually witha: glabrescent Grest igs jcc ww cece oes. cmcee BAO). 39 Leaflets elliptic, + symmetrical, very apex mucronulate (then crest on petal scales Bees ).on net (thencrestpdose)iie ssiq! tls. aalise! mest coun gid ... 40 Lobes of fruit much longer than high. Apex of leaflets abruptly narrowing 6. G. bicolor Lobes of fruit about as long as high. Apex of leaflets usually rather gradually nar- momecaeite, evelisut urol». i249) 26 ol dlame ausdunsade areub. 15. G. koelreuteria Leaflets without domatia, apex acute to acuminate, mucronulate. Crest of petal scales piahreus.iclavate:(Big36kK)m tacneiurd ema. Dares. £4680. 19. G. misimaensis Leaflets without or with one domatium, apex (acuminate to) cuspidate, not mucro- nulate. Crest of petal scales pilose, flat (Fig. 361) ................ 7. G. bijuga Disc uninterrupted (if in fruit check basal ring around several fruits; Fig. 35a) 42 Pesemutemmpted (Pie235e)q Jo-esce sunt! ose tte ee eae aE 49 Imm Fig. 38. Guioa Cay. Embryo types. — a. G. lentiscifolia Cav.; cotyledons superposed, apices almost not elongated. — b. G. pleuropteris (Blume) Radlk.; cotyledons secondarily collateral, apices elongated (a: Parks 16162; b: Currick 797). 556 Flora Malesiana ser. I, Vol. 11 (3) (1994) 42a. Leaves hirsute below. Apex of petal scales not broadened (Fig. 36b), crest absent or clavate and glabrous (Fig. 36h, k). Fruit of G. oligotricha 0.6-0.8 cm high (un- knowntontG. mollinscula). oo\. 20 celeb ie} care © 6 een Reed WAS See 43 . Leaves glabrous to sericeous (to subvillose) below; if subvillose (G. diplopetala) then apex of petal scales very broadened (Fig. 36a), crest if present flat and pilose (Fig:i36i), fruit. 0:7 =1.Sicmyhigh! 3 oie. Weis Set. «eT. eee 44 a. Leaflets 6.6-17.8 cm long; domatia absent or single. Crest on petal scales clavate (RigS6k)ye HS-5. eee t ete geek eee: eT. 20. G. molliuscula . Leaflets 4.1—9.1 cm long; domatia many. Crest on petal scales absent (Fig. 36h) 25. G. oligotricha . Crest on petal scales clavate (Fig. 36k). Lobes of fruit much longer than high (1.5 times). Leaflets ovate to elliptic, domatia absent or present. Stipe of fruit 1-2.5 mm LONE! ii ca sow Sok oie ecole oS AI ae EE, SO eee 45 . Crest on petal scales absent or flat (Fig. 36h, i). Lobes of fruit usually as long as high, if longer than leaflets elliptic, domatia absent, and stipe of fruit 2-3.3 mm lomerssersiad eae sete 2 ss Peed ERS AID ER, SE AE 46 . Leaflets ovate, domatia absent. Stipe of fruit c. 2.5 mm high, lobes not horizontally 38. G. scalariformis Leaflets elliptic, domatia absent to a single in at least a few leaflets. Stipe of fruit 1— Smnm: heh) lobesshonzontally 35 ies AR Eee eee 4. G. aryterifolia . Petals 0.5—4 by 0.3-2.2 mm, if small: apex of petal scales much broadened (Fig. 36a): EnuitstO:/ 2:2) byO7=128 mill. ek GE US Te ee 47 . Petals 0.8-1.2 by 0.3—0.4 mm, apex of scales not broadened (Fig. 36b). Fruit 0.7— OO byAO Se emp ia, Ae eae Re Levsises ot debeiod tere kel Pag ae 28. G. patentinervis . Leaflets (ovate to) elliptic; if ovate then apex of petal scales much broadened (Fig. 36a). Petal scales'0.3—2 mm, long (Figi36D).).. s:ee ake oarecee tee eee 48 . Leaflets ovate. Petal scales very short (Fig. 36g), 0.1—0.3 mm long, apex not broad- (eI d Gal aan ene anata we ae ae re ae Re PERL GES rc 6. 6 bono eae 5. G. asquamosa . Leaflets with (0 or 1 to) many domatia. Apex of petal scales (very) much broadened (Fiey36a)) Fruits 0:7-1.5 by 07-18 cree) 20g s one 10. G. diplopetala . Leaflets without domatia. Apex of petal scales not broadened (Fig. 36b). Fruits 1522.2 Dy 2a eC riba. 2c ae 2. so ees See sei eto 41. G. unguiculata . Leaflets ovate to elliptic, 2.9-24.2 by 1-8.9 cm, punctate or not, domatia absent or PEESCT ee cota shee Ue > = & SRRERERERCIOUe ieee on ro, ecto aes aee are ana 50 . Leaflets ovate, 10.9-30 by 3.8-13.3 cm, not punctate, domatia absent 12. G. grandifoliola . AD OIA ilayPATO, MANY” ates Wop. 5 a 55. + ws Raapemeteeeceepomee oy oceire cite hares 0 roneeaea- 0 Someone 51 : Domiatiatabsentionmone sy osc «a. ss.< ./ieppeeeua theses steeot: cor hopeless scene ene 52 . Crest on petal scales clavate, glabrous (Fig. 36k). Fruit 0.8—1.5 cm high; cotyledons (obliquely) superposed, sometimes apex of upper elongated (Fig. 38a) 8. G. comesperma . Crest on petal scales absent to usually present as a flat part (to a somewhat clavate part) of the scale apex, pilose (Fig. 36h-k). Fruit 1-2.2 cm high; cotyledons subcol- lateral, apices usually elongated (Fig. 38b) .............. 15. G. koelreuteria Adema, Leenhouts, Van Welzen — Sapindaceae 557 52a. Leaflets ovate to elliptic, with or without domatia. Fruit dehiscing completely, wall 4a. /}Domnatia swotomany isacatmpockets. .... . oo ns00s cman doors sans does ee 8 at suture less than 2 mm thick, pseudo-funicle on arillode (usually) present, if not or only partly then upper margin of fruit highly convex...................... 53 . Leaflets elliptic, without domatia. Fruit at most partly dehiscing, wall at suture up to 3.5 mm thick, upper margin of fruit straight, pseudo-funicle absent 9. G. contracta . Petal scales basally not auriculate (Fig. 36b), membranous margin indistinct; crest absent or clavate or part of bifid scale apex (Fig. 36h-k) .................. 54 . Petal scales basally auriculate (Fig. 36c), membranous margin distinct; crest a lin- Bananpendage (Figs 36)) cms. sae. cele.2miias.) aia 1. G. acuminata . Crest on petal scales absent or present, clavate or part of bifid scale apex, pilose eeerO Cis) 1 UES 605 8 FO) EN ORO 2 AY PCR es, ee 55 : ‘Creston petal scales clavate; glabrousi(Figs 36k)) is), dees. Wl gallate &: a7 . Leaflets ovate to elliptic, domatium absent or present. Scales of petals free (Fig. pba); Cotyledonsisubcollaterah (Fig: '\38b)«)...ic'-.cmedhgag. soled srallead.« 56 . Leaflets elliptic, domatium absent. Scales of petals folded margins (Fig. 36d). Co- tyledons presumably (obliquely) superposed (Fig. 38a) ..... 41. G. unguiculata . Leaflets elliptic; apex abruptly (acuminate to) cuspidate, very apex obtuse (to acute) 7. G. bijuga . Leaflets ovate (to elliptic); apex (obtuse to) usually gradually acuminate to caudate, very apex acute to usually mucronulate ................. 15. G. koelreuteria ePceneinciprioe 4s Aa NO NAMIeU eke Get JA log. Seto. 58 meats pyake CoP Tha R sr E Wee ee otals ers. (4804... a 23. G. normanbiensis . Leaflets subcoriaceous. Stipe of fruit 2-4 mm high; suture of fruit lobes flat to setiyiconver (Pig 3 7a)itiay oilers... een Aee).& 18. G. membranifolia Leaflets (sub)coriaceous. Stipe of fruit absent to up to 2 mm high; suture of fruit lobes usually highly convex, lobes almost touching (Fig. 37b) 37. G. rigidiuscula KEY TO THE SPECIES OF MALAY PENINSULA, SUMATRA, BORNEO, AND JAVA . Rachis of leaves slightly to distinctly winged (Fig. 33b, c)...............04. 2 Rchis;on leaves notiwinged: (Hip. 33a)i. o). Ueber dea hoe ob ad. eatin. eee 4 . Jugae (1—)2—7. Upper leaflets elliptic to obovate, lower surface densely greyish papillate, dull; domatiasmany.2 si ek devas ulate ciate oe ara bes 3 . Jugae 1-4. Upper leaflets elliptic; lower surface smooth (to somewhat papillate), rather shiny; domatia absent or a single ..................00005. 7. G. bijuga . Wings along rachis narrow, up to 3 mm broad. Apex of leaflets (obtuse to) abruptly acuminate (to cuspidate); lower surface with a convex, raised midrib, hairs oblique- ly hirsute (to sericeous), domatia many, pockets (to sacs; Fig. 34a, b, d) 30. G. pleuropteris . Wings along rachis always broad, up to 4 mm. Apex of leaflets gradually acuminate to cuspidate; lower surface with a hardly raised, flat midrib, hairs (very) sparsely sericeous, domatia many sacs (Fig. 34c)...............4. 34. G. pterorhachis Pematiaabsentiononé. sacecuiiee noth abso anise) a idginde asin 5 558 Flora Malesiana ser. I, Vol. 11 (3) (1994) Sa. 6a. Wa: 8a. 9a. la. . Leaflets papillate below, dull, greyish or whitish..............---.--5+-4--- 5 2a. Petals 1.8—3.8 mm long; scales with at most a hardly broadened apex (Fig. 36a); crest always present (Fig. 36i—-k). Disc interrupted (often almost uninterrupted; Fig. 35). Fruit lobes longer than high and usually sparsely sericeous or lobes about as longvas highrand ‘glabrovis 2p.) i UA RBI. eee 6 . Petals 0.5—4 mm long; scales with a (very) much broadened apex (Fig. 36a); crest usually absent (Fig. 36h). Disc uninterrupted (sometimes with a small slit; Fig. 35a, b). Fruit lobes usually longer than high, glabrous .......... 10. G. diplopetala Leaves 1—6-jugate. Leaflets ovate (to elliptic), asymmetrical; apex usually gradual- ly acuminate to caudate, very apex acute to usually mucronulate............. i . Leaves 1—4-jugate. Leaflets elliptic, usually symmetrical; apex abruptly (acumi- nate to) cuspidate, very apex usually obtuse (to acute), not mucronulate 7. G. bijuga Leaflets below smooth, shiny, glabrous. Ovary subhirsute. Fruits glabrous 15. G. koelreuteria Leaflets below papillate, dull, (slightly) shortly sericeous. Ovary densely hirsute. Fruits (very) sparsely sericeous, glabrescent ............... 35. G. pubescens Upper leaflets ovate to obovate; lower surface papillate, hirsute and with many pockets (to sacs) or (slightly) shortly sericeous and with many sacs. Petals (elliptic to) ob- ovate; apex of scales not to hardly broadened (Fig. 36b); crest (usually) present (Bigs 361219 ee ene ALO S96 La OL) 9 . Upper leaflets (ovate to) elliptic; lower surface usually smooth, glabrous (to slight- ly sericeous to subvillose), many sacs. Petals elliptic; scales with a (very) much broadened apex (Fig. 36a); crest usually absent (Fig. 36h)... 10. G. diplopetala Upper leaflets elliptic to obovate; apex (obtuse to) abruptly acuminate (to cuspi- date); lower surface (sub)sericeous to hirsute, domatia pockets (to sacs; Fig. 34a— €)aOvary subhirsute. Fruits glabrous 915%. 04 le. ee 30. G. pleuropteris Upper leaflets ovate (to elliptic); apex gradually acuminate to cuspidate; lower sur- face (slightly) shortly sericeous, domatia sacs (Fig. 34b). Ovary densely hirsute. Fruits (very) sparsely sericeous; glabrescent 272". S722 22. 35. G. pubescens KEY TO THE SPECIES OF THE PHILIPPINES Leaflets smooth below, without papillae, rather shiny................-.-.-. 2 Rachis not (to slightly) winged (Fig. 33a, b). Blade of petals obovate, gradually decusrent into the claw (Fig. 36)... 23st: 20. 1N9e 8 BOP. Le eee 3 . Rachis slightly winged (Fig. 33b). Blade of petals orbicular, abruptly clawed (Fig. SOR See eA ERT: X89 OO BL OEE LOS SER EGRLS BE 40. G. truncata . Apex of leaflets (obtuse to) acuminate to caudate. Scales of petals basically not auriculate (Fig. 36b), membranous margin along scales indistinct...........- = . Apex of leaflets (cuspidate to) caudate. Scales of petals basally auricled (Fig. 36c), membranous margin along scales distinct................... 1. G. acuminata . Rachis often (slightly) winged (Fig. 33b, c). Leaflets elliptic, usually symmetrical; apex abruptly (acuminate to) cuspidate; very apex obtuse (to acute); domatia absent OG URSA one owls e Hele He ENA re Oe es OE OI 2 RE 2s * 7. G. bijuga Adema, Leenhouts, Van Welzen — Sapindaceae 559 b. 5a: 6a. Ta. 10a. Rachis terete to flattened and broadened (to slightly winged; Fig. 33). Leaflets ovate (to elliptic), asymmetrical; apex (obtuse to) usually gradually acuminate to cau- date; very apex acute to usually mucronulate; domatia absent to many sacs (to pockets) 15. G. koelreuteria Rachis winged or not (Fig. 33). Leaflets ovate to obovate, (sub)coriaceous to very coriaceous, punctate or not; apex retuse to caudate; domatia absent to many. Inflo- rescence subglabrous to usually shortly brown sericeous to hirsute. Blade of petal obovate, gradually decurrent into claw (Fig. 36e); crest absent or part of bifid scale apex (to clavate; Fig. 36h—k) (N.B.: petals of G. palawanica and G. parvifoliola mmemervarh Tey hieeoeiinita: for Syeda woah how iwaanen (angdld oe 6 Rachis not winged (Fig. 33a). Leaflets ovate (to elliptic), coriaceous, punctate; doma- tia (absent to 1—many sacs to) many pockets. Inflorescence long golden (sericeous to) hirsute. Blade of petal orbicular, sharp transition with claw (Fig. 36f); crest peamate esti pitate «rion SOK) skh Sat, Da Bt as ae, bts 22. G. myriadenia een suoniusite) Fruits Glabrous A228 UP A) PO A i, Ee 7 . Ovary densely hirsute. Fruits sericeous, glabrescent ......... 35. G. pubescens Upper leaflets ovate to obovate; apex rounded to caudate; below (glabrous to) seri- ceous to hirsute, domatia absent to many pockets or sacs; venation usually conspic- uously raised below only, above usually concolorlous with lamina, densely to laxly sowemtate Pusually rather indistinct 4l>. 66) 4iind Se RE Se ee ee 8 Upper leaflets ovate; apex cuspidate to caudate; below sericeous, domatia a single (to many) sac(s); venation conspicuously raised on both sides, discoloured with famuina, deasely:reticulate: very distmet™ 0. ee ee se 36. G. reticulata . Upper leaflets ovate to obovate, 0.9-20 by 0.5-8.4 cm, if obovate either not punc- Bi orUase and Apex as ymunetricdl ss. ! fn. Jee coe sccav eee ee eed eee eae se e. Upper leaflets elliptic to obovate, 1.6-3.9 by 0.5—1.2 cm, slightly asymmetrical, LETT Sop tectge Oot Mate ot ete pee ice OG a etl ea Ne ae 27. G. parvifoliola . Upper leaflets ovate to obovate, if obovate then punctate, basally and apically asym- Gienacal, domatia iiany 7.455 eee So A ae ee ee eee ea ns ee 10 Upper leaflets elliptic to obovate, rather symmetrical, not punctate, domatia absent Sitio ee EU TERE TRE Le See Oe Ae 26. G. palawanica Rachis not (to slightly) winged (Fig. 33a, b). Upper leaflets ovate (to elliptic), sym- metrical to usually basally asymmetrical; apex (obtuse to) acuminate to caudate; below glabrous to sericeous to subhirsute (Mindanao only), usually at least some leaflets longer than 7.5 cm; nerves marginally looped and joined, sometimes (leaf- lets ovate) less, distinctly so in Tower half 2. <> se ads os. cs cee Oe 1] Rachis usually winged (Fig. 33b, c). Upper leaflets (elliptic to) obovate, apically and basally asymmetrical; apex (obtuse to) acute to acuminate (to cuspidate); be- low hirsute to (sub)sericeous, the (sub)sericeous leaflets always less than 7.5 cm long; nerves marginally looped and joined, but less distinctly so in the lower half 30. G. pleuropteris . Leaflets below somewhat differently coloured from above, brownish-greyish when dry, (usually) punctate; apex abruptly to gradually narrowing .............. 12 Leaflets below very differently coloured from above, almost whitish when dry, not punctate; apex gradually Harrowing... ... ss so ve caw ee Se eae 11. G. discolor 560 Flora Malesiana ser. I, Vol. 11 (3) (1994) 12a. Lobes of fruit about as long as high. Leaflets without domatia (then leaflets usually symmetrical) to many (then leaflets usually asymmetrical). Crest on petal scales (absent to) part of bifid scale apex (to clavate; Fig. 36h—-k)................. 13 b. Lobes of fruit much longer than high. Leaflets without or with | domatium, asym- metrical. Crest on petal scales clavate (Fig. 36k)...............- 6. G. bicolor 13a. Rachis often (slightly) winged (Fig. 33a, b). Leaflets elliptic, usually symmetrical; apex abruptly (acuminate to) cuspidate, very apex obtuse (to acute); domatia absent Oe MigSAcs Ra AAMAS TO A Te TT 7. G. bijuga b. Rachis terete to flattened and broadened (to slightly winged; Fig. 33a, b). Leaflets ovate (to elliptic) asymmetrical; apex (obtuse to) usually gradually acuminate to caudate, very apex acute to usually mucronulate; domatia absent to many sacs (to pockets) isicasg. sera aaa ee Se Sik GE Ee 15. G. koelreuteria KEY TO THE SPECIES OF CELEBES la. Axes and rachises, especially when young, hirsute. Lower surface of leaflets hir- Sule. .Wwithimany medere ct Plan dS). terpaapa) depo, ie aie ene 13. G. hirsuta b. Axes and rachises glabrous to very subsericeous. Lower surface of leaflets gla- broussred lands absent]... ....-.)-.5,. Jombanet- Tee ee 10. G. diplopetala KEY TO THE SPECIES OF THE MOLUCCAS la. Margin of leaflets entire or with few subapical teeth. Petals 5. Suture of fruits b. Margin of leaflets crenate. Petals 4. Suture of fruits very sharp 33. G. pteropoda 2a. Rachis slightly winged or not (Fig. 33a, b). Lower surface of the leaflets papillate, PlADTOUS LO SCRICCOUS, <5. sc ic ae wy sed oho sw he sr aeis OG «oO cpageis acne, ee 3 b. Rachis not winged (Fig. 33a). Lower surface of the leaflets not papillate, smooth, PADTOUS sey: oe aegyccuae Gach utes exes Gancaracic Actos) eens ecaer ae ico 4 3a. Rachis slightly winged (Fig. 33b). Lower surface of leaflets sericeous; domatia | to IMAM DOCK ta lIKC SACS oo paaess ooascan, ta accap cena cusps Sou aceeace as 16. G. malukuensis b. Rachis not winged (Fig. 33a). Lower surface of leaflets glabrous to sometimes very Sparsely sericeous-domatia. | (Ot) SaC(S). ste noe eee oe 2. G. acutifolia 4a. Petals 1.4-3 mm high; crest of scales well developed, clavate (Fig. 36k). Disc inter- rupted (Fig. 35c). Fruit 1.4-2.4 cm high by 1—3.6 cm broad 18. G. membranifolia b. Petals 0.8—1.2 mm high; crest absent (Fig. 36h). Disc uninterrupted (Fig. 35a). Fruit 0.7—0.9 cm high by 0 .7—1.3.. cm broad aos. 5 oop spans one Synch 28. G. patentinervis KEY TO THE SPECIES IN THE LESSER SUNDA ISLANDS Petals with very short scales, up to 0.3 mm long (Fig. 36g)........ 5. G. asquamosa Adema, Leenhouts, Van Welzen — Sapindaceae 561 3a. KEY TO THE SPECIES OF NEW GUINEA . Leaflets and inflorescences hirsute or velutinous. Rachis not winged (Fig. 33a) 2 . Leaflets and inflorescences glabrous, puberulous, tomentose, somewhat hirsute, or sericeous. If tomentose or somewhat hirsute then rachis winged (Fig. 33c) .... 4 . Lower surface of leaflets (smooth to) papillate, domatia (absent or | to) many. Crest of petal scales usually absent, seldom well developed (Fig. 36h, k). If leaflets smooth memMany domatia andno crest. (Fig..36h) —, 2 = 2). < scpscesce en da he oambsaed ee 3 . Lower surface of leaflets smooth, without papillae, domatia absent or a single sac. Crest of petal scales well developed (Fig. 36k) ............ 20. G. molliuscula Leaflets usually with a few subapical teeth; domatia many pockets. Inflorescences axillary, 1.2—6 cm long. Petals 0.7—1 by c. 0.3 mm; scales present (Fig. 36i), crest- less. Stipe of fruit 1.5—-2 mm high, slender (Fig. 37a) ........ 25. G. oligotricha . Leaflets entire; domatia absent to many pocket-like sacs. Inflorescences axillary to pseudoterminal, 1—26.6 cm long. Petals 0.9—2.3 by 0.5—1.6 mm; auricles present (Fig. 36d): crest usually absent (to developed). Stipe of fruit absent to up to 4 mm miele broadly obconical (Fig )37¢)) eioue chiles Jae? AO te 39. G. subsericea . Large inner sepals 0.8—3.2 by 0.9-3.5 mm. Petals (4 or) 5 (or 6). Sutures of fruit Bene pha COMpleterry atts 1) 36.31 NP Seog Bhetdonwlesthede: Qafiel i vce lhen Puls 5 . Large inner sepals 3-4 by 3—4.2 mm. Petals 4. Sutures of fruit very sharp; septa above attachment of funicle incomplete ................... 33. G. pteropoda a. Leaves 1—9-jugate, if all 1-jugate then leaflets usually longer than 7cm....... 6 Leaves all 1-jugate. Leaflets 1.1-6.3 cmlong .............. 29. G. pauciflora . Rachis winged, wing at least 0.5 mm broad (Fig. 33b, c) .........0. 02004... 7 Racks terete ito flattened, notywinged. (Pig33a) is. Soeew ks wxteolidoxittka ft 13 . Rachis slightly to broadly winged (Fig. 33b, c); if broadly winged then 2 to many BGmtitid PTESCHG755 «ua lina.|. see eel tins so Hastie a Ik) gai eee ee 8 Rachis broadly winged (wing more than 2 mm broad; Fig. 33c). Domatia absent (or Ste) Gs Seee Hie bob keenehiGi ks 38s sere tet nowealwigat 17. G. melanopoda . Heatlets densely. papillate below (dull, greyish): 2:s(2e42 00 e6\ee td. cdewdal lax 9 . Leaflets smooth below (rather shiny), no papillae ........................ 11 a. Leaflets 1—-2.4 by 0.4—0.9 cm; margin serrate or partly crenate in at least some leaf- lois; apex cmarginate toacute 743 tyeonte Jon aos’ cabot 4S. JES eaves) 10 . Leaflets 3.8—6.3 by 1—-1.7 cm; margin entire; apex acute....... 21. G. multijuga . Margin serrate in at least some leaflets; domatia 1, present in some leaflets 43. G. waigeoensis . Margin entire except for the usually crenate apex; domatia absent. 3. G. amabilis . Leaflets 0.8—6.5 by 0.5—2 cm; margin entire to usually (partly) crenate or serrate; domatia absent to many. Fruit blackish when dry, either 0.7—0.8 cm high with a slender stipe (Fig. 37a) or 1—2 cm high with a (rather) broadly obconical stipe (Fig. 37C) >». «. sab seochw staidoelel tigl'd - siubencask dads seeeietid awed ie de 12 . Leaflets 3.8—18.5 by 1.3—8.4 cm; margin entire; domatia 2 to many. Fruit reddish to reddish black when dry, 0.8—1.5 cm high, stipe slender (Fig. 37a) 8. G. comesperma Flora Malesiana ser. I, Vol. 11 (3) (1994) 14a. 20a. . Leaflets entire or usually slightly crenate; apex retuse to obtuse (to acute). Petals c. 34 byls2 mms Fruity 7 S19 emiacross ey. ee eee 32. G. pseudoamabilis Leaflets entire except for some subapical teeth; apex acuminate. Petals 0.2—1.1 by OME O02 Eruitt0/6=0/9 emracross fh Ae a. Be ee ee 42. G. venusta . Lower surface of leaflets glabrous or (sub)sericeous, domatia absent or a single sac (otonel or two pockets) tomany pocket-like'sacs” >. 2.4) 02 2.2) 2c) Seva 14 Lower surface of leaflets puberulous on midrib and basal nerves, domatia many POCKEES By -ricvaltpie Manet Hate RRM ene a eaee te eeen en see 14. G. hospita Leaflets below densely, long (greyish, dull) papillate, usually (sub)sericeous, if not sericeous then usually a single very large sac present (Fig. 34d) ............ 15 . Leaflets below (rather shiny) smooth (or at most very shortly papillate), glabrous (to very sparsely sericeous), sacs if present small (Fig. 34b) ............... Dll . Leaflets glabrous to (very sparsely) sericeous, domatia absent or a single small sac (or one or two pockets) to many pocket-like sacs (Fig. 34a, b).............. 16 . Leaflets glabrous except for a few sericeous hairs around the domatium, latter sin- gle, very large shighly domed (Fig. 34d) —. 0) 35.22 24. G. novobritannica PDiscinterrupted Gfin fruit, check several:fruits! Fig. 35¢)) 72 020 eee 17 § Discunmterupted:(tonwith asmallisht] Fie. 35a,D) 00. 2 ee eee eee 19 . Leaflets with 0-2 pocket domatia (Fig. 35a); if 2 pockets present then fruit blackish when'dry and stipe broadly obconicall (Figs 37) 225) 05. See. eee eee 18 Leaflets with 2 to many domatia, basally sacs (Fig. 35b) to upwards pockets. Fruit reddish to reddish black when dry, stipe slender (Fig. 37a)... 8. G. comesperma . Leaflets (sub)symmetrical, about 2.6-3 times as long as broad, coriaceous. Inflo- rescence subserceoust 2'.:: salle, Eh EAS 19. G. misimaensis Leaflets falcate, about 3.6—4.4 times as long as broad, coriaceous to very coria- CEOUS! IMtlOFESCENCE SEMICCOUS 2. ttt es eens ts WA ens 31. G. plurinervis . Leaves either 1—3(—5)-jugate or (3—)5—7-jugate. Leaflets (ovate to) elliptic; below glabrous to very slightly sericeous, domatia 0, 1 (or 2) sacs. Scales of petals free (except for the adnate lower part; Fig. 36a—c), often folded outwards ........ 20 . Leaves 1—3-jugate. Leaflets ovate (to elliptic) below (sub)sericeous, domatia ab- sent or 1 sac to many pocket-like sacs. Scales of petals present as inwardly folded, adnate auricles! (Figw36d) ete 10, 2S Tie eet En ees ke 39. G. subsericea Leaves 1—3(—5)-jugate; rachis not winged (Fig. 33a). Leaflets 2.8-23.5 cm long; lower surface glabrous, but sometimes very sparsely sericeous, especially on the midrib. Flower 1.5—2.5 mm in diam. Petals 0.3—1.8 mm long, claw absent or up to 02E0!4) mmrphish; scales O}1—1 mm long ....-. 522-3... . 2: 2. G. acutifolia . Leaves (3—)5—7-jugate; rachis not to very slightly winged (Fig. 33a, b). Leaflets 3.8—6.3 cm long; lower surface very sparsely sericeous. Flower c. 2.8 mm in diam. Petals 1.5—1.7 mm long, claw c. 0.3 mm high; scales c. 1 mm long 21. G. multijuga . Leaflets without or with | sac domatium. Fruit blackish when dry .......... yp . Leaflets with 2 to many domatia, basally sacs to upwards pockets. Fruits reddish to reddish blackewhemdryowiew. ans aoe: Hee wes. ele ee 8. G. comesperma . Some to all leaflets with a single (small) sac (Fig. 34b) ................... 23 VE DOMIAARAD SEM ak s.55 yt eccov aad weeecd 5) ccs hues ce heme oh tue esa gac eT @ tne tle creme 26 Adema, Leenhouts, Van Welzen — Sapindaceae 233° 29a. 1. Guioa acuminata Radlk., Philipp. J. Sc., Bot. 8 (1914) 463; Merr., Enum. Philip. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 1172; Welzen, Leiden Bot. Series 12 (1989) 171, f. 61. — Lectotype (Van Welzen 1989): BS (Ramos) 10916 (M holo; US), Phil- ippines. 563 Leaflets elliptic. Disc interrupted to uninterrupted (Fig. 35). Fruit 1.1—2.9 cm high; stipe absent or relatively short (c. 1/5—1/6 of fruit height), up to4 mm high... 24 Leaflets ovate. Disc interrupted (Fig. 35c). Fruit 1.2—1.5 cm high; stipe relatively high (c. 1/4 of fruit height), 3-3.5 mm high............ 23. G. normanbiensis . Disc interrupted (Fig. 35c). Stipe of fruit absent to up to 4 mm long; lobes about as long as high or higher than long (10-18 by 9-22 mm) .................... pe Disc uninterrupted (Fig. 35a). Stipe of fruit 1-1.5 mm long; lobes much longer than Bre S16) byiS- 12:5 a) eee. a ec. adios ght Ue 4. G. aryterifolia . Leaflets subcoriaceous. Fruit with a 2—4 mm high stipe; upper suture flat to slightly convex (Fig. 37a) 18. G. membranifolia Leaflets (sub)coriaceous. Fruit with a 0-2 mm long stipe; upper suture usually highly eonvex.(ebes almost touche: Fios37b)sicctesniviccsas 37. G. rigidiuscula MIME aCTCROVATCT AE Luctak L cotceteas SAAT. Ses i SE en et oe 27 Weer ee PCN es nrc a movant Sie woe S53 oa wane agaeim Gun. 5 wi a53ittvueas a aarp eee 28 . Leaflets 10.9-30 by 3.8—13.3 cm, index 2.1—2.9; upper surface usually with wax. Disc interrupted (Fig. 35c). Fruit wall 1-3 mm thick; stipe 1-1.5 mm high; lobes about as long as high, 12—16 by 10-14 mm 12. G. grandifoliola Leaflets 11.9—22.7 by 6.1—10.7 cm, index 1.8—2.1; upper surface without wax. Disc uninterrupted (Fig. 35a). Fruit wall less than | mm thick; stipe c. 2.5 mm high; lobes usually longer than broad, c. 17 by 12 mm.......... 38. G. scalariformis . Claw of petals either c. 0.4 mm high and then the crest of the scales well developed, clavate (Fig. 36k) or the claw c. 0.8 mm high and then the crest usually absent or present as part of a bifid scale apex (Fig. 36h, i). Disc (nearly) uninterrupted (Fig. 35a, b). Fruit wall rather thin (c. 1 mm thick); lobes much longer than high... 29 Claw of petals 0.2-0.4 mm high; crest well developed, clavate (Fig. 36k). Disc interrupted (Fig. 35c). Fruit wall very thick (up to 3.5 mm); lobes about as long 9. G. contracta Claw of petals c. 0.4 mm high; crest well developed, clavate (Fig. 36k). Disc unin- terrupted (Fig. 35a). Stipe of fruit 1-1.5 mm high........... 4. G. aryterifolia Claw of petals c. 0.8 mm high; crest usually absent or present as part of a bifid scale apex (Fig. 36h, i). Disc (nearly) uninterrupted (Fig. 35a, b). Stipe of fruit 2—3.3 mm high 41. G. unguiculata long. Leaflets opposite to subopposite, ovate to elliptic, somewhat falcate, 5.7—14.4 by 1.6—4.1 cm, index 2.64.2, asymmetrical, acroscopic side broad- er, coriaceous, punctate; base attenuate: margin entire, flat (to revolute); apex (cuspidate to) cau- date, mucronulate; upper surface glabrous; lower surface duller, not papillate, glabrous (to some Guioa perrottetii auct. non Radlk.: Sasaki, Cat. Gov. Herb. (1930) 328, p.p. (Ramos 10916). Tree, 6-8 m high, dbh up to 12.5 cm. Branch- lets sericeous when young; flowering twigs 2—4 mm thick. Leaves 1—3-jugate; rachis 1.9—-12 cm long, basally terete to upwards flattened, (sub)glabrous, petiole 1.5—8.2 cm long; petiolules up to 0.7 cm sparse hairs), domatia a single sac on basiscopic side in axil of second nerve; venation raised; nerves 0.4-1.9 cm apart, marginally looped and joined; veins laxly reticulate, indistinct. /nflorescences axillary, branching basally to mainly along the seri- ceous, flattened, 2.8-16.5 cm long axis; first order branches up to 7.7 cm long; cymules cincinnate, c. 3-flowered; bracts and bracteoles triangular, out- 564 side sericeous, inside glabrous; bracts 0.7—2 mm long; bracteoles 0.5—0.8 mm long; pedicels 3—4.4 mm long, sericeous except (sub)glabrous above ar- ticulation. Flowers 4—4.3 mm in diam., white. Se- pals 5, ovate, margin pilose, with glands, out- and inside glabrous; 2 outer smaller ones 1.5—2.3 by 1.2—2.6 mm; 3 inner larger ones 2.2—3.3 by 2.2— 3.7 mm, margin petaloid. Petals 5, obovate, 3.3— 3.7 by 1.2-1.3 mm, blade elliptic, gradually de- current into the 0.4—0.8 mm long claw, margin pi- lose, out- and inside glabrous, apex obtuse; scales 2.3—3.2 mm long, free, basally auriculate, mem- branous margin distinct; crest a linear appendage, (sub)glabrous. Disc interrupted. Stamens 8; fila- ment 2—5 mm long, pilose, especially basally; an- ther c. 0.5 mm long, glabrous. Pistil: ovary 0.4— 2.1 mm long, subhirsute; style and stigma 0.3—1.5 mm long. Fruits with 2 or 3 well developed lobes, c. 1.3 by 1.5 cm, completely dehiscent, rugose-ru- minate, glabrous, blackish when dry; stipe c. 4mm long, slender; margin blunt; lobes c. 8 by 8 mm, wall at suture less than 2 mm thick; septa com- plete. Seeds seen immature; arillode with pseudo- funicle. Distribution — Malesia: Philippines (Central & E Luzon, Polillo L.). Habitat & Ecology — Secondary forest; altitude sea level up to 100 m. FI. throughout the year. Note — Fruiting specimens of G. acuminata are hardly distinguishable from G. koelreuteria; they only differ in the relatively longer leaftip and the single instead of usually many domatia. 2. Guioa acutifolia Radlk., [Sapind. Holl.-Ind. (1879) 11, nom. nud.] Sitzungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 608; Bot. Jahrb. 56 (1921) 278; in Engl., Pflanzenr. 98 (1933) 1159; C.T. White, Contr. Arnold Arbor. 4 (1933) 60; S.T. Reynolds, Aus- trobaileya 2 (1984) 53; in Fl. Austral. 25 (1985) 48, f. 1ld-f, map 58; Welzen, Leiden Bot. Se- ries 12 (1989) 172, f. 62, 63. — [Cupania se- miglauca var. acutifolia F. Muell. ex Radlk., Sapind. Holl.-Ind. (1879) 11, nom. nud.] — Nephelium semiglaucum var. acutifolium F. Muell. ex Bailey & White, Bot. Bull. Queensl. Dep. Agr. 18 (1916) 9. — Lectotype (Van Welzen 1989): Dallachy s.n. (M holo; K, L, M, MEL, NSW), Australia. (Shrub to) tree, 2.5—25 m high, dbh 10—45 cm, spindly; outer bark nondescript, smooth to with small pustules and furrows, blotched variously grey to plum coloured, inner bark cream to pink to wine- red to light brown, with white fibrous vertical stripes; wood cream to light straw. Branchlets gla- Flora Malesiana ser. I, Vol. 11 (3) (1994) brous (to sparsely sericeous when young); flower- ing twigs I—5(-15) mm thick. Leaves 1—3(—6)-ju- gate; rachis 1-25 cm long, basally terete (to up- wards somewhat flattened), glabrous; petiole 0.5— 6.6(—8.8) cm long; petiolules up to 0.9 cm long. Leaflets opposite to alternate, (ovate to) elliptic (to slightly falcate), 2.8—23.5 by 1.2—8.7 cm, index 1.8— 4.7, usually symmetrical, subcoriaceous (to very coriaceous), usually punctate; base attenuate; mar- gin entire, flat; apex (obtuse to) acute to cuspidate (to caudate), usually mucronulate; upper surface glabrous (except for the sparsely pilose midrib), (thick waxy layer); lower surface glaucous, papil- late, glabrous (to very sparsely sericeous especial- ly on the midrib), domatia usually present in some to all leaflets, 1 (or 2) often small sacs on basi- scopic side in axil of second or third nerve (see note 3); venation on upper surface flat except for the basally raised midrib, on lower surface raised; nerves 0.2—3.9 cm apart, marginally looped and joined, (less so in lower part); veins laxly reticu- late. Inflorescences axillary (to pseudoterminal), unbranched to branching basally and along the terete, usually sericeous, 0.4-16.8 cm long axis; first order branches up to 6 cm long; cymules cincinnate to dichasial, |—7(—10)-flowered; bracts and bracteoles deltate, outside sericeous, inside glabrous; bracts (0.2—)0.4—1.4 mm long; bracteoles 0.2—0.9 mm long; pedicels 0.5—4 mm long, usual- ly sericeous except for the (sub)glabrous articu- late part. Flowers 1.5—2.5 mm in diam., sweet scent- ed. Sepals 5 (or 6), broadly ovate to orbicular, margin pilose, with glands, (outside sericeous), inside glabrous; 2 outer smaller ones 0.5—1.7 by 0.5-1.9 mm; 3 (or 4) inner larger ones 1—2.3 by 0.9-2.3 mm, margin petaloid. Petals 5, (rhombic to) obovate, 0.3—1.8 by 0.2-0.8 mm, white to tinged pinkish; claw 0—0.2(—0.4) mm high; margin pilose, outside and inside glabrous, apex obtuse to some- what fimbriate; scales 0.1—1 mm long, free, apex not broadened; crest absent (to clavate on slender short stipe, glabrous). Disc uninterrupted. Stamens 8: filaments 0.7—-3 mm long, completely pilose, especially basally; anthers 0.2-0.7 mm long, gla- brous to papillate, pink. Pistil: ovary 0.2—1.3 mm high, sparsely hirsute; stigma and style 0.1—0.7(— 2) mm long. Fruits with | or 2 (or 3) well devel- oped lobes, 0.5-1.2 by 0.8—2 cm, smooth to ru- gose-ruminate, glabrous, red when fresh, blackish when dry; stipe 0-3 mm high, broadly obconical to slender; margin blunt; lobes 5.S—12 by 5—8 mm, septa complete. Seeds obovoid, 4-8 by 3-5 mm; hilum 0.7—1.7 mm long. Distribution — Malesia: Moluccas, Irian Jaya (Vogelkop, Geelvinck Bay), Papua New Guinea (Western, Southern Highlands, Central Prov.); NE Adema, Leenhouts, Van Welzen — Sapindaceae 565 Australia: N Queensland (Cook, N Kennedy and Wide Bay District). Habitat & Ecology — Scarce to common under- storey tree; in disturbed rain forests, secondary bamboo forests, tall secondary montane forests, regrowth thickets, Araucaria vine thickets, along margins of forests, beach forests, water, mangroves, Melaleuca swamps. For vegetation types and soils in Australia see J.G. Tracey, Veget. Humid Trop. Region N. Queensl. (1982) 106. Altitude sea level up to 1800 m. Fl. (Mar.—)May-—Sept.(—Oct.); fr. Nov.—Jan. Notes — 1. Tracey (1982) records lowland and highland forms of G. acutifolia, but he did not ex- plain by what criteria these forms can be recog- nized. Note, however, that a specimen of G. come- sperma (Webb & Tracey 13251) was included in G. acutifolia. 2. Specimens found near salty water (along creeks, beaches, or on small islands) are sclero- phyllous with a thick waxy layer on the upper sur- face. The same phenomenon can be observed in specimens of G. bijuga from poor, sandy soil in Borneo. 3. Beccari FI 2810 is exceptional as the leaf- lets sometimes have a small subapical tooth, like G. oligotricha, and instead of a single sac several pockets are present. This specimen may belong to a new species, but as there is no other similar ma- terial, it is still included in G. acutifolia. 4. Several specimens in New Guinea possess very long leaflet apices, which make them look different from the more typical specimens of G. acutifolia. 3. Guioa amabilis Kanehira & Hatusima, Bot. Mag. Tokyo 57 (1943) 76, f. 11; Welzen, Lei- den Bot. Series 12 (1989) 176, f. 64. — Type: Kanehira & Hatusima 13999 (TI holo, n.v.; A), Irian Jaya. Shrub, 1-5 m high. Branchlets shortly sericeous, especially when young; flowering twigs |—3.5 mm thick. Leaves 3—S-jugate; rachis 0.6—5.4 cm long, slightly winged, wing at most | mm broad, slight- ly pilose, petiole 0.6-1.7 cm long. Leaflets sub- sessile, Opposite to alternate, elliptic to obovate, 1-1.9 by 0.5-0.9 cm, index 1.7—2.4, often asym- metrical, then acroscopic side broader, very coria- ceous, usually punctate; base attenuate; margin entire, except for the usually crenate apex, revo- lute; apex emarginate to acute, sometimes mucro- nulate; upper surface glabrous except for the some- times puberulous midrib, often wax; lower surface duller, papillate, sericeous, domatia absent; vena- tion on upper surface flat to slightly raised, raised below; nerves 0.1—0.4 cm apart, marginally looped and joined; veins densely reticulate, inconspicu- ous. Inflorescences axillary to pseudoterminal, unbranched to at most branching along the terete, sericeous, 0.6—9.7 cm long axis; first order branches up to 0.5 cm long; cymules cincinnate, 1- or 2- flowered; bracts and bracteoles triangular, outside sericeous, inside glabrous; bracts 0.5—1.3 mm long; bracteoles 0.3—0.6 mm long; pedicels 2—4 mm long, sericeous, articulate part less so. Flowers in bud; latter light green, tinged red. Sepals 5, ovate, mar- gin pilose, with glands, outside and inside glabrous, light green; 2 outer smaller ones 1.4—2 by 1.6—2.2 mm; 3 inner larger ones 2.2—3 by 2.2—-3 mm, mar- gin petaloid. Petals still immature, 5, clawed, white, margin pilose, outside and inside glabrous; scales still small. Disc uninterrupted. Stamens 8; filaments 1.7—-2 mm long, completely pilose, especially ba- sally, light green; anthers c. 0.6 mm long, glabrous, pink. Pistil: ovary c. 0.5 mm high, subhirsute; style and stigma c. 0.2 mm long. Fruits with 2 or 3 well developed lobes, 1—1.2 by 1.1—1.4 cm, smooth to slightly rugose, glabrous, brownish red when fresh, blackish when dry; stipe 1.5—2 mm high, broadly obconical; margin blunt; lobes 7—9 by 7-9 mm; septa complete. Seeds immature. Distribution — Malesia: Irian Jaya (Vogelkop). Habitat & Ecology — Scarce to common. Found at mountain forest edges and in occasionally burned scrubs. Soil: grey clay; altitude 2300-2500 m. FI. Jan. 4. Guioa aryterifolia Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 357; Bot. Jahrb. 56 (1921) 282; Baker in Rendle, J. Bot. 61, Suppl. (1923) 11; Radlk. in Engl., Pflanzenr. 98 (1933) 1174; P. Royen, Man. For. Trees Papua & New Guinea 2 (1964) 24, f. 11; Streimann, Pl. Upper Watut Watershed (1983) 169; Welzen, Leiden Bot. Series 12 (1989) 177, f. 65. — Lectotype (Van Welzen 1989): Forbes 870 (M holo: BM, MEL, P), Papua New Guinea. Shrub to tree, 5—20 m high, dbh up to 45 cm; bark grey, thin. Branchlets usually sericeous when young; flowering twigs 3—10 mm thick. Leaves 2— 4-jugate; rachis 4-21 cm long, terete, glabrous, petiole 2.6—7.3 cm long. Leaflets subsessile, op- posite to alternate, elliptic, 6.3—19.2 by 2.5—7.1 cm, index 2.2—3, usually rather symmetrical, (acroscop- ic side broader), coriaceous, usually punctate; base attenuate; margin entire, flat; apex acute to cuspi- date, often mucronulate; upper surface glabrous; lower surface duller, smooth, no papillae, glabrous (to sparsely sericeous), domatia absent or at least in some leaflets a single small sac on basiscopic side in axil of second nerve; venation on upper side 566 flat (to raised), raised on lower; nerves 0.3—2.9 cm apart, usually marginally very indistinctly looped and joined, especially in the lower part of the leaf- lets; veins laxly reticulate, often distinct. /nflores- cences axillary to ramiflorous, unbranched to branching basally and especially along the terete, sericeous, glabrescent, 3-13 cm long axis; first order branches up to 3.9 cm long; cymules cincin- nate, c. 2-flowered; bracts and bracteoles triangu- lar, outside sericeous, inside glabrous; bracts 0.7— 0.9 mm long; bracteoles 0.2—0.6 mm long; pedi- cels 3.3-5.5 mm long, sericeous. Flowers c. 3.2 mm in diam., sweet, faintly fragrant. Sepals 5, ovate, margin pilose, with glands, (outside sparse- ly pilose), inside glabrous; 2 outer smaller ones |— 1.7 by 1.2—2 mm; 3 inner larger ones 1.8—2.5 by 1.8—2.8 mm, margin petaloid. Petals 5, obovate, 1.8-1.9 by 0.6-0.9 mm, white; claw c. 0.4 mm high: margin pilose, apex acute, outside and inside sub- glabrous; scales c. | mm long, free; crest clavate, stipitate, apex lobed. Disc uninterrupted. Stamens 8; filaments 2.6—2.8 mm long, completely pilose, especially basally; anthers c. 0.5 mm long, gla- brous. Pistil: ovary c. 0.3 mm long, subhirsute; style and stigma c. 0.1 mm long. Fruits with 1-3 well developed lobes, 1.3—1.6 by 1.6—3.3 cm, rough to slightly rugose or ribbed, glabrous, black when dry; stipe 1—-1.5 mm high, slender; margin blunt; lobes 15-16 by 8—-12.5 mm, broadly spreading; wall thin to rather thick, not exceeding | mm; septa com- plete. Seeds globose to obovoid, c. 10 by 7 mm; arillode without or with a small pseudo-funicle, but always with a basal rim; hilum c. 1.5 mm long. Distribution — Malesia: Papua New Guinea (Central Prov.). Habitat & Ecology — Common. Found in low- land rain forest, the edge of the forest, and in sec- ondary forest.; altitude 50-1330 m. FI. Apr. Note — This species is part of the G. rigidiuscu- la-complex. The species can be distinguished by the venation with hardly joined nerves; the unin- terrupted disc; the fruits with a short slender stipe, a thin wall, and lobes which are longer than high and which spread widely. 5. Guioa asquamosa Welzen, Blumea 33 (1988) 411; Leiden Bot. Series 12 (1989) 179, f. 66. — Type: Metzner 227 (L holo), Timor. Guioa spec.: Verheijen, Dict. Manggarai Pl., Pac. Ling. ser. D, 43 (1982) 104. Tree, medium-sized. Branchlets usually seri- ceous when young; flowering twigs 1.5-3 mm thick. Leaves 2—4-jugate; rachis 4.6—19 cm long, terete to apically flattened above, subglabrous, petiole 2.8—8 cm jong; petiolules up to 0.9 cm long. Leaflets opposite (to alternate), ovate, 2.9-10.8 by Flora Malesiana ser. I, Vol. 11 (3) (1994) 0.93.3 cm, index 2.9—3.9, asymmetrical, acroscop- ic side broader, coriaceous, punctate; base attenu- ate; margin entire, flat; apex acuminate to cuspi- date, not mucronulate; upper surface glabrous, (wax); lower surface duller, smooth, no papillae, glabrous to very sparsely pilose, domatia (0 or) 1 to many sac(s), in axil of basiscopic second nerve to in axils of most nerves; venation on upper side flat to raised, raised below; nerves 0.3—1.6 cm apart, marginally looped and joined; veins laxly reticu- late, rather distinct. Inflorescences axillary, main- ly branching along the terete to somewhat flattened, subsericeous, 2.2—13.2 cm long axis; first order branches up to 4.2 cm long; cymules cincinnate, 2- or 3-flowered; bracts and bracteoles triangular, outside sericeous, inside subglabrous; bracts 0.7— 0.9 mm long; bracteoles 0.3—0.6 mm long; pedi- cels 3-6 mm long, subsericeous, articulate part less pilose. Flowers c. 4 mm in diam. Sepals 5, ovate, margin pilose, without glands, outside and inside glabrous; 2 outer smaller ones 1.3—1.8 by 1.3—1.8 mm; 3 inner larger ones 2.2—2.7 by 2.2—2.7 mm, margin petaloid. Petals 5, (elliptic to) obovate, 1.5— 2.5 by 1.1-1.7 mm; claw 0.1—0.2 mm high; mar- gin pilose, apex retuse to acute, outside and inside glabrous; scales free, very short, 0.1—0.3 mm high, apex not broadened; crest absent. Disc uninterrupt- ed. Stamens 8; filaments 1.4—3.5 mm long, pilose, especially basally; anthers 0.4—0.7 mm long, gla- brous. Pistil: ovary 0.5—1.5 mm high, subhirsute; style and stigma 0.2—1.5 mm long. Fruits with | or 2 well developed lobes, c. 1.4 by 1—1.4 cm, smooth, glabrous, blackish when dry; stipe c. 2 mm high, broadly obconical; margin blunt; lobes c. 9 by 9 mm. Seeds not mature. Distribution — Malesia: Lesser Sunda Islands (Flores, Timor). Habitat & Ecology — Altitude 800-1200 m. FI. May-June. 6. Guioa bicolor Merr., Philipp. J. Sc. 17 (1920) 279; Enum. Philipp Flow Pl. 2 (1923) 507; Radlk. in Engl., Pflanzenr. 98 (1933) 1170; Welzen, Leiden Bot. Series 12 (1989) 181, f. 67. — Type: BS (Ramos & Pascasio) 34487 (PNH}# holo; A, K, P), Philippines. Tree, 1-15 m high, dbh at least 4 cm; bark grey: wood white, hard. Branchlets sericeous when young, hairs brownish; flowering twigs 1-6 mm thick. Leaves 2—5-jugate; rachis 2.8—20.5 cm long, basally terete to apically flattened above, glabrous, petiole 2.1—9.1 cm long; petiolules up to | cm long. Leaflets opposite to subopposite, ovate to elliptic, 3.7-11.3 by 0.84.8 cm, index 2.4—4.6, asymmet- rical, not falcate, acroscopic side broader, coria- ceous to very coriaceous, punctate; base attenu- Adema, Leenhouts, Van Welzen — Sapindaceae ate; margin entire, flat to revolute; apex acuminate to caudate, usually abruptly narrowing (with a si- nus), usually mucronulate; upper surface glabrous (except the puberulous midrib); lower surface dull, papillate, glabrous to shortly subsericeous, doma- tia absent to a single small sac on basiscopic side in axil of second nerve; venation raised, above con- colorous with lamina; veins 0.2—2.3 cm apart, mar- ginally looped and joined, but often less distinctly so in lower part of leaflets; nerves densely to laxly reticulate, rather distinct. /nflorescences axillary, unbranched to branching basally and along the flat- tened, subsericeous, glabrescent, 1.9—14.4 cm long axis; first order branches up to 6.4 cm long; cy- mules cincinnate, c. 3-flowered; bracts and bracte- oles triangular, outside sericeous, inside glabrous; bracts 0.6—1.3 mm long; bracteoles 0.3—0.9 mm long; pedicels 2.5-4.9 mm long, sericeous, gla- brescent. Buds dark wine-red. Sepals 5, ovate, margin pilose, with glands, outside and inside (sub)glabrous, pink; 2 outer smaller ones 0.9—1.5 by 1.3-1.5 mm; 3 inner larger ones 2—3.5 by 1.8- 3.5 mm, margin petaloid. Petals 5, elliptic, 2.8— 3.2 by c. | mm, pinkish outside, white inside; blade elliptic, gradually decurrent into the c. 0.4 mm long claw, margin pilose, outside and inside glabrous, apex acute; scales c. 1.5 mm long, free, basally not auriculate, membranous margin distinct; crest a flat, pilose bifid part of scale apex when young, to clavate and glabrous when mature. Disc inter- rupted. Stamens 8; filaments c. 4 mm long, hir- sute, especially basally; anthers c. 0.7 by 0.6 mm, glabrous, light pink. Pistil: ovary c. 0.6 mm high, subhirsute; style and stigma c. 0.4 mm long. Fruits with 1-3 well developed lobes, c. 1.5 by 1.5—2.1 cm, smooth, glabrous, blackish when dry: stipe c. 3.5 mm high, slender; margin blunt; lobes c. 11 by 5 mm. Seeds obovoid, c. 9 by 5 mm; hilum c. 1.4 mm long. Distribution — Malesia: Philippines (Luzon, Mindanao, Sabtang I.). Habitat & Ecology — On ridges, in woods, cut forest. Soil: iron deposits present; altitude 130-850 m. Bud: Apr.; fr. May. 7. Guioa bijuga (Hiern) Radlk., Sapind. Holl.-Ind. (1879) 38; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 521, 611; in Engl., Pflanzenr. 98 (1933) 1165; Corner, Gard. Bull. Str. Settl. 10 (1939) 44; Wayside Trees (1940) 588; Meijer, Bot. News Bull. 9 (1967) 74; Briinig, Heidewald Sarawak Brunei (1968) 372; Corner, Gard. Bull. Sing., Suppl. 1 (1978) 153; Yap in Tree Fl. Malaya 4 (1989) 441; Welzen, Leiden Bot. Series 12 (1989) 183, f. 68, 69. — Cupania pleuropteris Blume var. 567 bijuga Hiern in Hook. f., Fl. Br. India | (1875) 677. — Guioa pleuropteris Radlk. var. bijuga King, J. As. Soc. Beng. 65, II (1896) 444; Rid- ley, Fl. Malay Penins. 1 (1922) 505. — Type: Wallich KD 8094 (K holo; BM, K), Malaya. [Guioa rubrofusca Radlk. ex Merr., Pl. Elm. Born. (1929) 175, nom. nud.; Radlk. in Engl., Pflan- zenr. 98 (1933) 1177; Masamune, Enum. Phan. Born. (1942) 427. Based on Elmer 21392.] See note 2. Tree, 3—30 m high, dbh 5 cm to | m; outer bark smooth to somewhat finely fissured, usually hard, usually whitish to (dark) grey with dark patches, also (red-)brown, inner bark yellow to red to brown, (fibrous); cambium white to yellow (to red); sap- wood white with growth rings to brown; heartwood pinkish brown. Branchlets usually glabrous, at most shortly sericeous when young, hairs brownish; flowering twigs 1—-7.5 mm thick. Leaves 1—4-ju- gate; rachis 0.8—16.8 cm long, usually flattened above, wing up to 2 mm broad, rachis at most pu- berulous, petiole 0.7—11 cm long. Leaflets subses- sile, opposite to subopposite (to alternate), ellip- tic, 2.1—-20 by 1.1-8.4 cm, index 1.2—4, usually symmetrical, subcoriaceous to very coriaceous, punctate; base sharply attenuate; margin entire, flat (to revolute); apex abruptly (acuminate to) cuspi- date, very apex obtuse (to acute), not mucronulate; upper surface glabrous to slightly sericeous; lower surface duller, smooth (to slightly papillate), gla- brous (to sparsely sericeous especially on the mid- rib), domatia absent or a single small sac on basi- scopic side in axil of second or third nerve (see note 3); venation on upper surface (slightly sunk- en to) flat to raised, concolorous with lamina, be- low raised; nerves 0.3—4.2 cm apart, marginally looped and joined, (less so in lower third of leaf- lets); veins laxly reticulate, rather indistinct. /n- florescences axillary (to pseudoterminal), (un- branched to) branching basally to especially along the terete to flattened, glabrous to pilose, 0.7—16.5 cm long axis; first order branches up to 7.4 cm long; cymules cincinnate, |—3(—6)-flowered; bracts and bracteoles deltate to triangular, outside sericeous, inside (sub)glabrous; bracts 0.5-1.9 mm long; bracteoles 0.2-1 mm long: pedicels 2.2-10 mm long, sericeous, (less so above articulation). Flowers 3.5—4.2 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside (sub)glabrous, green; 2 outer smaller ones 1—2.8 by 1—2.3 mm; 3 inner larger ones 1.4—3.4 by 1.2- 3.8 mm, margin petaloid. Petals 5, obovate, 1.8— 3.8 by 0.7-1.7 mm, white to yellow, blade ob- ovate, gradually decurrent into the 0.4—-1.2 mm high claw, margin pilose, outside and inside 568 Flora Malesiana ser. I, Vol. 11 (3) (1994) (sub)glabrous, apex emarginate to acute; scales (0.8—)1.2—2 mm long, free, basally without auri- cles, membranous margin indistinct, apex not to hardly broadened; crest a pilose flat part of the bi- fid scale apex. Disc interrupted. Stamens 8; fila- ments 1.6—5 mm long, pilose, especially basally; anthers 0.2—0.4 mm long, glabrous. Pistil: ovary 0.3-1.8 mm high, sparsely hirsute; style and stig- ma 0.2—2.5 mm long. Fruits with 1-3 well devel- oped lobes, 1.2—2.3 by 1.3—2.6 cm, smooth to some- what ribbed or rugose-ruminate, glabrous, black- ish when dry; stipe 2-5 mm high, rather slender; margin blunt; lobes 8.5—14 by 8-16 mm; wall less than 2 mm thick; septa complete. Seeds (globose to) obovoid, 8—9.8 by 7-7.8 mm; hilum 1.1—2 mm long; arillode with pseudo-funicle. Distribution — Thailand to Malesia: Peninsular Malaysia, Sumatra, Borneo, Philippines (Balabac I., Palawan). Habitat & Ecology — Often common. In lower and middle storey of primary and especially sec- ondary forest, savannah (kerangas), along roads, rivers, margin of forest, mangrove, in peat swamp, sometimes in cultivated fields. Soil: sandstone, podsolized white sand, ultrabasic, red-yellow loam; altitude sea level up to 1525 m. Fl. mainly (Nov.—) Feb.—Mar.(—Apr.), less so in Aug.—Sept.; fr. mainly Mar.—June, less so in Sept. and Oct. Notes — 1. Several specimens, growing on poor white sand, have smaller leaflets (3.2-14 by 1.5— 7.9 cm), which are very thick and scleromorphic; leaflets of specimens on richer soil are 3.9-19.6 by 1.8-8.4 cm, thinner, less scleromorphic. The same phenomenon can be observed in G. acutifo- lia. 2. One specimen is exceptional; S 32186: leaf- lets with many pockets, lower surface very papil- late and somewhat sericeous. 3. The difference between G. bijuga and G. pubescens is often vague on Borneo. Usually, G. bijuga has elliptic leaflets which are smooth and usually glabrous below; the fruits are glabrous too. Guioa pubescens usually has ovate leaflets, which are papillate below and sericeous, the fruits are glabrescent. On Borneo, G. bijuga can be papil- late and somewhat sericeous, while G. pubescens can have rather elliptic and more glabrous leaflets, especially in Sarawak and on the Kinabalu. Even there the more ovate shape of the leaflets and the few hairs remaining on the fruit indicate G. pubes- cens and the flattened above to winged rachis in- dicates G. bijuga. There are also anatomical dif- ferences: G. bijuga always has secretory idioblasts, while G. pubescens always lacks them. 8. Guioa comesperma Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1870) 357; Bot. Jahrb. 50 (1914) 77: 56 (1921) 281; in Engl., Pflanzenr. 98 (1933) 1173: Peekel, Fl. Bismarck-Arch. (1984) 337, f. 545: Welzen, Leiden Bot. Series 12 (1989) 188, f. 71. — Lectotype (Van Welzen 1989): MacGregor s.n., 1889, p.p. (M holo; MEL), New Guinea. Nephelium winterianum F.M. Bailey, Queensl. Agr. J. 3 (1898) 283. — Type: Bailey s.n. (BRI holo; K), New Guinea. Guioa rigidiuscula auct. non Radlk.: Hartley et al., Lloydia 36 (1973) 270. Guioa subsericea auct. non Radlk.: Streimann, PI. Upper Watut Watershed (1983) 169 (p.p.: NGF 9176, 14484). Guioa spec.: Streimann, Pl. Upper Watut Water- shed (1983) 169 (p.p.: NGF 14494). (Shrub to) tree, 2-18 m high, dbh 7—30 cm; outer bark smooth to finely vertically fissured to fluted, reddish brown to grey, usually patched, inner bark straw-coloured to pink to dark orange-red; sapwood white to pale pink, heartwood pinkish. Branchlets usually shortly sericeous when young, hairs brown- ish; flowering twigs 1.5—8 mm thick. Leaves 1—4- jugate; rachis 1.8-14.8 cm long, terete to winged, wing up to 3 mm broad, subglabrous, petiole 0.9— 6.6 cm long; petiolules up to 0.7 cm long. Leaflets opposite to alternate, ovate to elliptic, 3.8-18.5 by 1.3-8.4 cm, index 2—3.8, often asymmetrical, acro- scopic side broader, subcoriaceous, punctate; base attenuate; margin entire, flat; apex acuminate to cuspidate (to caudate), usually mucronulate; up- per surface glabrous (except midrib sparsely pu- berulous); lower surface duller, smooth (to papil- late), glabrous to very sparsely sericeous, domatia small, 2 to many in axils of nerves, basally sacs to apically pockets; venation on upper side (slightly sunken to) flat (to raised), raised below; nerves 0.2— 2.5(—3.5) cm apart, marginally looped and joined, (less distinctly so in lower part of leaflets); nerves laxly reticulate, often distinct. Inflorescences ax- illary to pseudoterminal, branching basally and along the terete to ribbed, subglabrous to sericeous, 1-17.6 cm long axis; first order branches up to 7.8 cm long; cymules cincinnate (to dichasial), 2-4(— 7)-flowered; bracts and bracteoles deltate to trian- gular, outside sericeous, inside glabrous; bracts 0.4— 1.3 mm long; bracteoles 0.2—0.6 mm long; pedi- cels 1.1-4.5 mm long, sericeous except subglabrous above articulation. Flowers 4-4.5 mm in diam. Sepals 5, ovate, margin pilose, with glands, out- side and inside glabrous, green to pinkish; 2 outer smaller ones 0.7—2 by 0.6—1.8 mm; 3 inner larger ones 1.8—3.2 by 1.7—3.5 mm, margin petaloid. Pet- Adema, Leenhouts, Van Welzen — Sapindaceae 569 als (4 or) 5, elliptic, 2.1—3.8 by 0.6—-1.7 mm, white to pale pink; blade gradually decurrent into the 0.3— 0.8 mm high claw, margin and less so outside pi- lose, inside (sub)glabrous, apex acute; scales 0.7— 2 mm long, free, basally not auriculate; crest cla- vate, long stiped, apex lobed, glabrous, yellow. Disc interrupted. Stamens 8; filaments 1.1—4.5 mm long, pilose, especially basally; anthers 0.3—0.8 mm long, sparsely pilose, (purplish) pink. Pistil: ovary 0.2— 2 mm long, subhirsute; style and stigma 0.1—2 mm long. Fruits with 1-3 well developed lobes, 0.8— 1.5 by 0.9-2 cm, smooth to rugosely ribbed, gla- brous, red when fresh, reddish to reddish black when dry; stipe 1.54.5 mm high, slender; margin blunt; lobes 7-11 by 5.5—11 mm; septa complete. Seeds obovoid, 6—8.5 by 4.3—6.5(—8) mm, glossy dark brown; hilum 0.8—1.4 mm long. Distribution — Malesia: Papua New Guinea (Southern Highlands, Western Highlands, Madang, Morobe, West & East New Britain, New Ireland, Manus, Northern, Milne Bay, Central Prov.) to NE Australia (Queensland: Cook Dist.). Habitat & Ecology — In savannah, secondary forest, lower montane forest on steep slopes or ridg- es, sometimes in primary lowland rain forest; along banks of rivers, lakes, mangrove, beach. Usually found in more open vegetations. Vegetation type in Australia: Semideciduous mesophyll vine for- est. Dominant plants in vegetation: Casuarina, Eucalyptus deglupta. Soil: alluvial, sand, limestone, rock, ultrabasic; altitude sea level up to 900 m. FI. (Apr.—)May—Oct.(—Dec.); fr. Jan.—Oct. Notes — 1. Guioa comesperma belongs to the G. rigidiuscula-complex. 2. A topocline can be found from the Central Province via Milne Bay towards the northern prov- inces. In Central Province the specimens have winged rachises, leaflets without papillae, and high- ly stipitate, small fruits with slender lobes. In Milne Bay the fruits are still the same, but the wing is seldom present and then usually very narrow. In Morobe Province and the other more northern and western provinces the specimens have no wings, the leaflets can be papillate, and the fruits are usu- ally larger (lobes broad and stipe small). 3. Distinguishing between G. subsericea and specimens of G. comesperma with large fruits and papillate, subsericeous leaflets is difficult when the specimens bear fruit. Then the only difference is the disc, which is interrupted in G. comesperma, and uninterrupted in G. subsericea. Flowering spec- imens are easier to separate, because the petals are very different, those of G. comesperma are large (2.1-3.8 mm long) with long, broad, free scales with big crests, while in G. subsericea the petals are much smaller (0.9—2.1 mm long), have infold- ed auricles as scales, and the crest is usually ab- sent but can be as well developed as in G. come- sperma. 9. Guioa contracta Radlk., Bot. Jahrb. 50 (1914) 77; 56 (1921) 283; in Engl., Pflanzenr. 98 (1933) 1174; Welzen, Leiden Bot. Series 12 (1989) 191, f. 72. — Type: Schlechter 18269 (B* holo; K, P), New Guinea. Guioa aryterifolia auct. non Radlk.: Hartley et al., Lloydia 36 (1973) 269. Guioa comesperma auct. non Radlk.: Hartley et al., Lloydia 36 (1973) 270 (p.p.: Hartley 10976, 12635). Tree, 5—27 m high, dbh 8-20 cm; outer bark smooth to faintly longitudinally fissured, light grey to reddish brown to dark blackish green, inner bark light straw to pale brown; wood soft, (straw-)white. Branchlets sericeous when young; flowering twigs 2.5-15 mm thick. Leaves 1—3-jugate; rachis 2.3- 19.2 cm long, terete, (sub)glabrous, petiole 2—8.5 cm long; petiolule up to 1.1 cm long. Leaflets sub- opposite to alternate, elliptic, 7.6—24.2 by 3-8.9 cm, index 2.1—3.3, + symmetrical, otherwise acro- scopic side broader, subcoriaceous, punctate; base attenuate; margin entire, flat; apex acuminate to cuspidate, very apex obtuse (to mucronulate); up- per surface glabrous; lower surface duller, smooth, no papillae, glabrous to often very sparsely seri- ceous, domatia absent; venation on upper side flat, raised on lower; nerves 0.3—3.3 cm apart, margin- ally looped and joined, less distinctly so in lower part of leaflets; veins laxly reticulate, rather indis- tinct. Inflorescences cauli- to ramiflorous (to axil- lary), branching basally to mainly along the terete, subsericeous, 1.3—18 cm long axis; first order branches up to 8.3 cm long; cymules cincinnate, (1—)3—4-flowered; bracts and bracteoles triangu- lar, outside hirsute, inside glabrous; bracts 0.5—1 mm long; bracteoles 0.2—0.5 mm long; pedicels 1.8—5 mm long, completely sericeous. Flowers 3.5— 4 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside glabrous, pale green; 2 outer smaller ones 0.7—1.4 by 1.1—2 mm; 3 inner larger ones 1.8—2.8 by 1.5—3 mm, margin petaloid. Petals 5, ovate to elliptic (to obovate), 1.7—2.8 by 0.5—1.3 mm, white; claw 0.2—0.4 mm high; mar- gin and less so outside pilose, inside glabrous, apex rounded to acute; scales 0.7—1.6 mm long, free, basally not auriculate; crest clavate, stipitate, apex lobed. Disc interrupted. Stamens 8; filaments |.2— 3.5 mm long, completely pilose, especially basal- ly pilose, white; anthers 0.4—0.6 mm long, glabrous to subpilose. Pistil: ovary 0.3-0.6 mm long, sub- hirsute; style and stigma 0.2—0.6 mm long. Fruits 570 with 1—3 well developed lobes, 1.4—1.9 by 1.3—2.4 cm, smooth to rugose, glabrous, not dehiscing to only (partly) dehiscing when completely ripe, red when fresh, blackish when dry; stipe 1-1.5 mm high, broadly obconical; margin blunt; lobes 11— 15 by 9-15 mm, widening towards the axis in trans- verse view; wall at suture up to 3.5 mm thick; septa complete. Seeds obovoid, c. 8 by 7.5 mm; arillode without a pseudo-funicle, basal rim present; hilum c. | mm long. Distribution — Malesia: Papua New Guinea (Madang, Morobe, Northern, and Central Prov.). Habitat & Ecology — In lowland and montane rain forest, secondary forest bordering oak forest, scrubby forest along creeks and mangrove, along roadsides; altitude sea level up to 1500 m. Fl. Aug.— Oct.; Fr. July. Many twigs hollow with ants. Note — Guioa contracta is part of the G. rigidius- cula-complex and it is characterized by its rather large, almost glabrous, elliptic leaflets without papillae and domatia, and its distinctive hardly lobed fruits. The lobes do not narrow towards the axis in transverse section, but widen; the wall is very thick (the fruits dehisce late and presumably only partly); the pseudo-funicle is reduced to a rim only. 10. Guioa diplopetala (Hassk.) Radlk., Sapind. Holl.-Ind. (1879) 88; Sitzungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 514, 521, 543, 610; Valeton, Bull. Inst. Bot. Buitenzorg. 15 (1902) 10; Koord. & Vale- ton, Bijdr. Booms. Java 9 (1903) 207; Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 63; Moll & Janss., Mikrogr. Holz. 2 (1911) 377; Koord., Exk. Fl. Java 2 (1912) 542; Radlk., Philipp. J. Sc., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 507; Pl. Elm. Born. (1929) 175; Radlk. in Engl., Pflanzenr. 98 (1933) 1162; Desch, Mal. For. Rec. 15 (1954) 526; Salvosa, Lex. Philipp. Pl. (1963) 104; Backer & Bakh. f., Fl. Java 2 (1965) 140; Welzen, Leiden Bot. Series 12 (1989) 197, f. 76, 77. — Cupania di- plopetala Hassk., Flora 25, 2, Beibl. (1842) 39; Cat. Hort. Bog. (1844) 224; Pl. Jav. Rar. (1848) 286. — Guioa diplopetala f. genuina Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 610, nom. il- leg.; in Engl., Pflanzenr. 98 (1933) 1162. — Type: not indicated, either Herbarium Hasskarl or Herbarium Bogoriense (n.v.), Java. Guioa squamosa Radlk., [Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 303, nom. nud.] Sapind. Holl.-Ind. (1879) 38; Sitzungsber. Math.-Phys. Cl. K6nig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 544, 609; Flora Malesiana ser. I, Vol. 11 (3) (1994) King, J. As. Soc. Beng. 65, II (1896) 444; Rid- ley, J. Str. Br. Roy. As. Soc. 33 (1900) 66; Bran- dis, Ind. Trees (1906) 186; Lecomte in FI]. Indo- Chine 1 (1912) 1025; Ridley, Fl. Malay Penins. 1 (1922) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 1161; Gagnep. in Fl. Indo-Chine, Suppl. 1 (1950) 981; Yap in Tree Fl. Malaya 4 (1989) 442. — [Sapindus squamosa Wall., non Roxb., Cat. (1847) 8097, nom. nud., nom. inval.] — Guioa squamosa f. genuina Radlk., Sitzungs- ber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 609, nom. illeg.; in Engl., Pflanzenr. 98 (1933) 1161. — Type: Wallich 8097 (K holo; A, BM, FI, P), Malay Peninsula. Cupania regularis Blume, Rumphia 3 (1847) 159; Walp., Ann. 2 (1851/2) 214; Filet, Plantk. Woordenb. ed. 2 (1888) 277, nr. 8115. — Gui- oa regularis Radlk., Sapind. Holl.-Ind. (1879) 12, 41; Masamune, Enum. Phan. Born. (1942) 427. — Guioa diplopetala f. borneensis Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 610; in Engl., Pflanzenr. 98 (1933) 1162. — Guioa diplo- petala var. borneensis Radlk., Bot. Jahrb. 49 (1913) 370. — Lectotype (Van Welzen 1989): Korthals s.n., no date (L, sh. 908.269-309), Borneo. Cupania minjalilen Blume, Rumphia 3 (1847) 162. — Guioa minjalilen Radlk., Sapind. Holl.-Ind. (1879) 10, 37; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 609. — Lectotype (Van Welzen 1989): Waitz s.n. (L holo; L, P), Java. Arytera karang Mig., Sumatra (1861) 510; Filet, Plantk. Woordenb. ed. 2 (1888) 151, nr. 3862b. — Type: Diepenhorst HB 2487 (U holo; BO), Sumatra. Cupania fuscidula Kurz, J. As. Soc. Beng. 41, II (1872) 302; ibid. 44, II (1875) 188, 189 (type); Hiern in Hook. f., Fl. Br. India 1 (1875) 676; Kurz, Fl. Burm. | (1877) 284. — Guioa fusci- dula Radlk., Sapind. Holl.-Ind. (1879) 38; Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 515, 609; King, J. As. Soc. Beng. 65, II (1896) 445; Brandis, Ind. Trees (1906) 186; Ridley, Fl. Malay Penins. 1 (1922) 506; Burk. & Hend., Gard. Bull. Str. Settl. 3 (1925) 363; Radlk. in Engl., Pflanzenr. 98 (1933) 1161; Corner, Gard. Bull. Str. Settl. 10 (1939) 44; Wayside Trees (1940) 588; Gagnep. in Fl. Indo-Chine, Suppl. 1 (1950) 979; Meijer, Bot. News Bull. 9 (1967) 75; Yap in Tree FI. Malaya 4 (1989) 442. — Type: Helfer 993 (K holo; A, L, M, P, W), Burma. Guioa squamosa Radlk. f. lineolata-punctata Radlk., Sitzungsber. Math.-Phys. Cl. Konig]. Adema, Leenhouts, Van Welzen — Sapindaceae Bayer. Akad. Wiss. Miinchen 9 (1879) 609; in Engl., Pflanzenr. 98 (1933) 1161. — Type: Helfer 983 (K holo; A, P), Burma. Guioa diplopetala f. microcarpa Radlk., Sitzungs- ber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 610; in Engl., Pflanzenr. 98 (1933) 1162. — Type: Beccari F1 2812 (FI holo, cited by Radlkofer as Beccari 6), Celebes. Guioa leptoneura Radlk., Sitzungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 611, 618; in Engl., Pflanzenr. 98 (1933) 1163. — Syntypes: Beccari FI 2807 (FI, cited by Radlkofer as Beccari 7”), Celebes, Kandari Prov., Lepo-lepo; Riedel s.n. (BO, K, W), Celebes. Guioa fuscidula (Kurz) Radlk. var. glabrescens King, J. As. Soc. Beng. 65, I (1896) 445; Rid- ley, Fl. Malay Penins. | (1922) 506. — Type: Scortechini 1714 (K holo), Malay Peninsula. Guioa microphylla Radlk., Rec. Bot. Surv. India 3 (1908) 354; Ridley, Fl. Malay Penins. | (1922) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 1161; Corner, Gard. Bull. Str. Settl. 10 (1939) 43. — Type: C. Curtis 1346 (K holo), Malay Peninsu- la. Guioa bullata Radlk. in Fedde, Rep. 18 (1922) 342; in Engl., Pflanzenr. 98 (1933) 1164; Masamune, Enum. Phan. Born. (1942) 426. — Lectotype (Welzen, 1989): Haviland 1003 (K holo; M), Borneo (cited by Radlkofer as Haviland 1063). Guioa sp. 1: Yap in Tree Fl. Malaya 4 (1989) 443. — Based on KEP 99960 (KEP, n.v., SAN), Malaysia, Kedah Peak; KEP FRI 3892 (K, L; KEP, n.v.), Malaysia. Guioa sp. 2: Yap in Tree Fl. Malaya 4 (1989) 443. — Based on KEP FRI 15969 (K, L; KEP, n.v.), Malaysia, Selangor, Fraser’s Hill; KEP FRI 19164 (K, KEP, L), Malaysia. ?Guioa sp. 3: Yap in Tree Fl. Malaya 4 (1989) 443. — Based on KEP 56953 (KEP, n.v.), Malaysia. Arytera montana auct. non Blume: Miq., Sumatra (1861) 510. Cupania glabrata auct. non Kurz: Hiern in Hook. f., Fl. Br. India 1 (1875) 676, p.p. (Wallich 8097, 8550). Cupania griffithiana auct. non Kurz: Kurz, J. As. Soc. Beng. 44, II (1875) 188, p.p. (Helfer 983); Fl. Burm. | (1877) 284. See Van Welzen (1989: 262) note 2 under G. pleuropteris for nomen- clature. Shrub to tree, 1-18.5 m high, dbh 3 cm to 1.3 m; outer bark smooth, grey-brown to grey-white, inner bark pink to pale brown; sapwood white to light yellow. Branchlets usually brown sericeous when young; flowering twigs 1.5-25 mm thick. 571 Leaves 1—9-jugate; rachis 2.3—33.5 cm long, terete to upwards flattened above, glabrous to subseri- ceous, petiole 1.4—13.5 cm long; petiolules 0—0.9 cm long. Leaflets opposite to alternate, (ovate to) elliptic, 2.3-24.3 by 0.8-7.8 cm, index (1.1—)2.5- 5.9, (slightly) especially basally asymmetrical, the acroscopic side broader, coriaceous, usually punc- tate; base attenuate; margin entire (to crenate), flat (to revolute); apex (obtuse to) acuminate to cuspi- date (to caudate), usually not mucronulate; upper surface usually glabrous to slightly sericeous; lower surface duller, smooth (to papillate), glabrous (to slightly sericeous to subvillose), domatia (0 or 1 to) many small sacs (to pockets), in axils of nerves; venation on upper surface (slightly sunken to) flat to raised, raised on lower side; nerves 0.2—3.7 cm apart, marginally looped and joined, (less distinct- ly so in lower part of leaflets); veins laxly reticu- late, usually indistinct. /nflorescences (ramiflorous to) axillary (to pseudoterminal), (unbranched to) branching basally and along the terete to slightly flattened, subsericeous (to subhirsute), 0.4—-18 cm long axis; first order branches up to 9 cm long; cymules cincinnate (to dichasial), 2—6-flowered: bracts and bracteoles deltate to triangular, outside sericeous, inside (sub)glabrous; bracts 0.5-2 mm long; bracteoles 0.2—0.9 mm long: pedicels 1.8— 7.3 mm long, completely sericeous. Flowers 3—4.5 mm in diam., without scent. Sepals 5, ovate, mar- gin and sometimes outside pilose, margin with glands, inside glabrous, green; 2 outer smaller ones 0.9-2.8 by 0.8-2.1 mm; 3 inner larger ones 1.4— 3.4 by 1.2-3.6 mm, margin petaloid, white. Petals 5, elliptic to obovate, 0.54 by 0.3—2.2 mm, white: claw 0.2—-1 mm high; margin pilose, outside and inside (sub)glabrous, apex rounded to acute; scales 0.3—2 mm long, free, apex (very) much broadened: crest usually absent to a pilose flat part of the bifid scale apex. Disc uninterrupted, yellow. Stamens 8; filaments |1.2—5 mm long, pilose, especially basal- ly, white; anthers 0.3—0.8 mm long, glabrous to slightly pilose, pink. Pistil: ovary 0.2—2 mm long, subhirsute, light green to white; style and stigma 0.1-2 mm long. Fruits with 1-3 well developed lobes, 0.7—1.5 by 0.7—1.8 cm, smooth to somewhat ribbed, glabrous, red when fresh, blackish when dry; stipe 2-5 mm high, slender; margin blunt; lobes 5-10 by 4-9.5 mm. Seeds obovoid, 5—9 by 4.1-— 7.3 mm, black; hilum 0.8—2 mm long. — Figs. 39, 40. Distribution — Burma, Thailand, Cambodia, Vietnam, Malesia: Malay Peninsula, Sumatra, Java, Borneo, Celebes. Habitat & Ecology — Regularly encountered, but not common. In primary and especially in second- ary forest, in heath forest, submontane forest, edge Flora Malesiana ser. I, Vol. 11 (3) (1994) 303 eC Fig. 39. Guioa diplopetala (Hassk.) Radlk. a. Habit; b. petal; c. fruit (a, c: Rahayu & Maskura 540; b: Beumée A765). of forest, along rivers, roads, seashore, in deserted cultivated fields, savannah (belukar). Soil: granit- ic sand, basalt, clay, loam on sandstone, limestone, marshy sand; altitude sea level up to 1700 m. FI. Sept.—Apr.; fr. Dec.—Apr. Said to be poisonous Uses — Boiled roots act against blennorrhea (suppurating inflamation of mucous membrames; Pételot, Pl. Medic. Cambodge, Laos, Viet-Nam I, 1952, 201). The wood is apparently usable for house construction (De Clercq, Nieuw Plantk. Woordenb., 1909, 251, nr. 1710), and is resistant against ter- mites and used as poles (Gagnep. in FI. Indo-Chine, Suppl. 1, 1950, 981). The wood of G. bijuga, rath- er similar to that of G. diplopetala, seems to be very vulnerable to insect attack. The arillode is presumably edible (Pételot, 1952). Notes — 1. Guioa diplopetala is a very wide- spread and variable species, with a peculiar, almost nN — ws) Adema, Leenhouts, Van Welzen — Sapindaceae Fig. 40. Guioa diplopetala (Hassk.) Radlk. Distribution map showing the change in leaflet form from 0.7). —a. ‘G. squamosa’ form; b. ‘G. microphylla’ form; c. ‘G. squamosa’ W to E Borneo (all leaflets c. - form; d. intermediate between the ‘G. sguamosa’ and ‘G. diplopetala’ forms; e. ‘G. squamosa’ form; f. up- per half: ‘G. fuscidula’ form, pilose; lower half: ‘G. fuscidula var. glabrescens’ form, glabrous; g. °G. bul- lata’ form; h. ‘G. diplopetala’ form (a: Geesink et al. 7688; b: Curtis 1346, c: Curtis 1041, d: Iboet 304; e: Put 3024; f: upper half Helfer 993, lower half Scortechini 1714; g: Haviland 2137; h: Rahayu & Mas- kura 540). 574 circular distribution of forms, see Fig. 40. On Bor- neo two distinguishable forms are present (Fig. 40g, h); specimens which link these two forms are found on the Malay Peninsula, Sumatra, and Java (see arrow in Fig. 40). The form ‘G. bullata’ has been described for the west point of Borneo (S Sarawak and SW Kalimantan; Fig. 40g shows an extreme form): the leaflets are broad, have a low leaf-in- dex, an asymmetrical base, and are slightly villose below. The same form has been described for the Malay Peninsula as ‘G. fuscidula’ (Fig. 40f upper half), and here the base can be even more asym- metrical, the indumentum is somewhat more hir- sute. The latter form is connected to ‘G. squamo- sa’ (Fig. 40a, c, e; synonym: G. cambodiana), and ‘G. microphylla’ with very small leaflets (Fig. 40b), through ‘G. fuscidula var. glabrescens’ (Fig. 40f, lower half), all from the Peninsula; ‘G. squamosa’ has smaller asymmetrical leaflets with a somewhat higher leaf-index and lacks the hirsute indumen- tum. Intermediate forms between ‘G. fuscidula/G. squamosa’ and typical G. diplopetala (syn.: G. regularis) are found on Sumatra (Fig. 40d). Typi- cal G. diplopetala is found on Sumatra, Java, E Borneo (the other form of Borneo!) and Celebes (Fig. 40h): the leaflets are rather symmetrical, long, narrow (high leaf-index), and lack hairs (sometimes the leaflets are very sparsely pilose below). The two forms on Borneo are spatially separated by the geologically old (dating from before the glacial periods) Lupar River system in W Borneo. This river is also a border for other species from differ- ent families, e.g., 57 species of Dipterocarpaceae occur east or west of it (Ashton, Ann. Missouri Bot. Gard. 64, 1977, 694-705). The presence of the cen- tral mountain range in Borneo may have prevent- ed a further mingling of both forms; however, few collections have been made in central Borneo and it is therefore possible that intermediates exist in Borneo itself. 2. From Java northwards to Borneo and Celebes the fruits gradually become somewhat smaller. 11. Guioa discolor Radlk. in Elmer, Leafl. Philipp. Bot..5 (4913) 1609; Philipp. J..Seé., Bot.,.8 (1914) 446; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 507; Radlk. in Engl., Pflanzenr. 98 (1933) 1168; Welzen, Leiden Bot. Series 12 (1989) 203, f. 78. — Lectotype (Van Welzen 1989): Elmer 7493 (M holo; A, BM, FI, K, L, NY, W), Philippines. Tree, 6-13 m high, dbh 15—20 cm; outer bark grey; wood white, hard. Branchlets sericeous when young; flowering twigs 3—8.5 mm in diam. Leaves 3—5-jugate; rachis 4.3—22 cm long, basally terete to upwards flattened, subglabrous, petiole 1.8—8.3 Flora Malesiana ser. I, Vol. 11 (3) (1994) cm long; petiolules up to 1 cm long. Leaflets op- posite to subopposite, ovate, falcate, 4.9—-14.4 by 1—3.6 cm, index 3.5—5, asymmetrical, acroscopic side broader, (very) coriaceous, not punctate; base attenuate; margin entire, flat to revolute; apex cus- pidate to caudate, gradually narrowing, usually mucronulate; upper surface glabrous; lower surface dull, densely whitish papillate, sericeous, domatia 0 or 1 small sac on basiscopic side in axil of sec- ond nerve; venation on upper surface flat (to raised), concolorous with lamina, below raised; nerves 0.2— 1.8 cm apart, marginally looped and joined; veins laxly reticulate, usually rather indistinct. /nflores- cences axillary, branching basally and along the usually flattened, brown sericeous, 1.3—9.5 cm long axis; first order branches up to 3.3 cm long; cy- mules cincinnate, c. 2-flowered; bracts and bracte- oles triangular, outside sericeous, inside subgla- brous; bracts 0.8—1.2 mm long; bracteoles 0.7—0.8 mm long; pedicels 1.8—4.5 mm long, sericeous except for the subglabrous articulate part. Flowers c. 4.2 mm in diam.; buds dark wine-red. Sepals 5, ovate, margin pilose, with glands, outside and in- side (sub)glabrous, green to pink; 2 outer smaller ones 1.2—1.3 by 1.1—1.3 mm; 3 inner larger ones 1.5—2.2 by 1.5-2.2 mm, margin petaloid. Petals 5, elliptic to obovate, 2.5—2.7 by 0.6—1 mm, white to pink; blade obovate, gradually decurrent into the 0.7—-1 mm long claw, margin pilose, outside and inside glabrous, apex obtuse to acute; scales 1.1— 1.7 mm long, free; crest a pilose flat part of the bifid scale apex to subglabrous and clavate. Disc interrupted. Stamens 8; filaments 2.8—3.7 mm long, pilose, especially basally; anthers c. 0.3 mm long, glabrous, pink. Pistil: ovary 0.4—0.7 mm long, sub- hirsute; style and stigma 0.4—0.5 mm long. Fruits immature. Distribution — Malesia: Philippines (Luzon, Samar). Habitat & Ecology — Primary dipterocarp for- est; altitude 90-850 m. Fl. May—June. 12. Guioa grandifoliola Welzen, Blumea 33 (1988) 412, pl. 4a, b; Leiden Bot. Series 12 (1989) 214, f. 83. — Type: NGF (Streimann) 45154 (L holo; BRI, K; LAE, n.v.), Papua New Gui- nea. Tree, 5-25 m high, dbh 5-30 cm; outer bark slightly pustular, grey to red-brown, inner red- orange; wood of moderate weight and hardness, cream. Branchlets shortly sericeous when young; flowering twigs 5-19 mm thick. Leaves 2- or 3- jugate; rachis 5.8-21 cm long, terete, glabrous, petiole 1.7—9.5 cm long. Leaflets subsessile, op- posite to subopposite, ovate, 10.9—30 by 3.8-13.3 cm, index 2.1—2.9, + symmetrical, otherwise acro- Adema, Leenhouts, Van Welzen — Sapindaceae scopic side broader, coriaceous, not punctate; base attenuate; margin entire, flat; apex acute to cuspi- date, mucronulate; upper surface glabrous, usual- ly wax; lower surface duller, smooth, no papillae, glabrous, domatia absent; venation on upper side flat (to raised), raised on lower; nerves 0.44.3 cm apart, marginally looped and joined, less distinct- ly so in lower part of leaflets; veins laxly reticu- late, indistinct. /nflorescences ramiflorous (to ax- illary), unbranched to branching basally and along the flattened to terete, (sub)sericeous, 0.7—12.7 cm long axis; first order branches up to 8.5 cm long; cymules cincinnate; bracts and bracteoles triangu- lar, outside hirsute, inside glabrous; bracts 0.9-1 mm long; bracteoles 0.3—0.6 mm long; pedicels 4— 5.6 mm long, sericeous except for the subglabrous articulate part. Flowers unknown. Sepals 5, ovate, margin pilose, outside and inside glabrous, light green; 2 outer smaller ones 0.8—1.7 by 1.5—1.8 mm, margin with glands; 3 inner larger ones 1.9—3 by 2-3 mm, margin petaloid, without glands. Petals unknown. Disc interrupted. Stamens unknown. Pis- til unknown. Fruits with 1|—3 well developed lobes, 1.2—1.6 by 2—2.7 cm, smooth to slightly rugose or ribbed, glabrous, red when fresh, black when dry; stipe 1-1.5 mm high, broadly obconical; margin blunt; lobes 12-16 by 10-14 mm; wall 1-3 mm thick near suture; septa complete. Seeds obovoid, c. 7.5 by 5.5 mm; pseudo-funicle often reduced in size to a basal rim on arillode; hilum c. 1.1 mm long. Distribution — Malesia: Papua New Guinea (Morobe, Northern Prov.). Habitat & Ecology — In lowland rain forest (Dip- terocarp-dominated), advanced secondary forest and gallery forest; altitude sea level up to 400 m. Fr. Aug. Note — Guioa grandifoliola is part of the G. ri- gidiuscula-complex and is characterized by its enor- mous, Ovate leaflets without indumentum and domatia. The fruits are like those of G. rigidiuscu- la, only smaller. 13. Guioa hirsuta Welzen, Blumea 33 (1988) 412, pl. la—c; Leiden Bot. Series 12 (1989) 216, f. 84. — Type: van Balgooy 3658 (L holo; K; BO, n.v.), S Celebes. Shrub to tree, 2-15 m high, dbh up to 19 cm; fluted; ornamental; outer bark brownish grey, not fissured, with many small elevated lenticels, inner bark cream-coloured; sapwood pale reddish cream, heartwood reddish ochre. Branchlets hirsute; flow- ering twigs 2.2-7 mm thick. Leaves 2—7-jugate; rachis 1.8—15.5 cm long, terete to upwards flattened above, hirsute, petiole 1.1—6.2 cm long. Leaflets subsessile, subopposite to alternate, (ovate to) el- X73 liptic, 3—10.7 by 1—3.5 cm, index 2.4—3.8, usually somewhat asymmetrical, acroscopic side broader, coriaceous, punctate; base attenuate; margin en- tire, flat (to revolute); apex acuminate to cuspidate, usually not mucronulate; upper surface hirsute; lower surface duller, smooth, no papillae, hirsute, small red erect glands abundant, domatia many pockets in axils of nerves; venation on upper side flat, raised on lower; nerves 0.3—1.6 cm apart, marginally looped and joined; veins laxly reticu- late, rather indistinct. /nflorescences axillary, branching basally and along the terete to somewhat flattened, hirsute, 1.5—17 cm long axis; first order branches up to 5 cm long; cymules cincinnate, 2- or 3-flowered; bracts and bracteoles triangular, outside sericeous, inside subglabrous; bracts 0.7— 1 mm long; bracteoles 0.3—0.5 mm long; pedicels 1.8—4 mm long, sericeous, less so in articulate part. Flowers 3.2—3.5 mm in diam., buds green. Sepals 5, ovate, margin and basally the outside pilose, margin with glands, inside glabrous; 2 outer smaller ones I—1.8 by I—1.8 mm; 3 inner larger ones 1.8— 2.8 by 1.4-3.2 mm, margin petaloid. Petals 5, el- liptic, 1-1.9 by 0.3—-0.7 mm; claw 0.4—0.8 mm high; margin pilose, outside and inside (sub)glabrous, apex obtuse; scales 0.9-1.3 mm long, free; crest absent (to slender pilose part of a bifid scale apex). Disc uninterrupted. Stamens 8; filaments 3.8—4.3 mm long, pilose, especially basally; anthers 0.4— 0.5 mm long, glabrous. Pistil: ovary 0.3—0.5 mm long, subhirsute; stigma and style 0.1—0.6 mm long. Fruits with 1-3 well developed lobes, 0.7—1.1 by 0.7—1.5 cm, smooth to sometimes somewhat ru- gose-ruminate, glabrous, red when fresh, blackish when dry; stipe c. 1.5 mm high, slender; margin blunt; lobes 5—8 by 6.5—8 mm. Seeds obovoid, 5.2— 5.3 by 2.9-4.2 mm, brown; hilum 1—1.1 mm long. Distribution — Malesia: Celebes. Habitat & Ecology — In swamp forest, disturbed forest, belukar, and along lakes. Vegetation usual- ly with Gleichenia, Imperata, Melastoma. Soil: usually ultrabasic, sometimes siliceous or clayey; altitude sea level up to 1200 m. FI. Apr.—July; Fr. Mar.—June. 14. Guioa hospita Radlk. [in Engl & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 346, nom. nud.], Bot. Jahrb. 56 (1921) 281; in Engl., Pflanzenr. 98 (1933) 1173; Welzen, Leiden Bot. Series 12 (1989) 218, f. 85. — Type: Expedition Royal Geographical Society of Australia s.n., Dec. 1890 (M holo), New Guinea. Tree? Branchlets puberulous when young; flow- ering twigs c. 5 mm thick. Leaves |- or 2-jugate: rachis 6.8—-16.5 cm long, terete, slightly puberu- lous, petiole 6.5—9.5 cm long. Leaflets subsessile, 576 Flora Malesiana ser. I, Vol. 11 (3) (1994) n,n opposite, ovate, 15.4-22.5 by 7.4-9 cm, index 2.1— 2.5, subbasally asymmetrical, acroscopic side broader, subcoriaceous, punctate; base attenuate; margin entire, flat; apex acuminate to cuspidate, mucronulate; upper surface puberulous on midrib and basal nerves; lower surface duller, smooth, no papillae, puberulous on venation, domatia many pockets in axils of nerves; venation on upper side flat, raised below; nerves 0.3—2.2 cm apart, mar- ginally looped and joined, less distinctly so in lower part of leaflets; veins laxly reticulate, rather incon- spicuous. /nfructescences axillary, branching along the terete, puberulous, c. 11 cm long axis; first or- der branches up to 4.4 cm long; bracts and bracte- oles caducous; pedicels 6—-6.5 mm long, puberu- lous. Flowers absent. Sepals and petals caducous. Disc interrupted. Stamens and pistil absent. Fruit immature, with | or 2 developing lobes, c. 1.8 by c. 2.8 cm, slightly ribbed, glabrous, blackish when dry; stipe c. 6 mm high, slightly obconical; margin blunt; lobes c. 16 by 7 mm; septa complete. Seeds unknown. Distribution — Malesia: Papua New Guinea (Gulf Prov.). Habitat & Ecology — Stems of specimen seen often swollen beneath the nodes, harbouring insect nests, probably those of ants. Note — This species is part of the G. rigidiuscu- la-complex and is distinct by its puberulous leaf- lets with many pockets, many and dense nerves, and very stipitate fruits with slender long lobes. 15. Guioa koelreuteria (Blanco) Merr., Sp. Blanc. (1918) 241; Brown, Min. Prod. Philipp. For. 3 (1921) 204; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 507; Radlk. in Engl., Pflanzenr. 98 (1933) 1172 (footnote); Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 300, f. 218; Welzen, Leiden Bot. Series 12 (1989) 219, f. 7b, 86. — Sapindus koelreuteria Blanco, Fl. Filip. (1837) 289 (‘kolreuteria’). — Koelreuteria arborea Blanco, Fl. Filip. ed. 2 (1845) 202. — Neo- type (Merrill 1918): Merrill 644 (PNH7# holo; BM, F, K, L, NY, P, W), Philippines. Hemigyrosa perrottetii Blume, Rumphia 3 (1847) 165; Walp., Ann. 2 (1851/2) 212; Gray, U.S. Expl. Exp. Bot. 1 (1854) 251; Mig., Fl. Ind. Bat. 1, 2 (1859) 568; Fern.-Vill., Nov. App. (1883) 349. — Guioa perrottetii Radlk., Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 273; Sapind. Holl.- Ind. (1879) 39; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 614: Vidal, Rev. Pl. Vasc. Filip. (1886) 95; Ceron, Cat. Pl. Herb. Manila (1892) 53; Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 63; Merr., Philipp. J. Sc. 1, Suppl. (1906) 87; Fl. Manila (1912) 306; Radlk., Philipp. J. Sc., Bot. 8 (1913) 446; in Engl., Pflanzenr. 98 (1933) 1172. — Lectotype (Van Welzen 1989): Perrottet s.n., no date (L holo; FI, L, P), Phil- ippines. Guioa salicifolia Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1608; Philipp. J. Sc., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 509; Radlk. in Engl., Pflanzenr. 98 (1933) 1165. — Type: Elmer 12286 (M holo; A, BM, F, FI, K, L, NY, P, U, W), Philippines. Guioa mindorensis Merr., Philipp. J. Sc. 20 (1922) 404; Enum. Philipp. Flow. Pl. 2 (1923) 508; Radlk. in Engl., Pflanzenr. 98 (1933) 1165. — Type: BS (Ramos) 39639 (PNH*# holo; A, K), Philippines. Guioa spec.: Vidal, Rev. Pl. Vasc.Filip. (1886) 95, p.p. (Vidal 1226, 1230). Quassia simaruba auct. non L.: Blanco., Fl. Filip. ed. 2 (1845) 247; Merr., Sp. Blanc. (1918) 241 (in syn.). Sapindus pubescens auct. non Zoll. & Mor.: Zoll. & Mor. in Mor., Syst. Verz. (1846) 22, p.p. (Perrottet s.n.). Cupania regularis auct. non Blume: Vidal, Sinop- sis (1883) 21, f. 34 b. Guioa rigidiuscula auct. non Radlk.: Vidal, Rev. Pl. Vasc. Filip. (1886) 95; Ceron, Cat. Pl. Herb. Manila (1892) 53. Guioa diplopetala auct. non Radlk.: Meijer, Bot. News Bull. 9 (1967) 75. Shrub to tree, 2-16 m high, dbh 2-50 cm; outer bark brown to dark greyish, smooth (to rough), in- ner pink to red-brown, fibrous; sapwood white to brownish, odourless, tasteless, rings slightly visi- ble. Branchlets especially sericeous (to hirsute) when young; flowering twigs |—7 mm thick. Leaves |-6-jugate; rachis 2.4-20.5 cm long, terete to flat- tened and broadened (to slightly winged), subgla- brous (to subhirsute), petiole 1-8 cm long; peti- olules 0-0.7 cm long. Leaflets usually subsessile, opposite to alternate, ovate (to elliptic), 2.8-17.8 by 0.9-5.4 cm, index 1.7—5.7, usually asymmetri- cal, acroscopic side broader, (very) coriaceous, usually punctate; base attenuate; margin entire, flat (to revolute); apex (obtuse to) usually gradually acuminate to caudate, very apex (acute to) mucro- nulate; upper surface glabrous to subhirsute; low- er surface duller, smooth (to slightly papillate), glabrous to subsericeous (to hirsute), domatia ab- sent to many small sacs (to pockets) on basiscopic side in axils of nerves; venation on upper surface (flat to) raised, usually concolorous with lamina, below raised; nerves 0.2—2.5 cm apart, marginally Adema, Leenhouts, Van Welzen — Sapindaceae looped and joined; veins laxly reticulate, usually distinct. Inflorescences axillary (to pseudotermi- nal), unbranched to branching basally and along the slightly flattened, brown (sub)sericeous (to hir- sute), 0.8—20.5 cm long axis; first order branches up to 9.8 cm long; cymules cincinnate (see note 4), 2-5(—8)-flowered; bracts and bracteoles trian- gular, outside sericeous, inside subglabrous; bracts 0.5—1.5 mm long; bracteoles 0.2—1 mm long; pedi- cels 1.4—-7 mm long, sericeous except for the usu- ally subglabrous articulate part. Flowers 3.24 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside (sub)glabrous, pink; 2 outer smaller ones 1—3.1 by 1—3.3 mm; 3 inner larger ones 1.5—3.8 by 1.2—-4 mm, margin petaloid. Pet- als 5, (elliptic to) obovate, 1.8—3.8 by 1-2.5 mm (see note 4), creamy white (to reddish); blade ob- ovate, gradually decurrent into the 0.4—1.3 mm high claw, margin (and outside) pilose, inside (sub)glabrous, apex retuse to acute; scales 0.92.1 mm long, free, basally not auriculate, membranous margin indistinct, apex not to hardly broadened: crest (absent to) a pilose, flat part of the bifid scale apex (to somewhat swollen and clavate). Disc in- terrupted. Stamens 8 (or 9); filaments 1.2-5 mm long, pilose, especially basally; anthers 0.3—0.6 mm long, glabrous to pilose. Pistil: ovary 0.2—2.4 mm long, subhirsute; style and stigma 0.1—2.7 mm long. Fruits with 1—3 well developed lobes, 1—2.2 by 1- 2.3 cm, completely dehiscent, smooth, glabrous, red when fresh, blackish when dry; stipe 2-5.5 mm high, slender; margin blunt; lobes 6.5—13 by 7-12 mm; septa complete. Seeds (globose to) obovoid, 5.3-8.5 by 5-7.5 mm; hilum 1-2.3 mm long; arillode with pseudo-funicle, edible. Distribution — Malesia: Borneo (islands NE of Sabah), Philippines. Habitat & Ecology - Common. In primary and especially in secondary forest, ridge forest, thick- ets, on heath, along the seashore, roads, streams, edges of plantations. Soil: sand, gravel, limestone, ultrabasic; altitude sea level up to 1350 m. FI. (Aug.—)Nov.—Feb.(—Mar.); fr. Mar.—Oct. Uses — The wood is used for agricultural im- plements and tool handles (Reyes in Desch, 1954). Oil extracted from the seeds can be used to cure certain skin diseases (Guerrero in Brown, Useful Pl. Philipp. 2, 1950, 363). Notes — 1. Guioa koelreuteria is rather variable as can be expected of a speceis whose range ex- tends over several islands. The leaflets differ much in size, Most are small, but some can become rath- er large. The leaflets are usually rather thin, but some have thick coriaceous leaflets. A rather dis- tinct form, found on Mindanao, Davao Prov., and occasionally on Guimaras Island, has leaflets which 577 always possess many domatia and short (combined with a few long) hairs sparsely all over the lower surface, while the inflorescences are patently pi- lose instead of (sub)sericeous. 2. The specimens which were described as G. salicifolia differ only slightly from ‘normal’ G. koelreuteria, the leaflets are more coriaceous and relatively narrower. 3. ‘Guioa mindorensis’ is also distinguishable. The characters mentioned by Merrill (small leaf- lets, winged rachis) can also be found among more typical specimens ot G. koelreuteria. On Mindoro two forms can be found, typical G. koelreuteria with leaflets without papillae and usually with a long cuspidate apex and ‘G. mindorensis’ with papillae and a short, rather obtuse apex. The latter form is presumably found at higher altitudes. 4. One exceptional specimen, ANU 1652 from Mindanao, Mt Apo, has dichasial instead of mon- ochasial cymes and the petals are almost completely reduced (the flowers are still young and may have opened prematurely because of drying). 16. Guioa malukuensis Welzen, Blumea 33 (1988) 418, pl. 5; Leiden Bot. Series 12 (1989) 227, f. 89. — Type: Kostermans 1206 (L holo; BRI, K, P: BO, n.v.), Moluccas. Tree, c. 13 m high, dbh up to 20 cm; bark grey; wood hard. Branchlets shortly sericeous, especial- ly when young; flowering twigs 3—3.5 mm in diam. Leaves 2-4-jugate; rachis 0.7—6 cm long, slightly winged, subsericeous, petiole 0.7—3 cm long. Leaf- lets subsessile, opposite to subopposite, elliptic, 3.4-7.2 by 1-2.2 cm, index 2.4—3.7, + symmetri- cal, coriaceous, punctate; base attenuate; margin entire, flat; apex acuminate, not mucronulate; up- per surface subsericeous, especially the venation; lower surface dull, papillate, sericeous, domatia | to many pocket-like sac(s) in axils of nerves; ve- nation on upper side flat, raised on lower; nerves 0.3-1.3 cm apart, marginally looped and joined; veins densely reticulate, distinct. /nflorescences axillary, branching basally and along the somewhat flattened, sericeous, 3.3—12.2 cm long axis; first order branches up to 3.3 cm long; cymules cincin- nate; bracts and bracteoles triangular, outside seri- ceous, inside glabrous; bracts c. 0.9 mm long: bracteoles 0.3—0.4 mm long; pedicels 3.5—3.8 mm long, sericeous except for the (sub)glabrous artic- ulate part. Flowers c. 3 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside gla- brous; 2 outer smaller ones c. 1.2 by 1.2 mm; 3 inner larger ones 1.8—2.2 by 1.8—2.2 mm, margin petaloid. Petals 5, elliptic, c. 1.4 by | mm, white; claw c. 0.2 mm high; margin and less so outside 578 pilose, inside (sub)glabrous, apex retuse to round- ed; scales inwardly folded auricles, c. 0.4 mm long; crest absent. Disc uninterrupted. Stamens 8; fila- ments c. 2.4 mm long, pilose, especially basally; anthers c. 0.4 mm long, glabrous. Pisti/ in young state. Fruits unknown. Distribution — Malesia: Moluccas (Morotai I.). Habitat & Ecology — Found at 1000 m altitude. Note — This species strongly resembles G. sub- sericea, but the latter has a wingless rachis and asymmetrical leaflets with a cuspidate to caudate apex instead of a winged rachis, symmetrical leaf- lets, and an acuminate apex. 17. Guioa melanopoda Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 514; Welzen, Leiden Bot. Series 12 (1989) 230, f. 91. — Type: Brass 13699 (A holo; BM, K, L), Irian Jaya. Tree, 8-15 m high. Branchlets glabrous except for some tomentose hairs on nodes; flowering twigs 3-6 mm thick. Leaves 3- or 4-jugate; rachis 3.2— 20.5 cm long, winged, wing 2.5—3 mm broad, sub- tomentose, petiole 2.7—7.7 cm long. Leaflets sub- sessile, subopposite to alternate, (ovate to) ellip- tic, 8-16.4 by 2.4-5.6 cm, index 2.7—3.5, slightly asymmetrical, acroscopic side broader, coriaceous, punctate; base attenuate; margin entire, flat; apex cuspidate, mucronulate; upper surface glabrous; lower surface duller, smooth, no papillae, glabrous to sparsely sericeous, domatia absent (to a single pocket on basiscopic side in axil of + second nerve); venation on upper side slightly sunken to flat, raised below; nerves 0.3—2.2 cm apart, marginally looped and joined, less distinctly so in lower part of leaf- lets; veins densely reticulate, rather conspicuous. Infructescences axillary (to pseudoterminal), branching basally and along the terete, subtomen- tose, 2.7—6 cm long axis; first order branches up to 4 cm long; bracts and bracteoles triangular, out- side sericeous, inside glabrous; bracts 0.7—-0.9 mm long; bracteoles 0.4-0.5 mm long; pedicels 1.8— 2.2 mm long, completely slightly sericeous. Flow- ers unknown. Sepals 5, ovate, margin pilose, with- out glands, outside and inside glabrous; 2 outer smaller ones 1—1.2 by 0.9—1.2 mm; 3 inner larger ones c. 1.5 by 1.5 mm, margin petaloid. Petals, see note. Disc uninterrupted. Stamens and pistil un- known. Fruits with 1 or 2 well developed lobes, 0.9-1 by 0.8-1.3 cm, rugose, glabrous, red when fresh, blackish when dry; stipe 1.5—2.5 mm high, broadly obconical; margin blunt; lobes 8-9 by 6— 6.5 mm; septa complete. Seeds obovoid, c. 6.5 by 5 mm; hilum c. 1.1 mm long. Distribution — Malesia: trian Jaya (Jayapura). Habitat & Ecology — Rain forest substage to seral rain forest of river banks; altitude 850-1200 Flora Malesiana ser. I, Vol. 11 (3) (1994) m. Fr. Feb.—Mar. Note — One petal has been found, but it is not known whether this is one of the well developed petals or the usually more reduced one between the two adjacent large sepals. The petal is ovate, c. 2.8 by 1 mm, claw c. 0.2 mm high, margin slightly pilose, outside and inside glabrous; scales c. 0.6 mm long, free; crest absent. 18. Guioa membranifolia Radlk., Sapind. Holl.- Ind. (1879) 11, 40; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 614 (typification); in Rech., Denkschr. K. Ak. Wiss. M.-N. KI. Wien 89 (1913) 573, t. 6a, f. 11; Bot. Jahrb. 56 (1921) 282; Lauterb., Bot. Jahrb. 62 (1929) 555; Radlk. in Engl., Pflanzenr. 98 (1933) 1174; Kanehira & Hatusi- ma, Bot. Mag. Tokyo 57 (1947) 77; Welzen, Leiden Bot. Series 12 (1989) 232, f. 92. — Type: Beccari PP 807 (FI holo, cited by Radlkofer as Beccari 9), Irian Jaya. Tree, 6-10 m high, dbh up to 10 cm; bark grey. Branchlets shortly sericeous when young; flower- ing twigs 2.5-8 mm long. Leaves 1—3-jugate; ra- chis 4.2-19.5 cm long, terete, glabrous, petiole 1.7— 6.4 cm long. Leaflets subsessile, opposite to alter- nate, elliptic, 3.5-16.2 by 1.5—6.4 cm, index 2.3— 3.8, usually slightly asymmetrical, acroscopic side broader, subcoriaceous, punctate; base attenuate; margin entire, flat; apex acute to cuspidate, mucro- nulate; upper surface glabrous; lower surface dull- er, smooth, no papillae, glabrous, domatia one small sac on basiscopic side in axil of + second nerve; venation on upper side flat, raised below; nerves (0.22.3 cm apart, marginally looped and joined, less distinctly so in lower part of leaflets; veins laxly to densely reticulate, rather inconspicuous. Inflorescences ramiflorous to axillary, branching basally and along the flattened to terete, sericeous, 1.9-12.3 cm long axis; first order branches up to 5.2 cm long; cymules cincinnate, 3- or 4-flowered; bracts and bracteoles triangular, outside hirsute, inside glabrous; bracts 0.5—0.8 mm long; bracte- oles 0.2-0.4 mm long; pedicels 2—5.7 mm long, sericeous except for the subglabrous articulate part. Flowers 3.5—4 mm in diam., fragrant. Sepals 5, ovate, margin pilose, with glands, outside and in- side glabrous; 2 outer smaller ones I—1.5 by 11.8 mm, 3 inner larger ones 1.4—3.2 by 1.4-3 mm, margin petaloid. Petals 5, elliptic, 1.43 by 0.7— 1.4 mm, white; claw 0.2—0.3 mm high; margin pi- lose, outside and inside subglabrous, apex acute; scales 0.8—2 mm long, free, basally not auriculate, membranous margin indistinct; crest clavate, stip- itate, apex lobed, pilose. Disc interrupted. Stamens 8; filaments 1.8—3.5 mm long, pilose, especially Adema, Leenhouts, Van Welzen — Sapindaceae basally; anthers 0.4—-0.8 mm long, glabrous. Pis- til: ovary 0.2-0.8 mm long, subhirsute; style and stigma 0.1—0.7 mm long. Fruits with 1-3 well de- veloped lobes, 1.4—2.4 by 1—3.6 cm, completely dehiscent, smooth to slightly rugose or ribbed, gla- brous, red when fresh, black when dry; stipe 2-4 mm high, slender; suture about flat to slightly con- vex; margin blunt; lobes 10.5—16 by 10-13 mm; septa complete. Seeds obovoid, 8.5—9.7 by 6.5—7.2 mm; hilum c. 1.7 mm long; arillode with pseudo- funicle. Distribution — Malesia: Moluccas (Halmahera, Morotai); Irian Jaya (Vogelkop, Geelvinck Bay, Jayapura). Habitat & Ecology — In fringing rain forest; al- titude 30-600 m. Fl. May—Aug.; fr. Mar.—May. Note — Guioa membranifolia is part of the G. rigidiuscula-complex. Typical for this species are the elliptic leaflets without papillae and hairs, but with a single sac. The fruit is always clearly stipi- tate and the upper margin of the lobes is at most slightly convex. 19. Guioa misimaensis Welzen, Blumea 33 (1988) 418, pl. 9; Leiden Bot. Series 12 (1989) 235, f. 94. — Type: Brass 2766] (L holo; K; LAE, n.v.), Papua New Guinea. Tree, up to 20 m high, dbh up to 30 cm. Branch- lets shortly sericeous when young; flowering twigs c. 4 mm thick. Leaves 1- or 2-jugate; rachis 4.2— 7.6 cm long, flattened above, subsericeous, peti- ole 2.7-4.7 cm long; petiolules up to 0.9 cm long. Leaflets subopposite, elliptic, 10.2-13.9 by 3.9- 4.7 cm, index 2.6—3, symmetrical, coriaceous, punctate; base attenuate; margin entire, flat; apex acute to acuminate, mucronulate; upper surface glabrous: lower surface dull, papillate, short seri- ceous, domatia absent; venation raised on both sides, concolorous with lamina; nerves 0.4—1.8 cm apart, marginally looped and joined; veins laxly reticulate, indistinct. Inflorescences axillary, branching along the flattened, subsericeous, 4.6— 13.7 cm long axis; first order branches up to 4.9 cm long; cymules cincinnate, c. 3-flowered; bracts and bracteoles triangular, outside subsericeous, inside glabrous; bracts c. 0.7 mm long; bracteoles c. 0.3 mm long; pedicels 2.5—3 mm long, subseri- ceous except for the (sub)glabrous articulate part. Flowers in bud. Sepals 5, ovate, margin pilose, outside and inside glabrous; 2 outer smaller ones 1-1.3 by c. 1.3 mm, margin with glands; 3 inner larger ones 1.8—2 by 2—2.2 mm, margin petaloid, without glands. Peta/s 5, immature, elliptic, blade gradually decurrent into the claw, margin pilose, outside and inside glabrous, apex acute, white; scales free; crest clavate, shortly stipitate, glabrous. 579 Disc interrupted. Stamens 8, immature; filaments pilose, especially basally; anthers glabrous. Pistil: ovary subhirsute. Fruits unknown. Distribution — Malesia: Papua New Guinea (Milne Bay Prov.: Misima L.). Habitat & Ecology - Common. In subcanopy of rain forest; altitude c. 150 m. Fr. Aug. Note — Guioa misimaensis is part of the G. ri- gidiuscula-complex. This species may be distin- guished by its symmetrical leaflets with papillae and short subsericeous hairs, its crested scales, and its interrupted disc. 20. Guioa molliuscula Radlk., Bot. Jahrb. 50 (1914) 76; 56 (1921) 280; in Engl., Pflanzenr. 98 (1933) 1163; Welzen, Leiden Bot. Series 12 (1989) 236, f. 95. — Type: Schlechter 19521 (By holo; K, P), Papua New Guinea. Tree, c. 10 m high. Branchlets velutinous; flow- ering twigs 2-4 mm thick. Leaves 1- or 2-jugate; rachis 2.2—9.5 cm long, terete, velutinous, petiole 1.1-4.8 cm long; petiolules up to 0.9 cm long. Leaf- lets usually subsessile, subopposite, ovate (to el- liptic), 6.6-17.8 by 3.4—8.2 cm, index 1.9-2.3, + symmetrical, coriaceous, punctate; base attenuate; margin entire, flat; apex acuminate to cuspidate, mucronulate; upper surface especially hirsute on the venation; lower surface duller, smooth, not pap- illate, hirsute, domatia 0 or | sac on basiscopic side in axil of second nerve; venation on upper side flat, raised on lower; nerves 0.4—2.5 cm apart, margin- ally looped and joined; veins laxly reticulate, in- distinct. Inflorescences axillary, hirsute, with very young buds. Sepals 5, margin with glands. Petals 5, elliptic, apex acute, white; scales free, apex not broadened; crest clavate, stipitate, glabrous. Disc uninterrupted. Stamens 8; filaments pilose; anthers glabrous. Pistil: ovary subhirsute. Fruits unknown. Distribution — Malesia: Papua New Guinea (Morobe Prov.). Habitat & Ecology — Understorey tree in allu- vial swamp; altitude c. 10 m. Buds in Jan. and May. Notes — 1. This species is part of the G. rigidius- cula-complex and is known only from two collec- tions. These look rather different at first sight as one has much larger and less pilose leaflets. Dis- tinctive of the species are the velutinous leaflets without papillae and the uninterrupted disc. 2. The shape of the leaflets, and even the hir- sute appearance, resemble G. subsericea, of which specimens with a hirsute indumentum are occasion- ally found. The absence of papillae and the type of petal (well crested scales instead of usually crest- less auricles) are characters not found in G. sub- sericea. 3. Guioa molliuscula also resembles G. oligo- 580 tricha; both have a hirsute indumentum, lack pa- pillae (some specimens of G. oligotricha except- ed), and have a uninterrupted disc. However, G. oligotricha has a different type of petal (smaller, less well developed scales, no crest), leaflets with many pockets as domatia instead of no domatia or a single sac, and often subapical teeth along the leaflet margins. 21. Guioa multijuga Welzen, Blumea 33 (1988) 418, pl. 7a, b; Leiden Bot. Series 12 (1989) 239, f. 97. — Type: BW (van der Sijde) 4061 (L holo; K, P; MAN, n.v.), Irian Jaya. Small tree, up to 5 m high. Branchlets shortly sericeous when young; flowering twigs 2.54.5 mm thick. Leaves (3—)5—7-jugate; rachis 7.3-14.7 cm long, terete, not to very slightly winged, subgla- brous to subsericeous, petiole 1.6—4.5 cm long; petiolules up to 0.6 cm long. Leaflets opposite to subopposite, (ovate to) elliptic, 3.8-6.3 by 1-1.7 cm, index 3.7—4.7, asymmetrical, acroscopic side broader, coriaceous, punctate; base attenuate; mar- gin entire, flat; apex acute to cuspidate, mucronu- late; upper surface glabrous; lower surface dull, papillate, very sparsely subsericeous, domatia at least in some leaflets a single small sac on basi- scopic side in axil of second nerve; venation on upper side flat, except for the raised midrib, raised below; nerves 0.2—0.9 cm apart, marginally looped and joined, (less distinctly so in lower part of leaf- lets); veins laxly reticulate, indistinct. Inflorescenc- es axillary, branching basally and along the terete, slightly sericeous, 0.5—6.5 cm long axis; first or- der branches up to 0.9 cm long; cymules cincin- nate, c. 2-flowered; bracts and bracteoles triangu- lar, outside pilose, inside glabrous; bracts c. | mm long; bracteoles c. 0.4 mm long; pedicels 2.1—2.5 mm long, sericeous except for the subglabrous ar- ticulate part. Flowers c. 2.8 mm in diam., smelling sweet. Sepals 5, ovate, margin pilose, outside and inside glabrous; 2 outer smaller ones 1.2—-1.5 by 0.8—-0.9 mm, margin with glands; 3 inner larger ones 1.5-1.8 by 1.3—-1.5 mm, margin petaloid, without glands. Petals 5, obovate, 1.5—1.7 by 0.3-0.5 mm, white; claw c. 0.3 mm high; margin densely pi- lose, outside and inside (sub)glabrous, apex acute; scales c. | mm long, free, folded outwards, apex not broadened; crest absent; petal between two adjacent large sepals not reduced. Disc uninterrupt- ed. Stamens 8; filaments c. 2.3 mm long, pilose, especially basally; anthers 0.3—0.4 mm long, sparsely pilose. Pistil: ovary c. 0.7 mm high, sub- hirsute; style and stigma c. 0.5 mm long. Fruits unknown. Distribution — Malesia: Irian Jaya (around Jayapura). Flora Malesiana ser. I, Vol. 11 (3) (1994) Habitat & Ecology — Rare; in old secondary forest and at edge rain forest-savannah on steep terrain. Soil: once recorded from clayey sand; alti- tude 100-185 m. Bud: Mar.; fl. Sept. Note — Guioa multijuga looks much like G. sub- sericea; see also note 3 under the latter. 22. Guioa myriadenia Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1610; Philipp. J. Sc., Bot. 8 (1914) 446; Merr., Enum. Philipp. Flow. PI. 2 (1923) 508; Radlk. in Engl., Pflanzenr. 98 (1933) 1168; Welzen, Leiden Bot. Series i2 (1989) 240, f. 98. — Lectotype (Welzen, 1989): Elmer 8704 (M holo; FI, K, L, NY, W), Philip- pines. Guioa falcata Radlk., Philipp. J. Sc., Bot. 8 (1914) 446, 461; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 507; Radlk. in Engl., Pflanzenr. 98 (1933) 1167. — Type: Elmer 5869 (M holo; K, NSW, NY, P), Philippines. Guioa obtusa Merr., Philipp. J. Sc., Bot. 12 (1917) 276; Enum. Philipp. Flow. Pl. 2 (1923) 508; Radlk. in Engl., Pflanzenr. 98 (1933) 1170. Type: BS (Ramos & Edafio) 26636 (PNH* holo; A, K, NY, P), Philippines. [Guioa ferruginea Merr. ex Salvosa, Lex. Philipp. Pl. (1963) 105, nom. nud., based on BS (Ra- mos) 42181.) Shrub to tree, 2-9 m high, dbh up to 6 cm; bark greyish white to mottled, hard; sapwood white, very hard. Branchlets golden (sericeous to) hirsute, es- pecially when young; flowering twigs 1.5—6 mm thick. Leaves 2-5-jugate; rachis 1.8—16.2 cm long, basally terete to upwards dorsoventrally flattened, subsericeous to hirsute, petiole 1-6.3 cm long; petiolules up to 0.9 cm long. Leaflets opposite, (to subopposite), ovate (to elliptic), often falcate, 2.5— 14.5 by 1-4.5 cm, index 2.3—-4.2, usually asym- metrical, acroscopic side broader, coriaceous, punc- tate; base attenuate; margin entire, (flat to) revo- lute; apex (rounded to) acuminate (to cuspidate), usually mucronulate; upper surface usually subseri- ceous to appressedly hirsute; lower surface dull, papillate, long sericeous to hirsute, domatia small, (1 to many sacs to) many pockets in axils of nerves; venation on upper side (slightly sunken to) flat (to raised), below raised; nerves 0.2—1.8 cm apart, marginally looped and joined; veins densely retic- ulate, distinct. Inflorescences axillary to pseudo- terminal, branching basally and along the flattened, densely golden (sericeous to) hirsute, 1.7—15.7 cm long axis; first order branches up to 7.6 cm long; cymules cincinnate, 2—4-flowered; bracts and bracteoles triangular, outside sericeous, inside sub- glabrous; bracts 1.5—2.5 mm long; bracteoles 0.5— 1.8 mm long; pedicels 2.8-8 mm long, sericeous, Adema, Leenhouts, Van Welzen — Sapindaceae 581 less so in articulate part. Flowers 3.5—4 mm in diam., not fragrant. Sepals 5, ovate, margin and outside pilose, margin with glands, inside glabrous, whitish; 2 outer smaller ones 1.6—2.5 by 1.1—2.4 mm; 3 inner larger ones 2—3.2 by 2—3.2 mm, mar- gin petaloid. Petals 5, obovate, 2.5—4 by 1.3-1.7 mm, white to red; blade orbicular, abruptly clawed, latter 0.8—2 mm high, margin pilose, outside and inside (sub)glabrous, apex retuse to obtuse; scales 1.2-1.7 mm long, free; crest clavate, shortly stipi- tate, apex lobed, subglabrous, yellow. Disc inter- rupted, greenish yellow. Stamens 8; filaments 2— 4.4 mm long, pilose, especially basally, white; an- thers 0.3—0.6 mm long, glabrous, red. Pistil: ovary 0.4-2.3 mm long, subhirsute; style and stigma 0. 1— 1.5 mm long. Fruits with 1-3 well developed lobes, 1.1-1.4 by 1.1—1.8 cm, smooth, glabrous, black- ish when dry; stipe 2.5—-4 mm high, slender; mar- gin blunt; lobes 8—9 by 7-10 mm. Seeds obovoid, c. 6.2 by 4.8 mm; hilum c. 1.9 mm long. Distribution — Malesia: Philippines (Luzon). Habitat & Ecology — In primary forest, forest- ed slopes, mossy forest, along pine crest; altitude 270-1700 m. Fl. Dec.—Apr.; fr. July. Notes — 1. This species can be distinguished by its ovate leaflets, reticulate venation, lower surface with sericeous to hirsute hairs, usually many pock- ets, punctation, sepals that are sericeous outside, and petals with a long claw, orbicular blade, and stipitate, clavate crests. Several forms are present, three of which are rather distinct. One form with leaflets with a sericeous lower surface occurs in the Benguet and Mountain Provinces. This form may be subdivided, some specimens have rather symmetrical leaflets, viz. G. myriadenia s.s., oth- ers have more falcate leaflets, viz. ‘G. falcata’. The second form, with hirsute indumentum, occurs in Rizal Province. Transitions between the two pre- vent the recognition of different species. The third form was known as G. obtusa (Tayabas Province) and has leaflets with a densely sericeous lower surface and obtuse apex. Only in the shorter apex it differs from the northern sericeous form with which it has a disjunct distribution (i.e.. Benguet and Mountain Prov. are north and Tayabas south of Rizal). 2. Some specimens of G. myriadenia look re- markably like the ‘G. lasiothyrsa’ form of G. pleu- ropteris. Guioa myriadenia has always a rather golden indumentum, not dull brown like G. pleu- ropteris, while the venation is denser and usually more distinct. The crest is also different, that of G. pleuropteris, if present, is part of a flat bifid scale apex, while that of G. myriadenia is clavate. The fruits of G. pleuropteris can be reddish when dry, those of G. myriadenia are black. 23. Guioa normanbiensis Welzen, Blumea 33 (1988) 418, pl. 6a—c; Leiden Bot. Series 12 (1989) 243, f. 99. — Type: Brass 25521] (L holo; K), Papua New Guinea. Tree, 7-20 m high, dbh at least 5 cm; bark brown; wood medium hard. Branchlets at most only sericeous when young; flowering twigs 3-13 mm thick. Leaves 2—4-jugate; rachis 4-17.7 cm long, terete, glabrous, petiole 3.5—7 cm long. Leaflets subsessile, opposite to alternate, ovate, 7—19.3 by 2.6—6.5 cm, index 2.7—3.2, + symmetrical, other- wise acroscopic side broader, coriaceous, (punc- tate); base attenuate; margin entire, flat; apex acu- minate to cuspidate, mucronulate; upper surface glabrous; lower surface duller, smooth, no papil- lae, glabrous (to very sparsely sericeous), domatia a single small sac on basiscopic side in axil of sec- ond nerve; venation on upper side flat (to raised), raised below; nerves 0.3—3.3 cm apart, marginally looped and joined, less distinctly so in lower part of leaflets; veins laxly reticulate, indistinct. /nflo- rescences ramiflorous to axillary, branching basally and along the flat to terete, subglabrous, 1.5—6.8 cm long axis; first order branches up to 2 cm long; cymules cincinnate, c. 3-flowered; bracts and bracteoles triangular, outside hirsute, inside gla- brous; bracts 0.7—1.1 mm long; bracteoles 0.3—0.8 mm long; pedicels 1.8—4 mm long, subglabrous except for the glabrous articulate part. Flowers c. 3.5 mm in diam. Sepals 5, ovate, margin pilose, outside and inside glabrous; 2 outer smaller ones 1—1.6 by 1.2—1.3 mm, margin with glands; 3 inner larger ones 1.9-2.8 by 1.7—2.7 mm, margin peta- loid, without glands. Petals 5, obovate, c. 2 by 0.6— 0.8 mm, white; claw c. 0.3 mm high; margin pi- lose, outside and inside (sub)glabrous, apex acute; scales 1.2—1.3 mm long, free, basally not auricu- late, membranous margin indistinct; crest clavate, stipitate, apex lobed, glabrous. Disc interrupted. Stamens 8; filaments 2.7—3.1 mm long, pilose, es- pecially basally; anthers c. 0.6 mm long, glabrous. Pistil: ovary c. 0.3 mm long, subhirsute; style and stigma c. 0.2 mm long. Fruits with 1—3 well devel- oped lobes, 1.2—1.5 by 1.3—2.2 cm, completely dehiscent, smooth to somewhat ribbed, glabrous, blackish when dry; stipe 3—3.5 mm high, slender; margin blunt; lobes 11-15 by 7.5—9.5 mm, septa complete. Seeds immature; arillode with pseudo- funicle. Distribution — Malesia: Papua New Guinea (Milne Bay Prov., Normanby I.). Habitat & Ecology — Frequently in lowland rain forest, along gullies; altitude 3-200 m. FI. Sept. Note — Guioa normanbiensis is part of the G. rigidiuscula-complex. It has ovate leaflets, (almost) 582 Flora Malesiana ser. I, Vol. 11 (3) (1994) without indumentum and without papillae, a sin- gle sac, and fruits with a long slender stipe and slender lobes. 24. Guioa novobritannica Welzen, Blumea 33 (1988) 419, pl. 17a, b; Leiden Bot. Series 12 (1989) 244, f. 100. — Type: NGF (Frodin) 26918 (L holo; BM, BRI, CANB; LAE, n.v.), Papua New Guinea. Tree, c. 20 m high, dbh up to 20 cm; buttresses absent; bark brown, moderately fissured, slightly scaly in places, red-brown; wood straw; odour and exudate absent. Branchlets sericeous when young; flowering twigs 2—2.5 mm thick. Leaves 1- or 2- jugate; rachis 1.2-4.3 cm long, flattened above, glabrous, petiole 1.3—2.7 cm long. Leaflets subses- sile, opposite to subopposite, elliptic, 4.2—4.9 by 1.1—1.7 cm, index 2.9-3.8, asymmetrical, acroscop- ic side broader, coriaceous, punctate; base attenu- ate; margin entire, flat; apex acuminate, mucronu- late; upper surface glabrous; lower surface dull, papillate, glabrous except for a few hairs near domatium, latter a single, large, highly domed sac on basiscopic side in axil of + second nerve; vena- tion on upper side flat, raised below; nerves 0.2— 0.8 cm apart, marginally looped and joined, less distinctly so in lower part of leaflets; veins dense- ly reticulate, distinct. Inflorescences axillary, branching along the flattened, subsericeous, 3-10 cm long axis; first order branches up to 4.8 cm long; cymules cincinnate, c. 2-flowered; bracts and bracteoles triangular, outside sericeous, inside gla- brous; bracts 0.7—0.8 mm long; bracteoles 0.2—0.4 mm long; pedicels c. 3 mm long, sericeous except for the (sub)glabrous articulate part. Flowers c. 3.5 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside glabrous; 2 outer small- er ones c. 1.5 by 1.7 mm; 3 inner larger ones 1.8— 2.5 by c. 1.8 mm, margin petaloid. Petals 5, ellip- tic, 3-3.2 by 1.2-1.3 mm, white; claw c. 0.8 mm high; margin pilose, outside and inside glabrous, apex obtuse to acute; scales 1.5—1.8 mm long, free; crest clavate, stipitate, apex lobed; petal between two adjacent large sepals not reduced in size. Disc interrupted. Stamens 8; filaments 1.7—1.8 mm long, pilose, especially basally; anthers c. 0.4 mm long, glabrous. Pistil: ovary c. 1.7 mm long, sub- hirsute; style and stigma c. 1.8 mm long. Fruits unknown. Distribution — Malesia: Papua New Guinea (W New Britain Prov.). Habitat & Ecology — In montane forest with dominant Casuarina rumphiana; altitude c. 800 m. Fl. May. Note — Guioa novobritannica belongs to the G. rigidiuscula-complex. Typical for this species are the small, falcate, papillate leaflets with a single large sac. 25. Guioa oligotricha Merr. & L.M. Perry, J. Ar- nold Arbor. 21 (1940) 512; Welzen, Leiden Bot. Series 12 (1989) 247, f. 102. — Type: Brass 8290 (A holo; BM, L), Papua New Guinea. Small tree, 5—7.5 m high, dbh c. 10 cm. Branch- lets hirsute when young; flowering twigs 2.5—11 mm thick. Leaves 1—3-jugate; rachis 2.1-8.5 cm long, terete, hirsute, petiole 1-2.7 cm long. Leaf- lets subsessile, opposite to alternate, ovate to el- liptic, 4.1-9.1 by 2-4 cm, index 2.1—2.5, symmet- rical, subcoriaceous, punctate; base acute; margin entire except usually for a few subapical teeth, flat to somewhat revolute; apex acuminate to cuspidate, sometimes mucronulate; upper surface glabrous to sparsely hirsute, midrib densely hirsute; lower sur- face duller, smooth to papillate, hirsute, domatia many pockets in axils of nerves; venation on up- per side slightly sunken (except for the raised mid- rib), raised below; nerves 0.3—1.3 cm apart, mar- ginally looped and joined, less distinctly so in lower third of leaflets; veins laxly scalariform to densely reticulate, rather distinct. Inflorescences axillary, unbranched to branching basally on the terete, hir- sute, 1.2—-6 cm long axis; first order branches up to 1.5 cm long; cymules cincinnate, 1- or 2-flowered; bracts and bracteoles narrowly triangular, outside hirsute, inside subglabrous; bracts c. | mm long; bracteoles 0.4—0.8 mm long; pedicels 1—3.2 mm long, hirsute except for the (sub)glabrous articu- late part. Flowers c. 3 mm in diam. Sepals 5, ovate, margin and outside pilose, margin with few glands, inside glabrous; 2 outer smaller ones 1—1.3 by 0.8— | mm; 3 inner larger ones 1.4—2.2 by 1.1—1.5 mm, margin petaloid. Petals 5, obovate, 0.7—1 by c. 0.3 mm, white; claw 0—0.3 mm high; margin and less so outside and inside hirsute, apex acute; scales 0.6-0.7 mm long, free, apex not broadened; crest absent. Disc uninterrupted. Stamens 8, filaments 1.1-1.8 mm long, pilose, especially basally; an- thers c. 0.3 mm long, glabrous. Pistil: ovary c. 0.3 mm long, subhirsute; style and stigma c. 0.2 mm long. Fruits with 1-3 well developed lobes, 0.6— 0.8 by 0.81.2 cm, smooth to rugose, glabrous, red when fresh, blackish when dry; stipe 1.5-2 mm high, slender; margin blunt; lobes 7.5—8 by 6-6.5 mm. Seeds obovoid, 5-6 by 3-4 mm; hilum c. 1 mm long. Distribution — Malesia: Irian Jaya (Southern Div.) and Papua New Guinea (Western Prov.). Habitat & Ecology — Common, in secondary forest, along edge of forest, river banks. Soil: re- corded from clayey soil; altitude 50-150 m. Fl. Mar.; fr. Aug.—Nov. Adema, Leenhouts, Van Welzen — Sapindaceae Note — See note 3 under G. molliuscula and under G. acutifolia for the differences with G. oli- gotricha. 26. Guioa palawanica Welzen, Blumea 33 (1988) 419, pl. 15a, b; Leiden Bot. Series 12 (1989) 250, f. 104. — Type: BS (Foxworthy) 697 (L holo; M), Philippines. Guioa glauca auct. non Radlk.: Radlk., Philipp. J. Se., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 507; Radlk. in Engl., Pflan- zenr. 98 (1933) 1171, p.p. (Philippine plants). Shrub to tree, 1-8 m high, dbh c. 15 cm. Branch- lets sericeous when young; flowering twigs 2.5— 4.5 mm thick. Leaves 1—3-jugate; rachis 1.1—9.2 cm long, dorsally flattened, subglabrous (to seri- ceous), petiole 1—3.3 cm long; petiolules up to | cm long. Leaflets opposite to subopposite, elliptic (to obovate), 2.5—7.6 by 1—2.5 cm, index 2.1-3.4, rather symmetrical, very coriaceous, not punctate; base attenuate; margin entire, revolute; apex round- ed to acuminate, very apex rounded to acute; up- per surface glabrous except for the basally puber- ulous midrib; lower surface dull, papillate, shortly sericeous, domatia absent or | small sac on basi- scopic side in axil of + second nerve; venation raised, concolorous with lamina; nerves 0.3—1.7 cm apart, marginally looped and joined; veins laxly reticulate, rather distinct. Inflorescences axillary, branching along the flattened, sericeous, glabres- cent, 1.2—9.8 cm long axis; first order branches up to 3.9 cm long; cymules cincinnate; bracts and bracteoles triangular, outside sericeous, inside gla- brous; bracts c. 1.3 mm long; bracteoles 0.3—0.8 mm long; pedicels 3.2-6.3 mm long, sericeous. Flowers unknown. Sepals 5, ovate, margin and outside sericeous, inside glabrous, glands along margin unknown; 2 outer smaller ones |.4—2.2 by 1.9—2 mm; 3 inner larger ones c. 2.3 by 2—2.2 mm, margin petaloid. Disc interrupted. Fruits with 1-3 well developed lobes, 1.2—1.4 by 1.1-1.9 cm, smooth to rugose-ruminate to rough, glabrous, blackish when dry; stipe 2-3 mm high, slender; margin blunt; lobes 8-10 by 7-10 mm. Seeds obovoid, 7.5—8.5 by 6.2—6.8 mm; hilum 1—1.3 mm long. Distribution — Malesia: Philippines (Palawan). Habitat & Ecology — In lowland forest on ul- trabasic rock, in stunted montane rain forest with many epiphytes, and along rivers; altitude 200-815 m. Fr. Nov.—Mar. 27. Guioa parvifoliola Merr., Philipp. J. Sc. 14 (1919) 417; Enum. Philipp. Flow. Pl. 2 (1923) 508; Radlk. in Engl., Pflanzenr. 98 (1933) 1170; Welzen, Leiden Bot. Series 12 (1989) 252, f. 583 105. — Type: BS (Ramos) 33187 (PNH* holo; A, K, P), Philippines. Tree? Branchlets brownish sericeous when young; flowering twigs c. 3 mm thick. Leaves 2— 3-jugate; rachis 0.7-4.8 cm long, dorsally flattened, subsericeous, petiole 1.1—1.3 cm long; petiolules up to 0.8 cm long. Leaflets opposite, elliptic to obovate, 1.6—3.9 by 0.5—1.2 cm, index c. 3.3, slight- ly asymmetrical, acroscopic side broader, very co- riaceous, punctate; base attenuate; margin entire, revolute; apex obtuse, very apex rounded; upper surface glabrous except for the puberulous mid- rib; lower surface dull, papillate, shortly subseri- ceous, domatia a small single sac on basiscopic side in axil of + second nerve; venation raised; nerves 0.2—0.7 cm apart, marginally looped and joined: veins densely reticulate, concolorous with lamina, distinct. Infructescences axillary, not branching; rachis somewhat flattened, 2.3—3.7 cm long, sub- sericeous; bracts and bracteoles triangular, outside sericeous, inside glabrous: bracts c. 2.2 mm long; bracteoles 1.3—1.6 mm long; pedicels c. 4 mm long, sericeous, less so in articulate part. Flowers un- known. Sepals 5, ovate, margin and outside seri- ceous, inside glabrous, glands along margin un- known; 2 outer smaller ones c. 2 by 1.8 mm; 3 in- ner larger ones c. 2.6 by 2.8 mm, margin petaloid. Disc interrupted. Fruits with 1-3 well developed lobes, c. 0.9 by 1.1-1.2 cm, smooth, glabrous, blackish when dry; stipe c. 1.5 mm high, slender; margin blunt; lobes c. 7 by 6 mm. Seeds obovoid, c. 5 by 5 mm; hilum c. 1.2 mm long. Distribution — Malesia: Philippines (Luzon: Ilocos Norte Prov.). Habitat & Ecology — Dry slopes; altitude c. 700 m. Fr. Aug. 28. Guioa patentinervis Radlk., Sapind. Holl.-Ind. (1879) 11, 40; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 610, 618; in Engl., Pflanzenr. 98 (1933) 1163; Welzen, Leiden Bot. Series 12 (1989) 253, f. 106. — Lectotype (Van Welzen 1989): de Vriese & Teijsmann s.n. (L holo; BO, K, L, M), Moluccas. Guioa spec.: Merr., Philipp. J. Sc., Bot. 11 (1916) 286. — Guioa multipunctata Radlk., Philipp. J. Sc., Bot. 12 (1917) 83; in Engl., Pflanzenr. 98 (1933) 1163. — Type: Robinson 1602 (M holo), Moluccas. Treelet, 5-8 m high. Branchlets sericeous when young; flowering twigs 3—6.5 mm thick. Leaves 2—5-jugate; rachis 4.4-20 cm long, terete, glabrous, petiole 2.2-6.6 cm long. Leaflets subsessile, op- posite to alternate, (ovate to) elliptic, 4.9-13.5 by 584 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1.4-4.6 cm, index 2.4—3.8, slightly asymmetrical, acroscopic side broader, coriaceous, punctate; base attenuate; margin entire (to with a tooth subapt- cally on either side of the margin), flat; apex acu- minate to cuspidate, usually not mucronulate; up- per surface glabrous; lower surface duller, smooth, no papillae, glabrous, domatia (0 or) 1 (to many) sacs to in axils of nerves; venation flat on upper side, raised on lower; nerves 0.3—3 cm apart, mar- ginally looped and joined; veins laxly reticulate, rather distinct, especially below. Inflorescences axillary, branching basally and along the terete to flattened, sericeous, 1.6-8.2 cm long axis; first order branches up to 3.1 cm long; cymules cincin- nate, c. 3-flowered; bracts and bracteoles triangu- lar, outside sericeous, inside subglabrous; bracts 0.5-0.7 mm long; bracteoles 0.2—0.3 mm long; pedicels 1.23.2 mm long, sericeous except for the subglabrous articulate part. Flowers c. 2 mm in diam. Sepals 5, ovate, margin pilose, outside and inside glabrous; 2 outer smaller ones 0.8—-1.9 by 0.8—1.3 mm, margin with few glands; 3 inner larg- er ones 1.3—2.5 by 1.3-1.9 mm, margin petaloid, without glands. Petals 5, very reduced, elliptic, 0.8— 1.2 by 0.3-0.4 mm; claw 0.2-0.3 mm high; mar- gin pilose, outside and inside glabrous, apex ob- tuse to acute; scales 0.3—0.4 mm long, free, apex very slender; crest absent. Disc uninterrupted. Sta- mens 8; filaments 1—2.3 mm long, pilose, especially basally; anthers c. 0.3 mm long, glabrous. Pistil: ovary 0.2—0.7 mm high, subhirsute; style and stig- ma 0.1—0.7 mm long. Fruits with 1-3 well devel- oped lobes, 0.7—0.9 by 0.7—1.3 cm, smooth to some- what rugose-ruminate, glabrous, red when fresh, blackish when dry; stipe c. 2 mm high, slender; margin blunt; lobes 5—7 by 4.5-5.5 mm. Seeds obovoid, 5—6.4 by 44.8 mm; hilum 0.7—1 mm long. Distribution — Malesia: Moluccas(Ambon, Buru, Ceram, Obi). Habitat & Ecology — In transition from coral sand beach to nickel-rich soil in open forest with very little undergrowth; altitude sea level up to 300 m. Fl. Jan.—Apr.; fr. Sept.—Nov. Uses — Wood is used for the construction of houses. 29. Guioa pauciflora Radlk., Bot. Jahrb. 56 (1921) 279: Nova Guinea 11 (1926) 183; in Engl., Pflanzenr. 98 (1933) 1160; Welzen, Leiden Bot. Series 12 (1989) 255, f. 107. — Type: Leder- mann 9026 (B+ holo; K, SING), Papua New Guinea. Shrub to tree, 3-25 m high. Branchlets seri- ceous, especially when young; flowering twigs c. 1.7 mm thick. Leaves 1-jugate; rachis 0.2—-1.3 cm long, terete, not to slightly winged, sericeous, pet- iole 0.2—1 cm long. Leaflets subsessile, opposite, elliptic to obovate, 1.1—6.3 by 0.5—2.6 cm, index 2.2-3.3, + symmetrical, coriaceous to very coria- ceous, usually punctate; base attenuate; margin entire, flat to revolute; apex retuse to acuminate, sometimes mucronulate; upper surface glabrous, (wax); lower surface dull, papillate, glabrous to very sparsely sericeous, domatia absent or a single small or large sac on basiscopic side in axil of second nerve; venation raised on both sides; nerves 0.2— 1.1 cm apart, marginally looped and joined; veins densely reticulate, very conspicuous, even the vein- lets. Inflorescences axillary, unbranched to branch- ing basally on the terete, sericeous, c. 3.9 cm long axis; cymules cincinnate, 2-flowered; bracts and bracteoles immature or caducous; pedicels not yet full-grown. Flowers in bud. Sepals and petals 5, immature, white. Disc uninterrupted. Stamens 8, immature. Pisti] not measured. Fruits with 2 or 3 well developed lobes, 1.1-1.2 by 1.1—-1.2 cm, ru- gose, glabrous, blackish when dry; stipe c. 3.5 mm high, rather broadly obconical; margin blunt; lobes c. 8 by 5 mm; septa complete. Seeds unknown. Distribution — Malesia: Irian Jaya (Snow Mts); Papua New Guinea (W & E Sepik Prov.). A seem- ingly somewhat disjunct distribution; the central area of New Guinea is very poorly collected, how- ever. Habitat & Ecology — Once recorded from lime- stone rocks on the crest of a steep slope. Presuma- bly also in cloud forest (the dried specimens are moss-covered); altitude 500-2500 m. Buds and young fr. Oct. Note — The three specimens studied are rather different. However, all have 1-jugate leaves, pap- illate leaflets with a very typical venation pattern, an uninterrupted disc, and rather small fruits with a broadly obconical stipe and slender lobes. 30. Guioa pleuropteris (Blume) Radlk., Sapind. Holl.-Ind. (1879) 10; Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 520, 610; Stapf, Trans. Linn. Soc. Bot. 4 (1894) 119, 142; King, J. As. Soc. Beng. 65, II (1896) 444; Ridley, J. Str. Br. Roy. As. Soc. 33 (1900) 66; Radlk. in Perkins, Fragm. FI. Philipp. 1 (1904) 63; Lecomte in Fl. Indo-Chine 1 (1912) 1024, f. 127: 1-6; Radlk., Philipp. J. Sc., Bot. 8 (1913) 446; Bot. Jahrb. 49 (1913) 370: Merr., Enum. Born. (1921) 361; Ridley, Fl. Malay Penins. 1 (1922) 505; Merr., Enum. Philipp. Flow. PI. 2 (1923) 508; Craib, Fl. Siam. Enum. | (1926) 332; Steen., Bull. Jard. Bot. Buitenzorg. II, 12 (1931) 172, 187, 189; Radlk. in Engl., Pflanzenr. 98 (1933) 1164; Adema, Leenhouts, Van Welzen — Sapindaceae Corner, Gard. Bull. Str. Settl. 10 (1939) 45; Gagnep. in FI. Indo-Chine, Suppl. 1 (1950) 981, f. 124: 17-26; Salvosa, Lex. Philipp. Pl. (1963) 105; Meijer, Bot. News Bull. 9 (1967) 74; Martin, Introd. Ethnobot. Cambodge (1971) 90; Corner, Gard. Bull. Sing., Suppl. 1 (1978) 153; Yap in Tree Fl. Malaya 4 (1989) 442; Welzen, Leiden Bot. Series 12 (1989) 257, f. 109, 110. — Cupania pleuropteris Blume, Rumphia 3 (1847) 158; Hiern in Hook. f., Fl. Br. India 1 (1875) 677; Fern.-Vill., Nov. App. (1880) 51, p.p.; (1883) 349; Ridley, Trans. Linn. Soc. Bot. 3 (1893) 289. — Lectotype (Van Welzen 1989): Korthals s.n. (L holo; L, W), Borneo. Cupania pleuropteris var. apiculata Hiern in Hook. f., Fl. Br. India 1 (1875) 677. — Cupania grif- fithiana Kurz, J. As. Soc. Beng. 44, II (1876) 188. — Guioa pleuropteris f. apiculata Radlk. in Schmidt, Bot. Tidsskr. 32 (1915) 315. — Lectotype (Van Welzen 1989): Maingay 442 (BM holo; A, K, L), Malay Peninsula. For no- menclature see Welzen (1989) 262, note 2. Guioa aptera Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 62; Merr., Philipp. J. Sc., Suppl. 1 (1906) 87; Radlk., Philipp. J. Sc., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 1166. — Syntypes: Warburg 11504, 13108 (n.v.), Philippines. Guioa lasiothyrsa Radlk. in Perkins, Fragm. FI. Philipp. 1 (1904) 63; Merr., Philipp. J. Sc., Suppl. 1 (1906) 87; Radlk., Philipp. J. Sc., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. PI. 2 (1923) 508; Radlk. in Engl., Pflanzenr. 98 (1933) 1166. — Type: Merrill 852 (PNH?# holo; A, K, NY, SING), Philippines. Guioa subapiculata Radlk. in Perkins, Fragm. FI. Philipp. 1 (1904) 64; Philipp. J. Sc., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 506; Radlk. in Engl., Pflanzenr. 98 (1933) 1166. — Type: Merrill 513 (PNH* holo; A, K, NY), Philippines. Guioa lasiothyrsa Radlk. f. elmeri Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1609. — Syntypes: Elmer 9315 (BM, F, FI, L, K, M, W), 9342 (A, BM, BO, FI, M, NY, W), Philip- pines. Guioa forbesii Bak. f., J. Bot. 62, Suppl. (1924) 26. — Type: Forbes 2617 (BM holo; FI, L, P), Sumatra. Shrub to tree, up to 30 m high, dbh 5—60 cm (to 2 m); no buttresses; outer bark smooth to some- times irregularly fluted, usually dark brown, often mottled with white or grey spots; inner bark white to dark brown, finely fibrous; cambium yellow to brown; sapwood finely grained, soft to hard, white to yellow, without odour or sap. Branchlets espe- cially hirsute (to sericeous) when young; flower- ing twigs 1-6 mm thick. Leaves (1—)2—5S(—7)-ju- gate; rachis 0.8—25.3 cm long, (terete to) upwards flattened, usually (slightly) winged, wing up to 3 mm broad, sericeous to hirsute, petiole 0.6—9.3 cm long. Leaflets usually subsessile, opposite to alter- nate, lower often ovate, upper (elliptic to) obovate, 0.9-18.7 by 0.5-8.3 cm, index 0.5—4.3, asymmet- rical, especially the base and the apex, acroscopic side broader, (sub)coriaceous, usually punctate; base (acute to cuneate to) attenuate; margin entire, flat (to revolute); apex (obtuse to) usually abrupt- ly acuminate (to cuspidate), often mucronulate; upper surface (glabrous to) sparsely sericeous to hirsute; lower surface dull, papillate, (sub)sericeous to usually hirsute, domatia many pockets (or sacs) in axils of nerves, raised; venation on upper sur- face (slightly sunken to) flat to raised, usually con- colorous with lamina, below raised; midrib below raised, convex; nerves 0.2—4.9 cm apart, margin- ally looped and joined, less distinctly so in lower half of leaflets; veins laxly reticulate, usually dis- tinct. Inflorescences axillary (to pseudoterminal), (unbranched to) branching basally and especially along the terete, usually brown hirsute, 0.5—21.6 cm long axis; first order branches up to 13.8 cm long; cymules cincinnate, 2—5-flowered; bracts and bracteoles (deltate to) triangular, outside sericeous, inside (sub)glabrous; bracts 0.7—-3.8 mm long; bracteoles 0.2—1.5 mm long; pedicels 1.1—-7 mm long, hirsute. Flowers 34.2 mm in diam., fragrant. Sepals 5, ovate, margin and less so outside pilose, margin with glands, inside (sub)glabrous, green to tinged reddish or whitish; 2 outer smaller ones 0.7— 2.8 by 0.6—2 mm; 3 inner larger ones 1.5—3.5 by 0.9-3.3 mm, margin petaloid. Petals 5, (elliptic to) obovate, |.3—3.5 by 0.7—2.2 mm, white; blade obo- vate, gradually decurrent into the 0.2—1 mm high claw, margin (and outside) pilose, inside glabrous, apex rounded (to acute); scales 0.8—2.2 mm long, free, apex not to hardly broadened; crest usually developed, a pilose flat part of the bifid scale apex, yellow. Disc (uninterrupted to) interrupted (gap often small). Stamens 8; filaments 1.3—5.1 mm long, pilose, especially basally, white; anthers 0.2—0.7 mm long, glabrous, pink. Pistil green; ovary 0.3— 2.8 mm long, subhirsute; style and stigma 0.1—2.2 mm long. Fruits with 1-3 well developed lobes, 1—-1.9 by 1-2.5 cm, smooth to somewhat ribbed to somewhat rugose-ruminate, glabrous, red when fresh, reddish (to blackish) when dry; stipe 2—5 mm high, slender; margin blunt; lobes 7.5—13 by 6.5- 13 mm. Seeds globose to obovoid, 5.5—9.7 by 5-8 mm; hilum 1—3 mm long; arillode yellow to or- 586 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 41. Guioa pleuropteris (Blume) Radlk. a. Habit; b. petal; c. fruit (a, c: Davidson 1325; b: Maxwell 81-34). ange, edible, sour. — Figs. 38b, 41, 42. Distribution — Burma, Cambodia (Kampot), Vietnam (Poulo-Condor, Hatien), Thailand, Male- sia: E coast of Malay Peninsula, Sumatra, Borneo, Philippines. Habitat & Ecology — (Rare to) very common, scattered in lower to middle storey of primary for- est, on flat to hilly to mountainous land, on dry to periodically flooded soils (e.g. swamps). In primary and especially secondary forest; edges of forests, plantations; in open landscapes like cultivated land and belukar; in mossy forest; along riverbanks, roads, sea. On Pulau Sekindan (Corner, Gard. Bull. Sing. 38, 1985, 19) in Terminalia-Barringtonia and Eugenia grandis formations. Soil: sandstone, brown stony soil, yellow sandy loam, black sand, ultra- basic alluvial deposits; altitude sea level up to 1800 m. Fl. Aug.—Mar.(—May); Philippines: Dec._May; fr. throughout the year, mainly Feb.—Apr. Uses — Wood is used for torches (Radlkofer, 1913); it is durable and elastic, but thin, therefore used in Indonesia as handles for axes and shafts of wagons and plows (Heyne, Nutt. Pl. Indon. ed. 3, 1950, 1000). The roots are used medically in NE Adema, Leenhouts, Van Welzen — Sapindaceae 587 Pahang (Malaysia) in the form of a decoction against fever and stomach ache. The name pokok serawan burang probably refers to a medical use as ‘serawan’ = sprue (Burk., Dict. Econ. Prod. Malay Penins. ed. 2, 1966, 1134). Also used to exterminate intestinal worms. Notes — 1. Guioa pleuropteris is a rather varia- ble species, and the very character to which it owns its name can be absent. The leaflets are usually obovate with a sharply decurrent short apex (see leaflet a and f in Fig. 42, but if the latter is elongat- ed the shape becomes more elliptic, as is found in forms from Sumatra, N Borneo and Philippines. The indumentum on the lower surface of the leaf- lets is usually dense, seldom sparse, usually ve- lutinous to occasionally sericeous; the hairs are usually long, but can be short. The domatia are pockets, but an occasional pocket-like sac is also found. The normal shape of the leaflets, found on the SE Asian mainland, most of Borneo and SE Phil- ippines, is represented by the leaflets a and f in the distribution map (Fig. 42). The Philippines have several distinct forms, which cannot be separated as species. Two clines can be found in the Philip- pines (see arrows on map, Fig. 42). One cline ranges from Palawan to Culion and Busuanga I. (Calami- an group). On Palawan the leaflets can be rather large and are usually densely velutinous on the low- er surface. On the other two islands the size of the leaflets decreases dramatically, but they still remain very velutinous, this form has been described as Fig. 42. Guioa pleuropteris (Blume) Radlk. Distribution map showing two transformation series of leaf- lets (all c. - thyrsa’ form; e. ‘G. aptera’ form; f. ‘G. pleuropteris’ form (a: Davidson 1325; b: Forbes 2617; ¢: Merrill 513; d: Elmer 10880; e: Elmer 9315; f: van Niel 4263). 0.7). —a. ‘G. pleuropteris’ form; b. ‘G. forbesii’ form; c. ‘G. subapiculata’ form, d. ‘G. lasio- 588 Flora Malesiana ser. I, Vol. 11 (3) (1994) ‘G. subapiculata’ (leaflet c). The forms on Pala- wan were described as G. pleuropteris (leaflet a, f) and the wingless specimens as ‘G. lasiothyrsa’ (leaflet d). The other cline is found from Borneo via Mindanao to Panay, Negros, Leyte, Samar to Luzon and finally to Mindoro. On Mindanao the situation is rather complex, the rather large-leaved form of Palawan is found, together with a some- what smaller form of the ‘typical’ G. pleuropteris (leaflet a and f) and a third form which was de- scribed as ‘G. aptera’ and ‘G. lasiothyrsa f. elmert’ (leaflet e). This latter form is mainly found on Lu- zon, it has very asymmetrical and small leaflets, with often (few) sericeous hairs and sometimes (partly) sacs instead of pockets on the lower sur- face. All intermediates among these forms are found. Morley & Flenley (in T.C. Whitmore: Bio- geograpical evolution of the Malay Archipelago, 1987, 50-59) show in a palaeogeographical recon- struction of the Sunda-Sahul region during the middle Pleistocene that a continuous landmass existed from N Borneo to Palawan and the Cala- mian group; this land mass is covered by one of the two clines; and another landmass existed from NE Borneo via Mindanao up to Luzon and Min- doro, the area occupied by the other cline. Proba- bly the clines are the result of dispersal accom- panied by phenological change. In Kalimantan Timur (Borneo) some specimens resemble G. pterorhachis, but the midrib on the lower surface of the leaflets is convex instead of flat and hardly raised. Specimens from Sumatra were described as ‘G. forbesii’ (leaflet b) because of the narrow, elliptic, rather than broad and obo- vate leaflets, which are densely velutinous below. 2. Merrill suggested that Sapindus guisian Blan- co is synonymous with G. pleuropteris. This deci- sion is incorrect, because of the large differences between the two. The full synonymy of Sapindus guisian and a discussion of the differences can be found in Welzen (1989) 263, note 3. 3. For the difference between the ’G. lasiothyr- sa’ form of G. pleuropteris and G. myriadenia see note 2 under the latter. 31. Guioa plurinervis Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 346; Bot. Jahrb. 56 (1921) 280 (typification); in Engl., Pflan- zenr. 98 (1933) 1169; Welzen, Leiden Bot. Series 12 (1989) 264, f. 111. — Type: Mac- Gregor s.n., 1890 (M holo; MEL), E Papua New Guinea. Treelet, 5-12 m high, dbh c. 8 cm; outer bark black, inner bark pinkish brown; sapwood cream, heartwood reddish brown. Branchlets sericeous when young; flowering twigs 3.5-5 mm thick. Leaves 2—5-jugate; rachis 4.3—15.2 cm long, some- what flattened above, sericeous, petiole 3—6.1 cm long; petiolules up to 1 cm long. Leaflets opposite to alternate, ovate, falcate, 5.9-13.2 by 1.5—3.3 cm, index 3.6—4.4, asymmetrical, acroscopic side broad- er, coriaceous to very coriaceous, not punctate; base attenuate; margin entire, flat; apex cuspidate to caudate, mucronulate; upper surface glabrous ex- cept for the puberulous midrib; lower surface dull, papillate, shortly sericeous, domatia 0—2 pockets, especially on basiscopic side in axil of second nerve; venation flat above, raised below; nerves (.2—1.3 cm apart, marginally looped and joined; veins laxly reticulate, indistinct. Inflorescences axillary to pseudoterminal, branching along the flattened to terete, sericeous, 3.1—15 cm long axis; first order branches up to 4.8 cm long; cymules cincinnate, c. 4-flowered; bracts and bracteoles tri- angular, outside sericeous, inside glabrous; bracts 0.9-1.1 mm long; bracteoles 0.5—0.6 mm long; pedicels 2.5—-3 mm long, sericeous except for the (sub)glabrous articulate part. Flowers c. 3 mm in diam. Sepals 5, ovate, margin (and outside) pilose, margin with glands, inside glabrous; 2 outer smaller ones 0.9—1.2 by 1—1.3 mm; 3 inner larger ones 1.8— 2.7 by 1.5-2.5 mm, margin petaloid. Petals 5, el- liptic, 3-3.2 by 1.2-1.5 mm, blade obovate, grad- ually decurrent into the c. 0.5 mm high claw, mar- gin pilose, outside and inside glabrous, apex acute; scales 1.7—1.8 mm long, free; crest clavate, stipi- tate, glabrous. Disc interrupted. Stamens 8; fila- ments c. 4.2 mm long, pilose, especially basally; anthers c. 0.7 mm long, pilose. Pistil: ovary c. 0.6 mm long, subhirsute; style and stigma c. 0.4 mm long. Fruits with 1-3 well developed lobes, 1—1.4 by 1-1.7 cm, slightly ribbed, glabrous, red when fresh, blackish when dry; stipe c. 2 mm high, broad- ly obconical, indistinct; margin blunt; lobes 8-9 by 7-9 mm; septa complete. Seeds obovoid, c. 9 by 6 mm; hilum c. 1.2 mm long. Distribution — Malesia: Papua New Guinea (Milne Bay Prov.: Rossel Island). Habitat & Ecology — Hill forest and secondary rain forest; altitude 10-50 m. FI. July; fr. Oct. Note — Guioa plurinervis belongs to the G. ri- gidiuscula-complex. It is characterized by its fal- cate leaflets, which are papillate below. 32. Guioa pseudoamabilis Welzen, Blumea 33 (1988) 419, pl. 13a—d; Leiden Bot. Series 12 (1989) 266, f. 112. — Type: Vinas & Bellamy 261 (L holo; A, CBG, K, LAE, UPNG, WEI), Papua New Guinea. Guioa venusta auct. non Radlk.; Hartley et al., Adema, Leenhouts, Van Welzen — Sapindaceae 589 aR EOE ESS ii 5 OEE A Fig. 43. Guioa pseudoamabilis Welzen. a. Habit; b. leaflet; c. petal; d. fruit (a, b: Saunders 802; c: Vinas & Bellamy 261; d: Hartley 12560). Lloydia 36 (1973) 270; Streimann, Pl. Upper Watut Watershed (1983) 169. Shrub to tree, 2-27 m high, dbh 1-60 cm; no buttresses; outer bark red to brown, smooth, inner bark straw-coloured; wood white. Branchlets seri- ceous to somewhat hirsute, especially when young; flowering twigs 1-4 mm thick. Leaves (1—)4-6- jugate; rachis 1.5—12 cm long, winged, wing up to | mm broad, subglabrous to sericeous, petiole 0.6— 2.2 cm long. Leaflets subsessile, opposite to alter- nate, ovate to obovate, 0.8—4.3 by 0.5—2 cm, index 1-2.9, asymmetrical, acroscopic side broader, sub- coriaceous, punctate; base attenuate; margin en- tire or usually slightly crenate, flat when crenate to revolute when entire; apex retuse to obtuse (to acute), mucronulate; upper surface (pilose, espe- cially the midrib); lower surface duller, smooth, no papillae, usually sericeous, domatia in at least some leaflets one to several large, hirsute sacs in axils of subbasal nerves; venation on upper side flat to slightly raised, raised on lower; nerves 0.1— 0.6 cm apart, marginally looped and joined; veins densely reticulate, very inconspicuous. /nflores- cences axillary to pseudoterminal, unbranched to branching along the terete, sericeous, 2.5—8 cm long axis; first order branches up to 3.1 cm long; cy- mules cincinnate, c. 2-flowered; bracts and bracte- oles triangular, outside sericeous, inside glabrous; bracts 0.4—-1.5 mm long; bracteoles 0.3-0.7 mm long; pedicels 3.3-6 mm long, sericeous, less so in articulate part. Flowers c. 4 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside glabrous, pinkish; 2 outer smaller ones 1.1— 1.8 by 1-2 mm; 3 inner larger ones 1.9—3.2 by 1.9- 2.6 mm. Petals 5, elliptic, c. 3.4 by 1.2 mm, pink- ish white; claw c. 0.5 mm long; margin pilose, outside and inside glabrous, apex acute; scales c. 0.7 mm long, free; crest stipitate, clavate, apex lobed. Disc uninterrupted. Stamens only seen in bud, 8; filaments pilose, especially basally; anthers glabrous. Pistil only seen in bud; ovary subhirsute. Fruits with 1-3 well developed lobes, 1.1—2 by 1.7— 1.9 cm, smooth (to slightly ribbed), glabrous, red when fresh, blackish when dry; stipe 2-3 mm high, rather broadly obconical; margin blunt; lobes 10- 590 12 by 6-10 mm; septa complete. Seeds globose to obovoid, 7-9 by 6—7 mm; hilum 1.8—2.9 mm long. — Fig. 43. Distribution — Malesia: Papua New Guinea (Enga, Eastern Highlands, and Morobe Prov.). Habitat & Ecology — Occasional in submontane, montane, and moss forest with low vegetation (once recorded to be dominated by Xanthomyrtus, Myr- taceae). Soil: once found on well-drained latosol in strong shade; altitude 1800-3300 m. FI. Jan.— Apr.; fr. Oct.—Dec. 33. Guioa pteropoda Radlk., Sapind. Holl.-Ind. (1879) 11, 41; Sitzungsber. Math.-Phys. Cl. KG6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 614 (typification); Bot. Jahrb. 56 (1921) 28; in Engl., Pflanzenr. 98 (1933) 1174; Welzen, Lei- den Bot. Series 12 (1989) 268, f. 113. — Type: Beccari it. sec. 16 (FI holo), New Guinea. Guioa crenifoliola Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 514. — Type: Brass 13082 (A holo; BM, L), Irian Jaya. See note. Tree, 13-20 m high, dbh 20-25 cm. Branchlets sericeous when very young; flowering twigs 3—4 mm thick. Leaves 2—7-jugate; rachis 1.9-17.3 cm long, terete, not to slightly winged, subglabrous, petiole 1.8—4.6 cm long. Leaflets subsessile, op- posite to alternate, elliptic, 3.5—-8.5 by 1.7—3.2 cm, index 2.1—3, asymmetrical, acroscopic side broad- er, (sub)coriaceous, not punctate; base attenuate; margin laxly crenate, slightly revolute; apex ob- tusely acuminate to caudate, usually mucronulate; upper surface glabrous, (wax); lower surface dull- er, smooth, no papillae, glabrous, domatia absent to several sacs, mainly on basiscopic side in axils of nerves; venation on upper side flat except for the raised midrib, slightly raised below, also the flat midrib; nerves 0.2—1 cm apart, marginally looped and joined; veins laxly reticulate, indistinct. Inflorescences axillary, unbranched to branching along the terete, subsericeous, 2.4—13 cm long axis; first order branches up to 5.3 cm long; cymules cincinnate, c. 4-flowered; bracts and bracteoles tri- angular, outside sericeous, inside glabrous; bracts (.8—1 mm long; bracteoles 0.3—0.5 mm long; pedi- cels 3.8-8 mm long, glabrous or very sparsely pi- lose. Flowers c. 4 mm in diam. Sepals 5, ovate, margin pilose, outside and inside glabrous; 2 outer smaller ones 1.2—2.2 by 1.8—2.3 mm, margin with glands; 3 inner larger ones 3—4 by 3—4.2 mm, mar- gin petaloid, without glands. Petals 4, obovate, 2.9— 3.1 by 1.5—2.1 mm, white; claw 0.2—0.3 mm high; margin subglabrous, outside and inside glabrous, apex truncate to rounded; scales 2—2.2 mm long, free; crest usually a pilose bifid scale apex or cla- vate, glabrous. Disc interrupted, very asymmetri- Flora Malesiana ser. I, Vol. 11 (3) (1994) cal. Stamens 8; filaments 2.3-4.6 mm long, only basally pilose; anthers 0.3—0.4 mm long, glabrous. Pistil: ovary 0.4—0.6 mm long, subhirsute; style and stigma c. 0.3 mm long. Fruits with 2 or 3 well de- veloped lobes, c. 2 by 1.4 cm, smooth, glabrous, red when fresh, blackish when dry; stipe indistinct, broadly obconical; margin very sharp; lobes c. 0.8 by 1.6 cm; septa above attachment of funicle in- complete. Seeds immature. Distribution — Malesia: S Moluccas (Ceram), Irian Jaya (Geelvink Bay, Jayapura). Habitat & Ecology — Common. In primary for- est along and in flood plains, in Agathis forest. Soil: clay, sand; altitude sea level up to 900 m. Fr. Mar. Note — Brass 13082, 13702 (‘G. crenifoliola’) are somewhat different from the other specimens in having leaves with a winged rachis and shorter, obtusely acute apices. 34. Guioa pterorhachis Welzen, Blumea 33 (1988) 419, pl. 12a, b; Leiden Bot. Series 12 (1989) 269, f. 114. — Type: Elmer 20268 (L holo; BM, F, K, M, NSW, P, U), Sabah. Guioa pleuropteris auct. non Radlk.; Merr., Pl. Elm. Born. (1929) 175. Shrub to tree, 3-16 m high, dbh 10 cm to 1.15 m; outer bark smooth, flaked, white to brown, soft, inner bark fibrous, pink to brown; cambium yel- low; sapwood white to brown. Branchlets glabrous, at most sparsely sericeous when young; flowering twigs 2-5 mm thick. Leaves 3—7-jugate; rachis winged, wing up to 4 mm broad, (sub)glabrous, petiole 1.1—12.2 cm long. Leaflets subsessile, op- posite to subopposite, elliptic (to obovate), 2.4— 14.6 by 1-4.6 cm, index 1.9—3.9, often slightly asymmetrical, especially basally, then acroscopic side broader, (sub)coriaceous, punctate; base acute to attenuate to cuneate; margin entire, flat; apex gradually acuminate to cuspidate, sometimes mucronulate; upper surface glabrous (to puberu- lous on the midrib and venation; lower surface dull, papillate, (very) sparsely, usually shortly sericeous, domatia many sunken sacs in axils of nerves; ve- nation on upper surface (slightly sunken to) flat to raised, raised on lower; midrib below hardly raised, flat; nerves 0.2—3.6 cm apart, marginally looped and joined, less so in lower half of leaflets; nerves laxly reticulate, rather indistinct. Inflorescences axillary (to pseudoterminal), (unbranched to) branching basally and along the terete, subseri- ceous, 1—20.8 cm long axis; first order branches up to 11.5 cm long; cymules cincinnate, 2—5-flow- ered; bracts and bracteoles deltate to triangular, outside sericeous, inside (sub)glabrous; bracts 0.7— 1.2 mm long; bracteoles 0.3—0.8 mm long; pedi- cels 2-7 mm long, sericeous. Flowers 3.5—4 mm Adema, Leenhouts, Van Welzen — Sapindaceae in diam. Sepals 5, ovate, margin (and outside) pi- lose, margin with glands, inside (sub)glabrous; 2 outer smaller ones 1—1.9 by 0.8—1.5 mm; 3 inner larger ones |.7—3 by 1.2—2.7 mm, margin petaloid. Petals 5, elliptic (to obovate), 1.1—2.8 by 0.8—1.6 mm, white; claw 0.2—0.5 mm high; margin pilose, outside and inside glabrous, apex + acute; scales 1.1—2 mm long, free; crest a pilose flat part of bi- fid scale apex. Disc interrupted to uninterrupted. Stamens 8; filaments 2-5 mm long, pilose, espe- cially basally; anthers c. 0.3 mm long, glabrous. Pistil: ovary 0.3—1 mm long, sparsely hirsute; style and stigma 0.1—1 mm long. Fruits with 1-3 well to) somewhat rugose-ruminate, glabrous, yellow to red when fresh, blackish when dry; stipe 4-7 mm high, slender; margin blunt; lobes 8.5—11 by 8-12 mm. Seeds obovoid, 7.5—10.5 by 6-9 mm; hilum 1.7—2 mm long; arillode edible, with yellow exu- date. Distribution — Malesia: Borneo (E Sabah: Sandakan & Tawau). Habitat & Ecology — In understorey of primary and especially secondary forest, along rivers and roadsides, on flat to undulating country. Soil: white, yellow, or black sands; altitude sea level up to 30 (—500) m. Fl. (July—)Nov.—Jan.; fr. Nov.—May. Uses — Firewood. Note — Guioa pterorhachis resembles G. pleu- ropteris closely; specimens of the latter from Ka- limantan Timur (Borneo) may differ only slightly from G. pterorhachis in having a raised and con- vex instead of a flat and hardly raised midrib on the lower surface of leaflets. 35. Guioa pubescens (Zoll. & Mor.) Radlk., Sit- zungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 302; Sapind. Holl.- Ind. (1879a) 10, 41 (lectotypification); Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879b) 543, 612; King, J. As. Soc. Beng. 65, II (1896) 445; Ridley, J. Str. Br. Roy. As. Soc. 33 (1900) 66; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 210; Koord., Exk. Fl. Java 2 (1912) 541; Radlk., Philipp. J. Sc., Bot. 8 (1913) 446; in Elmer, Leafl. Philipp. Bot. 5 (1913) 1616; Merr., Enum. Born. (1921) 361; Ridley, Fl. Malay Penins. | (1922) 506; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 509; RadIk. in Engl., Pflan- zenr. 98 (1933) 1169; Corner, Gard. Bull. Str. Settl. 10 (1939) 45; Wayside Trees (1940) 588, pl. 178; Desch, Mal. For. Rec. 15 (1954) 526; Backer & Bakh. f., Fl Java 2 (1965) 140; Burk., Dict. Econ. Prod. Malay Penins. ed. 2 (1966) 1134; Keng, Gard. Bull. Sing. 35 (1982) 90; 391 Yap in Tree Fl. Malaya 4 (1989) 442; Welzen, Leiden Bot. Series 12 (1989) 272, f. 115, 116. — Sapindus pubescens Zoll. & Mor. in Mor., Syst. Verz. (1846) 22, p.p. (Zollinger 1105). — Lectotype (Radlkofer 1879a): Zollinger 1105 (L holo; A, BM, FI, P), Java. Arytera silaka Miq., Sumatra (1861) 199, 510. — Type: Teijsmann HB 610 (U holo; BO), Suma- tra. Cupania pallidula Hiern in Hook. f., Fl. Br. India 1 (1875) 676; Ridley, Trans. Linn. Soc. Bot. 3 (1893) 289; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 509. — Syntypes: Griffith s.n. (K); Maingay s.n. (K), Malay Peninsula. Guioa diplopetala (Hassk.) Radlk. f. dentata Radlk., Sapind. Holl.-Ind. (1879) 88; Sitzungs- ber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 610; in Engl., Pflanzenr. 98 (1933) 1162. — Type: Teijsmann HB 3741 (BO holo), Sumatra. Guioa perrottetii auct. non Radlk.: Meijer, Bot. News Bull. 9 (1967) 75. Tree(let), 2-25 m high, dbh 1.3—85 cm; with- out buttresses; outer bark smooth, sometimes deep- ly fissured, hard, (greenish to) flaky greyish white or greyish brown to dark brown, inner bark yel- lowish white to reddish brown; cambium white; sapwood soft to hard, white to ochre; heartwood red. Branchlets sericeous when young, hairs brown; flowering twigs 1.5—6 mm thick. Leaves (1—)2-6- jugate; rachis 2.1—29.5 cm long, terete (to upwards slightly flattened), subsericeous, petiole 1.4—10.8 cm long; petiolules up to | cm long. Leaflets op- posite to alternate, ovate (to elliptic), often slight- ly falcate, 2.9-19.2 by 0.8—7.2 cm, index 1.3-5.5, especially basally asymmetrical, acroscopic side broader, coriaceous, not punctate; base attenuate; margin entire, flat (to revolute); apex gradually acuminate to cuspidate (to caudate), usually mucro- nulate; upper surface (glabrous to) sparsely short- ly sericeous, (wax); lower surface dull, papillate, (slightly) shortly sericeous, domatia (0 or) 1 small sac to many sacs in axils of (basal) nerves; vena- tion on upper surface (slightly sunken to) flat to raised, below usually raised, concolorous with lam- ina; nerves 0.3—3.3 cm apart, marginally looped and joined, (less so in lower third of leaflets); veins laxly reticulate, rather indistinct. /nflorescences axillary (to pseudoterminal), (unbranched to) branching basally to especially along the terete, brown sericeous, 1.4—24.2 cm long axis; first order branches up to 9.3 cm long; cymules cin- cinnate, 2—4-flowered; bracts and bracteoles deltate to triangular, outside sericeous, inside (sub)glabrous; bracts 0.6—1.8 mm long; bracteoles 592 Flora Malesiana ser. I, Vol. 11 (3) (1994) iE ————=E a (.2-1.2 mm long; pedicels 1.8-8 mm long, seri- ceous. Flowers 3.5—4.5 mm in diam.; buds green. Sepals 5, ovate, margin and often outside sericeous, margin with glands, inside (sub)glabrous, green to reddish; 2 outer smaller ones 1—2.8 by 0.8—2.1 mm; 3 inner larger ones 1.4—3.3 by 1.3—3.1 mm, mar- gin petaloid. Petals 5, obovate, 1.9-3.4 by 0.7-1.8 mm, white to yellow; blade obovate, gradually de- current into the 0.3—1.2 mm long claw, margin (and outside) pilose, inside glabrous, apex rounded (to acute); scales 1.1-2 mm long, free, apex not to hardly broadened; crest a pilose flat part of the bi- fid scale apex. Disc interrupted. Stamens 8, fila- ments 1.9—5.2 mm long, pilose, especially basally, white; anthers 0.2—0.4 mm long, glabrous, pink. Pistil: ovary 0.3-2 mm long, densely hirsute, yel- lowish green; style and stigma 0.1—2.5 mm long. Fruit with 1-3 well developed lobes, 1—-1.5 by 1— 1.9 cm, smooth (to somewhat ribbed), (very) sparse- ly sericeous, glabrescent, red when fresh, (reddish to) blackish when dry; stipe 1.5—5 mm high, slen- der: margin blunt; lobes 7-10 by 6.5—10 mm. Seeds globose to obovoid, 6.8—10 by 5.8—7.5 mm; hilum 1.1-2 mm long. Distribution — Malesia: W Malaysia, Singapore, Sumatra, Bangka, W Java (Djawa Barat), Karimun- djawa Island, Borneo (above equator), Philippines (Palawan). Habitat & Ecology — Rare to rather common in primary and especially secondary forest, kerangas, peat swamp forests, forest along sea. Soil: sand, sandy loam, sandstone, rocky soil, limestone, ul- trabasic, dry peat.; altitude sea level up to 1800 m. Fl. Singapore: July—Oct.; Java: Feb., Nov.; Borneo: Aug.—Feb.; Fr. Jan.—Apr. Uses — The wood is used as a construction tim- ber, in spite of the fact that it is brittle and the stem too narrow for sawing planks (Heyne, Nutt. PI. Indon. ed. 3, 1950, 1001); however, recorded di- ameters indicate the opposite. Note — See note 3 under G. bijuga. 36. Guioa reticulata Radlk., Philipp. J. Sc., Bot. 8 (1914) 446, 462; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 509; Radlk. in Engl., Pflanzenr. 98 (1933) 1168; Welzen, Leiden Bot. Series 12 (1989) 277, f. 118. — Lectotype (Van Welzen 1989): BS (Ramos) 7055 (M holo; NSW), Philippines. Guioa sulphurea Radlk., Philipp. J. Sc., Bot. 8 (1914) 446, 462; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 509; Radlk. in Engl., Pflanzenr. 98 (1933) 1167. — Type: FB (Alvarez) 22429 (M holo; F, L, NSW), Philippines. Tree, 7-8 m high, dbh c. 12.5 cm. Branchlets golden sericeous (to hirsute) when young; flower- ing twigs 2-4 mm thick. Leaves 2—5-jugate; rachis 2.8-17 cm long, basally terete, upwards flattened above, subglabrous, petiole 1.5—6.3 cm long; peti- olules up to 1 cm long. Leaflets opposite to alter- nate, ovate, falcate, 2.8-12 by 0.7—3 cm, index 3.5— 4.5, asymmetrical, acroscopic side broader, coria- ceous to very coriaceous, not punctate; base atten- uate; margin entire, flat to revolute; apex cuspi- date to caudate, usually mucronulate; upper sur- face glabrous to puberulous on the midrib; lower surface dull, papillate, short (to long) sericeous, domatia a single (to many) sac(s) on basiscopic side in axil of second nerve, small; venation con- spicuously raised, discolorous with lamina; nerves 0.3-2.1 cm apart, marginally looped and joined; veins densely reticulate, very distinct. Inflorescenc- es axillary, branching basally and along the usual- ly flattened, sericeous (to hirsute), 3.4-23.3 cm long axis; first order branches up to 11.2 cm long; cy- mules cincinnate (to dichasial), 2—5-flowered; bracts and bracteoles triangular, outside sericeous, inside glabrous; bracts 1-1.8 mm long; bracteoles (0.3-1.1 mm long; pedicels 2.3-5 mm long, seri- ceous except for the usually subglabrous articu- late part. Flowers 3.5-3.8 mm in diam. Sepals 5, ovate, margin pilose, with glands, (outside pilose), inside glabrous; 2 outer smaller ones 1.5—2.2 by 1.1-2.2 mm; 3 inner larger ones 2.2—2.8 by 1.6— 2.7 mm, margin petaloid. Petals 5, obovate, 2.8— 3.1 by 1-1.5 mm, blade obovate, gradually decur- rent into the 0.8—1.3 mm high claw, margin pilose, outside and inside (sub)glabrous, apex obtuse; scales 1.2-1.8 mm long, free; crest a pilose flat part of the bifid scale apex to clavate and subgla- brous. Disc interrupted. Stamens 8; filaments 1.7— 4 mm long, pilose, especially basally; anthers 0.4— 0.6 mm long, glabrous. Pistil: ovary 0.4—2 mm long, subhirsute; style and stigma 0.3—1.2 mm long. Fruits with 1 or 2 well developed lobes, 1—1.1 by 1.2-1.4 cm, smooth, glabrous, red when fresh, blackish when dry; stipe c. 2 mm high, slender; margin blunt; lobes c. 7.5 by 7 mm. Seeds obovoid, c. 7.2 by 5.8 mm; hilum c. 2 mm long. Distribution — Malesia: Philippines (Luzon). Habitat & Ecology — In secondary forest, on forested slopes. Fl. Jan.—Feb.; fr. May. Notes — 1. Several specimens have a patent, hir- sute indumentum instead of the more common ap- pressed sericeous hairs; they were described as a separate species (G. sulphurea). This phenomenon is also occasionally found in other species (e.g., G. chrysea, G. subsericea). 2. Jacobs 7983 is exceptional because the in- florescences, pedicels, and sepals are very dense- ly sericeous. Adema, Leenhouts, Van Welzen — Sapindaceae 37. Guioa rigidiuscula Radlk., Sapind. Holl.-Ind. (1879) 11, 41; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 614 (typification); ibid. 20 (1890) 361; K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1900) 420; Radlk., Bot. Jahrb. 56 (1921) 282; Baker in Rendle, J. Bot. 61, Suppl. (1923) 11; Laut., Bot. Jahrb. 62 (1929) 555; Radlk. in Engl., Pflanzenr. 98 (1933) 1173; Rehder, J. Arnold Arbor. 14 (1933) 63; P. Royen, Man. For. Trees Papua & New Guinea 2 (1964) 3, f. 1g; Welzen, Leiden Bot. Series 12 (1989) 281, f. 120. — Type: Beccari FI 2804 (FI holo, cited by Radlkofer as Beccari 8), New Guinea. Shrub to tree, 2-15 m high, dbh 10—24 cm; outer bark smooth, blotched variously grey to brown, inner bark brown, sap red; wood white to dark cream. Branchlets shortly sericeous when young; flowering twigs 2-6 mm thick. Leaves 1—4-jugate; rachis 2.2-17.2 cm long, terete (to somewhat flat- tened below jugae), subglabrous, petiole 1.3—8.2 cm long. Leaflets subsessile, opposite to alternate, elliptic, 3.6-17.1 by 1.2-6.3 cm, index 2.1-3.8, subsymmetrical, acroscopic side slightly broader, (sub)coriaceous, usually punctate; base attenuate; margin entire, flat; apex (acute to) acuminate to cuspidate (to caudate), usually mucronulate; up- per surface glabrous, (puberulous on midrib); lower surface duller, smooth, no papillae, subglabrous, domatia in at least some leaflets a single small sac on basiscopic side in axil of second nerve; vena- tion on upper side (slightly sunken to) flat (to raised), raised below; nerves 0.2—3.8 cm apart, marginally looped and joined, often less distinctly so in lower part of leaflets; veins laxly reticulate, indistinct. Inflorescences ramiflorous (to axillary), unbranched to branching basally and along the terete, sericeous, 1—13.3 cm long axis; first order branches up to 6.2 cm long; cymules cincinnate to dichasial, (1—)3—4(—5)-flowered; bracts and bracte- oles triangular, outside hirsute, inside glabrous; bracts 0.4—1 mm long; bracteoles 0.2—0.6 mm long; pedicels 1.9—5.5 mm long, sericeous except for the subsericeous articulate part. Flowers 3.5—4 mm in diam., slightly fragrant; buds green-white. Sepals 5, ovate, margin pilose, outside and inside glabrous, white; 2 outer smaller ones 0.7—2 by 0.8—2.3 mm, margin with glands; 3 inner larger ones 1.8—3.2 by 1.5-3 mm, margin petaloid, at most with few glands. Petals 5, ovate to obovate, 1.5—4.2 by 0.7— 1.6 mm, white; claw 0.2—0.5 mm high; margin and usually outside pilose, inside subglabrous, apex acute; scales 1-1.7 mm long, free, basally not au- riculate, margin not distinctly membranous; crest clavate, stipitate, apex lobed. Disc interrupted. Sta- mens 8; filaments 1.3—3.7 mm long, pilose, espe- cially basally, white; anthers 0.6—0.7 mm long, slightly pilose. Pistil: ovary 0.3—1.5 mm long, sub- hirsute; style and stigma 0.1—1.2 mm long. Fruits with 1—3 well developed lobes, i.1—2.9 by 1.1—3.5 cm, completely dehiscent, smooth to rugose to ru- gosely ribbed, glabrous, red when fresh, blackish when dry; stipe 0-2 mm high, slender; margin blunt; suture usually highly convex, lobes almost touching; lobes 10-18 by 9-22 mm; septa com- plete. Seeds obovoid, c. 8.7 by 5.2 mm; arillode without or with a small pseudo-funicle, but always with a basal rim; hiijum c. 1.3 mm long. Distribution — Malesia: Irian Jaya (Jayapura); Papua New Guinea (E. Sepik, Morobe, Milne Bay, Central, Gulf Prov.). Habitat & Ecology — In poor lowland rain for- est, gallery forest along creek, monsoon forest, garden regrowth, along edge of scrub and eucalypt savannah, road-sides; altitude sea level up to 450(— 935) m. Fl. Aug.—Sept.; fr. Jan. Note — Guioa rigidiuscula is part of the G. ri- gidiuscula-complex. Typical are the elliptic, smooth leaflets with a single sac and the barely stipitate fruits with highly convex upper sutures. 38. Guioa scalariformis Welzen, Blumea 33 (1988) 420, pl. 20a—c; Leiden Bot. Series 12 (1989) 283, f. 121. — Type: NGF (Sayers) 21576 (L holo; BM, CANB, U; LAE, n.v.), Papua New Guinea. Guioa spec.: Hartley et al., Lloydia 36 (1973) 270 (p.p.: Hartley 12459); Streimann, Pl. Upper Watut Watershed (1983) 169 (p.p.: Hartley 12459, NGF 21576). Shrub to tree, 3-10 m high, dbh up to 15 cm. Branchlets sericeous when young; flowering twigs 4-5.5 mm thick. Leaves |- or 2-jugate; rachis 9.3— 22.2 cm long, terete, glabrous, petiole 6—9.5 cm long. Leaflets subsessile, subopposite, ovate, | 1.9— 22.7 by 6.1—10.7 cm, index 1.8—2.1, subsymmetri- cal, acroscopic side somewhat broader, subcoria- ceous, not punctate; base attenuate; margin entire, flat; apex acuminate to cuspidate, often mucronu- late; upper surface glabrous; lower surface duller, smooth, no papillae, glabrous, domatia absent; ve- nation on upper side flat to raised, raised below; nerves 0.3—2.9 cm apart, marginally rather looped and joined; veins laxly reticulate to scalariform, very distinct. Inflorescences axillary, branching along the flattened to terete, sericeous, glabrescent, 6.1—13.5 cm long axis; first order branches up to 7.8 cm long; cymules cincinnate, c. 3-flowered;: bracts and bracteoles triangular, outside sericeous, inside glabrous; bracts 0.7—1 mm long; bracteoles 0.3—0.4 mm long; pedicels 4.2—5.8 mm long, seri- ceous except for the glabrous articulate part. Flow- 594 Flora Malesiana ser. I, Vol. 11 (3) (1994) ers c. 4 mm in diam. Sepals 5, ovate, margin pi- lose, with glands, outside and inside glabrous, greenish white; 2 outer smaller ones 1.2—1.9 by 1.8—2.2 mm; 3 larger inner ones 2.4—3.2 by 1.2- 2.5 mm, margin petaloid. Petals 5, obovate, 2.8—3 by 1.7—2 mm, white; claw c. 0.5 mm high; margin pilose, outside and inside glabrous, apex obtuse to acute; scales c. 1.8 mm long, free; crest clavate, stipitate, apex lobed; petal between two adjacent larger sepals not reduced in size. Disc uninterrupt- ed. Stamens 8: filaments 2.5—3.8 mm long, pilose, especially basally; anthers 0.6—0.7 mm long, gla- brous. Pistil: ovary c. 0.4 mm long, subhirsute; style and stigma c. 0.2 mm long. Fruits with 2 or 3 well developed lobes, c. 1.8 by 2.3-2.9 cm, slightly ribbed, glabrous, red when fresh, blackish when dry; stipe c. 2.5 mm high, slender; margin blunt; wall less than | mm thick at suture; lobes c. 17 by 12 mm; septa complete. Seeds globose, c. 11.2 by 9.8 mm; hilum c. 3 mm long. Distribution — Malesia: Papua New Guinea (Morobe Province: near Wagau). Habitat & Ecology — In primary mid-mountain forest; altitude 1500-1700 m. FI. Dec. Note — This species is part of the G. rigidiuscu- la-complex. It is distinctive in having large, broad leaflets with distinct venation, large petals, and an uninterrupted disc. 39. Guioa subsericea Radlk., Bot. Jahrb. 56 (1921) 277; in Engl., Pflanzenr. 98 (1933) 1158; Strei- mann, Pl. Upper Watut Watershed (1983) 169 (p.p.: Hartley 11835); Welzen, Leiden Bot. Series 12 (1989) 287, f. 123. — Lectotype (Van Welzen 1989): Ledermann 10005 (L; K), NE New Guinea. Guioa molliuscula auct. non Radlk.: Hartley et al., Lloydia 36 (1973) 270. Guioa dasyantha auct. non Radlk.: Streimann, PI. Upper Watut Watershed (1983) 169. Shrub to tree, 3-30 m high, dbh 6—50 cm; outer bark usually smooth to longitudinally fissured or finely pustulated, greenish to greyish to reddish brown, inner bark white to red-brown, no exudate; sapwood white, (rings prominent), heartwood brown. Branchlets shortly sericeous (to hirsute), especially when young; flowering twigs 1.5-7 mm thick. Leaves 1—3-jugate; rachis 1.2—20 cm long, terete to somewhat flattened below the jugae, sub- sericeous (to hirsute), petiole 0.9—9 cm long; peti- olules up to 0.9 cm long. Leaflets opposite to sub- opposite (to alternate), ovate (to elliptic), 3.5—18.3 by 0.9-9.3 cm, index 1.9-4.3, usually very asym- metrical, acroscopic side broader, coriaceous to very coriaceous, usually punctate; base attenuate; margin entire, flat (to revolute), apex gradually (acuminate to) cuspidate to caudate, mucronulate; upper surface densely pilose when young to sub- sericeous when older, (wax); lower surface dull, papillate, (sub)sericeous (to hirsute), domatia ab- sent to many small pocket-like sacs in axils of nerves; venation on upper side (slightly sunken to) flat (to raised), raised on lower; nerves 0.2—2.8 (—4.2) cm apart, marginally looped and joined, less distinctly so in lower part of leaflets; veins densely to laxly reticulate, usually distinct. Inflorescences axillary to pseudoterminal, branching basally and along the terete, sericeous (to hirsute), 1-26.6 cm long axis; first order branches up to 10.5 cm long; cymules cincinnate (to dichasial), 2—4(—6)-flow- ered; bracts and bracteoles triangular, outside seri- ceous, inside glabrous; bracts 0.5—2.2 mm long; bracteoles 0.2—1.1 mm long; pedicels (1.2—)2.2— 3.6(-6) mm long, sericeous except for the (sub) glabrous articulate part. Flowers c. 3 mm in diam. Sepals 5, ovate, margin pilose, with glands, out- side and inside glabrous, light to midgreen; 2 outer smaller ones 0.6—1.8 by 0.6—2.3 mm; 3 inner larger ones 0.8—3 by 1-3 mm, margin petaloid. Petals 5, rhombic to elliptic to obovate, 0.9-2.3 by 0.5—1.6 mm, white to yellow; claw 0.2—0.3 mm high; mar- gin and less so outside pilose, inside (sub)glabrous, apex retuse to rounded; scales inwardly folded au- ricles, 0.3-1.2 mm high; crest absent (to clavate, shortly stipitate). Disc uninterrupted. Stamens 8; filaments 1.2—2.8 mm long, pilose, especially ba- sally, white; anthers 0.4—-0.6 mm long, glabrous to sparsely pilose, light pink to purplish red. Pistil: ovary 0.3—1.2 mm long, subhirsute, midgreen; style and stigma 0.2—1.3 mm long. Fruits with 1-3 well developed lobes, 0.9-1.6 by 0.9—2.4 cm, smooth to rugose to slightly ribbed, glabrous, red when fresh, usually blackish when dry; stipe 0-4 mm high, broadly obconical, indistinct; margin blunt; lobes 7-17 by 5-11 mm; septa complete. Seeds (globose to) obovoid, 5.8—7.8 by 3-8 mm; hilum 1—1.2 mm long; arillode orange. — Fig. 44. Distribution — Malesia: Irian Jaya (Vogelkop, Jayapura); Papua New Guinea (W & E Sepik, Enga, Western Highlands, Chimbu, Southern Highlands, Morobe, Central, Milne Bay Provinces). Habitat & Ecology — Occasional to fairly com- mon in understorey (to subcanopy trees) in lower to midmontane primary forest, in secondary forest and along edges of wood, water and roads; often on steep slopes and ridges. Forests often dominat- ed by Castanopsis, Casuarina, Nothofagus, and Pandanus. Soil: detritus mud, sand, laterite, stony clay or loam; wet; shade light to medium; altitude (70—)1000—3000 m. FI. Feb.—July; fr. throughout the year. Adema, Leenhouts, Van Welzen — Sapindaceae 595 Imm Imm imm Fig. 44. Guioa subsericea Radlk. a. Habit with E New Guinean form of leaflets; b. leaflet of W New Guinean form; c. petal; d. fruit (a, d: ANU (Flenley) 2772; b: BW 4076; c: NGF 11042}. 596 Notes — 1. Guioa subsericea 1s distinct because of its long-tipped leaflets that are sericeous and papillate below, an uninterrupted disc, and the ba- sically broadly obconical fruits. The species is rath- er diverse, with some distinctive forms which are connected by intermediates. Most obvious is the topocline between W and E New Guinea, although within W as well as E New Guinea there are rea- sonably uniform groups. Central New Guinea forms the transition zone. Plants in E New Guinea have larger leaflets (3.5-18.3 by 1.4-8.3 cm versus 3.6— 9.8 by 0.9-3 cm), with usually longer hairs, and they have larger fruits (0.9-1.6 by 1—2.4 cm ver- sus 0.9-1.2 by 0.91.4 cm). In Morobe and W High- land Provinces (Papua New Guinea) several spec- imens possess leaflets which are, in a dried condi- tion, very coriaceous and very silvery on both sur- faces of the leaflets. Guioa subsericea has a sericeous indumentum, but some specimens are very hirsute and, at first sight, look like a different species. This phenome- non also occurs in other species of Guioa, e.g., G. pleuropteris, G. chrysea, G. villosa. 2. Guioa multijuga, from Irian Jaya, is remark- ably similar to G. subsericea from the same area; however, G. multijuga always has leaves with more than 3 jugae and the leaflets are almost glabrous below instead of sericeous. Furthermore, the pet- als have very different scales, those of G. subseri- cea are inwardly folded auricles, those of G. mul- tijuga are free scales folded outwards. 3. The differences with G. molliuscula, G. malukuensis, and G. comesperma are discussed in notes under these species. 40. Guioa truncata Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1611; Philipp. J. Sc., Bot. 8 (1913) 446; Merr., Enum. Philipp. Flow. PI. 2 (1923) 509; Radlk. in Engl., Pflanzenr. 98 (1933) 1171; Welzen, Leiden Bot. Series 12 (1989) 292, f. 125. — Type: Elmer 11219 (M holo; A, BM, F, FI, K, L, NY, U, W), Philip- pines. Tree, 5-7 m high, dbh 10-15 cm; bark dull brown, smooth; sapwood whitish, rather hard, odourless, tasteless, heartwood reddish brown. Branchlets sericeous when young; flowering twigs 2-3 mm thick. Leaves 2—3-jugate; rachis 2-10 cm long, basally terete to upwards flattened above, slightly winged, glabrous, petiole 1.9—3.8 cm long. Leaflets subsessile, opposite to subopposite, ellip- tic, 3.4-7.6 by 1.3—2.6 cm, index 2.6—3, asymmet- rical, acroscopic side broader, coriaceous, punc- tate; base attenuate; margin entire, flat; apex acu- minate, mucronulate; upper surface glabrous (ex- cept for the puberulous midrib); lower surface dull- Flora Malesiana ser. I, Vol. 11 (3) (1994) er, smooth, no papillae, glabrous to slightly pilose near domatia, latter a single to many sacs in axils of nerves; venation raised; nerves 0.4—1.6 cm apart, marginally looped and joined; veins laxly reticu- late, distinct. Inflorescences axillary, branching basally to usually along the flattened, subsericeous, 4.3-15.4 cm long; first order branches up to 4.2 cm long; cymules cincinnate, c. 3-flowered; bracts and bracteoles triangular, outside sericeous, inside glabrous; bracts c. 0.8 mm long; bracteoles 0.3— 0.7 mm long; pedicels c. 3.3 mm long, sericeous except for the subglabrous articulate part. Flowers c. 3 mm in diam. Sepals 5, ovate, margin pilose, with glands, outside and inside glabrous; 2 outer smaller ones 1.4—1.6 by 1—1.3 mm; 3 inner larger ones 2—3.1 by 1.8-2.6 mm, margin petaloid. Pet- als 5, obovate, c. 2.1 by 1.4 mm, white; blade or- bicular, abruptly clawed; latter c. 0.7 mm high; margin pilose, apex rounded; scales c. 0.8 mm long, free: crest a flat subglabrous part of the slightly bifid scale apex. Disc interrupted. Stamens 8; fila- ments 2.9-3.1 mm long, pilose, especially basal- ly; anthers c. 0.5 mm long, glabrous. Pistil: ovary c. 0.7 mm long, subhirsute; style and stigma c. 0.2 mm long. Fruits immature. Distribution — Malesia: Philippines (Mindanao). Habitat & Ecology — In dense moist forests and mossy forests; altitude 1300-2300 m. FI. Mar. Note — PNH 118626 (G. koelreuteria) looks like G. truncata; but its venation is not as distinct, the leaflets are narrower, and do not possess large sac- like domatia. Guioa koelreuteria has at most one large sac, not several, as G. truncata usually has. 41. Guioa unguiculata Welzen, Blumea 33 (1988) 420, pl. 18a—c; Leiden Bot. Series 12 (1989) 294, f. 126. — Type: Vink 16407 (L holo; BRI, K, NSW, P; LAE, n.v.), Papua New Guinea. Guioa membranifolia auct. non Radlk.: Hartley et al., Lloydia 36 (1973) 270. Tree(let), 5-20 m high; dbh up to 15 cm; outer bark smooth, greyish green to dark grey, inner bark straw coloured; wood straw coloured. Branchlets sericeous when young; flowering twigs 3-7 mm thick. Leaves 1—3-jugate; rachis 2—15.8 cm long, terete, glabrous, petiole 1.4-6.9 cm long. Leaflets subsessile, subopposite to alternate, elliptic, 6.3— 17 by 1.9-6.5 cm, index 2.3-3.7, slightly asym- metrical, acroscopic side broader, (sub)coriaceous, usually punctate; base attenuate; margin entire, flat; apex acuminate to caudate, mucronulate; upper surface glabrous; lower surface duller, smooth, no papillae, glabrous except for a few hairs on vena- tion (to sparsely sericeous), domatia absent; vena- tion on upper side flat, raised below; nerves 0.3— 2.9 cm apart, marginally looped and joined, less Adema, Leenhouts, Van Welzen — Sapindaceae distinctly so in lower part of leaflets; veins laxly reticulate, rather indistinct. /nflorescences axillary to ramiflorous, branching basally and along the flat- tened to terete, subsericeous, 2.2—8.5 cm long axis; first order branches up to 2.3 cm long; cymules cincinnate, 2—4-flowered; bracts and bracteoles tri- angular, outside sericeous, inside glabrous; bracts 0.5—1 mm long; bracteoles 0.6—0.7 mm long; pedi- cels 2.8-6.2 mm long, sericeous except for the gla- brous articulate part. Flowers c. 4 mm in diam. Sepals 5, ovate, margin pilose, with glands, out- side and inside glabrous, light green; 2 outer smaller ones 1.2—2 by 1.5—2 mm; 3 larger inner ones 2- 2.8 by 1.6—2.9 mm, margin petaloid. Petals 5, obo- vate, 2—3.2 by 1-1.5 mm, white; claw c. 0.8 mm high; margin pilose, outside and inside subpilose, apex rounded, somewhat fimbriate; scales 1.2—1.5 mm long, resembling folded margins, without ba- sal auricles, without a distinct membranous mar- gin; crest absent (to some scales with a bifid apex); petal between two adjacent larger sepals not re- duced in size. Disc (nearly) uninterrupted, ochre- green. Stamens 8; filaments 2—3 mm long, pilose, especially basally, white; anthers 0.4—0.6 mm long, glabrous, light pink. Pistil green; ovary 0.4—1.5 mm long, subhirsute; style and stigma 0.2—1.5 mm long. Fruits with 1-3 well developed lobes, 1.5—2.2 by 2—2.7 cm, completely dehiscent, smooth to slight- ly rugose, black when dry; stipe 2—3.3 mm high, slender; margin blunt; wall less than | mm thick; lobes 13-18 by 8.5—12 mm; septa complete. Seeds not full-grown. Distribution — Malesia: Papua New Guinea (Western Highlands, Madang, Morobe Prov.). Habitat & Ecology — In open forest, oak forest, old secondary forest; altitude 1200-1900 m. FI. Aug.—Sep. Fruits eaten by birds. Note — Guioa unguiculata belongs to the G. ri- gidiuscula-complex. It has long-clawed petals with folded, apically bifid scales; a (nearly) uninterrupt- ed disc; and leaflets which are smooth, without papillae or domatia on the lower side, and with a strongly raised venation. 42. Guioa venusta Radlk., Sapind. Holl.-Ind. (1879) 11, 40; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 609; Bot. Jahrb. 56 (1921) 280; in Engl., Pflan- zenr. 98 (1933) 1160; Welzen, Leiden Bot. Series 12 (1989) 295, f. 127. — Type: Beccari FI 2813 (Fl holo, cited by Radlkofer as Bec- cari 5"), New Guinea. Small tree, 2—2.5 m high. Branchlets sericeous when young; flowering twigs 2.5—3 mm thick. Leaves 4—9-jugate; rachis 1.5—12.2 cm long, slight- ly winged, wing up to at most 0.5 mm broad, sub- sb | sericeous, petiole 1.3—2.4 cm long. Leaflets sub- sessile, opposite to alternate, elliptic, 1.8—-6.5 by 0.6—2 cm, index 2.2—3.3, asymmetrical, acroscop- ic side broader, subcoriaceous, punctate; base at- tenuate to cuneate; margin entire except for some subapical teeth, flat to slightly revolute; apex acu- minate, mucronulate; upper surface glabrous ex- cept for the basally puberulous midrib; lower sur- face duller, smooth, no papillae, glabrous to slightly sericeous, domatia many pockets in axils of nerves; venation on upper side slightly sunken to flat, raised below; nerves 0.2—1.2 cm apart, marginally looped and joined; veins laxly to densely reticulate, rath- er distinct. /nflorescences axillary, unbranched to branching basally and along the terete, pilose, 1.9— 5.4 cm long axis; first order branches up to 3.4 cm long; cymules cincinnate, c. 4-flowered; bracts and bracteoles triangular, outside sericeous, inside gla- brous; bracts 0.3—0.6 mm long; bracteoles 0.3—0.4 mm long; pedicels 0.8—3 mm long, sericeous; less so in the articulate part. Flowers in bud. Sepals 5, ovate, margin pilose, with glands, outside and in- side glabrous; 2 outer smaller ones 1.1—1.5 by 0.7— 1.1 mm; 3 inner larger ones 1.4—1.8 by 1.3—1.5 mm, margin petaloid. Petals 5, immature, 0.2—1.1 by 0.1—0.2 mm, no distinct claw yet, margin pilose, outside and inside glabrous, white; scales slightly developed, free; crest (still?) absent. Disc uninter- rupted. Stamens 8; filaments |.7—2.2 mm long, pi- lose in lower half; anthers c. 0.3 mm long, gla- brous. Pistil: ovary c. 0.2 mm long, subhirsute; style and stigma c. 0.1 mm long. Fruits with | or 2 well developed lobes, 0.7—0.8 by 0.6—0.9 cm, smooth to slightly rugose, glabrous, blackish when dry; stipe 2—2.2 mm high, slender; margin blunt; lobes 4—6 by 3.8-4.3 mm; septa complete. Seeds imma- ture. Distribution — Malesia: Irian Jaya (Schouten Island: Biak, and Japen Island). Habitat & Ecology — On flat terrain, in second- ary scrubby vegetation; altitude c. 60 m. FI. Apr. Note — The type of G. venusta, from Japen Is- land, is somewhat different from the specimens from Biak. It has smaller (1.8—3.9 by 0.6—1.2 cm versus 2.9—6.5 by 0.6—2 cm), more numerous pairs of leaflets (6-9 versus 4 or 5), and the petiole is more densely sericeous. 43. Guioa waigeoensis Welzen, Blumea 33 (1988) 420, pl. 19a, b; Leiden Bot. Series 12 (1989) 300, f. 129. — Type: van Royen 5409 (L holo), W Irian Jaya. Arytera xerocarpa auct. non Adelb.: P. Royen, Nova Guinea 5 (1960) 60. Small tree, c. 4 m high, dbh c. 5 cm, Branchlets sericeous, especially when young; flowering twigs 598 c. 1 mm thick. Leaves 3- or 4-jugate; rachis 0.4— 3.7 cm long, flattened above, slightly winged (wing less than 1 mm broad), subsericeous, petiole 0.4— 0.8 cm long. Leaflets subsessile, opposite to sub- opposite, elliptic, 1.3—2.4 by 0.4—0.6 cm, index 3.3— 4, rather symmetrical, coriaceous, punctate; base attenuate; margin at least in some leaflets slightly serrate, revolute: apex acute, mucronulate; upper surface subsericeous; lower surface dull, papillate, shortly sericeous, domatia in at least some leaflets a single, small sac on basiscopic side in axil of sec- ond nerve; venation flat to slightly raised on lower side; nerves 0.2—0.4 cm apart, marginally looped and joined; veins densely reticulate, indistinct. /n- fructescences axillary, not or at most branching along rachis; latter flattened, 1.2—2 cm long, seri- ceous; first order branches up to 0.6 cm long; bracts and bracteoles triangular, outside sericeous, inside glabrous; bracts 0.6—0.7 mm long; bracteoles c. 0.3 mm long; pedicels c. 5.8 mm long, sericeous ex- cept for the glabrous articulate part. Flowers white. Sepals 5, ovate, margin pilose, glands unknown, outside and inside glabrous; 2 outer smaller ones c. 1.8 by 1.3-1.9 mm; 3 inner larger ones 2—2.4 by 1.7—2.4 mm, margin petaloid. Petals unknown. Disc uninterrupted. Stamens and pistil unknown. Fruits with 1—3 well developed lobes (see note), c. 1.2 by 1.2 cm, smooth to slightly rugose, glabrous, black- ish when dry; stipe c. 3 mm high, rather slender; margin blunt; lobes c. 8 by 7 mm; septa complete. Seeds still immature. Flora Malesiana ser. I, Vol. 11 (3) (1994) Distribution — Malesia: W Irian Jaya (Waigeo Island). Habitat & Ecology — Relatively common tree, in hilly xerophytic vegetation; altitude c. 300 m. Note — One fruit had 4 lobes instead of 3, which is unique in Guioa. DUBIOUS NAME Guioa elegans Radlk., Bot. Jahrb. 56 (1920) 280; in Engl., Pfianzenr. 98 (1933) 1162; Welzen, Leiden Bot. Series 12 (1989) 301. Type: Schultze Jena 337 (B+ holo), Papua New Gui- nea. Note — Known only from the type, which is lost, and it cannot be identified by its description. Plants identified as G. elegans by Hartley et al. (Lloydia 36, 1973, 270) are Toechima erythrocarpum sub- sp. papuanum. Typical for G. elegans should be the small leaflets (6-8 by 1.5—2 cm) with revolute and sinuate margins, and the uninterrupted disc. The small leaflets with sinuate margins may be like those of G. amabilis, G. pseudoamabilis, or G. pteropoda (the latter a species also described by Radlkofer and therefore hardly likely to be identi- cal to G. elegans). The first two have revolute mar- gins and an uninterrupted disc; however, both are mountain species, and, moreover, G. amabilis is only found near the Anggi Lakes in the Vogelkop Peninsula (Irian Jaya). HARPULLIA (P.W. Leenhouts & M. Vente) Harpullia Roxb. [Hort. Beng. (1814) 86, nom. nud.] FI. Ind. 2 (1824) 441; Radlk. in Engl., Pflanzenr. 98 (1933-1934) 1433; Leenh. & Vente, Blumea 28 (1982) 3S Reynolds in FI. Austral. 25 (1985) 38; Yap in Tree Fl. Malaya 4 (1989) 443. — Type species: Harpullia cupanioides Roxb. [Donatophorus Zipp. ex Macklot, Bijdr. Natuurk. Wetensch. 5 (1830) 181, nom. nud. — Thanatophorus Zipp. ex Walp., Ann. Bot. Syst. 2 (1851/52) 213, nom. inval. in syn. — Indicated type species: Donatophorus erythrospermus Zipp. ex Macklot (= Har- pullia cupanioides Roxb.).] Otonychium Blume, Rumphia 3 (1847) 179. — Type species: Otonychium imbricatum Blume [= Harpullia arborea (Blanco) Radlk. |]. Blancoa Blume, Rumphia 3 (1847) 181, nom. illeg., non Lindl. (1840). — Type species: Blancoa arborea (Blanco) Blume [= Harpullia arborea (Blanco) Radlk. ]. Streptostigma Thwaites, Hook. J. Bot. Kew Gard. Misc. 6 (1854) 298. — Type species: Streptostigma viridiflorum Thwaites [= Harpullia arborea (Blanco) Radlk.]. ?Apiocarpos Montrouz., Mém. Acad. Roy. Sci. Lyon, Sect. Sci., sér. 2, 10 (1860) 190. — Adema, Leenhouts, Van Welzen — Sapindaceae 599 Type species: Apiocarpos moguinii Montrouz. [= ?Harpullia austrocaledonica Bail- lon]. Shrubs to medium-sized trees, dioecious. /ndumentum solitary and stellate tufts of simple hairs (and glandular hairs); no glandular scales. Leaves paripinnate, 1—9-jugate, without pseudostipules; (petiole and rachis winged). Leaflets alternate (or opposite), not papillate beneath; margin entire (Malesian species). /nflorescences axillary, (pseudoter- minal), to truly terminal, solitary, or rami- and/or cauliflorous and often tufted, usually thyrses; bracts and bracteoles usually caducous. Flowers unisexual, actinomorphic. Se- pals 5, free, imbricate, equal or the outer two sometimes slightly smaller, not petaloid, not ciliate, (glandular hairs mainly along the margin), entire. Petals 5, longer than the sepals, distinctly clawed with a pair of auricles above the claw, or sessile with a broad or narrow base and without auricles and scales, entire. Disc uninterrupted (to divided into 5 lobes), without appendages. Stamens 5—8, in male flowers exserted; filaments glabrous; anthers basally attached, base cleft for up to 1/5, dehiscence latero-introrse. Pistil 2- or 3- (or 4-)locular; ovary sessile or short-stalked, hairy; ovules | or 2 per locule; style apical, shorter to much longer than the ovary, slender, often hooked and the upper part twisted, lower part hairy, with stigmatic lines usually till slightly above the base. Fruits loculicid- al capsules, usually short-stipitate, not winged, 2- or 3-lobed, the lobes erect to spread- ing, inflated, rounded; wall pergamentaceous to woody. Seeds with a thin-crustaceous testa; arillode restricted to a narrow annular sarcotesta around the hilum, or composed of a basal sarcotestal part, covering half the seed, and an upper, free arilloidal part, reaching to near the apex, the arillode entire and without appendages; hilum covering less than 1/6 of the seed. — Figs. 45-47. Distribution — 26 species, occurring from Sri Lanka and India through SE China and Malesia to Australia (Northern Territory, Queensland, and New South Wales to c. 32° 30' S), New Caledonia, and Tonga. See Avé in Van Balgooy (ed.), Pacific Plant Areas 4 (1984) ZIOR Lo: Habitat & Ecology — Mainly substage or lower storey trees of primary and some- times secondary rain forests, sometimes growing in low or open forest or savannahs and in shrubberies on coastal dunes; altitude from sea level up to 2000 m. The seeds are probably mainly dispersed by birds, possibly also by mammals and lizards. Pollen morphology — See Muller, Blumea 31 (1985) 161-218. Note — For evolutionary notes see Muller, Blumea 31 (1985) 161—218 and Leenh., Blumea 31 (1985) 219-234. KEY TO'THE SPECIES hatikeat rachis‘and petiole notowineed Oy b2..29U M001 Gl als PORT EIS 2 baieeas rath and petiole winged 2... ov sesh en soa oe 16. H. rhachiptera 2a. Petals not clawed, without auricles, thin-fleshy. Arillode completely or nearly com- pretely.envelopmeg. the seed... «.. 9304 JOA AG AIO PAU R Ea 8 3 b. Petals clawed and auricled, membranous. Arillode restricted to a ring around the MRPOMSICUE SIC cag rcs cree ee oe en a oo hee pe a ee i ae 1. H. arborea BassStamens (FORMS OULU OT, 004pViotiob Oud Bea AS. OU Ae heats elm il 4 Bahan T ie ob os kde vecde ens sees dei inl ann RPaes 6 600 Flora Malesiana ser. I, Vol. 11 (3) (1994) alt Teeaviesliair yi yen axph aN ee = ef SLPERTSRI. NM GP AIUL2Alene Fale eve Ph Sct on 5 Se AVES SIADTOUS co sencuzuce aacenpoeaPagao o oer eb eokgcr her SE oaspemetnac 7c 10. H. longipetala a. Midrib above (in dried leaves) sunken, nerves flat to sunken. Fruit lobes erect, com- pletelyeumited*@ver S25 Cli, eer piera uate teins vet ste etree 7. H. giganteacapsula _ Midrib and nerves above (in dried leaves) slightly raised. Fruit lobes widely spread- ie Panel y MMI Le CLOVCT AT MINN ein renee aes neue alae 12. H. oococca . Inflorescences ramiflorous, though usually initially in the leaf axils .......... 7 Inflorescences axillary to (rarely) terminal). 5.0.0 eosin sae Joe ee eee 13 a. Leaflets herbaceous to chartaceous, usually hairy. Sepals up to 7 mm long .... 8 Leaflets coriaceous, glabrous. Sepals 7-8 mm long.......... 13. H. peekeliana . Inflorescences short, up to about 4 cm long, not or hardly branched .......... 5) Inflorescences longer than c. 4 cm, often repeatedly branched.............. 10 a. Twigs slender, 2-7 mm thick. Fruits leathery, inside glabrous .. 15. H. ramiflora . Twigs stout, 8-15 mm thick. Fruits woody, usually hairy inside . 4. H. cauliflora ER RUME MAINS WOOR sj a: rctselae ccs d s

    3 12 . Petiolules 0-4 mm long. Inflorescences exclusively ramiflorous/cauliflorous, up to DS ORGimmLOMe eee es tL aed MOOREA os tenis ene tg orn aetna one 14. H. petiolaris . Leaflets abruptly acuminate. Axillary inflorescences solitary. Fruits 2.2—2.5 cm wide and about as high, nardly,Stipitatess 5 spo jecsgi cusses 9) ipo 11 H. myrmecophila . Leaflets gradually acuminate. Axillary inflorescences clustered like the rami- and/ or cauliflorous ones. Fruits 2.5—3.4 cm wide, c. 1.5 times as wide as high, distinctly Ait Oy UE {Wasps Sk SR ask Oh Pa NP ord rn a Aue 17. H. solomonensis a. Fruit lobes spreading, usually united for less than] cm ................... 14 Fruit lobes erect, united for more than 1 cm, hence the fruit ovoid to obovoid . 16 » Leaflets acuminate, Hill and low montane $5... 222-255 3... ya eee bs » Leatlets obtuse to roundeds Lowlanduts An.2e. wee 2. ae ccs 9. H. leptococca . Leaflets glabrous to sparsely hairy on midrib and nerves on both sides 3. H. carrii . Leaflets on midrib and nerves densely puberulous on both sides, beneath also sparsely SOOM VeIMSeandeVeIMletS Az tes foes ee ye tee ays ee 12. H. oococca . Vegetative parts, at least when young, distinctly and + densely hairy ........ 17 . Twigs glabrous except for the terminal bud, leaves glabrous or at most very sparse- lyshainy*on-axess midribyand nerves {os ... lee ane ee = 6. H. cupanioides . Twigs rather slender, 3-10 mm thick. Leaves up to 5-jugate. Leaflets distinctly pet- 1olulatesnenvies ADOVE a= TalseG sce. i. soca t Se re ce ce ee eb cue Sree ene 18 . Twigs rather stout, 9-15 mm thick. Leaves 5—9-jugate. Leaflets subsessile, the nerves sitchtlyasunkem abOVG =. .)252 6.045405 s50 + Tee eh easter eee 5. H. crustacea Ss lntloneseencestencet, 105lloicmilone eq) .es ee. EO DG OE Bee 19 Minilorescences pendulous, up to /O;emilones ; Pease ene ae 18. H. vaga . Leaflets at most thin-puberulous on midrib and nerves on both sides 2. H. camptoneura . Leaflets hirsute on both sides, most densely so on he midrib and on the nerves on fhe mippensiderte, ete CAO. ROY. Oks tek err aed pitta nn rae 8. H. hirsuta Adema, Leenhouts, Van Welzen — Sapindaceae 601 Subgenus Otonychium Harpullia subg. Otonychium (Blume) Radlk., Sapind. Holl.-Ind. (1879) 52; in Engl., Pflan- zenr. 98 (1934) 1438: Leenh. & Vente, Blumea 28 (1982) 9. — Otonychium Blume, Rumphia 3 (1847) 179. — Harpullia sect. Euotonychium Radlk., Sapind. Holl.-Ind. (1879) 52, nom. inval.; in Engl., Pflanzenr. 98 (1934) 1439. — Type species: Otony- chium imbricatum Blume [= Harpullia arborea (Blanco) Radlk. ]. Blancoa Blume, Rumphia 3 (1847) 181. — Type species: Blancoa arborea (Blanco) Blume [= Harpullia arborea (Blanco) Radlk.}. Streptostigma Thwaites, Hook. J. Bot. Kew Gard. Misc. 6 (1854) 298. — Type species: Streptostigma viridiflorum Thwaites [= Harpullia arborea (Blanco) Radlk.]. Harpullia sect. Otonychidium Radlk., Sapind. Holl.-Ind. (1879) 53; in Engl., Pflanzenr. 98 (1934) 1438. — Type species: Harpullia pendula Planch. ex F. Muell. Sepals deciduous. Petals clawed and with a pair of infolded auricles just above the claw, membrabous. Style straight. Arillode restricted to a narrow sarcotestal ring around the hilum. Distribution — 2 species, one restricted to Australia. 1. Harpullia arborea (Blanco) Radlk., Sitzungs- ber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 16 (1887) 404; Lecomte in FI. Indo- Chine | (1912) 1022; Le Renard, Ann. Sci. Nat. Bot. LX, 17 (1913) 373; Merr., Sp. Blanc. (1918) 243; Enum. Philipp. Flow. Pl. 2 (1923) 515; Craib, Fl. Siam. Enum. | (1926) 335; Radlk. in Engl., Pflanzenr. 98 (1934) 1456; Gagnep. in Fl. Indo-Chine, Suppl. 1 (1950) 954, f. 119: 18— 21; Backer & Bakh. f., Fl. Java 2 (1965) 142; Leenh. & Vente, Blumea 28 (1982) 11; S.T. Reynolds in FI. Austral. 25 (1985) 44, map 51; Yap in Tree Fl. Malaya 4 (1989) 444. — Ptelea arborea Blanco, FI. Filip. (1837) 63. — Ser- ingia lanceolata Blanco, Fl. Filip. ed. 2 (1845) 45, nom. illeg., non Steetz (1848); ed. 3, 1 (1877) 85. — Blancoa arborea Blume, Rumphia 3 (1847) 181, comb. illeg. — Neotype (Leen- houts & Vente 1982): Merrill Sp. Blancoanae 339 (A holo; BM, BO, K, L, NSW, P, US, W), Philippines. Otonychium imbricatum Blume, Rumphia 3 (1847) 180. — Harpullia imbricata Thwaites, Enum. Pl. Zeyl. (1858) 56; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 236; Atlas 1 (1913) pl. 142: f-n. — Syntypes: Korthals s.n. (L), Cen- tral Sumatra; Kiihl & van Hasselt s.n. (L), Java. ?Harpullia blancoi Fern.-Vill. in Mercado, Libro Medic. (1880) 4. — Type: unknown. Harpullia pedicellaris Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 279; in Engl., Pflanzenr. 98 (1934) 1454. — Type: C. Hartmann s.n., 1887 (M holo; MEL), SE New Guinea. Harpullia glanduligera Radlk. in Fedde, Rep. 20 (1924) 41; in Engl., Pflanzenr. 98 (1934) 1459. — Type: Ledermann 6760 (B* holo; M), NE New Guinea. Harpullia arborea var. megalocarpa Merr., Philipp. J. Sc. 27 (1925) 35. — Type: Loher 13273 (PNH* holo), Philippines. Harpullia tomentosa Ridley, Kew Bull. (1933) 192; Radlk. in Engl., Pflanzenr. 98 (1934) 1462. — Type: Haviland 2232 (K holo), Sarawak. Harpullia sphaeroloboa Radlk. in Engl., Pflanzenr. 98 (1934) 1454. — Type: Bogor Hort. Bot. 111.K.19a (M holo; BO, BRI, U), Moluccas. Harpullia cupanioides auct. non Roxb.: Fern.- Vill. in Blanco, FI. Filip. ed. 3, Nov. App. (1880) 53, p.p.; Radlk., Meded. Rijks-Herb. 22 (1914) 20 as for Elbert 3261; in Engl., Pflanzenr. 98 (1934) 1444, as for Celebes. Harpullia sp.: Ceron, Cat. Pl. Herb. Manila (1892) 55 as for Vidal 2524 & 2526. Harpullia sp. A: Koord., Meded. Lands Planten- tuin 12 (1894) 33. Arytera litoralis auct. non Blume: Koord., Minah. (1898) 402. (Shrub or) tree, up to 33 m high, dbh up to 60 cm, but usually much smaller. Young parts + dense- ly hirsute, hairs rather long; (on lower side of mid- rib, in inflorescences, and sepals also with longer glandular hairs). Twigs 3.5—9 mm thick. Leaves 2- 6-jugate; petiole 4.5-15 cm long; petiolules 3-8 mm long; all axes hairy, glabrescent. Leaflets ovate to elliptic, 5.5—30 by 2-10 cm, index 1.5—3.5, her- baceous; base oblique with the upper half cordate 602 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 45. Harpullia arborea (Blanco) Radlk. Habit (PNH 12482). or both sides acute, or symmetrical and acute to rounded; apex acute to rounded (to acuminate), acumen usually short, acute; above glabrous but for the midrib, beneath glabrous or sparsely hairy on midrib and very sparsely so on the nerves; mid- rib above flat to slightly raised; nerves 0.75—2.25 cm apart, above flat; (intersecondary nerves incon- spicuous). /nflorescences axillary to rami- or cau- liflorous, hairy, ramified only near the base into several axes of about the same length, up to 17 cm long, or with a distinct, up to 35 cm (in fruit up to 60 cm) long main axis with short branches; pedi- Adema, Leenhouts, Van Welzen — Sapindaceae 603 cels in fruit 12-30 mm long. Sepals all equal, ovate to obovate, 5—10.5 by 3-5 mm, (some scattered glandular hairs especially along the margin). Per- als: claw 3—7 mm long, blade obovate-oblong, 8— 17 by 3-10 mm, white, outside glabrous or claw and lower half of blade mainly in the centre sparsely hairy, margin often ciliate mainly in the basal part, inside often sparsely hairy. Disc sparsely to densely hairy. Stamens 5(—7); filaments 10-17 mm long; anthers 2—2.5 mm long. Piszil 2(—4)-locular; style 14-17 mm long; ovules | or 2 per locule. Fruits 9— 31 by 27-65 mm; stipe up to 4.5(—7) mm high; lobes spreading, slender ellipsoid to globular; out- side prominently veined to smooth, red, fairly densely to sparsely hairy; wall thin, chartaceous to woody; inside reddish, sparsely hairy to glabrous. Seeds | or 2 per locule, black, mahogany-brown, or dark-purple; sarcotesta up to 2.5 mm wide, orange. — Figs. 45, 46a—f. Distribution — Sri Lanka, the Deccan Peninsula from Maharashtra to the south, Assam, Thailand, S Vietnam, throughout Malesia, from the Solomon Islands to Samoa and Tonga, and N Queensland. Habitat & Ecology — On ridges, slopes, and plains, in ravines and sometimes along or in swamps, on river banks, or along the seacoast; on fertile soil, clay, loam, or sand, in limestone areas or on volcanic soils. Usually in well-drained pri- mary rain forest, but also in secondary forests, in lowland, hill, lower and middle montane forest, also in Araucaria forest; exceptionally in open vegeta- tion; sea level up to 1200 m altitude. Fl. and fr. throughout the year. Uses — The bark is used as a fish poison. A watery exudate of the bark and sometimes the fruits is used for washing, to keep away leeches, or is drunk to allay pain. For a description of the tim- ber, see p. 427. The oil pressed out of the seeds is used as an anti-rheumatic. See Brown, Useful PI. Philipp. 2 (1950) 363; Desch, Mal. For. Rec. 15 (1954) 528 (timber). Notes — 1. Though H. arborea is a variable spe- cies with many distinct local forms it is not possi- ble to subdivide it. The most conspicuous differ- ence is between the W Malesian population, up to and including Java and Borneo, with 2 ovules and thus 2 seeds per locule, and the E Malesian ones with usually only 1 ovule and | seed per locule. However, there is a wide overlap in the Philippines, and while the western material is rather uniform the number of ovules and seeds varies in the east. 2. Harpullia arborea seems to be scarce in W Malesia, whereas it is common in the eastern part. Subgenus Harpullia Harpullia subg. Harpullia: Leenh. & Vente, Blumea 28 (1982) 15. — Harpullia Roxb., Fl. Ind. 2 (1824) 441. — Harpullia subg. Euharpullia Radlk., Sapind. Holl.-Ind. (1879) 52, nom. inval.; in Engl., Pflanzenr. 98 (1934) 1435. — Type species: Harpullia cupanioides Roxb. Donatophorus Zipp. ex Macklot, Bijdr. Natuurk. Wetensch. 5 (1830) 181. — Harpullia sect. Thanatophorus Radlk., Sapind. Holl.-Ind. (1879) 52, nom. inval.; in Engl., Pflan- zenr. 98 (1934) 1436. — Type species: Donatophorus erythrospermus Zipp. ex Macklot [= Harpullia cupanioides Roxb.}. ?Apiocarpos Montrouz., Mém. Acad. Roy. Sci. Lyon, Sect. Sci. sér. 2, 10 (1860) 190. — Type species: Apiocarpos moguinii Montrouz. [= ?Harpullia austrocaledonica Bail- lon]. Harpullia sect. Harpulliastrum Baillon, Adansonia 11 (1874) 241; Radlk. in Engl., Pflan- zenr. 98 (1934) 1438. — Type species: Harpullia austrocaledonica Baillon. Sepals nearly always persistent in fruit. Petals sessile with a broad or narrowed base, without auricles, thin-fleshy to rarely membranous. Style bent or hooked once or twice. Seeds with a complex arillode composed of a sarcotestal part and a free arillode. Distribution — 24 species, as widely distributed as the genus with the exception of Sri Lanka, the New Hebrides, Fiji, Samoa, and Tonga. 604 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 46. Harpullia Roxb. Petals, ovary, infructescence, fruits, seeds, and embryos. — H. arborea (Blan- co) Radlk. a. Petal; b. ovary; c. seed, lateral; d. ibid., from below; e. embryo, dorsal; f. ibid., lateral. — H. cupanioides Roxb. g. Petal; h. ovary. — H. cauliflora K. Schum. & Laut. i. Fruit; j. seed. — H. gigan- teacapsula Vente. k. Fruit. — H. leptococca Radlk. |. Fruit. — H. ramiflora Radlk. m. Infructescence (a, b: van Beusekom et al. 3862; c, d: SAN 31225; e, f: Merrill 2843: g, h: Waterhouse 36B; i: Hoogland & Craven 10551; j: Brass 13802; k: LAE 62090; |: NGF 12367; m: Pullen 6609). Adema, Leenhouts, Van Welzen — Sapindaceae 605 2. Harpullia camptoneura Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 360; Radlk. in Engl., Pflan- zenr. 98 (1934) 1446; Leenh. & Vente, Blumea 28 (1982) 32. — Type: Warburg s.n., 1881 (B+ holo; M), NE New Guinea. Harpullia crustacea auct. non Radlk.: K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1900) 424 as for Lauterbach 601. Shrub or treelet, up to 4.5 m high. Twigs, leaf axes, and inflorescences densely, shortly, brown- ish puberulous, usually early glabrescent except for the inflorescences. Twigs 3.5—6 mm thick. Leaves 2—4-jugate; petiole 4-8 cm long; petiolules 2-8 mm long. Leaflets ovate (lowermost leaflets) to ellip- tic, 7-21 by 3.5—8 cm, index 1.75-3.25, stiff-per- gamentaceous; base symmetrical (to slightly ob- lique), acute to rounded, not or slightly decurrent; apex acute, fairly abruptly (to hardly acuminate), acumen fairly short, rounded to acute; above (gla- brous to) thin-puberulous on midrib and nerves, beneath with some scattered hair tufts on base of midrib only (to thin-puberulous on midrib and nerves); midrib above slightly raised to basally nearly flat; nerves 1.25—3 cm apart, above slightly raised; intersecondary nerves variably developed. Inflorescences axillary (to pseudoterminal), soli- tary, 1.5—15 cm long, sparsely and laxly branched, few- (to 1-)flowered; pedicels in flower c. 4 mm long. Only male flowers known. Sepals elliptic, c. 6 by 4 mm, persistent. Petals linear-lanceolate, c. 8 by 1.5 mm, white, glabrous. Disc uninterrupted, velvety. Stamens 5; filaments c. 5 mm long; an- thers c. 2 mm long. Pistil 2-locular; ovules 1 per locule. Fruits transverse-ellipsoid, 20—25 by 25- 30 mm, to subglobular, c. 2 cm in diam., the lobes erect, strongly bulging; stipe slightly hollowed, c. 1.5 mm long; apex obtuse; wall thin, hard, outside granulate and in old fruits coarsely reticulately veined, bright red, glabrous, inside glabrous or with scattered tufts of few hairs. Seeds black with a pink to bright red arillode. Distribution — Malesia: NE New Guinea. Habitat & Ecology — Primary and secondary rain forests, also Castanopsis-oak forest; 300-1500 m altitude. Fl. Jan., Oct.; fr. Jan., Mar., July. The fruits are said to be eaten by birds. 3. Harpullia carrii Leenh. in Leenh. & Vente, Blumea 28 (1982) 35. — Type: Carr 12443 (L holo; K, SING), SE New Guinea. Shrub or treelet, |.8—2.4 m high. Twigs and leaf axes thin-puberulous, glabrescent, inflorescences and infructescences short-velvety. 7wigs 3-5 mm thick. Leaves 1—S-jugate; petiole 4-10 cm long; petiolules 3—9 mm long. Leaflets elliptic, 5.5—23 by 2.5—9.5 cm, index 1.75—3, pergamentaceous; base symmetrical (or oblique), acute to rounded, only slightly decurrent; apex acuminate, acumen short to long, slender to broad, rounded; glabrous to sparsely hairy on midrib (and nerves); midrib above slightly raised to flat; nerves 1.25—3 cm apart, slightly raised above; intersecondary nerves fre- quent, variably developed. /nflorescences restrict- ed to the uppermost leaf axil (to the upper two ax- ils), solitary or few together, 4.5—12 cm long, hardly to repeatedly branched from the base onwards; pedicels in fruit c. 3 mm long. Sepals elliptic to broad-ovate, 3.5—S by 2-3 mm, persistent under the fruit. Petals glabrous (known in bud only). Disc uninterrupted, velvety. Stamens 5; anthers c. 1.2 mm long. Pistil 2- or 3-locular; style 2.5—3 mm long; ovules | per locule. Fruits with widely spread- ing ellipsoid lobes, c. 1.5 by 1.25 cm, the central axis 6-7 mm high ; stipe c. 3 mm high; wall thin- woody, outside pustular or coarsely veined, orange, glabrous, inside glabrous. Distribution — Malesia: Papua New Guinea (Central Prov., near Port Moresby). Habitat & Ecology — In primary and secondary forest; on ridges and steep rocky hillside, in the savannah in gallery forest; 300-400(—1200) m al- titude. FI., fr. May. Note — Harpullia carrii is distinctly closely al- lied to H. leptococca and occurs in the same re- stricted area. However, it has a distinctly different ecology and occurs at a higher altitude. 4. Harpullia cauliflora K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1900) 424; Radlk., Bot. Jahrb. 56 (1920) 312, p.p. (W New Guinea col- lections are H. ramiflora); in Engl., Pflanzenr. 98 (1934) 1440, p.p. (ditto); Merr. & L.M. Per- ry, J. Arnold Arbor. 21 (1940) 525; Leenh. & Vente, Blumea 28 (1982) 40. — Type: Kersting s.n. in herb. Lauterbach 2411 (B¥ holo; M, WRSL), NE New Guinea. Harpullia peekeliana auct. non Melch.: Kanehira & Hatusima, Bot. Mag. Tokyo 57 (1943) 79. (Shrub to) tree, up to 20 m high, dbh up to 20 cm. Twigs, leaf axes, inflorescences, and infruct- escences rather densely to sparsely puberulous, glabrescent to glabrous. Twigs 6-15 mm thick. Leaves 3—5-jugate; petiole 8-17 cm long; petiolules 4-12 mm long. Leaflets (ovate to) elliptic (to obo- vate), 2.5-33.5 by 5-13 cm, index 1.6—3, perga- mentaceous to chartaceous; base symmetrical, acute, and gradually long-decurrent (especially upper leaflets) to oblique, rounded, and abruptly shortly attenuate (lower ones); apex tapering to abruptly acuminate, acumen straight to slightly 606 Flora Malesiana ser. I, Vol. 11 (3) (1994) falcate, short to long, with rounded tip; on midrib and often on nerves densely to sparsely puberu- lous, especially beneath glabrescent; midrib above slightly raised; nerves 1.5—5.5 cm apart, slightly raised above; intersecondary nerves few to many, usually short and inconspicuous. /nflorescences axillary, rami-, and cauliflorous, in one or two groups of up to 3.5 cm long rachises, these not or hardly branched; pedicels in fruit 6-13 mm long. Sepals oblong-obovate to elliptic, 5—7 by 2.5—4.5 mm, outer 2 sometimes slightly smaller, persistent. Petals oblong, c. 7 by 3 mm or more, white, gla- brous. Disc short-velvety. Stamens 5; filaments c. 4 mm long or more; anthers c. 3 mm long. Pistil: 2-locular; ovules | per locule. Fruits transversely broad-ellipsoid to subglobular (or subtrapezoid), 1.5—2.5 by 1.8-3.2 cm, base obtuse to rounded to sometimes nearly truncate; stipe up to 3.5 mm long; apex rounded to emarginate; wall up to 1 mm thick, woody, outside granulate with scattered prominent pustules and prominulous irregular ridges, red, gla- brous or sparsely puberulous, inside glabrous to densely, shortly tufted-hairy. Seeds black or dark brown with an orange to red arillode. — Fig. 46i, j. Distribution — Malesia: New Guinea. Habitat & Ecology — Understorey of primary rain forest on alluvial plains; soil rocky clay or a mixture of sand, clay, and peat; 50-700 m altitude. Fl. Mar., Apr., July; fr. Mar., June, Oct., Nov. 5. Harpullia crustacea Radlk. in K. Schum. & Hollr., Fl. Kais. Wilh. Land (1889) 67; in Engl., Pflanzenr. 98 (1934) 1442; Leenh. & Vente, Blumea 28 (1982) 31. — Type: Hollrung 549 (M holo; BO, K, MEL), NE New Guinea. Harpullia cupanioides auct. non Roxb.: Hartley et al., Lloydia 36 (1973) 270, p.p. Tree, up to 24 m high, dbh up to 22.5 cm. Young parts, leaf axes, inflorescences, and infructescenc- es sparsely to moderately densely short hirsute. Twigs 9-15 mm thick. Leaves up to 1 m long, 5—9- jugate; petiole 4-23 cm long; petiolules 0-3 mm long. Leaflets elliptic, 6-31 by 3-13 cm, index 1.8— 3.4, papyraceous to thin-pergamentaceous; base symmetrical (or + oblique), acute to rounded (to subcordate); apex (rounded and hardly apiculate to) tapering acuminate, acumen short to long, acute to rounded; above densely puberulous on the mid- rib, more sparsely so on the nerves (to very sparsely all over), beneath thin puberulous on midrib and nerves (and on veins); midrib above slightly raised; nerves 0.5—2.3 cm apart, sunken above; intersec- ondary nerves many and often strongly developed, making the nervation sometimes irregular; veins hardly raised at both sides. Inflorescences axillary, pendulous, 30-60 cm long, simple; pedicels in fruit 4-6 mm long. Flowers fragrant. Sepals broad-ovate, 4.5-6.5 by 4.2-4.5 mm, persistent. Petals oblan- ceolate, 7—9 by 2.5—3.5 mm, white (to cream or yellowish green), (outside basally slightly hairy). Disc uninterrupted, 5-lobed, velvety. Stamens 5; filaments 3—5.5 mm long; anthers 2.5—3 mm long. Pistil 2-locular; style 3.5—4 mm long; ovules | per locule. Fruits transversely flattened broad-ellipsoid, c. 2 by 3-3.5 cm; base slightly cordate to truncate, abruptly narrowed into a |—2 mm long stipe; apex (slightly) emarginate; wall woody, outside promi- nently reticulate or pustulate, orange red, sparsely to moderately densely hairy, inside with a rather dense indumentum of longer tufted hairs. Distribution — Malesia: E New Guinea. Habitat & Ecology — In and along rain forest; on slopes, river banks, banks of mangrove creeks, also in swamps; on clay on volcanic rocks; sea level up to 480 m altitude. Fl. Feb./Mar., July, Sept., Nov./ Dec.; fr. Apr., Dec. 6. Harpullia cupanioides Roxb., [Hort. Beng. (1814) 86, nom. nud. (‘cuponioides’)] Fl. Ind. 2 (1824) 442; Migq., Fl. Ind. Bat. 1, 2 (1859) 570; Radlk., Sapind. Holl.-Ind. (1879) 94; Fern.- Vill. in Blanco, FI. Filip. ed. 3, Nov. App. (1880) 53 (p.p. = H. arborea); King, J. As. Soc. Beng. 65, II (1896) 451; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 239; Lecomte in FI. Indo- Chine 1 (1912) 1022, f. 126: 6-8; Koord. & Valeton, Atlas 1 (1913) pl. 142: a—-e; Radlk., Meded. Rijks-Herb. 22 (1914) 20 (Elbert 3261 = H. arborea); Bot. Jahrb. 56 (1920) 313 (ma- terial from W New Guinea = H. arborea); Merr., Enum. Philipp. Flow. Pl. 2 (1923) 516 (Ramos 30150 = H. ramiflora); Craib, Fl. Siam. Enum. 1 (1926) 335; Docters van Leeuwen, Zoocecidia (1926) 337, f. 609 & 610; Radlk. in Engl., Pflan- zenr. 98 (1934) 1444 (material from Celebes = H. arborea); Gagnep. in Fl. Indo-Chine, Suppl. 1 (1950) 954; Backer & Bakh. f., Fl. Java 2 (1965) 142; Hartley et al., Lloydia 36 (1973) 270 (p.p. = H. arborea); Leenh. & Vente, Blu- mea 28 (1982) 26; Yap in Tree Fl. Malaya 4 (1989) 444. — Type: Roxburgh s.n., 1813 (A, BO iso), Bangladesh. Ay-Assa Rumph., Herb. Amb. Auct. (1755) 20, nom. inval.; see Hassk., Abh. Naturf. Ges. Halle 9, 2 (1866) 329; Merr., Int. Rumph. (1917) 509. Tina rupestris Blume, Bijdr. (1825) 235; Span., Linnaea 15 (1841) 181. — Cupania rupestris Cambess., Mém. Mus. Hist. Nat. 18 (1829) 29. — Cupania blumei Steud., Nomencl. ed. 2, 1 (1840) 453, nom. illeg. — Harpullia rupestris Blume, Rumphia 3 (1847) 175; Migq., FI. Ind. Bat. 1, 2 (1859) 570; Radlk., Sapind. Holl.-Ind. Adema, Leenhouts, Van Welzen — Sapindaceae (1879) 50, 94. — Lectotype (Blume 1847): Blume 1625 (L holo), Java. {Donatophorus erythrospermus Zipp. ex Macklot, Bijdr. Natuurk. Wetensch. 5 (1830) 181, nom. nud.}. Harpullia confusa Blume, Rumphia 3 (1847) 176; Ridley, Fl. Malay Penins. 1 (1922) 510; Burk., Dict. Econ. Prod. Malay Penins. (1935) 1128. — Syntypes: Blume s.n. (L), W Java; Reinwardt gs.n. (L), E Java, Juti Kalangan. Harpullia fraxinifolia Blume, Rumphia 3 (1847) 177; Radlk. in Engl., Pflanzenr. 98 (1934) 1442. — Type: Spanoghe s.n. (L holo; M, P), Timor. Harpullia fruticosa Blume, Rumphia 3 (1847) 179; Radlk. in Engl., Pflanzenr. 98 (1934) 1443. — Type: Zippelius 168 (L holo), W New Guinea. Harpullia juglandifolia Blume, Rumphia 3 (1847) 177. — Syntypes: Korthals s.n. (L), Borneo; Korthals s.n. (L), SE Borneo, Dusun; Muller s.n. (L), Borneo; Anonymous s.n. (L), Borneo. Harpullia juglandifolia Blume var. multiflora Blume, Rumphia 3 (1847) 177. — Type: Jung- huhn s.n. (L holo), Sumatra. Harpullia thanatophora Blume, Rumphia 3 (1847) 178; Gresh., Meded. Lands Plantentuin 10 (1893) 44; Radlk. in Engl., Pflanzenr. 98 (1934) 1440. — Harpullia cupanioides Roxb. var. la- tifolia Miq., Fl. Ind. Bat. 1, 2 (1859) 571. — Syntypes: Zippelius 134a (A, L, S), 185b (L), W New Guinea. Harpullia macrocalyx Radlk., Philipp. J. Sc., Bot. 8 (1914) 473; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 516; Radlk. in Engl., Pflanzenr. 98 (1934) 1441. — Type: Loher 5891 (M holo), Philippines. Harpullia obscura Radlk., Bot. Jahrb. 56 (1920) 314; in Engl., Pflanzenr. 98 (1934) 1448. — Syntypes: Ledermann 12494, 12968 (B+, M), both NE New Guinea. Harpullia longithyrsifera Kanehira & Hatusima, Bot. Mag. Tokyo 57 (1943) 78, f. 12. — Type: Kanehira & Hatusima 14234 (A, BO iso), W Irian Jaya. Harpullia sp.: Ceron, Cat. Pl. Herb. Manila (1892) 55 as for Vidal 2525 & 2527. Walsura villosa auct. non Wall.: Ridley, Fl. Malay Penins. | (1922) 412, as for coll. Burn-Mur- doch trom Gombak, Selangor. Harpullia reticulata auct non Radlk.: Radlk., Nova Guinea 14 (1926) 185; in Engl., Pflanzenr. 98 (1934) 1443, as for both collections by Feuille- teau de Bruyn. (Shrub to) tree; up to 20(—40) m high, dbh up to 40(—100) cm; (buttresses up to 2 m high and 2 m spreading). 7wigs 2-10 mm thick; usually gla- 607 brous but for the terminal bud (to all young parts up to and including the infructescences densely appressedly short-hairy and usually early glabres- cent). Leaves (1—)3—6(—7)-jugate; petiole up to 20 cm long; petiolules 2—12 mm long; axes glabrous (or sparsely puberulous). Leaflets ovate (lower ones) to elliptic to obovate (upper ones), 5-36 by 2—15 cm, index 1.5—4, herbaceous to chartaceous; base symmetrical to oblique, acute or the broader apical half (rarely both halves) rounded, (not or) only slightly decurrent; apex acute to rounded to gradually to abruptly acuminate, acumen short to rather long, rounded or retuse (to acute); glabrous or very sparsely hairy on midrib and below on nerves; midrib above usually slightly raised to ba- sally sunken, (or raised or sunken over its whole length); nerves 0.75—3 cm apart, above slightly sunken; intersecondary nerves feeble (or absent). Inflorescences axillary (to pseudoterminal to truly terminal), erect or pendulous, infructescences more often + pendulous, solitary, 5-85 cm long, simple (or with a strong branch near the base), usually sparsely and + widely branched, often + few-flow- ered, the upper parts usually tardily hairy, for the rest glabrescent; pedicels in fruit 3—7.5 mm long; bracts small, simple (or ternate) leaves. Flowers fragrant. Sepals elliptic to suborbicular, 3—6 by 2.5— 4.5 mm, persistent in fruit. Petals oblong-obovate to oblanceolate, 5—10 by 2—3 mm, white to creamy (yellow, greenish, pink), glabrous. Disc complete, low, short-velvety. Stamens 5 (or 6); filaments 2.5— 3.5 mm long, white; anthers 1.5—3.5 mm long, yel- lowish white, greyish yellow, or dark mauve. Pis- til 2-locular; ovary light green, yellow, or reddish brown; style 1.75—6.5 mm long, light green, stig- ma whitish; ovules | per locule (to 2 in one of the locules). Fruits slightly kidney-shaped, transversely ellipsoid, broadly ovoid, obovoid, or globular, 12— 20 by 12—32.5 mm; base rounded to truncate (to slightly concave), stipe 1—3 mm long; apex slight- ly concave to obtuse-angular, apiculate; wall leath- ery to woody, outside and inside red, outside vari- ably hairy, early to late glabrescent, inside glabrous to very laxly rather long hairy or glandular hairy. Seeds shining brown to black with a bright glossy red arillode. — Fig. 46g, h. Distribution — Southern China, Assam, the An- daman Islands, Bangladesh, Burma, Thailand, Indo- China, throughout Malesia, and the Northern Ter- ritory of Australia (Entrance I., Port Essington). Habitat & Ecology — Primary and secondary rain forest, more rarely in light forest, teak forest, tidal forest, scrub, or on open places; often on slopes and ridges, on river banks and along ravines, also on flats and along the beach, usually on dry (to swampy) soil; rocky, sand, clay, or loam, usually 608 Flora Malesiana ser. I, Vol. 11 (3) (1994) on limestone on fertile volcanic soils; sea level up to 1200(—1800) m altitude. Fl. (Jan.—)Apr.—July(— Dec.); fr. (Jan.—)Apr.—Oct.(—Dec.). Uses — The wood is used for charcoal and as firewood; the bark is used as a fish poison. Notes — 1. Harpullia cupanioides is best dis- tinguished from its allies by the combination of glabrous vegetative parts (except for the terminal buds) and exclusively axillary inflorescences. Noth- withstanding its rather wide distribution it is not very variable. However, there is a gradual shift from a mainly western ‘cupanioides’ type towards an eastern ‘thanatophora’ type, the latter being char- acterized by fewer, larger, and broader leaflets with more distant nerves, by less hairy inflorescences, by smaller flowers, and by bigger fruits that are earlier glabrescent. 2. The type of H. fruticosa from West New Guinea is the only specimen in which the leaflets are quite densely hairy along the complete length of the midrib. 3. The two collections made by Feuilleteau de Bruyn from Schouten Island differ in their thick coriaceous leaflets with very pronounced and dense reticulation. 4. A truly terminal inflorescence is known only from Verheijen 3241, from Flores. 5. A somewhat doubtful collection is PNH (Su- lit) 10113 from Mindanao. It differs mainly in its unusually big fruits, which are up to 25 by 45 mm. This may represent a separate species, though in the H. cupanioides alliance. 7. Harpullia giganteacapsula Vente in Leenh. & Vente, Blumea 28 (1982) 17. — Type: NGF (Streimann & Kairo) 27614 (L holo; BRI, K), NE New Guinea. Tree, up to 20 m high, dbh up to 80 cm. Young parts with a short, dense, brown indumentum, gla- brescent. Twigs 6-21 mm thick. Leaves (2—)4—5- jugate; petiole 7.5—35 cm long; petiolules 2-10 mm long; axes hairy, glabrescent. Leaflets elliptic to obovate (upper ones), 7.5—26 by 4-11 cm, index 1.7—3, herbaceous to thin-pergamentaceous; base symmetrical to slightly oblique, acute to rounded, short-decurrent; apex rather abruptly (or gradual- ly) acuminate, acumen short to moderately long, acute to rounded; above slightly hairy on midrib (and nerves), beneath sparsely to densely hairy on midrib and nerves; midrib above sunken; nerves 1.2-4.3 cm apart, above flat or sunken; intersec- ondary nerves few, usually feeble. Infructescences axillary, 10-33 cm long, with few up to 6 cm long branches, hairy; bracts lanceolate, up to 2.5 cm long; pedicels in fruit up to 18 mm long. Flowers only known from remnants under the fruit. Sepals broad-elliptic to suborbicular, 6-8 by 4.5—7 mm, persistent or caducous. Disc uninterrupted, hairy. Stamens 7 or 8 (scars). Pistil 2-locular; style c. 4.5 mm long; ovules 2 per locule, the lower one with a long thin funicle, probably abortive. Fruits trans- versely broad-ellipsoid, c. 35 by 58 mm; base emar- ginate, stipe 1-2 mm long; apex emarginate to trun- cate; lobes erect, strongly inflated; wall thin woody, outside minutely pustulate and sulcate, densely hairy, glabrescent, inside very sparsely hairy. Seeds 1 per locule. — Fig. 46k. Distribution — Malesia: E New Guinea. Habitat & Ecology — In forests on ridges and slopes; limestone; 1350-1850 m altitude. Fr. Apr.— June, Oct. 8. Harpullia hirsuta Radlk., Nova Guinea 8 (1912) 618; in Engl., Pflanzenr. 98 (1934) 1450; Leenh. & Vente, Blumea 28 (1982) 33. — Type: von Rémer 980 (L holo; M), SW New Guinea. Tree. Twigs and leaf axes brownish hirsute. Twigs c. 3 mm thick. Leaves 2- or 3-jugate; petiole 5-7 cm long; petiolules 4-6 mm long. Leaflets el- liptic, 11-20 by 5—6.5 cm, index 2—3, pergamenta- ceous; base symmetrical or slightly oblique, acute and decurrent; apex acuminate, acumen long, fal- cate, with acute tip; above densely hirsute on mid- rib and nerves, more sparsely so all over, beneath hairy on midrib, sparsely so all over; midrib and nerves slightly raised above; nerves 1.5—2.5 cm apart; intersecondary nerves few. Inflorescences axillary, probably unbranched, 1.5 cm long (or more?). Flowers known from female buds only. Sepals elliptic. Petals thin, not (yet?) clawed, not auriculate, glabrous. Disc densely long-hairy. Sta- mens 5. Pistil 2-locular; ovary densely hairy; ovules 1 per locule. Fruits unknown. Distribution — Malesia: SW New Guinea (only known from the type); a second doubtful collec- tion from the Vogelkop Peninsula. Note — The present species is very incomplete- ly known. It may be closest to H. camptoneura. 9. Harpullia leptococca Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 278; in Engl., Pflanzenr. 98 (1934) 1450; Leenh. & Vente, Blumea 28 (1982) 34. — Type: Chalmers s.n., 1880 (M holo; MEL), SE New Guinea. Harpullia camptoneura auct. non Radlk.: Rehder, J. Arnold Arbor. 14 (1933) 64. Erect bushy tree, up to 9 m high. Young parts, leaf axes, inflorescences, and infructescences densely, appressedly, minutely hairy. Twigs 3.5—7 mm thick. Leaves 2- or 3(—5)-jugate; petiole 2.5-8 Adema, Leenhouts, Van Welzen — Sapindaceae 609 cm long; petiolules 2—5(—7) mm long. Leaflets el- liptic to obovate, 4.5—15 by 2.5—8 cm, index 1.5— 3, chartaceous; base symmetrical to slightly ob- lique, rounded to acute and usually (abruptly) short- decurrent; apex rounded to obtuse(to slightly emar- ginate); glabrous or very sparsely hairy on the mid- rib below; midrib above slightly sunken to slightly raised; nerves 0.9—2(—2.6) cm apart, about equally prominent on both surfaces; intersecondary nerves rather many, usually rather feeble. /nflorescences axillary, pseudoterminal, or terminal, 8.5-30 cm long, solitary and simple or with 2 equally strong basal branches (to with some more higher up); pedi- cels in flower and fruit 2—4 mm long. Sepals obo- vate to elliptic to orbicular, 4—5.5 by 3-4 mm, per- sistent. Petals oblong to lanceolate to oblanceo- late, 6-7.5 by 2—2.5 mm, creamy green to creamy white, glabrous (or in the same flower some slightly hairy). Disc uninterrupted, short-velvety. Stamens 5(—7). Pistil 2-locular; style 2.5—5 mm long; ovules | per locule (to 2 in one of the locules). Fruits with widely spreading, obovoid to ellipsoid lobes, 5—6 by 27-34 mm; base cordate to obtuse, stipe up to 2 mm long; wall pergamentaceous, outside smooth with faint longitudinal nerves, orange to red, sparse- ly short-hairy, inside very sparsely short-hairy. Seeds black with a red arillode. — Fig. 46 1. Distribution — Malesia: SE New Guinea (Cen- tral Prov., mainly around Port Moresby and on Yule I.). Habitat & Ecology — In and along coastal scrub, between strand vegetation and grassland, on savan- nah, in the undergrowth of semi-deciduous forest; along the shore, behind the mangrove, on tidal flats, also on sandhills and ridges; sea level up to 60 m altitude. Fl. Apr., Aug.; fr. Apr., July—Sept., Nov. 10. Harpullia longipetala Leenh. in Leenh. & Vente, Blumea 28 (1982) 18. — Type: Carr 11498 (L holo; K, SING), SE New Guinea. Shrub or tree, up to 18 m high, dbh up to 20 cm. Glabrous but for shortly dark- to orange-brown velvety buds and inflorescences. Twigs 3-14 mm thick. Leaves 2—4-jugate; petiole 4-15 cm long; petiolules 5S—10 mm long. Leaflets + elliptic, 12.5— 32 by 4-14 cm, index 2-4, thin- to stiff-pergamen- taceous, chartaceous, or coriaceous; base symmet- rical (to oblique), acute (to rounded lower), usual- ly slightly decurrent; apex rounded to gradually to abruptly acuminate, acumen short (to long), round- ed to acute; midrib usually slightly raised above; nerves |—5 cm apart, above slightly to hardly raised; intersecondary nerves absent (to well developed). Inflorescences axillary, rami-, and cauliflorous, in dense groups, 2—2.5 cm long, axes racemoid, few- flowered; pedicels in fruit c. 1 cm long. Sepals: outer roundish, c. 6 by 5 mm, inner broad-elliptic, c. 7 by 5 mm; persistent in fruit. Petals oblanceo- late, c. 15 by 4 mm, white, cream, or very pale mauve, glabrous. Disc velvety. Stamens 8, white; filaments up to 9 mm long; anthers c. 2.5 mm long. Pistil 2-locular; style c. 4.5 mm long; ovules 2 per locule (to 1 in one of the locules). Fruits broad- cordate, c. 18 by 22-25 mm, widest at or above the middle, orange to red; base + truncate to broadly rounded, stipe c. 2 mm long; wall leathery, outside densely short-velvety, + glabrescent, inside very sparsely hairy to glabrous. Seeds usually | per loc- ule, arillode bright red. Distribution — Malesia: E New Guinea. Habitat & Ecology — Rain forest, sometimes dry monsoon forest, swamp forest, or secondary for- est; on river banks, hill sides, in gullies, also in coastal savannahs; usually below 100 m (up to 1650 m) altitude. Fl. Nov._Feb.(—Apr., June, July, Sept.); fr. July—Sept. (Nov., Mar.). 11. Harpullia myrmecophila Merr. & L.M. Per- ry, J. Arnold Arbor. 21 (1940) 526; Leenh. & Vente, Blumea 28 (1982) 39. — Type: Brass 13414 (A holo; BO, L), NW New Guinea. Treelet, up to 4 m high. All young parts, includ- ing twigs, leaf axes, and infructescences, moder- ately to (latter) densely puberulous, glabrescent. Twigs 6—7 mm thick. Leaves 3- or 4-jugate; peti- ole 8.5—-11.5 cm long; petiolules 5-7 mm long. Leaflets elliptic, 14.5—24.5 by 7.5—9 cm, index 1.9— 2.8, chartaceous; base symmetrical or in upper leaf- lets oblique, acute and gradually, to rounded and abruptly (lowermost leaflets) decurrent; apex + abruptly acuminate, acumen rather long, straight (to falcate), the tip rounded; midrib and nerves above fairly densely, beneath sparsely puberulous; midrib and nerves above slightly raised; nerves 2.2— 3.5 cm apart; intersecondary nerves few, short. /n- florescences axillary and solitary or ramiflorous and in clusters of c. 3, not to sparsely branched, 6—15 cm long; pedicels in flower and fruit 49 mm long. Sepals ovate to elliptic, 4.5—5 by 3-4 mm, persist- ent. Petals oblanceolate, c. 10 by 3.2 mm, green- ish white, glabrous. Disc uninterrupted, short-vel- vety. Stamens 5; filaments c. 2.5 mm long; anthers 3.5-4 mm long. Pistil 2-locular; ovules | per loc- ule. Fruits broad-ellipsoid, 24-26 by 22-25 mm, rounded at base and apex, stipe c. 0.5 mm long; wall thin, woody, outside granulate and irregular- ly pustulate, red, very sparsely minutely hairy, in- side glabrous. Seeds with a yellow arillode. Distribution — Malesia: NW New Guinea (Iden- burg R. near Bernhard Camp). Habitat & Ecology — Rain forest, on stream bank; altitude c. 850 m. FI., fr. Mar. 610 Flora Malesiana ser. I, Vol. 11 (3) (1994) 12. Harpullia oococca Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 278; in Engl., Pflanzenr. 98 (1934) 1449: Leenh. & Vente, Blumea 28 (1982) 35. — Type: Sayer s.n., 1887 (M holo; MEL), SE New Guinea. Harpullia hirsuta auct. non Radlk.: Kanehira & Hatusima, Bot. Mag. Tokyo 57 (1943) 77. Straggling tree, c. 6 m high. Twigs and leaf axes sparsely puberulous, infructescences short-velvety. Twigs 2 mm thick (or more?). Leaves 3—(to more?)- jugate; petiole c. 2.5 cm long; petiolules 7-10 mm long. Leaflets elliptic, 10-30 by 5.5—11 cm, index 2-3, pergamentaceous; base symmetrical to slightly oblique, acute and slightly decurrent; apex rather abruptly acuminate, acumen short, acute; densely puberulous on midrib and nerves, more sparsely so on veins beneath; midrib slightly raised above; nerves |.5—4 cm apart, hardly raised above; some intersecondary nerves strongly developed. /nfruct- escences (position unknown) probably solitary, probably no more than 5—6 cm long, with some strong patent branches near the base; pedicels c. 7.5 mm long. Sepals elliptic, c. 5 by 3 mm, per- sistent. Disc velvety. Fruits with 2 widely spread- ing ellipsoid lobes of c. 1.5 by 1 cm, central axis 4-7 mm high with ac. 3 mm long stipe and crowned by ac. 5 mm long style; wall thin-woody, outside pustular, orange, thinly puberulous, probably gla- brescent, inside glabrous. Seeds 1 per locule, or- ange with a deep carmine arillode. Distribution — Malesia: New Guinea (SE, Cen- tral Prov., once collected; 2 doubtful collections from the West: Nassau Mts and near Nabire). Habitat & Ecology — Altitude c. 600 m. 13. Harpullia peekeliana Melch., Notizbl. Bot. Gart. Berlin-Dahlem 10 (1928) 279; Radlk. in Engl., Pflanzenr. 98 (1934) 1461; Leenh. & Vente, Blumea 28 (1982) 44. — Type: Peekel 987 (B+ holo), Papua New Guinea. Tree, c. 12 m high. Vegetative parts glabrous, inflorescences olive-brown tomentellous. Leaves 2-4-jugate, 42-50 cm long; petiolules 5-15 mm long. Leaflets elliptic or oblong-elliptic, 18—40 by 11—16 cm, coriaceous; base symmetrical, short-at- tenuate; apex acuminate; midrib and nerves above not or hardly raised. Inflorescences rami- and cau- liflorous, 2-10 cm long, paniculate; pedicels in flower 10—12 mm long. Only male flowers known. Sepals broad-elliptic, 7-8 by c. 5 mm. Petals c. 12 by 3.5 mm, narrowed at base, not auriculate, fleshy, glabrous. Disc low, tomentellous. Stamens 5, c. 9 mm long, filaments and anthers of about the same length. Pistil and fruits unknown. Distribution — Malesia: New Ireland (Lamekot to Panamangaf); only known from the type. Habitat & Ecology — Altitude c. 200 m. Fl. May. Note — Harpullia peekeliana is only known from Melchior’s original description: the type was lost in the Berlin fire and apparently there are no du- plicates in other herbaria. There is hardly any doubt that it belongs to the alliance of H. ramiflora, where it seems nearest to H. solomonensis. Comparison is difficult as only male flowers have been described from H. peekeliana, while only fruits and the re- mains of the female flowers under these are known for H. solomonensis. The two specimens from the Bismarck Archipelago, referred to H. solomonen- sis, agree fairly well vegetatively with H. peekeli- ana, but their infructescences are axillary. More- over, both are from coastal habitats, whereas Peekel collected his material at 200 m altitude. More col- lections from the Bismarck Archipelago are a pre- requisite to establish the position of H. peekeliana and its relationship to 4. solomonensis. 14. Harpullia petiolaris Radlk., Bot. Jahrb. 56 (1920) 315; in Engl., Pflanzenr. 98 (1934) 1450; Leenh. & Vente, Blumea 28 (1982) 41. — Lectotype (Leenhouts & Vente 1982): Le- dermann 8814 (B* holo; M), NE New Guinea. Harpullis cupanioides auct. non Roxb.: Holth. & Lam, Blumea 5 (1942) 208. Tree, up to 6(—20) m high, dbh c. 4 cm. Twigs, leaf axes, inflorescences, and infructescences (very) densely set with patent, short or long, brown hairs or minutely tomentose, + early glabrescent. Twigs (5—)10—13 mm thick. Leaves 4—6(—7)-jugate; peti- ole 10—27(—33) cm long; petiolules 0-4 mm long. Leaflets ovate to elliptic, 9-44 by 3.5—15 cm, in- dex 2.1—3.6, herbaceous to thin-pergamentaceous; base symmetrical and subcordate to rounded (to oblique with one half acute, the other rounded); apex gradually (or fairly abruptly) acuminate, acu- men short to rather long with acute tip; glabrous to (sparsely above to) densely hairy on midrib and nerves, sparsely on veins below; midrib above + raised; nerves 1—3.3 cm apart, above slightly sunken (to slightly raised); intersecondary nerves often strongly developed. Inflorescences cauliflorous, in groups of 1—5 rachises, 22-50 cm long, simple or with few widely spaced, erecto-patent, up to 7 cm long branches mainly in the basal half, lax; cymes scattered along rachis and branches; pedicels in fruit 5-8 mm long. Flowers without smell. Sepals broad- elliptic, 4-6.5 by 3.2-4.5 mm: in fruit persistent or caducous. Petals oblanceolate, (7—)11—12.5 by 2.5-4 mm, white, glabrous (to ciliate at base). Disc low, 5-angular, short-velvety. Stamens 5; filaments 2—5.5 mm long; anthers 3-4.75 mm long. Pistil 2- Adema, Leenhouts, Van Welzen — Sapindaceae 611 locular; style c. 6 mm long; ovules | per locule. Fruits transversely ellipsoid, 16—20 by 26-28 mm; base slightly cordate to rounded, stipe 1-2 mm long; apex slightly emarginate or obtuse; wall woody, outside irregularly pustulate, red, puberulous to pubescent, glabrescent, inside glabrous. Seeds dark brown to black with a yellow to red arillode. Distribution — Malesia: Borneo, Moluccas, and New Guinea. KEY TO THE SUBSPECIES AND VARIETIES la. Twigs, leaves, inflorescences, and infructes- cences thin-puberulous. Nerves spaced... 2 (a. subsp. moluccana) b. Twigs, leaf axes, midrib, and nerves on lower side of leaflets, and inflorescences ferrugine- ous-velvety. Nervation rather dense d. subsp. petiolaris a. Sepals persistent in fruit. ¢. var. moluccana b. Sepals caducous short after fertilization b. var. decidens a. subsp. moluccana Leenh. in Leenh. & Vente, Blumea 28 (1982) 42. — Type: De Vogel 4102 (L holo), Moluccas. For characters see the key. Distribution — Malesia: Borneo and Moluccas. b. var. decidens Vente in Leenh. & Vente, Blumea 28 (1982) 43. — Type: De Vogel 3077 (L holo), Moluccas. For characters see the key. Distribution — Malesia: Moluccas (Halmahera). Habitat & Ecology — Rather dense primary for- est, with very little undergrowth on plane or slight slope; soil clay; altitude 10-20 m. FI., fr. Oct. c. var. moluccana — Leenh. & Vente, Blumea 28 (1982) 43. For characters see the key. Distribution — Malesia: Borneo and Moluccas (Batjan [.). Habitat & Ecology — Rather dense primary for- est with little undergrowth on dry soil, clay with limestone boulders; altitude up to 500 m. Fr. May. d. subsp. petiolaris — Leenh. & Vente, Blumea 28 (1982) 42. For characters see the key. Distribution — Malesia: New Guinea. Habitat & Ecology — Primary and secondary forests, Fagaceous and marsh forest; altitude up to 1400 m. Fl. Mar., May, Sept., Nov.; fr. Oct. Uses — Sap is used for poisoning fish. 15. Harpullia ramiflora Radlk., Sapind. Holl.- Ind. (1879) 15, 54; in Engl., Pflanzenr. 98 (1934) 1439; S.T. Reynolds, Austrobaileya | (1981) 415, f. 29g; Leenh. & Vente, Blumea 28 (1982) 38; S.T. Reynolds in Fl. Austral. 25 (1985) 42, map 48. — Type: Beccari herb. 2822 (FI holo), Aru. Islands. Harpullia angustifolia Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 599; in Engl., Pflanzenr. 98 (1934) 1439. — Type: d’Albertis s.n. in herb. Beccari 2892 (FI holo), SE New Guinea. Harpullia aeruginosa Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 278; in Engl., Pflanzenr. 98 (1934) 1448. — Type: Chalmers s.n., 1885 (M holo), SE New Guinea. Harpullia weinlandii K. Schum. in K. Schum. & Laut., Nachtr. Fl. Schutzgeb. Siidsee (1905) 310; Radlk. in Engl., Pflanzenr. 98 (1934) 1449. — Type: Weinland 258 (B¥ holo; BRI, L, M, SING, US, WRSL), NE New Guinea. Harpullia reticulata Radlk., Bot. Jahrb. 56 (1920) 313; in Engl., Pflanzenr. 98 (1934) 1443 (type only; further collections = H. cupanioides). — Type: Branderhorst 439 (M holo; BO), SW New Guinea. Harpullia cupanioides auct. non Roxb.: Koord., Nova Guinea 8 (1909) 171; Radlk., Bot. Jahrb. 56 (1920) 313, as for W New Guinea; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 516, as for Ramos 30150; Rehder, J. Arnold Arbor. 14 (1933) 64. Harpullia cauliflora auct. non K. Schum. & Laut.: Radlk., Nova Guinea 8 (1912) 618; Bot. Jahrb. 56 (1920) 312, as for W New Guinea; in Engl., Pflanzenr. 98 (1934) 1440, ditto. Harpullia camptoneura auct. non Radlk.: Hartley et al., Lloydia 36 (1973) 270. Bushy to sparsely branched shrub or tree, up to 15 m high, dbh up to 15 cm. Twigs and leaf axes, or only buds, inflorescences, and infructescences sparsely to densely variably hairy, + glabrescent. Twigs 3-11 mm thick. Leaves (1—)3—4(—7)-jugate: petiole (2—)5—14(—18) cm long; petiolules (2—)4— 8(-11) mm long. Leaflets ovate to elliptic, (+ fal- cate), (4—)8—20(—31) by (1.5—)3.5—8(—15) cm, in- dex (1.5—)2—4, herbaceous to chartaceous, + bul- late if thin; base symmetrical (to oblique), acute to (especially in the lower leaflets) rounded, not or variably decurrent; apex acute to (in obovate leaf- lets) rounded to acuminate, the acumen short to long, straight to strongly curved, acute to round- ed, (mucronulate); glabrous to (sparsely to) densely hairy on midrib and nerves, sparsely between nerves below; midrib basally slightly sunken to apically slightly raised to completely raised; nerves 0.5—6 cm apart, above flat to raised; intersecondary nerves 612 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1com i Ve ssenaer} ate Fig. 47. Harpullia rhachiptera Radlk. Leaf (LAE 51779). Adema, Leenhouts, Van Welzen — Sapindaceae 613 hardly to fairly strongly developed. /nflorescences initially axillary, after leaf-drop rami- to cauli- florous, at first solitary (or some together), later clustered, (in fruit pendent), 1—20(—40) cm long, simple or sparsely branched, mainly in the lower part, few-flowered; pedicels in fruit 4-10 mm long. Flowers fragrant. Sepals ovate (outer) to elliptic (or inner orbicular), 3.5—6 by 2.5—4.5 mm, persist- ent. Petals (lanceolate or) oblanceolate to obovate, 6-13.5 by (1.5—)2—3 mm, white to creamy, gla- brous. Disc uninterrupted, low, short-velvety (or thin-puberulous). Stamens 5; filaments 5—6.5 mm long; anthers 2.25—3.5 mm long. Pistil 2-locular; style (2—)4.5—7 mm long; ovules | per locule. Fruits transversely ellipsoid to broadly heart-shaped (or obtriangular), | 1—16(—22) by 16-25 mm; base trun- cate to rounded (to angular), stipe 1—2(—3) mm long: apex emarginate (or truncate to rounded); wall per- gamentaceous to leathery, outside minutely warty or coarsely reticulately veined (or smooth), orange to bright red, fairly densely puberulous to short- velvety, inside orange, glabrous or very sparsely hairy. Seeds black with a shining orange arillode. — Fig. 46m. Distribution — Malesia: Philippines (Catandu- anas), Moluccas (Halmahera), New Guinea: Queensland. Habitat & Ecology — Mainly in the undergrowth or primary rain forest and in young secondary for- est, in gallery forests, coastal scrub, monsoon for- est, periodically flooded forest, and sago swamp; on river banks, in gulleys, on steep slopes and ridg- es; on stony soil, loam, or clay; altitude sea level up to 200(—1950) m. FI., fr. Feb.—Sept.(—Nov.). 16. Harpullia rhachiptera Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 278; in Engl., Pflanzenr. 98 (1934) 1440; Verdc., Kew Bull. 32 (1977) 223; Leenh. & Vente, Blumea 28 (1982) 23. — Type: Bauerlen 191 = Capt. Everett’s Exp. s.n. (M holo; MEL), SE New Guinea. [Affonsea pteropoda Kosterm., Adansonia n.s. 6 (1966) 371, pl. 5, nom. inval. — Lectotype (Verdcourt 1977): d’Albertis herb. 3544 & 3544a (FI), SE New Guinea. ] Shrub, less than | m high, to treelet up to 3 m high. Young parts, inflorescences, and infructes- cences densely, appressedly, shortly, brown hairy. Twigs 4—6 mm thick. Leaves 3- or 4-jugate; petiole 8—11 cm long, winged except for the basal 1/4-1/2, wings 2—3.5 mm wide; rachis winged, wings 4.5-6 mm wide, widest beneath leaflet; petiolules 0-3 mm long; axes hairy. Leaflets ovate to elliptic, 7.5—25 by 2.5—10 cm, index 2—3.75, her- baceous; base symmetrical, narrowly rounded to subcordate; apex gradually to rather abruptly acu- minate, acumen short to rather long, straight, with rounded tip; midrib and nerves very sparsely to densely minutely hairy, below also with glandular hairs; midrib above slightly raised; nerves 1.5—2.5 cm apart, slightly raised above; intersecondary nerves few, faint. Inflorescences axillary, simple or slightly branched, up to 3.5 cm long, few-flow- ered; pedicels in fruit 3-4 mm long. Female flow- ers unknown. Sepals elliptic or (outer) ovate, 4.5— 5.5 by 2.5—3 mm, in fruit semipersistent. Petals oblanceolate, c. 9 by 2 mm, white, glabrous. Disc annular, puberulous. Stamens 5 or 6; filaments c. 8 mm long; anthers 2.3—2.5 mm long. Pistil 2-locu- lar; style c. 3.5 mm long; ovules | per locule. Fruits broad-ellipsoid to subglobular, 14-22 by 18—20 mm, red, base obtuse to truncate, short-stipitate, apex (sub)emarginate; wall thin coriaceous, out- side prominently coarsely reticulate and pustulate, pubescent, inside glabrous. Seeds black with a yel- low to orange arillode. — Fig. 47. Distribution — Malesia: New Guinea ( Vogelkop Peninsula, Western Prov.). Habitat & Ecology — In undergrowth of forest, also in swamp forest, on ridges and river banks: altitude 45-90 m. FI. Aug.; fr. Apr—June, Aug. Note — The present species seems on the one hand to be allied with the Australian H. fruticosa group, especially with H. rhyticarpa, and on the other hand with H. cupanioides. 17. Harpullia solomonensis Vente in Leenh. & Vente, Blumea 28 (1982) 36. — Type: Kajew- ski 1968 (BRI holo; A, BISH, BO, P), Bou- gainville Island. Tree, up to 18 m high, dbh up to 25 cm. Twigs and leaf axes subglabrous, infructescences thin- puberulous. 7wigs 9-11 mm thick. Leaves 3-jugate, up to 75 cm long; petiole 9.5—18 cm long; peti- olules (6—)9—12 mm long. Leaflets ovate to ellip- tic, 11-42 by 6-15.5 cm, index 1.8—2.7, herbaceous to pergamentaceous; base symmetrical, acute or rounded, short-decurrent; apex tapering-acuminate, acumen long; above sparsely puberulous on mid- rib (and nerves), beneath glabrous; midrib and nerves above flat to slightly raised, nerves 2—3.5 cm apart; intersecondary nerves few. /nflorescences axillary to cauliflorous, densely clustered, the ra- chises branched at base, the axillary ones up to 4.5 cm long, those on the trunk up to 22 cm long; pedi- cels 8-12 mm long. Flowers without or with a strong smell, greenish- to golden-yellow; known only from remnants under the fruit. Sepals (sub)orbicular to elliptic, (3—)4—6 by 3—5 mm, per- sistent. Disc short-velvety. Stamens 5. Pistil 2-loc- ular; style c. 6 mm long; ovules | per locule. Fruits 614 Flora Malesiana ser. I, Vol. 11 (3) (1994) with erect, ellipsoid lobes, (17—)23—24(—30) by (25—)28—34 mm; base broadly rounded, stipe c. 3 mm long; apex slightly emarginate; wall slightly woody, outside granulate, yellow to orange, varia- bly minutely hairy, inside glabrous. Distribution — Solomon Islands, possibly also in Malesial Bismarck Archipelago. Habitat & Ecclogy — Well-drained primary rain forest on ridge tops, hill sides, and in deep valleys; altitude 150—250(—900) m. FI. Apr., June, Nov.; fr. Mar.—Apr., June—July. Notes — 1. Harpullia solomonensis differs from its nearest related species, H. ramiflora, particu- larly in its inflorescences, which, when axillary, are short, dense clusters developing from both ax- illary buds, in its usually longer pedicels, and in its larger fruits with a thicker and rather woody wall. 2. Two collections from the Bismarck Archi- pelago, NGF 45722 from New Britain and NGF 29839 from New Ireland, may belong to this spe- cies. However, the nearly unknown H. peekeliana was described from New Ireland and seems to be closest to H. solomonensis. This problem can be solved only when more material from the Bismarck Archipelago becomes available. 18. Harpullia vaga Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 526; Leenh. & Vente, Blu- mea 28 (1982) 30. — Type: Kajewski 2544 (A holo; BISH, BM, BO, BRI, L, P, SING), Solo- mon Islands. Tree, up to 18 m high, dbh up to 48 cm. Jndu- mentum dense, short, yellowish brown, and hirsute. Twigs 4—10 mm thick, sometimes hardly more than the terminal bud hairy. Leaves (2- to) 3—-5-jugate; petiole 4-14 cm long; petiolules 3-8(—10) mm long; axes fairly densely hairy, glabrescent. Leaflets ovate to elliptic, 4.75-27 by 2.5-8.5 cm, index 2-4.5, chartaceous to papyraceous; base oblique to sym- metrical, acute to (lower) obtuse to rounded, at- tenuate; apex gradually acuminate, acumen long, and acute (or short and rounded); above (glabrous to) sparsely hairy on midrib (and nerves), beneath subglabrous to thinly hairy (to densely tomentel- lous) all over, most densely on midrib and nerves; midrib above slightly raised to slightly sunken, nerves 1—2.5 cm apart, slightly raised above; in- tersecondary nerves few and usually feeble. Inflo- rescences axillary, solitary, pendulous, 10—37(—70) cm long, subspicate to thyrsoid; pedicels in fruit 5-10 mm long. Sepals ovate, 6-8 by 5—6 mm, per- sistent in fruit. Petals glabrous. Disc uninterrupt- ed, puberulous. Stamens 5. Pistil 2-locular; ovules 1 or 2 per locule. Fruits transversely broad-ellip- soid, kidney-shaped, broad-ovoid, or subglobular, up to 3 by 4.5 cm; base obtuse or rounded, stipe c. 2.5 mm long; apex rounded or sub-emarginate; wall woody, 1—1.5 mm thick, outside orange, coarsely granulate (to with rather big pustules), subglabrous, inside glabrous. Distribution — Queensland, the Solomon Islands, and Malesia: New Ireland (?). Habitat & Ecology — In well-drained primary or rarely secondary rain forest on slopes or ridge; altitude sea level up to 200(—1200) m. Fl. May, July—Sept., Dec.; fr. Feb.—Mar., June-July, Dec. Note — NGF 46042 from the Namatanai Sub- dist., New Ireland, may belong to this species. The above description is based on material from the Solomons; NGF 46042 differs in the following points: leaflets relatively broad, up to 20 by 9 cm, with the nerves widely spaced (up to 3.5 cm apart); old female flowers with relatively large sepals (8 by 5 mm). Its fruits are unknown, this makes it the more difficult to place this specimen with cer- tainty. EXCLUDED Otonychium retusum Mig., Fl. Ind. Bat. 1, 2 (1859) 572 = Rhus taitensis Guillemin, Ann. Sci. Nat. Bot. II, 7 (1837) 361 (Anacardiaceae). See Ding Hou in Fl. Males. I, 8 (1978) 537-538. JAGERA (P.W. Leenhouts) Jagera Blume, Rumphia 3 (1847) 155; Radlk. in Engl., Pflanzenr. 98 (1933) 1238-1243; Adema, Blumea 37 (1993) 195. — Type species: Jagera javanica (Blume) Blume ex Kalkman [= Jagera speciosa Blume]. Trees, often + pachycaulous, or shrubs, monoecious. Indumentum of solitary simple hairs; no glandular scales. Twigs often strongly ribbed, with a thick pith and a very thin Adema, Leenhouts, Van Welzen — Sapindaceae 615 wood cylinder; terminal bud protected by bud scales. Leaves often verticillate, some- times opposite or spirally arranged, paripinnate, 4—40-jugate, without pseudo-stipules; petiole base swollen and beneath deeply 3-lobed, leaving a very characteristic scar, nei- ther petiole nor rachis winged. Leaflets alternate to opposite, below with naked glands; base at the basiscopic side narrow, cuneate, at the acroscopic side broad and rounded; margin serrate to dentate (entire in an Australian variety of J. pseudorhus); midrib prom- inent above, nerves and veins raised on both sides, nerves ending in the marginal teeth. Inflorescences in the axils of bracts or normal leaves, together pseudoterminal, thyrses. Flowers unisexual, monoecious, regular. Sepals 5, slightly connate at base, narrowly imbricate, all equal, margin sometimes slightly petaloid, outside hairy, the margin (partly glandular) ciliolate, inside slightly hairy to glabrous, entire, in fruit persistent and slight- ly reflexed. Petals 5, as long as or slightly longer than the calyx, shortly clawed to sessile, glabrous or outside at the base and sometimes in the centre, and along the margin slightly hairy, entire, inside either auricled or with two, sometimes with one, erect woolly scale(s), exceptionally (an Australian subspecies of J. javanica) with a crest. Disc entire, lobed, without appendages, glabrous. Stamens 8 (7-10), in male flowers exserted, spreading; filaments at least slightly hairy at base (to completely hairy); anthers obovate, ciliate (or glabrous). Ovary sessile, hairy, 3-locular; style apical, about as long as the ovary, hairy, with 3 swollen stigmatic lines. Ovules 1 per locule. Fruits sessile, loculicidal, slightly warty, densely pilose with stiff, irritating hairs, inside also densely pilose. Seeds with a swollen annular sarcotesta around the hilum. — Fig. 48. Distribution — Two species in the Moluccas, New Guinea, and Australia (Queens- land and northern New South Wales). Habitat — Understorey tree of different forest types. KEY TO THE SPEGIES la. Twigs 10-35 mm thick, with 10-12 deep grooves. Leaves mostly in whorls of up to 6, sometimes in a dense spiral, 13—40-jugate .................. 1. J. javanica b. Twigs 3—5 mm thick, with 6 slight grooves. Leaves arranged in an irregular spiral, Somecmes opposites 4—1 7s gate: wea... ode tae sels Eke tees 2. J. pseudorhus 1. Jagera javanica (Blume) Blume ex Kalkman, Blumea 7 (1953) 470; S.T. Reynolds in Fl. Aus- tral. 25 (1985) 67. — Garuga javanica Blume, Bijdr. (1826) 1165. — Jagera speciosa Blume, Rumphia 3 (1847) 155, nom. illeg.; Miq., Fl. Ind. Bat. 1, 2 (1859) 564. — Type: ?Korthals S27. \(L2), [Papaja litorea boeronensis Attehu dicta Rumph., Herb. Amb. | (1741) 150, pl. 53, f. 2. Sapindus serrata Roxb., Hort. Beng. (1814) 88, nom. nudum, inval.; Fl. Ind. ed. Carey (1832) 284. — Jagera roxburghii Blume, Rumphia 3 (1847) 155, nom. illeg. — Jagera serrata (Roxb.) Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Akad. Wiss. Miinchen 8 (1878) 303; Sapind. Holl.-Ind. (1879) 36; in Engl., Pflan- zenr. 98 (1933) 1241; S.T. Reynolds, Austrobai- leya 1 (1983) 411, f. 28E. — Jagera serrata (Roxb.) Radlk. f. genuina Radlk., Bot. Jahrb. 56 (1920) 297, nom. illeg. — Type: Roxburgh s.n., Moluccas. Jagera serrata (Roxb.) Radlk. f. fulvinervis Radlk., Bot. Jahrb. 56 (1920) 297. — Type: Forbes 750 (FI, L), New Guinea. Jagera macrophylla Radlk., Bot. Jahrb. 56 (1920) 297, f. 3; in Engl., Pflanzenr. 98 (1933) 1243, f. 37. — Syntypes: Moszkowski 34] (M), Ledermann 10759, 10842, all New Gui- nea. The species is divided into two subspecies of which only one occurs in Malesia, the other one (subsp. australiana Leenh.) is restricted to N Aus- tralia. 616 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 48. Jagera Blume. Habit, leaflet and fruit. — J. javanica (Blume) Blume ex Kalkman subsp. javani- ca. a. Habit, leafy part; b. ibid., topmost part with inflorescences; c. fruit. — J. pseudorhus (A. Rich.) Radlk. d. Leaflet (a, b: Kostermans & Soegeng 70; c: Hoogland 4910; d: Brass 6437). subsp. javanica rather smooth or sometimes pustular, greyish. Twigs 1-3.5 cm thick, with 10 or 12 deep grooves and a Not or sometimes sparsely branched pachycau- conspicuous sequence of zones with scars of bud lous tree, up to c. 12, rarely to 25 mhigh, dbh up _ scales, zones with 5 or 6 whorled leaf scars sepa- to 20(—50) cm, sometimes with low buttresses; bark —_ rated by up to4cm long internodes, and zones with Adema, Leenhouts, Van Welzen — Sapindaceae scars of inflorescences axillary to scars of bracts; usually densely hairy. Leaves mostly whorled, 13— 40-jugate, 35-120 cm long; petiole 5—28 cm long, 3-9 mm thick, strongly 3-lobed below at the base: axes tomentellous; petiolules 1-3 mm long. Leaf- lets 2.5—20 by 1-6 cm, index 2.5—5, variably per- gamentaceous, sometimes papyraceous or coria- ceous, above on the midrib variably hairy, very sparsely so on the nerves, beneath hairy all over, mainly on midrib and nerves only, with 0 to many naked glands usually in 2 rows along the midrib; margin serrate to crenate-dentate; apex acute to acuminate. /nflorescences up to 1 m long, widely laxly branched, many-flowered, the flowers in near- ly sessile, compact, few-flowered dichasia or in compact monochasia, densely hairy: pedicels 1-3 mm long, pendent in fruit. Sepals 1—-1.8 by 0.8- 1.5 mm. Petals ovate to obovate, 1-2 by 0.6—1.5 mm, inside with 2 scales or sometimes with 2 auricles. Stamens: filaments up to 3 mm long, partly woolly, at least near the base; anthers 1.3—1.8 by 0.8—1 mm, ciliate. Fruits obovoid to globular, 2.5 by 2.cm, pericarp hard, c. 2 mm thick. — Fig. 48a-c. Distribution — Malesia: Moluccas and New Guinea. Habitat & Ecology — Mostly an understorey tree, sometimes a canopy tree of primary and second- ary rain forest, sometimes of swamp or savannah forest, often on slopes and ridges, sometimes on flat and river banks, sometimes periodically flood- ed, both on (limestone) rocks and on clay; mostly below 100 m altitude, sometimes up to 1050 m. Fl. mainly Feb.—July; fr. May—Dec. Notes — 1. Unfortunately, the type material is labelled ‘Java’, hence Blume’s specific epithet ‘javanica’. Later on he realized that this material came from the Bogor Botanic Garden and had been introduced from the Moluccas (see Blume 1847). 2. Several characters show rather wide varia- tion. Forma fi/vinervis shows the extremes of some mostly independently varying characters. The den- sity of the indumentum is especially variable; the most glabrous forms occur in the Moluccas, but not all Moluccan collections are glabrous. 2. Jagera pseudorhus (A. Rich.) Radlk., Sapind. Holl.-Ind. (1879) 37; in Engl., Pflanzenr. 98 (1933) 1240; Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 522; Francis, Austral. Rain- 617 for. Trees ed. 2 (1951) 256, f. 149-151; S.T. Reynolds, Austrobaileya 1 (1983) 409, f. 25A, 28B-D; Stanley & Ross, Fl. SE Queensl. 1 (1983) 509, f. 79K; S.T. Reynolds in Fl. Aus- tral. 25 (1985) 67, f. 13E, F. — Cupania pseu- dorhus A. Rich. in Urv., Voy. Astrolabe 2, Bot. (1834) 34, pl. 14. — Type: Fraser s.n., Aus- tralia. Jagera pseudorhus f. pilosiuscula Radlk., Sitzungs- ber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 621. — Lectotype (S.T. Rey- nolds 1983): Wawra 628, Queensland. The species is divided into two varieties: var. pseudorhus, with two forms: f. pseudorhus (S Queensland, northern New South Wales) and f. pilosiuscula Radlk., and var. integerrima S.T. Rey- nolds (Atherton Tableland). Only f. pilosiuscula, described below, occurs in Malesia. Trees, up to 8, rarely up to 18 m high, dbh up to 12 cm; bark smooth or slightly pustular, patchy grey and brown. Twigs 3—5 mm thick, with 6 slight grooves, gradually becoming terete, sparsely hairy. Leaves arranged in an irregular spiral, upper ones sometimes opposite, 4—17-jugate, 6.5—32 cm long: petiole 2—7 cm long, 1—2 mm thick, below at base with 2 slight grooves; axes variably hairy; petiolules 1-2 mm long. Leaflets 4-11 by 1.25—3.5 cm, in- dex 2-5, thin- to stiff-pergamentaceous, above on the midrib slightly hairy, beneath glabrous but for a few hairs on the midrib, with 0 to many naked glands in | row along the midrib; margin distantly serrate; apex tapering acuminate, acute. /nflores- cences 4.5—20 cm long, sparsely hairy: branches up to 5 cm long, bearing several-flowered dicha- sia; pedicels 1.5—2 mm long. Sepals 1—1.8 by 0.6— 1 mm. Petals ovate to orbicular, 1.4—2 by 1-1.5 mm, inside with one or two scale(s). Stamens: fil- aments 2.5—3 mm long, sparsely to densely long- hairy; anthers 1.5—2 by 0.8—0.9 mm, glabrous or slightly ciliate. Fruits obovoid, 1.75 by 1.25 cm; pericarp hard, c. 1 mm thick. — Fig. 48d. Distribution — Queensland (south to Fraser Is- land); in Malesia: New Guinea (Irian Jaya: around Merauke; Papua New Guinea: Western Prov.). Habitat & Ecology — In rain, monsoon, and sa- vannah forest, on banks of rivers and lakes: from sea level to 30 m altitude. Fl. mainly Aug., Sept.: fr. Aug. 618 Flora Malesiana ser. I, Vol. 11 (3) (1994) LEPIDEREMA (A.M. Schot) Lepiderema Radlk., Sapind. Holl.-Ind. (1879) 99; in Engl., Pflanzenr. 98 (1933) 1216; S.T. Reynolds, Austrobaileya 1 (1982) 488; in Fl. Austral. 25 (1985) 52; Schot, Blu- mea 36 (1991) 235. — Type species: Lepiderema papuana Radlk. (Small) trees. Indumentum of solitary hairs; (some) scale hairs present in inflores- cences. Branchlets terete, more or less straight, pith narrow. Leaves paripinnate, pseu- dostipules absent; rachis above grooved, not winged. Leaflets opposite to subopposite, elliptic to narrowly elliptic, symmetrical except for the base, not punctate; base attenuate (to cuneate), oblique; margin entire, slightly curved; apex cuspidate, very apex obtuse, not mucronulate; both surfaces smooth, domatia absent below; venation raised to almost flat above, raised below; veins densely reticulate. Inflorescences pseudoterminal or axil- lary or ramiflorous thyrses, branches with scale-like hairs; cymules 1—7-flowered, dicha- sial. Bracts and bracteoles caducous. Pedicels sparsely hairy, glabrescent. Sepals 5, pet- aloid, connate at base, biseriate. Petals 5; claw very small to absent; scales absent. Disc uninterrupted. Stamens 8; filaments filiform; anthers basifixed, dehiscence latrorse. Ova- ry 3-locular, one ovule per locule; style pyramidal with three longitudinal stigmatic grooves. Fruits capsular, obovoid, loculicidal, not winged, stipe slender, outside and inside gla- brous. Seeds obovoid, covered by an arillode; the latter open at apical end, without basal pseudofunicle. — Fig. 49. Distribution — 8 species: 6 in Australia, 2 in Malesia: New Guinea. Note — Only two specimens of Lepiderema are known from New Guinea, both being the types of the two Malesian species. KEY FO THE SPECIES la. Rachis of leaf shortly hirsute; midrib of leaflets sericeous on both sides; marginal IGOMSMNGISHNEt oucert. soe kts ot aa... sot ee solar er 1. L. melanorrhachis b. Rachis of leaf glabrous; leaflets, including midrib, glabrous; marginal loops dis- (TS os co 5 ata alt tite eke ce PAPER PRERE TPL ek Wt tee SSS 2. L. papuana 1. Lepiderema melanorrhachis Merr. & L.M. like hairs, black; part above abscission zone c. 1 Perry, J. Arnold Arbor. 21 (1940) 521; P. Roy- — mm long. Flowers unknown. Sepals elliptic to sub- en, Man. For. Trees Papua & New Guinea 2 orbicular, glabrous, outer two c. 1 by 0.75 mm, (1964) f. 1k; Schot, Blumea 36 (1991) 237. — inner three c. 1.5 by | mm. Fruits obovoid, 10-12 Type: Brass 7432 (A holo; L), New Guinea. by 9-10 mm, not lobed, rough; stipe 3-4 mm long; wall thin. Seeds obovoid, c. 4.5 by 3 mm, only Tree, tall, slender. Flowering branchlets 5-9 mm young ones seen. — Fig. 49. thick, slightly rough, very sparsely hirsute, black. Distribution — Malesia: Papua New Guinea (Fly Leaves 3—5S-jugate; peduncle 2.2—7.8 cm long; ra- River). chis 4.5—-14.2 cm long, shortly hirsute. Leaflets el- Habitat & Ecology — In forest canopy. Fr. Aug. liptic to narrowly elliptic, 4.7—14.3 by 2.4—5.7 cm; both surfaces glabrous except for the sericeous 2. Lepiderema papuana Radlk., Sapind. Holl.- midrib; venation almost flat above; nerves curv- Ind. (1879) 100; in Engl., Pflanzenr. 98 (1933) ing towards margin, indistinctly joined. /nflores- 1217; P. Royen, Man. For. Trees Papua & New cences axillary to ramiflorous; rachis 4—9.5 cm Guinea 2 (1964) 28; Schot, Blumea 36 (1991) long, hirsute; cymules (1—)3—7-flowered; pedicel 238. — Type: Teijsmann HB 14250 (M holo), c. 2 mm long, glabrous to sparsely hairy with scale- Irian Jaya. Adema, Leenhouts, Van Welzen — Sapindaceae 619 Fig. 49. Lepiderema melanorrhachis Merr. & L.M. Perry. a. Habit; b. fruit (a, b: Brass 7432). 620 Flora Malesiana ser. I, Vol. 11 (3) (1994) (Presumably) small tree. Flowering branchlets c. 6 mm thick, glabrous. Leaves 3—S-jugate; pe- duncle 5.4—13.7 cm long; rachis 5.3—11.8 cm long, glabrous. Leaflets elliptic to narrowly elliptic, 4.6— 16.5 by 2.2—5.8 cm; both surfaces glabrous; vena- tion raised above; nerves looped and joined near the margin. Inflorescences pseudoterminal; rachis 3.8—6.9 cm long, shortly hirsute, branching at base; cymules 1|-flowered; pedicel c. 1.75 mm long, short- ly, sparsely hairy with simple and with scale hairs; part above abscission zone c. 1 mm long. Flowers 2.5-3 by c. 1.5 mm, probably only female ones seen. Sepals elliptic to suborbicular, glabrous, outer two c. | by 0.75 mm, inner three c. 2 by 1.25 mm. Petals 5, elliptic, c. 1.75—2 by 0.5—1.25 mm, gla- brous to sparsely hairy; claw less than 0.1 mm long, hairy at base; apex rounded. Stamens 8, filaments 0.5—0.75 mm long, densely hairy at least at base; anthers c. | mm long, mainly hairy on connective. Ovary c. 1.5—2 mm long, glabrous; style and stig- mac. 2 mm long. Fruits and seeds unknown. Distribution — Malesia: Irian Jaya (Misool). LEPIDOPETALUM (P.C. van Welzen) Lepidopetalum Blume, Rumphia 3 (1847) 171; Radlk., Bot. Jahrb. 56 (1920) 56; in Engl., Pflanzenr. 98 (1933) 1316; S.T. Reynolds in Fl. Austral. 25 (1985) 87; Welzen et al., Blumea 36 (1992) 439. — Type species: Lepidopetalum perrottetii (Cambess.) Blume. Lachnopetalum Turcz., Bull. Soc. Nat. Mosc. 21 (1848) 571. — Type species: Lachno- petalum glabrum Turcz. [= Lepidopetalum perrottetii (Cambess.) Blume]. Trees; bark greyish brown, slightly fissured and/or with lenticels; indumentum of simple hairs only. Branchlets smooth to slightly grooved, sericeous when young. Leaves paripin- nate, pulvinate at base, no pseudo-stipules or wings; petiolules present as pulvinus. Leaf- lets subopposite to opposite, basal ones ovate, upper ones elliptic (to obovate), slightly asymmetric, acroscopic side broader, thin, not punctate; base usually cuneate, asymmet- ric; margin entire, flat; both surfaces smooth, subglabrous except for at least the subseri- ceous basal part of the midrib, domatia as (a few) glabrescent hair tufts below; venation raised, especially below, nerves marginally looped, veins reticulate. Inflorescences ram- iflorous to axillary to pseudoterminal, reduced thyrses with usually 3 cymules per node; rachis (sub)sericeous; cymules basally dichasial to apically cincinnate, those with male flowers often with an exceptional number of sepals, petals, and stamens in first devel- oped flowers. Bracts and bracteoles: mainly margin and outside sericeous. Pedicels with glandular hairs besides simple ones, articulated. Flowers actinomorphic, yellowish white. Sepals (sub)equal, basally united; tube disc-shaped, c. 0.3 mm long; blades triangular (to ovate). Petals: claw c. 0.1 mm high; blade triangular; scales united into one, larger than blade, margin irregularly crenate, pilose; crests usually absent (to present). Disc circular, smooth, glabrous, enveloping base of stamens, without appendages. Stamens (7—)8(—10); filaments especially basally pilose; anthers basifixed, dehiscence latrorse, pilose. Pistil: ovary flat, 2- (or 3-)locular; ovules one per locule; style and stigma united, flat, triangu- lar, stigmatic part folded outwards like overhanging roof. Fruit an obovoid, loculicidally dehiscing capsule, with usually 1 developed seed, flat when 2-locular, smooth to some- what rough, outside glabrous, margin sharp, almost slightly winged, inside glabrous to pilose, red; stipitate or not; wall thin, coriaceous to woody, at most 2 mm thick. Seeds ellipsoid, triangular in transverse view, base straight to oblique, smooth, shiny, black; sarcotesta glabrous, covering seed only basally to almost completely, orange, testa thin, without radicle pocket on the inside. — Figs. 50, 51. Adema, Leenhouts, Van Welzen — Sapindaceae 621 Distribution — 6 species from India (Andaman and Nicobar Islands; | non-endemic species) to NE Australia and the Solomon Islands (also 1 non-endemic species each); all species occurring in Malesia, mainly endemic. Habitat & Ecology — Trees in primary and secondary forest, on waste land, along rivers and roads; sea level up to 1200 m altitude. Note — The phylogeny and historical biogeography of Lepidopetalum are described in Van Welzen et al. (1992). KEY TO THE SPECIES (based on flowering characters) Caution: the differences in floral characters among the species are only slight. Therefore geography is used as an additional ‘character’; however, due to insufficient collecting, the geographical ranges of some species may be larger than described in the key. Pet ALTO PCCUCCES POSE tees ee he kee we ee me ee. See 2 b. Upper part of pedicels glabrous except for glandular hairs .................. i) 2a. Disc as high as broad or broader than high (flat). Philippines, New Guinea, Solo- MES ei ire. Hoes 1a ie nxceetent . . A. ng SSR. Be ee ene 3 b. Disc higher than broad. Nicobar Islands, Sumatra............ 3. L. montanum 3a. Ovary (sub)glabrous (to pilose); if pilose then disc usually with slits. Petals 0.5—2.2 mm high, crests often present as ribs or as flat scales. New Guinea, Solomons . 4 b. Ovary pilose; disc without slits. Petals 0.4—1 mm high, crests absent. Philippines 4. L. perrottetii 4a. Disc as high as broad, without slits. Crests usually present on petal scales as ribs or as flat appendages. Ovary (sub)glabrous. N coast of New Guinea: E Geelvink Bay teplaGano Province acs kriee ss 2c eh. 2 es Oe bY ee 2. L. micans b. Disc flat to as high as broad, usually with slits. Crests usually absent, at most present as ribs. Ovary subglabrous to pilose. New Guinea: Peninsula to S coast to Vogelkop 6. L. xylocarpum 5a. Stamens 8, some flowers of a cymule, especially the first flowering male ones, with 9 or more. Solomons and New Guinea (absent in Morobe Prov.) ............. 6 b. Stamens 8. New Guinea: Morobe Province ............. 1. L. fructoglabrum 6a. Inflorescences up to 17 cm long, axillary to pseudoterminal. Each inflorescence with at least a few big bracts 1—3.3 mm long. Solomons and N New Guinea: Jayapura to Morobe Province to Bougainville .................. 5. L. subdichotomum b. Inflorescences up to 8.5 cm long, usually ramiflorous on thin twigs to axillary to pseudoterminal. Inflorescences with only small bracts 0.7—1.3 mm long. New Guinea: Peninsula to:S coast taVogelkopues... . . /ovun! AN iain Fh: 6. L. xylocarpum KEY TO THE SPECIES (based on fruit characters) Ban Fie PU OGENIOSEMS tp pee Seif gc 5 <5 LOUK Ai ete aga «4 Gh a Naan cig ie 0 ea 2 b.’ Prart'plabrows inside... S20. Pi. «s . «cos Sa Ee 1. L. fructoglabrum 622 Flora Malesiana ser. I, Vol. 11 (3) (1994) 2a. Sarcotesta present only basally around hilum...........-...---+0 +++ seeees 3 b. Sarcotesta covering seed except for (part of) dorsal side............--...--. 5 3a. Upper part of pedicels sericeous. Sarcotesta with well developed obpyramidal basal OUtenOwthy-4K). AAO Le ne EEN PEPSI Hehe RE. t ie «2 EN A 4 b. Upper part of pedicels glabrous except for some glandular hairs. Sarcotesta with inconspicuous obpyramidal basal outgrowth ........... 5. L. subdichotomum Aa shri 3. 2=4.5, by, li —2-4 Cisne ceo + or hetenetns ee > ener Bee 3. L. montanum bag kit 3—225 DysOsS il Oicimienhs eth. «eta aes te eels ck 4. L. perrottetii 5a. Fruit long pilose inside, hairs c. 1.5 mm long. Sarcotesta covering seed except for small dorsal triangular part (Fig. 50D)............-.---.++++-:: 2. L. micans b. Fruit short pilose inside, hairs c. 0.5 mm long. Sarcotesta covering seed except for donsalsside (Hig. S02) ican cpt -cyeropory- 1. Lepidopetalum fructoglabrum Welzen in Welzen et al., Blumea 36 (1992) 455, f. 7. — Type: Hartley 9948 (L holo; A, BRI, CANB, K), Papua New Guinea. Lepidopetalum subdichotomum auct. non Radlk.: S.T. Reynolds in Fl. Austral. 25 (1985) 87, f. 16, 21k, map 111. Tree, 10-17 m high, dbh 7.5—50 cm; outer bark blotched light and dark grey to dark brown, smooth 6. L. xylocarpum to finely tessellated to rather warty, inner bark pink- ish to reddish brown, hard; wood cream to dark straw, moderately hard and heavy, heartwood red- dish brown. Flowering branchlets 1|—4 mm in diam. Leaves 3- or 4-jugate; rachis 2.7-14.5 cm long, terete, subsericeous. Leaflets 3.2-17.5 by 1.4-6.1 cm; apex (acuminate to) caudate, very apex emar- ginate to rounded; upper surface glaucous when dry. Inflorescences mainly axillary to pseudoter- minal, axes branching in axil and along rachis, all — icm tem icm Fig. 50. Lepidopetalum Blume. Fruits and seeds. — L. micans Laut. & K. Schum. a. Fruit; b. seed. — L. perrottettii (Cambess.) Blume. c. Fruit; d. seed. — L. subdichotomum Radlk. e. Fruit. — L. monta- num (Blume) Radlk. f. Fruit. — L. xylocarpum Radlk. g. Fruit (a, b: LAE 50083; c, d: FB 20524; e: BSIP 10242; f: King’s coll. 7; g: NGF 27507). Adema, Leenhouts, Van Welzen — Sapindaceae up to 7.5 cm long. Bracts and bracteoles triangu- lar; bracts 0.6—1 mm long; bracteoles 0.3—0.8 mm long. Pedicels subglabrous, upper part up to 5.7 mm long, glabrous except for some glandular hairs. Flowers 3.54.3 mm in diam. Sepals 5; lobes 1.3— 1.8 by 0.7—1.1 mm, subglabrous. Petals 5; blade 0.8—1 by 0.7—1.3 mm, subpilose; scale 0.7—1.3 by 1—].6 mm; crest absent (to weli developed central ribs). Disc uninterrupted, c. 0.6 by 0.5 mm, broad- er than high. Stamens 8; filaments white, in male flowers 2.7—3.3 mm long, in female flowers 0.5— 1.5 mm long; anthers 0.8—1.1 by 0.5—0.8 mm, yel- lowish. Pistil: ovary in male flowers c. 0.3 mm high, up to 2.6 mm high in female flowers, 2-locular, subglabrous; style and stigma in male flowers c. 0.1 mm high, in female flowers up to 1.3 mm high. Fruits 2—2.2 by 1.2—1.4 cm, inside glabrous; stipe absent. Seeds 1.1—1.3 by c. 0.9 cm, base straight; sarcotesta only around hilum, forming slight ob- pyramidal outgrowth below seed; hilum 2-4 mm long. Distribution — Malesia: NE Papua New Guinea (Morobe Prov.). Habitat & Ecology — Found in second storey of primary and secondary rain forest, periodically flooded forest, and along streams and roads; 30— 900 m altitude. Fl. Aug.—Nov.; fr. Nov.—Feb. Chemical compounds — Hartley et al. (Lloydia 36, 1973, 270) report L. fructoglabrum (as L. hebe- cladum) to be devoid of alkaloids in bark and leaves. 2. Lepidopetalum micans Laut. & K. Schum. in K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1901) 423; Welzen et al., Blumea 36 (1992) 456, f. 2a, 3c, 7. — Type: Lauterbach 2840 (B+ holo; BM, K, M), Papua New Guinea. Lepidopetalum hebecladum auct. non Radlk.; Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 269; K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1901) 422; Radlk., Bot. Jahrb. 56 (1920) 307; Nova Guinea 14 (1926) 184; in Engl., Pflan- zenr. 98 (1933) 1319. In all references Hollrung 341] excluded; see note 1. (Shrub to) tree, 2-30 m high, 10-85 cm dbh; outer bark whitish to greyish brown, smooth to somewhat rough with numerous pustular lenticels, thin, hard, inner bark cream to orange to straw brown; wood white to brown, heartwood reddish brown. Flowering branchlets 1.5—2.5(—11) mm in diam. Leaves (2—)3—4(-5S)-jugate; rachis 1.7—-14cm long, flat above, especially upper side sericeous. Leaflets 2-21 by 1-7 cm; apex acuminate to cus- pidate, very apex emarginate to rounded. /nflores- cences axillary to pseudoterminal, usually not 623 branching, rachis up to 8 cm long. Bracts and bracteoles triangular; bracts 1.1—-1.6 mm long; bracteoles 0.3—1 mm long. Pedicels puberulous, upper part 1.8 (flower)—10 (fruit) mm long. Flow- ers 3.3-6.3 mm in diam. Sepals 5; lobes 1.2—3.3 by 0.7—2.3 mm, margin pilose. Petals 5; blade 0.5— 2 by 0.8—2 mm, margin pilose, outside subglabrous; scale sometimes divided into two, 0.5—1.5 by 0.8— 2.2 mm; crest (absent to) well developed central ribs to 2 small flat appendages. Disc uninterrupt- ed, c. 0.5 mm high, as high as broad. Stamens (7—)8(—10); filaments in male flowers 1—3.3 mm long, in female flowers 0.8—1.3 mm long; anthers 0.8-1.2 by 0.6-1 mm. Pistil: ovary in male flowers 0.3—-0.4 mm high, up to 2.5 mm high in female flowers, 2- (or occasionally 3-)locular, sub- glabrous; style and stigma in male flowers c. 0.1 mm high, in female flowers up to 1.2 mm high. Fruits 1.6—2.2 by 1—1.6 cm, inside densely hirsute with hairs up to 1.5 mm long; stipe absent to 2.5 mm high. Seeds 1.2-1.7 by 0.7-1.2 cm, base straight; sarcotesta covering seed almost completely except for small dorsal triangular spot; hilum 1.2 4 mm long. — Fig. 50a, b. Distribution — Malesia: Along north coast of New Guinea from E Geelvink Bay (Irian Jaya) to Madang Prov. (Papua New Guinea). Habitat & Ecology — Found in under- to middle storey of primary forest, secondary forest, swamp forest, and along rivers and roads. Soil: limestone, sandy clay. From sea level up to 425 m altitude. Fl. June—Nov.; fr. July—April. Notes — 1. The type specimen of L. hebecla- dum (Hollrung 341) has always been interpreted incorrectly. This specimen is conspecific with L. subdichotomum. All other specimens, always iden- tified as L. hebecladum, are L. micans. 2. Lepidopetalum xylocarpum strongly tesem- bles L. micans in the western part of its distribu- tion area. See note 3 under L. micans. 3. Lepidopetalum montanum (Blume) Radlk., Sapind. Holl.-Ind. (1879) 14; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 510, 535, 623; in Engl., Pflanzenr. 98 (1933) 1320; Welzen et al., Blu- mea 36 (1992) 457, f. 2b, 7. — Arytera mon- tana Blume, Rumphia 3 (June 1847) 171. — Ratonia montana Fern.-Vill., Nov. App. (1880) 51, nom. illeg. — Type: Korthals s.n. (L holo), Sumatra (probably Junghuhn 4/ in L and de Vriese 18 in L are the same specimen). {[Connarus ? jackianus Wall., Cat. (after 22 Oct. 1847) 8552, nom. nud.] — Cupania jackiana Hiern in Hook. f., Fl. Br. India 1 (1875) 678. — Lepidopetalum jackianum Radlk., Sapind. Holl.- 624 Flora Malesiana ser. I, Vol. 11 (3) (1994) Ind. (1879) 45; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 535, 623; in Engl., Pflanzenr. 98 (1933) 1318. — Type: Jack s.n. (K holo; M), Nicobar Is. Tree, 5-16 m high. Flowering branchlets 2-4 mm in diam. Leaves 2—4-jugate; rachis 1.9-14.7 cm long, terete, subsericeous. Leaflets 4.5—21 by 1.5-7.5 cm; apex acuminate, very apex rounded (Nicobar Islands) to almost acute (Sumatra). //- florescences axillary to pseudoterminal, axes branching in axil or along rachis, all up to 11 cm long. Bracts and bracteoles triangular to long-tri- angular; bracts 0.8—-2 mm long; bracteoles 0.3-1.6 mm long. Pedicels sericeous, upper part 1.2—6.3 mm long. Flowers 2.8-4.2 mm in diam. Sepals 5; lobes 1-2.8 by 0.8—-1.4 mm, mainly outside seri- ceous. Petals 5; blade triangular, 0.6—1.2 by 0.5— 1.1 mm, completely pilose; scale 0.8—1 by 0.8-1.5 mm; crests absent. Disc uninterrupted, 0.5—0.7 mm high, margin often revolute, higher than broad. Sta- mens 8; filaments in male flowers 1.2—3 mm long, yellow; anthers in male flowers 0.7—-1.2 by 0.6— 0.8 mm, pink to reddish purple. Pistil: ovary in male flowers 0.2—0.5 mm high, 2-locular, subgla- brous; style and stigma in male flowers 0.05—0.1 mm high. Fruits 3.2-4.5 by 1.7—2.4 cm, inside densely hirsute with hairs 1-1.5 mm long; stipe 3— 5 mm high. Seeds 1.9-2.2 by 1.3-1.7 cm, base oblique; sarcotesta covering seed at base, present around hilum, forming obpyramidal outgrowth below seed; hilum 7—8 mm long. — Fig. 50f. Distribution — S Andaman and Nicobar Islands; Malesia: N Sumatra. Habitat & Ecology — In primary forest, along rivers and roads; sea level up to 1000 m altitude. Rare to locally common. FI. and fr. seemingly the whole year round. The seeds are eaten by Calenas nicobarica and other frugivorous pigeons (Prain, Proc. As. Soc. Beng. 1891, 167). 4. Lepidopetalum perrottetii (Cambess.) Blume, Rumphia 3 (1847) 172; Miq., FI. Ind. Bat. 1, 2 (1859) 569; Radlk., Sapind. Holl.-Ind. (1879) 15, 46, 92; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 535, 622; Radlk. in Engl., Pflanzenr. 98 (1933) 1317; Welzen et al., Blumea 36 (1992) 458, f. 2c, 3a, 7. — Cupania perrottetii Cambess., Mém. Mus. Hist. Nat. Paris 18 (1829) 45. — Type: Perrot- tet s.n. (P holo; L, P), Philippines. Lachnopetalum glabrum Turcz., Bull. Soc. Nat. Moscou 21 (1848) 572; Miq., Fl. Ind. Bat. 1, 2 (1859) 557. — Ratonia lachnopetala Turcz., Bull. Soc. Nat. Moscou 36 (1863, n.v.) 586, nom. illeg.. — Type: Cuming 1169 (holo un- known; iso L, M, MEL, NY, P), Philippines. Cupania ? richii A. Gray, U.S. Expl. Exped. Bot. 1 (1854) 257; Merr., Philipp. J. Sc., Bot. 3 (1908) 79. — Type: U.S. Expl. Exped. s.n. (A holo), Philippines. Shrub to tree, 2-17 m high, dbh 6—25 cm. Flow- ering branchlets 3-5 mm in diam. Leaves 2—4- jugate; rachis 1.6-17.3 cm long, terete, (sub)seri- ceous. Leaflets 2.5-24 by 1.2-10 cm; apex acumi- nate, very apex (emarginate to) rounded. /nflores- cences axillary with a tendency towards ramiflory, branching mainly in axil or along rachis, latter up to 7 cm long. Bracts and bracteoles triangular to long-triangular; bracts 1.1—2 mm long; bracteoles (.6-1 mm long. Pedicels sericeous, upper part 2-8 mm long. Flowers c. 4 mm in diam. Sepals 5 (or 6); lobes 0.8-2.3 by 0.6-1.5 mm, mainly outside sericeous. Petals 5; blade triangular, 0.4—1 by 1— 1.6 mm, completely pilose; scale 0.8—-1.7 by 1.2— 1.8 mm; crests absent. Disc uninterrupted, c. 0.4 mm high, broader than high. Stamens 8; filaments in male flowers 1.2—3.3 mm long, in female flow- ers 0.4-1.8 mm; anthers in male flowers 0.6—0.9 by 0.5—0.8 mm, in female flowers 0.4—0.7 by 0.5- 0.8 mm. Pistil: ovary in male flowers 0.3—0.4 mm high, in female flowers 1.82.5 mm high, 2-locu- lar, hirsute; style and stigma in male flowers 0.05— (0.1 mm long, in female flowers 0.9—1.8 mm long. Fruits 1.3-2.5 by 0.8—1.6 cm, inside densely hir- sute with hairs c. 0.5 mm long; stipe absent (to 2 mm high). Seeds 6-12 by 5—9.5 mm, base oblique; sarcotesta only around hilum, forming obpyrami- dal outgrowth below seed; hilum 0.8—1.1 mm long. — Fig. 50c, d. Distribution — Malesia: Philippines (mainly E and N). Habitat & Ecology — Found along roadsides, in secondary forest, on waste land, and along forest edges; sea level up to 1200 m altitude. Probably two flowering seasons: Jan.—Mar.(—Apr.) and May— June; fr. (Feb—)March—June and Sept—Nov.(—Feb.). Note — Clinal variation exists from Mindanao to Luzon, the leaflets and fruits decrease in size. 5. Lepidopetalum subdichotomum Radlk. in K. Schum. & Hollr., Fl. Kais. Wilh. Land (1889) 67; Sitzungsber. Math.-Phys. Cl. K6nigl. Bay- er. Akad. Wiss. Miinchen 20 (1890) 269; Bot. Jahrb. 56 (1920) 307; in Engl., Pflanzenr. 98 (1933) 1319; S.T. Reynolds in Fl. Austral. 25 (1985) 87, f. 16, 21k, map 111; Welzen et al., Blumea 36 (1992) 460, f. 1, 2d, 7. — Type: Hollrung 387 (B+ holo; K, L, M, MEL, P), Pa- pua New Guinea, Kaiser Wilhelmsland, Hatz- feldhafen. Adema, Leenhouts, Van Welzen — Sapindaceae 625 Fig. 51. Lepidopetalum subdichotomum Radlk. a. Habit; b. hairtuft domatia; c. male flower; d. petal with scale from outside; e. female flower after pollination (a—d: Schodde & Craven 4092; e: BSIP 14557). 626 Flora Malesiana ser. I, Vol. 11 (3) (1994) Lepidopetalum hebecladum Radlk. in K. Schum. & Holir., Fl. Kais. Wilh. Land (1889) 67; Sit- zungsber. Math.-Phys. Cl. Konig. Bayer. Akad. Wiss. Miinchen 20 (1890) 269; Bot. Jahrb. 56 (1920) 307; in Engl., Pflanzenr. 98 (1933) 1319. — All references only include the type: Holl- rung 341 (Bt holo; K, P), Papua New Guinea, Kaiser Wilhelmsland, Hatzfeldhafen. See note I. Tree, 3-20 m high, girth 15 cm to 1 m; outer bark light to dark brown to grey, smooth to slight- ly fissured to scaly, soft, inner bark pinkish to mid- dle brown; wood white to reddish browa, (soft to) hard. Flowering branchlets 2-4 mm in diam. Leaves 3- or 4-jugate; rachis 4—15.2 cm long, terete, subsericeous. Leaflets 3.6-19 by 1.3—8.5 cm; apex acuminate, very apex rounded. Inflorescences ax- illary to pseudoterminal, mainly branching along rachis, sometimes also in axil, rachis up to 17 cm long. Bracts and bracteoles triangular to at least a few obovate to leaf-like; bracts 1—-3.3 mm long, bracteoles 0.4—0.7 mm long. Pedicels subglabrous, upper part 1.3 (flower)—12 (fruit) mm long, gla- brous except for some glandular hairs. Flowers 3— 5.5 mm in diam., fragrant. Sepals 5; lobes 0.8—2.3 by 0.8-1.6 mm, only margin subpilose. Petals 5; blade 0.7—1.6 by 0.8—-1.6 mm, mainly margin sub- pilose; scale 0.5—1.3 by 1.1—2.5 mm, crest absent to well developed central ribs to 2 small append- ages. Disc uninterrupted, 0.5—0.8 mm high, as high as broad. Stamens 8 ox at least in some flowers 9; filaments in male flowers 2.2—2.7 mm long, in fe- male flowers 0.8—1.7 mm long; anthers 0.7—1.2 by 0.6-1 mm. Pistil: ovary in male flowers 0.4—0.5 mm high, up to 2.5 mm high in female flowers, 2- (or 3-)locular, subglabrous; style and stigma in male flowers c. 0.1 mm high, in female flowers up to 1.3 mm high. Fruits 1.6-3.1 by 1-2.5 cm, inside densely hirsute with hairs 0.5—0.8 mm short; stipe 1-6 mm high. Seeds 1.3-1.7 by 0.9-1.1 cm, base straight; sarcotesta only around hilum, forming slight obpyramidal outgrowth below seed; hilum c. 1.5 mm long. — Figs. 50e, 51. Distribution — Solomon Islands to Malesia: New Guinea (Irian Jaya: Jayapura; Papua New Guinea: W Sepik to Morobe Provinces, New Britain, New Ireland, Bougainville). Habitat & Ecology — Found mainly in second- ary forest, also in primary forest, littoral forest, and grassland. At least found on sandy clay. Occasion- ally common. From sea level up to 330 m altitude. Fl. and fr. throughout the year. Uses — Wood is used in house construction (Bou- gainville). Notes — 1. Lepidopetalum hebecladum has al- ways been wrongly interpreted. The type specimen is conspecific with L. subdichotomum, while all other specimens have to be included in L. micans. 2. A geographical cline exists, the eastern spec- imens (Solomon Islands) have larger fruits than the western ones (New Guinea). 6. Lepidopetalum xylocarpum Radlk., Sitzungs- ber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 269; Bot. Jahrb. 56 (1920) 307; in Engl., Pflanzenr. 98 (1933) 1319; Welzen et al., Blumea 36 (1992) 461, f. 3b, 7. — Lec- totype (Van Welzen 1992): Forbes 379 (M holo; BM, MEL), Papua New Guinea, base of Owen Stanley's Ranges, Sogeri region. Tree, 4-22 m high; dbh 5—25 cm; (complex many-footed buttress system); outer bark blotched pale green and greyish to grey-brown to dark brown with many round pustular lenticels, inner bark straw to pinkish brown, shortly fibred; wood straw, turn- ing pale redbrown to centre, moderately hard and heavy, splits very readily, tendency to rings. Flow- ering branchlets 1-3 mm in diam. Leaves 2—4-ju- gate; rachis 1.7-16 cm long, flat above, especially upper side sericeous. Leaflets 3—22.5 by 1.5-7.5 cm; apex (slightly) acuminate (to cuspidate), very apex emarginate to rounded (to mucronulate); above often glaucous when dry. /nflorescences usu- ally ramiflorous on thin twigs to axillary to pseu- doterminal, at most branching along rachis, latter up to 8.5 cm long. Bracts and bracteoles triangu- lar; bracts 0.7—1.3 mm long, broad; bracteoles 0.4— 0.5 mm long. Pedicels (sub)glabrous except for glandular hairs to puberulous, upper part 2.3 (flow- er)—9 (fruit) mm long. Flowers 3.5—5 mm in diam. Sepals 5 (see note 1); lobes 1.2—2.9 by 1-2.2 mm, margin and outside pilose. Petals 5 (see note 1); blade 0.5—2.2 by 0.7-2 mm, margin pilose, out- side subglabrous; scale (divided into two) 0.4-1.3 by 1-2 mm; crest absent (to central ribs). Disc of- ten with slits, usually 5-lobed, flat, 0.3—-0.5 mm high, usually broader than high to as broad as high, pale green to yellow. Stamens 8(—10; see note 1); filaments white, in male flowers 1.8—3.3 mm long, in female flowers 0.7—1.6 mm long; anthers 0.7— 1.3 by 0.4-1.2 mm, brown. Pistil: ovary in male flowers c. 0.4 mm high, up to 2.5 mm high in fe- male flowers, 2- (or 3-)locular, subglabrous to pi- lose; style and stigma in male flowers up to 0.1 mm high, in female flowers up to 1.5 mm high. Fruits 1.7-3.6 by 1.2—2 cm, inside densely hirsute with hairs c. 0.5 mm short; stipe absent to 7 mm high. Seeds 1-1.9 by 0.71.1 cm, base straight; sarcotesta covering seed for greater part except for dorsal side; hilum 3—5.5 mm long. — Fig. 50g. Adema, Leenhouts, Van Welzen — Sapindaceae Distribution — NE Australia to Malesia: New Guinea (Irian Jaya: Vogelkop, Fakfak, Mimika, Digul; Papua New Guinea: Western, Gulf, Central, Milne Bay, and Northern Provinces). See also notes 2 and 3. Habitat & Ecology — Growing in understorey of primary , secondary , lower montane, and gal- lery forest, and along rivers. Soil: alluvial, lava. limestone, loam. From sea level up to 700 m alti- tude. Fl. Sept.—Apr.; fr. Jan.—Aug.(—Nov.). Uses — In the Vogelkop the wood is used to start fires (BW (Moll) 12782). Notes — 1. Flowers, usually male, at the end of an inflorescence often show aberrant numbers of sepals (6), petals (6 or 7), and anthers (9 or 10). 627 2. There is geographical variation, with the east- ern specimens having larger fruits than the west- ern ones. 3. Specimens from the western part of the dis- tribution area (Aet 287, Bauerlen 449, bb 32673, BW 12782, Everill 449, Jacobs 9132, 9200, NGF 35311, and Versteeg 1356) strongly resemble L. micans. The fruits are almost similar in shape, the only differences being the short hairs inside the fruit and the sarcotesta which does not cover the dorsal side of the seeds (in L. micans the hairs are long and the sarcotesta almost completely covers the seed except for a small dorsal triangle). Flow- ering specimens can be separated only if the pedi- cel is glabrous (it is always pilose in L. micans). LEPISANTHES (P.W. Leenhouts) Lepisanthes Blume, Bijdr. (1825) 237; Radlk. in Engl., Pflanzenr. 98 (1932) 726; Leenh., Blumea 17 (1969) 33-91. — Lepisanthes Blume sect. Eulepisanthes Radlk., Sapind. Holl.-Ind. (1879) 34, nom. illeg. — Lepisanthes Blume subg. & sect. Lepisanthes — Type species: Lepisanthes montana Blume [= Lepisanthes tetraphylla (Vahl) Radlk.]. Erioglossum Blume, Bijdr. (1825) 229; Radlk. in Engl., Pflanzenr. 98 (1932) 692. — Lepisanthes subg. Erioglossum (Blume) Leenh., Blumea 17 (1969) 60, 81. — Type species: Erioglossum edule Blume [= Lepisanthes rubiginosa (Roxb.) Leenh.]. Aphania Blume, Bijdr. (1825) 236; Radlk. in Engl., Pflanzenr. 98 (1932) 699. — Apha- nia Blume sect. Euaphania Radlk., Sapind. Holl.-Ind. (1879) 69, nom. illeg. — Lepi- santhes Blume subg. Aphania (Blume) Leenh., Blumea 17 (1969) 60, 83. — Type species: Aphania montana Blume [= Lepisanthes senegalensis (Poir.) Leenh.]. Otophora Blume, Rumphia 3 (1847) 142; Radlk. in Engl., Pflanzenr. 98 (1932) 753. — Otophora Blume sect. Euotophora Radlk., Sapind. Holl.-Ind. (1879) 86, nom. illeg. — Otolepis Turcz. sect. Otophora (Blume) Kuntze in Post & Kuntze, Lex. Phan. (1903) 408. — Lepisanthes Blume subg. Otophora (Blume) Leenh., Blumea 17 (1969) 62, 70. — Lepisanthes Blume sect. Otophora (Blume) Leenh., Blumea 17 (1969) 71. — Lectotype species (Leenhouts 1969): Otophora amoena Blume [= Lepisanthes amoe- na (Blume) Leenh.]}. Otophora Blume subg. or sect. Pseudophora Blume, Rumphia 3 (1847) 142, nom. inval. — Otophora Blume sect. Pseudotophora Radlk., Sapind. Holl.-Ind. (1879) 85. — Otolepis Turcz. sect. Pseudotophora (Radlk.) Kuntze in Post & Kuntze, Lex. Phan. (1903) 408, nom. illeg. — Lepisanthes Blume sect. Pseudotophora (Radlk.) Leenh., Blumea 17 (1969) 75. — Lectotype species (Leenhouts 1969): Otophora fruticosa Blume [= Lepisanthes fruticosa (Blume) Leenh. ]. Otolepis Turcz., Bull. Soc. Nat. Mosc. 21 (1848) 572. — Type species: Otolepis nigres- cens Turcz. (= Lepisanthes fruticosa (Blume) Leenh.). Hebecoccus Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 246; in Engl., Pflanzenr. 98 (1932) 719. — Lepisanthes Blume sect. Hebecoc- 628 Flora Malesiana ser. I, Vol. 11 (3) (1994) cus (Radlk.) Leenh., Blumea 17 (1969) 60, 67. — Type species: Hebecoccus ferru- gineus Radlk. [= Lepisanthes ferruginea (Radlk.) Leenh.]. Otophora Blume sect. Anomotophora Radlk., Sapind. Holl.-Ind. (1879) 85. — Otolepis Turez. sect. Anomotophora (Radlk.) Kuntze in Post & Kuntze, Lex. Phan. (1903) 408. — Lepisanthes Blume sect. Anomotophora (Radlk.) Leenh., Blumea 17 (1969) 79. — Type species: Otophora ramiflora Radlk. |= Lepisanthes ramiflora (Radlk.) Leenh.]. For a more complete list of synonyms and references, see Leenhouts (1969). Trees or shrubs, exceptionally lianas; mostly monoecious. /ndumentum of solitary, simple hairs; no glandular scales. Leaves spirally arranged, pari- or imparipinnate, some- times simple, 1- to more than 40-jugate, petiole and/or rachis winged or not, with or without pseudo-stipules. Leaflets opposite or alternate, not papillose beneath, margin entire. Inflorescences terminal, axillary, rami-, or cauliflorous, rarely sticky. Flowers unisexual, actinomorphic. Sepals 4 or 5 (rarely 3 or 6), free, imbricate, outer PLA (vied i) mostly distinctly smaller, mostly at least inner ones partly petaloid, entire or partly den- ticulate. Petals 4 or 5 (rarely 3, 6, or 7), shorter to longer than sepals, mostly distinctly clawed; scale mostly well developed, sometimes only represented by a hairy rim or a pair of small auricles, crested or not. Disc interrupted or not, mostly slightly lobed, without appendage. Stamens mostly c. 8 (4-18), in male flowers not to distinctly exserted; fila- ments nearly always hairy mostly either the base or the apex excepted; anthers hairy or glabrous. Ovary sessile to short-stipitate, lobed or not, 2- or 3(—4)-celled; style apical, about as long as the ovary or stigma sessile; stigma globular or dome-shaped, slightly lobed. Ovules 1 per cell, subbasal to median, placenta with an obturator. Fruits sessile to short-stipitate, not to distinctly lobed, drupaceous; outside smooth (or slightly warty), hairy to glabrous; pericarp thin- (or thick-)fleshy, inside hairy or glabrous; septa mostly complete, sometimes interrupted or represented merely by a rib. Seeds with a shining brown to black, glabrous or sometimes hairy testa; without arillode or sarcotesta; hilum basal, small. — Figs. 52-56. Distribution — 24 species in tropical Africa, Madagascar, S and SE Asia from Sri Lanka to Hainan, Malesia, and NW Australia. Habitat & Ecology — Mainly under everwet, sometimes also (or even mainly) in seasonal conditions. Most species are treelets or shrubs (or trees up to 20 m high) of the middle and lower storeys of primary and secondary forests, probably especially on open places, escarpments, steep slopes, river banks, forest edges, clearings, etc., and of more open secondary vegetations; altitude from sea level to c. 2000 m. The flowers have either a greenish calyx and creamy to white petals longer than the calyx, or (subg. Otophora and Aphania) a pinkish to red calyx not exceeded by the petals. They are small but are aggregated in small to large inflorescences, and are sweet-scented, and are probably pollinated by insects. The fruits of at least those species with a fleshy pericarp will be dispersed by animals. Taxonomy — Several genera were combined into Lepisanthes by Leenhouts (I.c.). He subdivided the genus as follows (in brackets the number of species in Malesia): Subgenus Lepisanthes Section Lepisanthes (1) Section Hebecoccus (2) Adema, Leenhouts, Van Welzen — Sapindaceae 629 Subgenus Otophora Section Otophora (4) Section Pseudotophora (2) Section Anomotophora (2) Subgenus Erioglossum (2) Subgenus Aphania (3) Uses — A few species have some value as ornamental trees. The wood of several species is used, but only few reach a sufficient size to have value as timber trees. Differ- ent parts of several species are of medicinal value. The fruits of some species are eaten, but only L. fruticosa and L. alata are planted for their fruits. Note — Outside Malesia petals may be absent and the ovary may be |-celled. KEY TO THE SPECIES 2. ESS SSS GS 2 So) SE an So rece oe. > ic 2 b. Pseudo-stipules (a pair of leaflets attached at the base of the petiole) present... 9 Bae Neither petiole Wor TACHIS WINGED oa a wi a a so a ee = Se eo See oe anaes 3 PE INCE ANG/OF TACIHS WINGER Stree abs See we Sk mee wale 0 15. L. mixta 3a. Sepals (sub)glabrous, as long as the petals.................0 20.202 eee eee, + bo Sepals sericeous, shorter than the petals... ....2...2 22). 000. cae ee ee 5 4a. Midrib angular beneath; nerves up to | cm apart. Inflorescences solitary, simple, up Satocmilone Pio Lye 2 CUE IY AE Sia. 28 14. L. dictyophylla b. Midrib rounded beneath, nerves at least 1 cm apart. Inflorescences either solitary and widely branched, or simple and fascicled, up to 60 cm long 16. L. senegalensis 5a. Fruits distinctly lobed, lobes spreading, subglabrous ..........-....-.--+--- 6 b. Fruits not or slightly, rarely distinctly lobed, lobes erect, densely hairy ....... 7 6a. Inflorescences axillary, sometimes together pseudoterminal, up to 5 cm long, hard- ly branched. Ovary subglabrous ..............------: 12. L. membranifolia b. Inflorescences terminal and in the upper leaf axils, 25-50 cm long, widely branched. wary censelwhaingts:s. sink s Geet JA e208 wee kk eee: 13. L. rubiginosa 7a. Leaflets mostly with glandular-pitted warts. Petal scales not crested. Dry fruit coarsely TEE EY a | 0 i RR ER A ar le OSI aera oe hae oe Fiat BL ile Sat ot SR ee 8 b. Leaflets without glandular-pitted warts. Petal scales mostly crested. Dry fruit smooth 1. L. tetraphylla 8a. Leaves more than 3-jugate. Fruits hardly lobed ..............-.. 2. L. faleata b. Leaves (1—)2(—3)-jugate. Fruits distinctly lobed.............. 3. L. ferruginea a ECUGIC AUG FAGHIS WINGER cco c nh Sos. Sing ene Cee ete eee Le epee ee ae eee 10 b. Petiole never, rachis very rarely winged ......... 2.2.60. e eee eee eee ene ees 1] 10a. Ovary 3- (or 4-)celled. Twigs and leaves glabrous. Leaflets narrow, usually +~7 times as long as wide, rarely more than 20 by 4 cm, base acute or sometimes round- 2. pa Ri OR TUN AES SO a eh WAS Wg Wk te ac ate RON 10. L. alata b. Ovary 2-celled. Twigs and leaves sparsely hairy. Leaflets broader, 3-5 times as long as wide, up to 33 by 10.5 cm, base cordate ............. 11. L. ramiflora lla. Fruits distinctly lobed, lobes spreading. .............-6 000s ee eee eee ee eee 12 630 Flora Malesiana ser. I, Vol. 11 (3) (1994) b. Fruits not or slightly, rarely distinctly lobed, lobes + erect ................. 14 12a. Leaves imparipinnate. Fruits 3-lobed ......... 2.0.50. s eee ee ee 13 b. Leaves paripinnate. Fruits nearly always 2-lobed .......... 16. L. senegalensis 13a. Leaves 7-14-jugate. Twigs up to | cm in diam., tomentose . 6. L. kinabaluensis b. Leaves (15—)30—40-jugate. Twigs 1.5-2 cm in diam., glabrous .. 7. L. multijuga 14a. Leaflets with pitted warts looking like small white scales. Leaves mostly impari- pinnate, (3—)7—42-jugate. Inflorescences terminal and axillary ............. 15 b. Leaflets densely finely pitted underneath. Leaves paripinnate or with a reduced ter- minal leaflet, 1—8(—14)-jugate. Inflorescences rarely terminal .............. 16 {San Neration open’)! 7a TA BEE aniees ee SNE cs SNRs 4. L. amoena be SNesvation closéd? Acre act SASAN OSE AY BEY GILES RE Te oe 5. L. divaricata Ga) Intlorescence and sepals densely hairy 7. 22.5. ..0...- 0/32 eae 8. L. bengalan Db dnilorescence and'sepals’clabrousae ssh ee...) ee ee ee 9. L. fruticosa Subgenus Lepisanthes Lepisanthes Blume — For a complete synonymy see Leenhouts (1969: 62). Leaves paripinnate, petiole and rachis not winged, without (outside Malesia excep- tionally with) pseudo-stipules. Leaflets opposite to alternate, nervation open to closed. Outer sepals sericeous outside. Petals longer than sepals, outside at least partly seri- ceous, scale crested or not. Disc glabrous or hairy. Stamens (4—)8(—18); filaments longer than anthers. Ovary 2- or 3-celled, densely hairy to subglabrous. Fruits slightly, rarely distinctly lobed, densely hairy, rarely glabrous, septa never interrupted. Distribution — S and SE Asia from Sri Lanka to Hainan, and Malesia. Section Lepisanthes Lepisanthes Blume — Lepisanthes Blume sect. Eulepisanthes Radlk. — For a complete synonymy see Leenhouts (1969: 62). Shrubs or trees. Leaves glabrous or at least petiole, rachis, and petiolules hairy. Leaf- lets without glandular-pitted warts, apex not mucronulate, midrib sunken to prominulous above, rounded to angular beneath. Inflorescences mostly axillary or rami- to cauliflorous, rarely terminal, solitary or fascicled, simple or (mostly sparsely) branched. Petal scales mostly crested. Fruits smooth or sometimes warty, outside densely hairy (to glabrous), pericarp thin, inside hairy or glabrous. Distribution — As the subgenus; two species, one of which (L. andamanica) endemic in the Andaman Islands. 1. Lepisanthes tetraphylla (Vahl) Radlk., Sit- 39, 63. — Sapindus tetraphylla Vahl, Symb. 3 zungsber. Math.Phys. Cl. Konigl. Bayer. Akad. (1794) 54. — Type: Koenig s.n. (herb. Vahl, C), Wiss. Miinchen 8 (1878) 276; Ridley, Fl. Ma- India. lay Penins. 5 (1925) 301; Craib, Fl. Siam. Enum. — Lepisanthes montana Blume, Bijdr. (1825) 238; 1 (1926) 327; Radlk. in Engl., Pflanzenr. 98 Hiern in Hook. f., Fl. Br. India 1 (1875) 679; (1932) 743, f. 15; Gagnep. in Fl. Indo-Chine, Koord. & Valeton, Bijdr. Booms. Java 9 (1903) Suppl.! (1950) 947; Leenh., Blumea 17 (1969) 165; Atlas 1 (1913) t. 131; Radlk. in Engl., Adema, Leenhouts, Van Welzen — Sapindaceae 1om Fig. 52. Lepisanthes Blume. Fruits. — a. L. senegalensis (Poir.) Leenh. — b. L. fruticosa (Roxb.) Leenh. —c. L. rubiginosa (Roxb.) Leenh. — d. L. ferruginea (Radlk.) Leenh. —e. L. alata (Blume) Leenh. — f. L. falcata (Radlk.) Leenh. subsp. borneensis (Leenh.) Leenh. — g. L. multijuga (Hook. f.) Leenh. — h. L. tetraphylla (Vahl) Radlk. (a: Kostermans 60; b: Nooteboom 1323; c: Kartawinata 683; d: SFN 21731; e: S 40142; ft: SMHI 618 (Podzorski); g: Kokawa & Hotta 2329; h: SAN 25591). Pflanzenr. 98 (1932) 732; Adelb., Blumea 6 (1948) 323; Backer & Bakh. f., Fl. Java 2 (1965) 135. — Type: Blume 676 (L), W Java. Leptsanthes sessiliflora Blume, Rumphia 3 (1847) 153; Boerl., Hand. 1 (1890) 271 (‘sessilifolia’); Radlk. in Engl., Pflanzenr. 98 (1932) 734. — Type: Blume 3100 (L), W Java. Lepisanthes angustifolia Blume, Rumphia 3 (1847) 154; Radlk. in Engl., Pflanzenr. 98 (1932) 735; Adelb., Blumea 6 (1948) 323. — Type: Korthals s.n. (L), W Java. Hemigyrosa longifolia Hiern in Hook. f., Fl. Br. Ind. 1 (1875) 671. — Lepisanthes longifolia (Hiern) Radlk., Sitzungsber. Math.-Phys. Cl. KGnigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 276; Ridley, Fl. Malay Penins. 1 (1922) 494; Radlk. in Engl., Pflanzenr. 98 (1932) 749. — Anomosanthes longifolia (Hiern) Pierre, Fl. Coch. (1895) t. 327 text. — Syntypes: Griffith KD 994 (M), Maingay KD 446 (L, M), both Malay Peninsula. Lepisanthes cuneata Hiern in Hook. f., Fl. Br. In- dia 1 (1875) 680; Ridley, Fl. Malay Penins. | (1922) 493; Radlk. in Engl., Pflanzenr. 98 (1932) 737. — Type: Porter s.n. (M), Malay Peninsula. Lepisanthes eriolepis Radlk., Sapind. Holl.-Ind. (1879) 36; Merr., Enum. Philipp. Flow. Pl. 2 632 Flora Malesiana ser. I, Vol. 11 (3) (1994) (1923) 500; Radlk. in Engl., Pflanzenr. 98 (1932) 738. — Type: Cuming 785 (A, FI, K, M), Philippines, Luzon. Lepisanthes schizolepis Radlk., Sapind. Holl.-Ind. (1879) 87, in Perkins, Fragm. Fl. Philipp. | (1904) 60; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 500; Radlk. in Engl., Pflanzenr. 98 (1932) 739. — Lepisanthes schizolepis Radlk. f. genuina Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 60, nom. illeg.; in Engl., Pflanzenr. 98 (1932) 740. — Type: Barthe s.n. (M, P), Phil- ippines, Luzon. Ostodes appendiculata Hook. f., Fl. Br. India 5 (1887) 401. —Lepisanthes kunstleri King, J. As. Soc. Beng. 65, II (1896) 427, nom. illeg.; Rid- ley, Fl. Malay Penins. 1 (1922) 493; Radlk. in Engl., Pflanzenr. 98 (1932) 746. — Lepisan- thes appendiculata (Hook. f.) Symington, Kew Bull. (1937) 320. — Type: King’s coll. 4634 (K), Malay Peninsula. Lepisanthes scortechinii King, J. As. Soc. Beng. 65, II (1896) 429; Ridley, Fl. Malay Penins. | (1922) 493; Radlk. in Engl., Pflanzenr. 98 (1932) 750. — Type: Scortechini 2090 (CAL, K, M), Malay Peninsula. Lepisanthes blumeana Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 168; Adelb., Blumea 6 (1948) 323; Backer & Bakh. f., Fl. Java 2 (1965) 135. — Type: Koorders 7406 (L, M), W Java. Lepisanthes viridis Radlk., Philipp. J. Sc., Bot. 8 (1914) 454; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 500; Radlk. in Engl., Pflanzenr. 98 (1932) 735. — Type: FB 9266 (M), Philippines, Mindanao. Lepisanthes acutissima Radlk., Philipp. J. Sc. 20 (1922) 675; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 499; Radlk. in Engl., Pflanzenr. 98 (1932) 737. — Type: Merrill 9564 (M), Philip- pines, Palawan. Lepisanthes macrocarpa Radlk., Philipp. J. Sc. 20 (1922) 657; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 500; Radlk. in Engl., Pflanzenr. 98 (1932) 738. — Type: BS 19460 (M), Philippines, Luzon. Aglaia chartacea Kosterm., Reinwardtia 7 (1966) 261, f. 14. — Type: Van Steenis 6455 (BO), Sumatra. For a more complete list of synonyms and refer- ences, see Leenhouts (1969). Mostly a shrub or treelet, sometimes a tree up to 22.5 m high, dbh up to 60 cm. Twigs terete (to angular), up to 2.5 cm in diam., glabrous or varia- bly fulvous to ferrugineous hairy, mostly early gla- brescent, usually at first dark (purplish) brown, later on often greyish brown, mostly conspicuously len- ticellate. Leaves 2—10-jugate, axial parts glabrous or variably hairy; petioles 5—SO cm long; petiolules 2-25 mm. Leaflets lanceolate to elliptic, ovate to obovate, 7-55 by 2—10(—20) cm, chartaceous (per- gamentaceous), glabrous or variably hairy; base slightly (or strongly) oblique to equal-sided, acute to rounded (or subcordate); apex tapering to abrupt- ly acuminate, acumen short to long, obtuse to acute, nerves variable, at least the upper ones (rarely all) looped and joined or exceptionally connected by an intramarginal vein, above prominulous or some- times sunken, beneath prominulous to prominent, intercalary veins variable, veins and veinlets fine- ly reticulate. Inflorescences 2-70 cm long, varia- bly, mostly shortly and densely hairy, hardly to long peduncled, the main branches often racemous; cymes patent, sessile or up to 0.5 cm long stalked, up to c. 7-flowered; bracts ovate-lanceolate to sub- ulate, up to 5 mm long, rarely ovate and up to 7 by 6 mm; pedicels up to 6 mm. Flowers white, some- times greenish white, cream, or pink, sweet-scent- ed. Sepals: outer ovate to suborbicular, 1.2—7 by 1.2-6 mm, densely (sometimes partly glandular- )ciliolate, inside glabrous (to sparsely hairy); in- ner 3 elliptic to orbicular, ovate to obovate, 2.2— 6.2 by 1.8-6 mm, margin mostly narrow (to broad) petaloid, outside glabrous, crenulate to fimbriate- denticulate, indument otherwise like outer sepals. Petals 2.2-10 mm long, claw 0.5—2(—4) mm long, blade elliptic to oblong to ovate, 1.2-4 mm wide, more or less dentate in the upper part, outside most- ly up to 2/3 sericeous, rarely subglabrous, margin glabrous or below insertion of scale sparsely to densely woolly-ciliate, rarely rest of the blade (part- ly glandular-)ciliate, inside glabrous or the claw (rarely also the base of the blade) sparsely hairy; scale mostly well-developed (rarely a hairy rim only), entire to deeply 2—4-lobed, glabrous, cili- ate, or inside sparsely to densely woolly, without or more often with a crest which may vary from a small wart (then often present only in some of the petals of a flower) to a deeply bilobed duplication of the scale or to 2 brushes. Stamens: filaments 1.5-6.5 mm, sparsely to densely woolly, mostly mainly in the upper, rarely in the lower half; an- thers (broad-)elliptic to oblong, ovate to obovate, (0.82.5 mm, connective broad and obtuse (rarely narrow and/or pointed), woolly to glabrous. Ova- ry and lower part of style densely hairy (to subgla- brous). Fruits green when young, later yellowish, grey, or greyish pink (ripe?), flattened ellipsoid, shortly obovoid, or subglobular, slightly lobed, the lobes rarely carinate, 1.5-5 cm in diam. Seeds brown, testa papery, probably partly fleshy when fresh. — Fig. 52h. Distribution — SE Asia from Sri Lanka and the Deccan Peninsula to Hainan; Malesia: Sumatra, Adema, Leenhouts, Van Welzen — Sapindaceae 633 Malay Peninsula, Borneo, Java (western half only, incl. also Nusa Kembangan), Philippines, N Cele- bes (one collection), Timor, and New Guinea. Habitat & Ecology — Understorey of primary and secondary forests, also along forest edges, on river banks, and in more open country, as well on dry land as in swampy localities or periodically inundated places, or even in streams, on all kinds of soils; up to 1200 m altitude. Fl. Jan.—Dec.; fr. Nov.—July. Uses — The wood of some forms is heavy, hard, and close-grained, and good for furniture and for turning. One specimen from the Malay Peninsula was said to be used in the preparation of dart poi- son. See also Burkill, Dict. Econ. Pl. Malay Penins. (1935) 1332, sub L. kunstleri. Chromosomes — 2n = 26: Sarkar et al. in Love, Taxon 26 (1976) 636, 649. Notes — 1. Lepisanthes tetraphylla is closest to L. andamanica King (S Andaman) which differs only by the shifting of the lower pair of leaflets to the very base of the leaf, simulating a pair of stip- ules. 2. Lepisanthes tetraphylla in the sense accept- ed here is a very polymorphous species. An exten- sive survey of the many forms distinguishable on a local scale can be found in Leenhouts, Blumea 17 (1969) 39-52. In some parts of the area (Malay Peninsula, Philippines) most of the specimens can easily be placed in distinct forms, in other parts the limits are vague, and in some parts (Borneo, Sumatra) new collections may not match any of the forms provisionally distinguished. Some of the more important local forms are: Malay Peninsula: ‘longifolia’: large leaflets (18-38 by 5.5—9 cm); large (c. 8 mm long) ferru- gineous-hairy flowers; and warty fruits. — ‘race 21’: indumentum of dense, short, patent, soft hairs intermingled with c. 5 mm long, stiff, irritating hairs; drooping catkin-like inflorescences with broad-ovate, large (5 by 5-6 mm) bracts. — ‘hir- ta’: indumentum beautiful reddish, yellowish, or greyish brown velvety; large (19-39 by 7-17 cm) leaflets, with many straight nerves, hispid all over the lower surface, the hairs being on minute warts; and relatively large (4 by 4 mm) broad-ovate bracts. — ‘cuneata’: glabrous twigs; sepals not petaloid; ovary 2-celled. — ‘montana’ like ‘cuneata’, but with midrib sunken above and ovary 3-celled. Sumatra: ‘lamponga’ is the predominant form, much resembling ‘montana’ but usually with 4 petals, an interrupted disc and densely hairy seeds. — ‘cuneata’ (but with 3-celled ovary) and ‘hirta’ are known from the E Coast, ‘montana’ from Pa- lembang. Java: ‘montana’ but lacking any crest to the petal scale. — ‘heterolepis’ like ‘montana’, but with at least some of the petal scales + crested. (The de- limitation between these two forms is highly arti- ficial.) Timor: All material from Timor represents ‘pal- lens’. Borneo: The main forms from Borneo are ‘mon- tana’ and ‘heterolepis’. Philippines: The predominant form is ‘schizo- lepis’. Section Hebecoccus Lepisanthes sect. Hebecoccus (Radlk.) Leenh., Blumea 17 (1969) 60, 67 (see there for complete synonymy). — Hebecoccus Radlk. Trees. Leaves glabrous. Leaflets mostly with scattered glandular-pitted warts on both surfaces, apex mucronulate, midrib prominulous above, rounded beneath. /nflorescences in Malesian species terminal and in the upper leaf axils, spreadingly but sparsely branched. Petals 5, scale in Malesian species not crested. Disc entire. Fruits not or slightly lobed, smooth but with thick, fleshy pericarp which is more or less strongly wrinkled when dry, outside thinly to densely hairy, inside glabrous. Seeds glabrous. Distribution — 5 species, 3 in continental Asia from Sri Lanka to Indo-China, the other 2 in Malesia. Pflanzenr. 98 (1932) 721. — Lectotype (Leen- houts 1969): FB 1263] (L, M), Philippines, Leyte. Hebecoccus inaequalis Radlk., Philipp. J. Sc., Bot. 2. Lepisanthes falcata (Radlk.) Leenh., Blumea 17 (1969) 69. — Hebecoccus falcatus Radlk., Philipp. J. Sc., Bot. 8 (1914) 453; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 499; Radlk. in Engl., 634 8 (1914) 453; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 499; Radlk. in Engl., Pflanzenr. 98 (1932) 721. — Type: FB 6459 (L, M), Philip- pines, Cebu. Lepisanthes aphanococca Leenh., Blumea 17 (1969) 67. — Aphanococcus celebicus Radlk. in Durand, Ind. Gen. (1888) 74; in Engl., Pflan- zenr. (8 (1932) 723. — Type: Riedel s.n. (K, M), Celebes. Lepisanthes borneensis Leenh., Blumea 17 (1969) 68. — Type: Chew, Corner & Stainton RSNB 2936 (BO, L, SAR), Sabah. Tree, up to 22 m high, dbh up to 40 cm. Branch- lets terete, 1—-1.5 cm in diam., greyish or silvery brown, when young sparsely yellowish brown to- mentose to glabrous. Leaves 4—10-jugate; petiole terete to slightly flattened above, swollen at base, 7-25 cm, light yellowish green; petiolules grooved above, swollen, 4-15 mm. Leaflets subopposite to alternate, elliptic to ovate or oblong to lanceolate, widest at or below the middle, not or slightly fal- cate, 10-28 by 3-10 cm, index 2—3.5, thin- to stiff- chartaceous, shining bright green above, dull green beneath, glandular-pitted warts rather numerous on both sides or scattered on lower surface only; base equal-sided to oblique, acute to rounded and at- tenuate, or subcordate; apex acute to tapering acu- minate; nerves c. 15 per side, 1.5—3 cm apart, an- gle to midrib 60—85° spreading, slightly to more curved, looped and joined at some distance from the margin or not, some intercalary veins some- times as strongly developed as the nerves. Jnflo- rescences 20-70 cm long, minutely brown tomen- tose; branches up to 40 cm long, patent to oblique erect; cymes condensed, several-flowered, stalks c. 5 mm long; bracts narrowly triangular, c. 3 mm long; pedicels c. 1 mm. Sepals ovate to obovate to suborbicular, 34.5 by 24.5 mm. Petals 4—7 mm long, claw 0.75—2.5 mm long, blade elliptic to sub- orbicular, 2.3-5 mm long, outside claw and base of blade glabrous or sericeous, claw (and basal 1/5 of the blade) densely long-ciliate, inside glabrous or claw sparsely hairy; scale short and broad, en- tire, bilobed, or 3-dentate, densely bearded. Sta- mens: filaments 2-3 mm; anthers ovate, 1—1.5 mm. Fruits not or 3-lobed, often only | part developed, but sterile cells not visible, 1.8—2.5 by 1.8—3.5 cm, outside densely shortly brown velvety, corrugat- ed. Seeds globular-obovoid, 15 by 8-10 mm, hilum basal, small, circular. Distribution — Malesia: Borneo, Philippines, Celebes. Habitat & Ecology — Primary mountain forests, rocky banks of streams, forested steep slopes and ridges; altitude 60-1500 m. Fl. Nov.; fr. Mar. Flora Malesiana ser. I, Vol. 11 (3) (1994) KEY TO THE SUBSPECIES la. All nerves looped and joined ........... 2 b. Only the nerves in the upper 1/3—1/4 of the leaflet distinctly looped and joined b. subsp. borneensis 2a. Leaflets c. 25 by 7 cm, not falcate, base not OUNGUE. 2AM. Be Ae Ae c. subsp. celebica b. Leaflets up to 16 by 6 cm, slightly falcate, base ODMGUC™ vais iam so HF a. subsp. falcata a. subsp. falcata Leaves 6—10-jugate. Leaflets 10-22 by 3-7 cm, slightly falcate, nervation closed, nerves looped and joined at some distance from the margin, many glandular-pitted warts on both sides. Fruits not lobed, usually only 1 part developed, c. 2 by 1.8 cm, wrinkled. Distribution — Malesia: Philippines (Palawan, Luzon, Leyte, Cebu). Habitat & Ecology — Forested slopes and ridg- es; altitude up to 320 m. FI. Nov.; fr. Mar. Uses — Good timber. b. subsp. borneensis (Leenh.) Leenh., Blumea 18 (1970) 429. — Lepisanthes borneensis Leenh. Leaves 4—6-jugate. Leaflets 14-28 by 4-10 cm, not or slightly falcate, glandular-pitted warts scat- tered on lower surface, base equal-sided or oblique, nervation mostly open, only the nerves in the up- per 1/3—1/4 distinctly looped and joined at several mm from the margin. Fruits slightly 3-lobed, c. 2.5 by 3.5 cm, not rarely 1 or 2 cells not developed, finely wrinkled. — Fig. 52f. Distribution — Malesia: Borneo, Philippines (Palawan). Habitat & Ecology — Primary mountain forests and on rocky banks of streams; altitude 60-1500 m. Fl. Mar., June, Sept., Nov.; fr. Mar., May. c. subsp. celebica (Radlk.) Leenh., Blumea 18 (1970) 429. — Aphanococcus celebicus Radlk. in Durand — Lepisanthes aphanococca Leenh. Leaves at least 3-jugate (only upper part of leaf known). Leaflets 25-27 by 7—7.5 cm, not falcate, base not oblique, nervation closed, nerves looped and joined at some distance from the margin. Fruits not lobed, c. 1.8 by 2 cm, minutely wrinkled. Distribution — Malesia: Celebes (once collect- ed: Gorontalo). Habitat & Ecology — Fr. June. 3. Lepisanthes ferruginea (Radlk.) Leenh., Blu- mea 17 (1969) 69.— Hebecoccus ferrugineus Adema, Leenhouts, Van Welzen — Sapindaceae Radlk., Sitzungsber. Math.-Phys. Ci. Konig]. Bayer. Akad. Wiss. Miinchen 8 (1878) 301; Sapind. Holl.-Ind. (1879) 19, 22, 56, 68; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 162; Backer, Schoolfl. Java (1911) 262; Koord. & Valeton, Atlas (1913) f. 134; Radlk. in Engl., Pflanzenr. 98 (1932) 720; Backer & Bakh. f., Fl. Java 2 (1965) 134. — Type: Zollinger 3459 (FI, L), Java. Sapindus laurifolia auct. non Vahl: Teijsm. & Binn., Cat. Hort. Bog. (1866) 215, p.p. Tree, up to 20 m high, dbh up to 60 cm. Branch- lets terete, 6 mm in diam., slightly ribbed, greyish to yellowish brown, when young rather densely minutely hairy, glabrescent. Leaves (1—)2(—3)-ju- gate; petiole slightly flattened above, rounded be- neath, swollen at base, 5—9 cm long; petiolules deeply sulcate above, slightly swollen, 3—5(—9) mm. Leaflets mostly opposite, oblong-ovate to oblong to lanceolate, 10-20 by 4—7.5 cm, chartaceous, brownish to greenish above, greenish beneath, glan- dular-pitted warts very scarce; base equal-sided, obtuse to acute; apex slightly tapering acuminate, acute to obtuse; nerves |—3 cm apart, angle to mid- rib 45—80°, usually slightly curved, looped and joined at some distance from the margin, some- times straight and not joined but for the few upper ones, some intercalary veins strongly developed, veins and veinlets finely reticulate. /nflorescences 635 up to 45 cm long, ferrugineous to fulvous velvety, glabrescent; pedicels 2-4 mm long, deeply 3- grooved. Sepals in vivo bright green, outside ful- vous-sericeous, margin glandular-ciliolate, inside glabrous, outer 2 oblong-ovate, 2.8 by 2 mm, in- ner 3 broadly obovate, 3.5 by 2.5 mm. Petals obo- vate-cuneate, 4.5 by 2.8 mm, white, hardly clawed, outside in the basal half densely appressed long- hairy, the margin except for the upper 1/3 long- ciliate; scale erect, slightly more than half as long as the petal, laterally partly adnate to its margin, the free part deltoid to bilobed, densely ciliate. Disc glabrous. Stamens slightly exserted; filaments 3.5 mm, densely long-hairy; anthers ovate, apiculate, 1 mm, glabrous. Ovary 1.5 mm, style conical, | mm, stigma faintly 3-lobed. Fruits slightly 3-lobed, mostly only | lobe developed, this globular, c. 2.5 cm in diam., densely ferrugineous (when fresh or- ange to brown) velvety; pericarp in vivo 3 mm thick, orange. Seeds slightly flattened ellipsoid, 1.8 by 1.2 by 1 cm, testa shining, papery, hilum elliptic, 5 by 2 mm. — Fig. 52d. Distribution — Malesia: Sumatra (once collect- ed: East Coast, Sibolangit), Malay Peninsula (Pe- rak: near G. Bubu; Tioman I.), Java (few: Bantam, Preanger, Pasuruan). Habitat & Ecology — Primary rain forest, up to c. 450 m altitude. Fl. Feb.—Apr.; fr. Apr.—June, Aug. Uses — Timber used for construction. Subgenus Otophora Lepisanthes subg. Otophora (Blume) Leenh., Blumea 17 (1969) 60, 70 (see there for a complete synonymy). — Otophora Blume. Leaves pari- or imparipinnate, petiole and rachis winged or not, with pseudo-stipules. Leaflets opposite to alternate, nervation open to closed. Petals shorter than sepals, out- side mostly glabrous, scale not crested. Disc entire, glabrous or hairy. Stamens 5—10; filaments shorter than anthers. Ovary 2- or 3(—4)-celled (outside Malesia also 1-celled), often glabrous. Fruits mostly not or only slightly lobed, sometimes parted, glabrous or thin-hairy, septa often more or less interrupted. Distribution — SE Asia from Burma (Mergui) and Hainan eastwards to the Moluccas. Section Otophora Lepisanthes sect. Otophora (Blume) Leenh., Blumea 17 (1969) 60, 71 (see there for a complete synonymy). — Otophora Blume sect. Euotophora Radlk. Twigs glabrous or glabrescent. Leaves usually imparipinnate, often many-jugate, hairy or glabrous; petiole and rachis not winged. Leaflets especially above with pitted warts 636 Flora Malesiana ser. I, Vol. 11 (3) (1994) resembling small white scales, often hairy, midrib beneath usually rounded. /nflores- cences terminal and axillary, mostly pyramidal, hairy or glabrous. Sepals 5 (or 6), mostly (sub)glabrous. Petals (4 or) 5, outside often hairy; scale either represented by two auri- cles, or lobed, ciliolate. Anthers hairy. Ovary 2- or 3-celled, hairy or glabrous. Fruits smooth to scurfy, hairy or glabrous; septa complete. Distribution — 5 species, | in Hainan, the others in West Malesia as far as Borneo and Timor. 4. Lepisanthes amoena (Hassk.) Leenh., Blumea 17 (1969) 71. — Melicocca amoena Hassk., Flora 25, 2 (1842) Beibl. 39 (‘Melicoccus amoe- nus’); Tijd. Nat. Gesch. Phys. 10 (1843) 139; Aanteek. Nut Java (1845) 17. — Schleichera amoena (Hassk.) Walp., Rep. 5 (1845) 366. — Otophora amoena (Hassk.) Blume, Rumphia 3 (1847) 142; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 172; Backer, Schoolfl. Java (1911) 263; Radlk. in Engl., Pflanzenr. 98 (1932) 771; Backer & Bakh. f., Fl. Java 2 (1965) 135. — Otolepis amoena (Hassk.) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Type: Reinwardt s.n. (L), Java. Otophora spectabilis Blume, Rumphia 3 (1847) 142: Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 171; Atlas 1 (1913) f. 130; Radlk. in Engl., Pflanzenr. 98 (1932) 772. — Capura spectabilis (Blume) Teijsm. & Binn., Cat. Hort. Bog. (1866) 214. — Otolepis spectabilis (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Type: Anonymous s.n. (L), Java. Otophora spectabilis Blume var. pubicosta Blume, Rumphia 3 (1847) 143. — Syntypes: Zippel s.n., Anonymous s.n., both Java. Otophora confinis Blume, Rumphia 3 (1847) 143. — Type: Korthals s.n. (L), E Borneo. Otophora imbricata Blume, Rumphia 3 ( 1847) 144; Ridley, Fl. Malay Penins. 1 (1922) 495; Radlk. in Engl., Pflanzenr. 98 (1932) 773. — Otolepis imbricata (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Syntype: Korthals s.n. (L), Bor- neo. Otophora pubescens Blume, Rumphia 3 (1847) 145; Radlk. in Engl., Pflanzenr. 98 (1932) 770. — Otolepis pubescens (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Syntypes: Korthals s.n. (L), Borneo; S. Miiller s.n. (L), S Borneo. Otophora cordigera Radlk., Sapind. Holl.-Ind. (1879) 85; in Engl., Pflanzenr. 98 (1932) 770. — Otolepis cordigera Kuntze, Rev. Gen. PI. 1 (1891) 144. — Type: Beccari PB 3359 (FI, M), Borneo. Otophora styligera Radlk. in Engl., Pflanzenr. 98 (1932) 774. — Type: Native collector 1784 (A, M), Borneo. Tree, up to 10 m high, dbh up to 15 cm, or shrub up to 6 m. Twigs 0.8—1(—1.2) cm in diam., brown, when young sparsely, rarely rather densely, short- ly fulvous hairy, mostly early glabrescent. Leaves imparipinnate, 7—42-jugate, up to 90 cm long, usu- ally glabrous to thin-tomentose on axial parts, sometimes axial parts densely fulvous-tomentose, rarely moreover hirsute, glabrescent; flush from white over pink to salmon or coral-red; petiole terete to flattened above, 1-9 cm long; pseudo-stip- ules orbicular, ovate, or transversely elliptic, usu- ally oblique, 1-6 by 0.8—6 cm, base truncate to deeply cordate, apex rounded, obtuse, acute, or shortly and broadly acuminate, some very oblique pseudo-stipules with a second more lateral apex, penni- or retinerved; pseudo-stipules usually con- nected with the normal leaflets by a series of inter- grades. Leaflets opposite to alternate, (sub)sessile, linear or sometimes ovate-lanceolate, 7—22.5 by 1.2—5 cm, index 3.5—7, thin-chartaceous or perga- mentaceous, greenish grey to dark brown above, yellowish- to red-brown beneath, glabrous to the midrib densely fulvous- to ferrugineous-tomentose above, sparsely hirsute beneath, rarely thinly hairy all over the lower surface, often on both surfaces or only above with scattered, minute, glandular- pitted warts; base oblique or not, obtuse to subcor- date; apex obtuse to tapering acuminate, acumen short to long, obtuse to acute; nerves 0.5—2 cm apart, angle to midrib 65—85°, curved, only the upper ones looped and joined at some distance from the margin. Inflorescences terminal and axillary, pyramidal, up to 60 cm long, sparsely to densely short fulvous-hairy, the axial parts in vivo often reddish, rachis and main branches sharply 3-angu- lar; branches nearly transverse to ascending, up to 30 cm long, the lower ones often with some short, spreading branches; cymes short-stalked to sessile, several- to 1-flowered; pedicels 2-4 mm. Flowers mostly reported to be white, sometimes creamy to yellow or pink to red, not scented. Sepals red, out- side thinly appressed short-hairy to glabrous, in- side glabrous to sparsely appressed short-hairy in the basal half, usually sparsely partly glandular ciliolate mainly towards the base, outer two ovate to oblong, 1.5—3.5 by 1.2—2 mm, inner 3 + orbicu- Adema, Leenhouts, Van Welzen — Sapindaceae 637 Fig. 53. Lepisanthes amoena (Hassk.) Leenh. a. Leaf; b. detail of lower surface of leaflet; c. fruit (a, b: Danser 5474, c: Wirawan 63). 638 Flora Malesiana ser. I, Vol. 11 (3) (1994) lar, 2.5—4 by 2—3.5 mm. Petals white or red, usual- ly up to 2/3 mm clawed, outside very sparsely to densely appressed long-hairy, inside variably hairy, claw and at least base of blade densely to very sparsely ciliate, blade subdeltoid to suborbicular, up to 1.5 by 1.2 mm, at base with 2 incurved, + connected lobes. Disc glabrous, orange or yellow. Stamens 7-9; filaments white or reddish; anthers 1.5-1.8 mm, densely hairy, yellowish. Ovary 3- celled, densely velvety to nearly glabrous, cream to reddish; style 1.5 mm. Fruits slightly 3-lobed, widest in or above the middle, 2—2.5 by 2.2—2.8 cm (fresh 2.5 by 3 cm), apiculate by the style base, (sometimes hardly) scurfy, thinly short-hairy to glabrous, in vivo light green, spotted brown when unripe, to brownish or purple when ripe, pulp yel- lowish to white. Seeds oblique-ellipsoid, hilum orbicular to lanceolate, small. — Fig. 53. Distribution — Malesia: Sumatra, Malay Penin- sula (Pahang, Selangor), Borneo, W Java, Timor (one collection). Habitat & Ecology — Primary (or secondary) forests, scrub, or bamboo forests, often along riv- ers, in swamp forests, or even in periodically in- undated localities, but also on dry-land, as well on acid as on basic soils, on loam or clay as well as on sand, at 0-400(—1650) m altitude. FI., fr. Jan.— Dec. The often large pseudo-stipules, appressed to the twigs, are used as shelter by ants. Uses — In Java sometimes planted as an orna- mental tree. The bark as well as the young leaves contain saponin and are applied against ulcers (Bor- neo) or an extract of the bark is used against hoarse- ness. The wood is reported to be very hard and was used for making hooks to catch crocodiles. The fruitpulp is sweet and edible. See Heyne, Nutt. PI. Indon. ed. 3 (1950) 990; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1613; Jansen et al. in Ver- heij & Coronel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991) 343. Notes — 1. Lepisanthes amoena occupies a cen- tral position in sect. Otophora and is more or less closely related to all the other species. It is itself a variable species but, apart from one race in Su- matra East Coast, no clearly delimited infraspecif- ic taxa can be distinguished. That race, represent- ed by Lérzing 4164, 5219 and 11988, all from around Sibolangit, differs from the above descrip- tion constantly in a few characters: twigs 1.5 cm thick; leaves 40-50-jugate, 1-2 m long (from young, unbranched treelets), leaflets 12-13 by 1.8- 2.2 cm, index 6—7, above midrib densely fulvous- tomentose, beneath mainly on the midrib thin-hir- sute, base slightly cordate, nearly equal, apex long acute-acuminate, nerves rather dense, 0.5—1 cm apart, all looped and joined at some distance from the margin; calyx light green to whitish; blade of petals 2.8-3 by 2.5 mm, scale sometimes hardly developed, glabrous; anthers glabrous; fruits dense- ly ferrugineous velvety, rarely glabrescent. What taxonomic status should be given to this race is not yet clear. It is doubtless nearest to what was described as Otophora pubescens. 2. Other variability is only somewhat discon- tinuous. Three ‘forms’ are more or less distinguish- able, clearest in Borneo where they are also geo- graphically more or less restricted. Outside of Bor- neo, the differences are often less clear, however. These races are: ‘amoena’: leaves rather many-ju- gate, leaflets not very narrow, linear, often greyish green above, reddish brown beneath. Throughout the area. — ‘imbricata’: leaves few-jugate, leaf- lets large, relatively wide, the sides not parallel, obtuse at base, brown. The series of intergrades between pseudo-stipules and leaflets on which Otophora imbricata was based, is exceptional. E Borneo and Java. — ‘pubescens’: leaves many- jugate, leaflets small and narrow, linear, cordate at base, mostly dark brown above, midrib usually hirsute beneath. W Borneo and Sumatra. 5. Lepisanthes divaricata (Radlk.) Leenh., Blu- mea 17 (1969) 72. — Otophora divaricata Radlk. in Fedde, Rep. 18 (1922) 338; in Engl., Pflanzenr. 98 (1932) 758. — Type: Anonymous s.n. (M, SAR), Sarawak. Otophora pyramidalis Radlk. in Fedde, Rep. 18 (1922) 338; in Engl., Pflanzenr. 98 (1932) 759. — Type: Native collector 375 (M), Sarawak. Otophora lunduensis Radlk. in Fedde, Rep. 18 (1922) 339; in Engl., Pflanzenr. 98 (1932) 769. — Type: Foxworthy 17 (M), Sarawak. Otophora macrocarpa Ridley, Kew Bull. (1933) 190; Radlk. in Engl., Pflanzenr. 98 (1934) 1494. — Type: Haviland 67 (K, SAR), Sarawak. Small tree, up to 7 m high, or shrub. Twigs 3— 13 mm in diam., brown, glabrous to rather densely shortly ferrugineous hairy and glabrescent. Leaves pari- or imparipinnate, (3—)7—11(—16)-jugate, 25— 40 cm long, the axial parts ferrugineous or fulvous short-hairy, glabrescent; petiole + terete, 2—6(—17) cm; pseudo-stipules suborbicular to elliptic, often slightly oblique, (0.4—)1—1.8 by (0.3—)1.2—2.2 cm, cordate to cuneate at base, rounded to acute at apex, reti-, palmati-, or penninerved. Leaflets opposite to alternate, sessile or with up to 2 mm long peti- olules, elliptic to lanceolate, 7-20 by 2—7 cm, in- dex 2.2—4, stiff-chartaceous, mostly greenish grey to blackish above, light to dark brown beneath, gla- brous or the midrib ferrugineous tomentose on both sides, mainly beneath with pitted warts; base sub- Adema, Leenhouts, Van Welzen — Sapindaceae cordate or rounded to cuneate, oblique, in termi- nal leaflet long attenuate; apex tapering acuminate, acute to obtuse; nerves 1—2.2 cm apart, angle to midrib c. 75°, nearly straight, distinctly looped and joined at some distance from the margin. /nflores- cences terminal and in the upper leaf-axils, pyram- idal, up to more than 40 cm long, glabrous to fulvous short-hairy, repeatedly branched; main branches patent, up to 25 cm long; cymes short- stalked to sessile, few- to many-flowered; pedicels slender, 1-1.2 mm. Flowers (orange-)yellow. Se- pals sparsely (outer) to densely (inner), partly glan- dular ciliolate, outer two broad-ovate to obovate, 1.8—2.2 by 1.2 mm, inner suborbicular to ovate, 1.8-3.5 by 1.8—-2.2 mm. Petals distinctly clawed, completely long-hairy, blade transversely semi-el- liptic, 0.7—1.5 by 1.2—2 mm, auricled at base. Disc glabrous or fairly densely short-hairy. Stamens 5— 8; anthers | mm, rather densely woolly. Ovary 2- or 3-celled, glabrous to thinly appressed-hairy; stig- ma sessile or on a short style. Fruits slightly 2- or 3-lobed, 2—3 by 2—3 cm, hardly pointed, smooth and glabrous, yellow to pale brown. Seeds obovoid, hilum elliptic, 3 mm long. Distribution — Malesia: Borneo (Sarawak: near Kuching). Habitat & Ecology — Disturbed kerangas or swamp forest behind the mangrove, at low altitude (up to 400 m). Fl. Oct.—July; fr. Oct.—Jan. The following two forms can be distinguished: a. forma divaricata — Otophora divaricata Radlk. — Otophora macrocarpa Ridley. Leaves imparipinnate, 6- or more-jugate; pseu- do-stipules orbicular, at least 8 by 8 mm, sessile, base rounded to cordate, never penninerved. Leaf- lets sessile, index 3-4, up to 4 cm wide, with par- allel sides. Ovary 2-celled. — Fig. 54. b. forma lunduensis (Radlk.) Leenh., Blumea 17 (1969) 72. — Otophora lunduensis Radlk. Leaves paripinnate, up to 6-jugate; pseudo-stip- ules elliptic, up to 8 by 4 mm, subsessile, base cuneate to subcordate, mostly penninerved. Leaf- lets short-stalked, index 2.5—3, up to 7 cm wide, the sides not parallel. Ovary 3-celled. Notes — 1. The species seems closest to L. amoe- na. 2. A few specimens are intermediate between the two forms. Native collector 375, the type of Otophora pyramidalis, comes near f. lunduensis but has the ovary 2-celled; Daun (Haviland) 917 and Daun (Haviland) s.n. (herb. SING 23156) look like f. divaricata but have a 3-celled ovary and small 639 pseudo-stipules, whereas Daun (Haviland) s.n., moreover, lacks the terminal leaflet. 6. Lepisanthes kinabaluensis Leenh., Blumea 17 (1969) 73. — Type: Chew & Corner RSNB 4998 (K, L), Sabah. Tree, up to 10 m high, or shrub. Twigs 0.8-1 cm in diam., dark purplish to greyish brown, dense- ly and shortly fulvous-tomentose, glabrescent. Leaves imparipinnate, 7—14-jugate, up to 75 cm long, axial parts hairy like the young twigs; peti- ole terete, flattened towards the base, S—13 cm long; pseudo-stipules orbicular to broad-ovate, 2—5 by 2.5—4 cm, deeply cordate at base, rounded or ob- tuse to short-acuminate at apex, penni- to reti- nerved, distinctly set off from the normal leaflets. Leaflets mostly about opposite, sessile or with up to 2 mm long petiolules, lanceolate to oblong, par- allel-sided, 12—20 by 3.5—5 cm, index 3-5, perga- mentaceous, mostly greyish green above, greenish beneath, glabrous or above on midrib shortly ful- vous-tomentose, on both faces with scattered, glan- dular-pitted warts; base oblique to equal-sided, cuneate or lower half rounded; apex tapering acu- minate, acumen long, acute or obtuse; nerves 1—2 cm apart, angle to midrib 70—75°, straight to slightly curved, more or less distinctly looped and joined near or at some distance from the margin. /nflo- rescences terminal, pyramidal, c. 25 cm high, red, densely short-hairy to subglabrous; branches pat- ent, short-stalked to sessile; cymes several-flow- ered; pedicels up to 5 mm long. Sepals deep rose red, sparsely ciliolate in the apical part, outer 2 ovate, 3.5—5 by 2.5—3 mm, inner 3 elliptic to or- bicular, 4-5 by 2.5—5 mm, especially the inner ones rather thin and probably for the greater part peta- loid, sometimes more or less grading to the petals. Petals 4, 0.5 mm clawed, outside pink, towards the base more red, inside whitish, claw and base of blade ciliate, upper part sparsely ciliolate, further glabrous or outside appressed hairy, plate wide- rhomboid, 2.5 by 3.5 mm, scale bilobed or the lat- eral parts reflexed, the centre erect and hood- shaped, 2/5 as long as the blade. Disc glabrous or short-hairy. Stamens 7-10; anthers 2 mm. Ovary 3-celled, 3-angular-obovoid, 3-lobed, glabrous: stigma sessile, dome-shaped. Fruits 3-lobed, of- ten 1 or 2 parts not developed, then style nearly basal, lobes spreading, ellipsoid, 3.2 by 2.2 cm, scabrous, glabrous, reported brown, yellow, orange, or purplish-red when ripe. — Fig. 55. Distribution — Malesia: Borneo (Mt Kinabalu, Crocker Range). Habitat & Ecology — In forest between 1200 and 2000 m altitude. Fl. Mar.—Apr-.; fr. Jan., Mar.— Apr., June. 640 Flora Malesiana ser. I, Vol. 11 (3) (1994) JH YOs77 Fig. 54. Lepisanthes divaricata Leenh. f. divaricata. a. Habit; b. female flower; c. pistil, longitudinal section; d. petal from inside; e. sepal from inside; f. staminode from outside; g. fruit (a-f: Brunig S 17527; g: Rehal 13015). Adema, Leenhouts, Van Welzen — Sapindaceae 641 Fig. 55. Lepisanthes kinabaluensis Leenh. a. Habit; b. fruit; c. female flower; d. pistil; e. petal from in- side; f. stamen; g. seed, longitudinal section (a, b: RSNB 4998; c—g: RSNB 4944). 642 Flora Malesiana ser. I, Vol. 11 (3) (1994) Note — Apparently nearest to L. multijuga, and like that species different from L. amoena mainly by the deeply 3-lobed fruits. 7. Lepisanthes multijuga (Hook. f.) Leenh., Blu- mea 17 (1969) 73. — Nephelium multijuga Hook. f., Trans. Linn. Soc. 23 (1860) 164. — Capura multijuga Hook. f. ex Radlk., Sapind. Holl.-Ind. (1879) 11, nom. inval. — Otophora muitijuga (Hook. f.) Merr., Enum. Born. PI. (1921) 358; Radlk. in Engl., Pflanzenr. 98 (1932) 774. — Type: Motley s.n., Borneo. Otophora tricocca Radlk. in Merr., Pl. Elm. Born. (1929) 174, nom. nud. — Syntypes: E/mer 20010, 20200 (both A, BO, L, M, SING), N Borneo. Otophora imbricata auct. non Blume: Radlk., Sa- pind. Holl.-Ind. (1879) 82, p.p. Tree or shrub, up to 12 m high, dbh up to 10 em. Twigs 1.5—2 cm in diam., blackish to reddish brown, glabrous. Leaves imparipinnate, (15—)30— 40-jugate, 80-90 cm to more than | m long, axial parts fulvous-tomentose, glabrescent; petiole terete to flattened above, 3—6(—8) cm long; pseudo-stip- ules | or 2 pairs, ovate to suborbicular, 2.5—7.5 by 2.5—5 cm, base cordate, apex acute, penni- to reti- nerved, more or less connected with the normal leaflets by intergrades. Leaflets opposite to alter- nate, (sub)sessile, linear-lanceolate, up to 18 by 3 cm, index 6—7, chartaceous, blackish brown to greenish grey above, medium to dark brown be- neath, midrib above densely fulvous-tomentose, beneath very sparsely hairy to glabrous, above densely, beneath more sparsely covered with glan- dular-pitted warts; base rounded to subcordate, oblique; apex tapering acuminate, obtuse to acute; nerves 1—1.5 cm apart, angle to midrib 65—85°, slightly curved, vaguely looped and joined at a dis- tance from the margin. Inflorescences terminal, broad-thyrsoid, 25-30 cm high, subglabrous, sparsely branched; branches obliquely patent, up to 45(—60) cm long, narrowly thyrsoid; cymes scat- tered, short-stalked to sessile, condensed, few-flow- ered; pedicels 0.5—4 mm. Flowers red. Sepals out- side very sparsely appressed short-hairy to gla- brous, sparsely glandular-ciliolate, outer 2 elliptic to ovate, 2.5—3 by 1.8-2.5 mm, inner 3 orbicular, 3-4 by 2.5—4 mm, the latter with a broad petaloid margin. Petals 0.5 mm clawed, rather densely long ciliate, outside sparsely hairy at the base or gla- brous, blade broad-ovate, subtruncate at base, 2 by 2.5 mm; scale slightly bilobed, about 1/3 as high as the blade, slightly recurved, ciliate. Disc gla- brous. Stamens (6—)8; anthers 1.5—1.8 mm, long- ciliate to completely woolly. Ovary 3-celled, deeply grooved, glabrous; style conical. Fruits 3-lobed (if only 1 part is developed style near the base above the scars of the dropped sterile lobes), lobes spread- ing, oblique-ellipsoid, 22 by 13 mm, scabrous, gla- brous, in vivo chocolate-coloured or yellow or or- ange. Seeds hazelnut-shaped, 12 by 9 mm. — Fig. 52g. Distribution — Malesia: Borneo (Sabah, Labuan I): Habitat & Ecology — In primary and secondary forest, level land, at low altitude. Fl. July, Sept., Novy.; fr. Jan., Sept., Nov., Dec. Notes — 1. Hooker f. in Benth. & Hook. f., Gen. Pl. 1 (1862) 405, included Nephelium multijugum in the genus Capura, but without making the com- bination. The name Capura multijuga was first used by Radlkofer, Sapind. Holl.-Ind. (1879) 11, but since it was placed under the genus Otophora, it was not accepted by him, and therefore is invalid. 2. The present species hardly differs from L. amoena except in the deeply 3-lobed fruits, a char- acter, however, which seems to be of importance in this relationship. It shares this character with L. kinabaluensis. Section Pseudotophora Lepisanthes sect. Pseudotophora (Radlk.) Leenh., Blumea 17 (1969) 60, 75 (see there for a complete synonymy). — Otophora subg. vel sect. Pseudophora Blume, Rumphia 3 (1847) 142, nom. inval. — Otophora sect. Pseudotophora Radlk., Sapind. Holl.-Ind (1879) 85. Twigs glabrous or sometimes hairy. Leaves mostly paripinnate, 1—8(—14)-jugate, mostly glabrous; petiole and rachis only very rarely winged. Leaflets nearly always densely fine- ly pitted underneath, exceptionally with pitted warts, often glabrous, midrib beneath an- gular or more rarely rounded. Inflorescences rarely terminal, nearly always axillary, rami- or cauliflorous, solitary or few together, simple or branched, hairy or glabrous. Sepals (3 Adema, Leenhouts, Van Welzen — Sapindaceae 643 or) 4 or 5, outside hairy or glabrous. Petals 4 or 5, outside glabrous or hairy; scale often faint, entire, hairy. Stamens 5—8(—10); anthers hairy or glabrous. Ovary 2- or 3(—4)-celled, mostly glabrous. Fruits rarely lobed, smooth, glabrous, septa mostly interrupted to near- ly fully reduced. Distribution — 2 species, in SE Asia and Malesia as far east as the Moluccas. 8. Lepisanthes bengalan Leenh., Blumea 17 (1969) 75. — Type: Kostermans 4889 (L, SING), NE Borneo. Tree, up to 12 m high, dbh up to 40 cm, or shrub. Twigs 1.2 cm in diam., shining olive-brown, slightly pustular lenticellate, glabrous. Leaves imparipin- nate with a reduced terminal leaflet, 4—6-jugate, glabrous; petiole terete, swollen at base, 4-9 cm long; rachis in the lower part terete, central part angular above, upper part flat with lateral ribs to marginate beneath insertion of leaflets; petiolules 1 mm; pseudo-stipules suborbicular, c. 4 by 5 cm, sessile, cordate at base, rounded at apex, palmati- to retinerved. Leaflets subopposite, oblong, up to 17-19 by 5-7 cm, index c. 3.5, stiff-chartaceous, greyish green above, light- to red-brown beneath, above with scattered gland-warts, beneath densely finely pitted; base slightly oblique, narrowly round- ed; apex obtuse to narrowly rounded; midrib be- neath acute, nerves 1.5—3 cm apart, angle to mid- rib c. 65°, slightly curved, at least the upper ones distinctly looped and joined near the margin. /n- florescences axillary, 30—35 cm long, rachis sub- glabrous, widely branched; branches long, oblique- erect, densely fulvous-puberulous, some branched again; cymes sessile, few- to 1-flowered; pedicels rather thick, 2 mm long. Sepals 5, outer elliptic, 4.5 by 2.5 mm, inner suborbicular, 3.5 by 3 mm, all with petaloid margin, dark red, outside densely fulvous-puberulous, margin densely ciliolate, in- side glabrous. Petals 5, sessile, elliptic, 3 by 2.5 mm, thick-fleshy, outside basally rather densely ap- pressed short fulvous hairy, margin ciliolate; scale a narrow, reflexed, ciliolate strip. Stamens 8; an- thers 1.5 mm, glabrous (in female flowers ciliola- te along the sides). Ovary 3-celled, ciliate along the sides; pistillode densely hairy. /nfructescences and fruits not observed; fruit described as dark purplish. — Fig. 56. Distribution — Malesia: NE Borneo. Habitat & Ecology — On ridges and in forest, on basic loam; altitudes up to 200 m. FI. May; fr. June, Aug. 9. Lepisanthes fruticosa (Roxb.) Leenh., Blumea 17 (1969) 76. — Sapindus fruticosa Roxb., {Hort. Beng. (1814) 29, nom. nud.] FI. Ind. ed. Carey (1832) 283. — Otophora fruticosa (Roxb.) Blume, Rumphia 3 (1847) 142; Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 329; Sapind. Holl.-Ind. (1879) 31; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 174; Backer, Schoolfl. Java (1911) 264; Merr., Fl. Manila (1912) 305; Sp. Blanc. (1918) 239; Enum. Philipp. Flow. Pl. 2 (1923) 500; Radlk. in Engl., Pflanzenr. 98 (1932) 759; Backer & Bakh. f., Fl. Java 2 (1965) 135. — Otolepis fruticosa (Roxb.) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Capura fruticosa (Roxb.) Vidal in Ceron, Cat. Pl. Herb. Manila (1892) 54. — Type: Roxburgh s.n., Hort. Bot. Calcutta (introduced from the Moluccas). Sapindus baccata Blanco, FI. Filip. (1837) 290. — Koelreuteria edulis Blanco, Fl. Filip. ed. 2 (1845) 202, nom. illeg.; ed. 3, 2 (1878) 14. — Otophora blancoi Blume, Rumphia 3 (1847) 142, nom. illeg. — Neotype: Merrill Sp. Blanc. 374 (A, BO, L), Philippines, Palawan. Otophora erythrocalyx Hiern in Hook. f., Fl. Br. India 1 (1875) 680; Radlk. in Engl., Pflanzenr. 98 (1932) 769. — Ofolepis erythrocalyx (Hiern) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Type: Maingay KD 477 (M), Malay Peninsula. Otolepis sessilis King, J. As. Soc. Beng. 65, II (1896) 430; Craib, Fl. Siam. Enum. | (1926) 328; Radlk. in Engl., Pflanzenr. 98 (1932) 763. — Syntypes: King’s collector 2460 (M), 5043 (M drawing), Malay Peninsula. Otophora resecta Radlk., Rec. Bot. Surv. India 3 (1907) 346; Ridley, Fl. Malay Penins. 1 (1922) 495; Hend., Gard. Bull. Str. Settl. 4 (1928) 243; Radlk. in Engl., Pflanzenr. 98 (1932) 766. — Type: Ridley 6948 (M, SING), Malay Peninsu- la. Otophora glandulosa Radlk. [in Merr., Pl. Elm. Born. (1929) 174, nom. nud.] in Engl., Pflan- zenr. 98 (1932) 763. — Type: Elmer 20128 (A, BO, L, M, SING), N Borneo. Otophora acuminata Radlk. in Engl., Pflanzenr. 98 (1932) 769. —Type: Hallier 91] (L, M), W In- donesian Borneo. Otophora latifolia Ridley, Kew Bull. (1933) 190, nom. illeg.; Radlk. in Engl., Pflanzenr. 98 (1934) 1494. — Otophora glandulosa Radlk. ex Rid- ley, Kew Bull. (1933) 490, nom. illeg., non Radlk. (1932). — Type: Creagh s.n. (K), N Borneo. 644 Flora Malesiana ser. I, Vol. 11 (3) (1994) Ses we yi ( Wea — XN \ | a : A | Fig. 56. Lepisanthes bengalan Leenh. a. Leaf; b. detail of lower surface of leaflet (a, b: Kostermans 4889). Adema, Leenhouts, Van Welzen — Sapindaceae 645 For a more complete list of synonyms and refer- ences, see Leenhouts (1969). Shrub or tree, 1.5—10(—15) m high, dbh 2-15 em. Twigs 2.5—20 mm in diam., red when young, later variably brown to silvery grey, smooth or len- ticellate, glabrous or sometimes variably fulvous- hairy and glabrescent. Leaves without or sometimes with a rather strongly reduced terminal leaflet, 1— 8(—14)-jugate, 25 cm to more than | m long, most- ly glabrous; petiole terete or sometimes flattened, 0.5—32 cm long; rachis terete to laterally flattened, in the upper part often marginate to exceptionally narrowly winged; petiolules 0-30 mm; pseudo-stip- ules very rarely absent, mostly persistent, ovate, obovate, elliptic, or orbicular, sometimes very ob- lique, 0.2—10 by 0.1—10 cm, base cordate to ob- tuse, apex obtuse to rounded or exceptionally fur- cate, reti- or palmatinerved, rarely penninerved. Leaflets opposite to alternate, ovate- to obovate- oblong to narrowly lanceolate, 9-40 by 2—12 cm, index 2-9, thin-chartaceous to coriaceous, often greyish above, brown beneath, above with scattered sunken glands, beneath sparsely to densely finely pitted or exceptionally with minute pitted warts on both faces, glabrous or sometimes beneath hairy mainly on the midrib; base oblique or not, subcor- date to acute, mostly attenuate, in oblique leaflets lower half sometimes rounded, upper cuneate; apex obtuse (rarely acute) to acuminate, acumen short, broad, and obtuse to long, slender, and acute, some- times mucronate; midrib beneath acute (to round- ed), nerves 0.8—5 cm apart, angle to midrib 35— 75°, straight to curved, none to all looped and joined. Inflorescences (terminal or) axillary to rami- or cauliflorous, solitary or (if cauliflorous) some- times some together, simple or branched either with some to several ascending long branches from near the base or all over and pyramidal, up to 75 cm long, glabrous; cymes (sub)sessile (rarely up to 1 cm stalked), few- to several-flowered, in the upper part often flowers solitary, sometimes sticky (race ‘glandulosa’); pedicels filiform, 0.3—1(—1.5) cm long. Flowers scentless. Sepals 4—5, outer 2 some- times smaller, elliptic, orbicular, or obovate, 2-4 by 1.5-3 mm, dark red (rarely yellow to white), margin, especially of the inner ones, petaloid, crenulate to fimbriate-ciliolate, glabrous or very sparsely glandular-ciliolate. Petals 4—5, short- clawed, blade broad-ovate or elliptic to obovate, 1.5—3 by 1-2 mm, dark red (rarely yellow to white), glabrous or rarely claw ciliate or outside hairy; ei- ther 2 auricles or | small and reflexed scale, scale mm, yellow to white, glabrous or hairy. Ovary 2- or 3(—4)-celled, glabrous; stigma (sub)sessile, slightly lobed. /nfructescences with patent, most- ly slender, up to 1.5 cm long pedicels. Fruits ovoid, ellipsoid, subglobular, or transversely ellipsoid, rarely distinctly lobed, 1—3 by 0.6—2 by 0.5—2 cm (fresh up to 4 cm in diam.), apparently white when young, dark red to blackish when ripe; fruit-wall thin, apparently fleshy when fresh; septum rarely complete, usually interrupted to (mostly) reduced to a rib all around. Seeds mostly 2, subglobular to semi-ellipsoid, flattened on the axial side, 8—23 by 6-18 by 4-18 mm, hilum orbicular to lanceolate, up to 6 by 3-4 mm. — Fig. 52b. Distribution — Indo-China, Lower Burma, Thai- land, and Malesia: Sumatra (Indragiri, one collec- tion), Malay Peninsula, Borneo (not known from the southern part), Java (only Central Java, few collections, Semarang and Solo, possibly not in- digenous), Philippines, Celebes, Lesser Sunda Is- lands (Dompo I. near Sumbawa: one old collec- tion, wild?), Moluccas (Talaud, Ternate, Bacan, Ambon). Habitat & Ecology — In primary and secondary vegetations, probably mainly in open places in the forest, along the edges, on ridges, along rivers, swamps, and the beach, also in logged areas, aban- doned plantations, and grasslands, on dry to swampy, rich as well as poor, clayey as well as sandy, acid as well as basic soils, from sea level up to 600(—1400) m altitude. Fl., fr. mainly Dec.—May. Fruits sweet, eaten by birds, wild pigs and deer. Uses — In Malaya, the roots are medicinally used. The wood is hard, durable, and heavy and is used for house-building in Malacca. Sometimes cultivated because of its edible fruits. See Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1614: Jansen et al. in Verheij & Coronel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991) 343; Ochse, Indische Vruchten (1927) f.126. Notes — 1. Lepisanthes fruticosa is a rather var- iable species. Though it is impossible to subdivide it into well delimited infraspecific taxa, often some races are locally clearly distinguishable. In the Malay Peninsula there are two races: ‘erythrocalyx’, characterized by distinctly stalked leaflets, S-merous flowers, and completely 3-celled fruits; and ‘resecta’ (including also O. sessilis), with (sub)sessile leaflets, 4-merous flowers, and incompletely 2-celled fruits. In Borneo there are three main races, ‘acumi- nata’, ‘fruticosa’, and ‘glandulosa’, the latter hav- ing 2 ecotypes. Race ‘acuminata’ is characterized by acute leaflets, rami- or cauliflorous inflorescenc- es, 5-merous flowers, completely 3-celled fruits; it is restricted to the Kapuas basin. Race ‘frutico- sa’ has obtuse leaflets, axillary or sometimes ter- minal inflorescences, 4-merous flowers, and very 646 Flora Malesiana ser. I, Vol. 11 (3) (1994) incompletely 2-celled fruits; itis apparently most- ly restricted to N Borneo. Race ‘glandulosa’ is characterized by obtuse leaflets, axillary and rami- or cauliflorous inflorescences, 5-merous flowers, densely short-hairy anthers, and very incomplete- ly 2-celled fruits. The lowland form is rather wide- spread, a mountain form, mainly restricted to Mt Kinabalu, differs in the more slender petiole and rachis (the latter may be narrowly winged), small- er and more caducous pseudo-stipules, narrower leaflets with a longer attenuate apex, and the mid- rib more often rounded beneath. Outside Borneo, ‘fruticosa’ in the same strict sense is known from Java and the Lesser Sunda Islands, and it is identical with some material from the Philippines, Celebes, and the Moluccas. In the Philippines and E Malesia, however, it seems im- possible to distinguish between races, as several characters seem to vary independently. This is true not only of the vegetative parts, but the inflores- cences may be axillary, rami-, or cauliflorous, the flowers are mostly 4-merous, but 5-merous flow- ers may be found in the same inflorescences, and both flower types have a 2-celled, but sometimes a 3-celled pistil. 2. Meijer 4238, the only collection known from Sumatra, seems to belong to the present species, but differs mainly by the narrow (1—1.5 mm wide) wings along petiole and rachis. Sarawak For. Dept. S 18404 is distinctive in its completely 3-merous flowers and glandular pitted warts on both surfac- es of the leaflets; furthermore, it strongly resem- bles ‘acuminata’ in particular and that is why it was included here. Endert 2922 (NE Borneo), ap- parently also the present species, is especially re- markable because of its very large (12 by 12 cm), coriaceous pseudo-stipules. Section Anomotophora Lepisanthes sect. Anomotophora (Radlk.) Leenh., Blumea 17 (1969) 60, 79 (see there for complete synonymy). — Otophora Blume sect. Anomotophora Radlk., Sapind. Holl.- Ind. (1879) 85. Leaves pari- or (more rarely) imparipinnate, 3—6(—13)-jugate, hairy or glabrous, pet- iole and rachis winged. Leaflets densely finely pitted underneath, glabrous or hairy, mid- rib beneath angular or rounded. Inflorescences axillary, rami- or cauliflorous, solitary or fascicled, simple or slightly branched, glabrous. Sepals 4 or 5 (or 6), outside glabrous. Petals 4 or 5 (or 6), outside nearly always glabrous; scale entire to bilobed, glabrous or hairy. Stamens 5-8; anthers (sub)glabrous. Ovary 2- or 3- (or 4-)celled, glabrous. Fruits not or slightly lobed, smooth, glabrous, septa mostly complete. Distribution — 3 species, one in Indo-China, two in W Malesia. 10. Lepisanthes alata (Blume) Leenh., Blumea 17 (1969) 80. — Otophora alata Blume, Rumphia 3 (1847) 145; Schoolfl. Java (1911) 264; Koord., Exk. Fl. Java 2 (1912) 538; Koord. & Valeton, Atlas 1 (1913) t. 129; Radlk. in Engl., Pflanzenr. 98 (1932) 768; Backer & Bakh. f., FI. Java 2 (1965) 135. — Capura alata (Blume) Teijsm. & Binn., Cat. Hort. Bog. (1866) 214. — Otolepis alata (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Lectotype (Leenhouts 1969): Korthals s.n. (L), S Borneo. Otophora edulis C.E.C. Fischer, Kew Bull. (1932) 178; Radlk. in Engl., Pflanzenr. 98 (1934) 1493. — Type: Orolfo 1319 (K), N Borneo. Tree, up to 15 m high, dbh up to 30 cm, or shrub. Twigs 1.2—0.8(-1.5) cm in diam., blackish brown to grey, glabrous. Leaves paripinnate, 3—5(—13)- jugate, 20-45 cm (to more than | m long), gla- brous; petiole angular above, exceptionally flat to rounded, 1.5—7.5(—25) cm long; wings of petiole and rachis 3—8 mm wide; pseudo-stipules oblique- ovate, 1—3(—8.5) by 0.8—2.2(—5) cm, base deeply cordate, apex obtuse, sometimes acuminate, pen- ninerved. Leaflets opposite (to alternate), sessile or petiolules up to 2 mm long, lanceolate (excep- tionally linear-lanceolate or oblong to obovate- oblong, lower sometimes ovate-lanceolate), 10- 20(-45) by 1.8-4(-8) cm, index (3—)4—7(-14), thin- chartaceous, above greyish to blackish brown, be- neath brown to greyish green; base mostly hardly oblique, acute and sometimes tapering, in oblique leaflets one half or both halves rounded; apex ta- pering long acuminate, acute; midrib beneath most- Adema, Leenhouts, Van Welzen — Sapindaceae ly acute, nerves 1—1.5(—2) cm apart, angle to mid- rib 55—70°, curved, at least those in the upper half of the leaflet looped and joined near the margin. Inflorescence often drooping, in vivo often purple to reddish brown, apparently mostly unisexual, though male and female ones on the same trees; male: usually axillary, narrowly thyrsoid, un- branched, 20—25(—40) cm long, with scattered, ses- sile fascicles of 3—5(—7) flowers on 2—7 mm long pedicels; female: either in the lower leaf-axils, or rami- or cauliflorous, at least near the base with some branches about equalling the rachis, up to 45 cm long, with scattered, mostly solitary flowers on 6—13 mm long pedicels. Flowers in vivo dark wine- red to purple. Sepals obovate-orbicular, 2.5—4 by 2—3 mm, outer slightly smaller than inner, inner partly petaloid with crenulate margin, all sparsely glandular-ciliolate. Petals sessile (in L. edulis claw 0.5 mm), sparsely, partly glandular, ciliolate, gla- brous or rarely sparsely appressed short-hairy out- side in the basal half (L. edulis); blade suborbicu- lar, up to 4 by 3 mm (L. edulis 2 by 2.5 mm), scale erect, slightly hood-shaped, 1/3-1/5 as long as blade, glabrous (L. edulis densely ciliolate). Disc glabrous, pink. Stamens 8; anthers 1.8 mm, sub- glabrous, yellow with white pollen. Ovary ellip- soid, 3- (or 4-)celled, pale mauve; style very short, stigma dome-shaped to flat, slightly 3-lobed, white. Infructescences with patent, up to 2 cm long, slen- der pedicels. Fruits shortly stipitate, trigonous- obovoid, 2.5—4 by 2.2—3 cm, apiculate, apparently dark brownish purple to nearly black when ripe, pulp rather thick, fleshy, white. Seeds ellipsoid, up to 2.5 by 1.5 cm, hilum rhomboid, 6 by 5 mm. — Fig. 52e. Distribution — Malesia: Malay Peninsula (Jo- hore, one collection), Borneo, and Java (probably only naturalized). Habitat & Ecology — In and along forests, on river banks, etc.; on clay; altitude up to 100(—450) m. Fl. Aug.—May; fr. Aug.—Apr. Ants may live un- der the stipules [see Schimper, Pfl. Geogr. Physi- ol. Grundlage (1898) 165-166, f. 87]. Uses — The fruits and possibly also the seeds are eaten; in Sumatra, Java, and Borneo sometimes grown as a fruit tree. See Heyne, Nutt. Pl. Indon. ed. 3 (1950) 990; Jansen et al. in Verheij & Coro- nel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991) 343. Note — Lepisanthes alata is as a whole a rather uniform species. Only a few specimens from Bor- neo deviate considerably from the mean in some characters. Among these Endert 1702 and Jaheri 1693 are both characterized by rather large (15— 23 by 5—7.5 cm), relatively wide (index c. 3), dis- tinctly obovate leaflets with rather spaced nerves 647 (1.5—2 cm apart). Hotta 12660 from Brunei differs in its many-jugate leaf (13 pairs of leaflets), peti- ole and rachis rounded or flat above, very large stipules (8.5 by 5 cm), and very long and narrow leaflets (33 by 2.2 cm, index c.14). Orolfo 1319 from Sabah, the type of Otophora edulis, differs in its rather narrow leaflets (index 8) with nearly perpendicular nerves, petals which are distinctly clawed and are hairy outside and on the scale, and especially in its large fruits (c. 4 by 3 cm) with a thick, fleshy pulp (the wall of the ovary is already exceptionally thick). However, one should keep in mind that fleshy fruits are nearly always collected unripe and Orolfo 1319 may be exceptional in this respect; furthermore, from the label it is not clear whether it was taken from a wild specimen or pos- sibly some cultivar. 11. Lepisanthes ramiflora (Radlk.) Leenh., Blu- mea 17 (1969) 81. — Otophora ramiflora Radlk., Sapind. Holl.-Ind. (1879) 32, 85; in Engl., Pflanzenr. 98 (1932) 758. — Otolepis ramiflora (Radlk.) Kuntze, Rev. Gen. Pl. 1 (1891) 144. — Lectotype (Leenhouts 1969): Beccari PB 364 (FI), Sarawak. Small tree up to 22.5 m high and 2 cm in diam., or shrub. Twigs 5—7 mm in diam., yellowish to red- dish brown, densely pustular-lenticellate, sparsely hirsute, early glabrescent. Leaves pari- (rarely im- pari-)pinnate, (3—)4—6-jugate, 40-SO cm; petiole and rachis sparsely hairy to subglabrous; petiole terete, 6-18 cm long; wings of petiole and rachis up to 5 mm wide; pseudo-stipules ovate, 0.8—3.5 by 0.5—2 cm, base cordate, apex acute to rounded, penninerved. Leaflets (sub)opposite, sessile, oblong (sometimes slightly obovate), up to 33 by 10.5 cm, index 3—5, chartaceous, sometimes slightly bullate, grey or brown above to greenish beneath, midrib beneath at least near the base sparsely hirsute; base oblique, narrowed, cordate; apex tapering acumi- nate, acute; midrib rounded beneath, nerves 1—2.5 cm apart, angle to midrib 65—70°, more or less curved, at least those in the upper half distinctly looped and joined at some distance from the mar- gin. Inflorescences cauliflorous, several to many fascicled on knobs on the stem, simple or hardly branched, racemoid, c. 1.5 cm long, flowers few; pedicels patent, 1.5—5 mm long, slender. Flowers 4-merous. Sepals pale pink, outer 2 elliptic, 2.5 by 2 mm, with petaloid margin, inner broad-ovate, 3 by 2 mm, for the greater part petaloid. Petals white, sparsely long ciliate near the base, 0.4 mm long clawed, blade semi-transversely ellipsoid, 1 by 1.8 mm; scale broad, rounded to bilobed, c. 0.3—0.5 as high as the blade, slightly hairy. Dise pale yellow, 648 Flora Malesiana ser. I, Vol. 11 (3) (1994) glabrous. Stamens 5-7; anthers | mm, glabrous, cream-coloured. Ovary suborbicular, flattened, 2- celled, pale yellowish; style short, stigma dome- shaped, slightly bilobed, white. Fruits transverse- ly ellipsoid, slightly bilobed, 1 by 1.5 cm, crimson to dark purple, incompletely 2-celled. Seeds ellip- soid. Distribution — Malesia: Borneo (Sarawak: around Kuching). Habitat & Ecology — In primary and secondary forest on sandstone; altitude 0-100 m. FI. Jan.— Feb., July—Sept., Nov.; fr. Jan., Oct., Nov. Uses — The fruits are eaten. Note — Closest to L. amplifolia (Pierre) Leenh., from S Vietnam, which has 5-merous flowers (ac- cording to Gagnepain), infructescences up to 40 cm long, widely branched, and completely 2-celled fruits. Subgenus Erioglossum Lepisanthes subg. Erioglossum (Blume) Leenh., Blumea 17 (1969) 60, 81 (see there for complete synonymy). — Erioglossum Blume. Leaves paripinnate, neither petiole nor rachis winged, without pseudo-stipules. Leaf- lets (sub)opposite, nervation open. Petals longer than sepals, outside subglabrous, scale crested. Disc interrupted, glabrous. Stamens 8; filaments longer than anthers. Ovary 3- celled, very sparsely to densely hairy. Fruits parted, (sub)glabrous, septa complete. Distribution — 2 species, in SE Asia, Malesia, and NW Australia. 12. Lepisanthes membranifolia (Radlk.) Radlk., Bot. Jahrb. 56 (1920) 252; in Engl., Pflanzenr. 98 (1932) 746; Leenh., Blumea 17 (1969) 81. — Erioglossum membranifolium Radlk., Sa- pind. Holl.-Ind. (1879) 17, 55. — Lectotype (Leenh. 1969): Beccari PP 317 (FI, M), New Guinea. Treelet. Twigs terete, c. 6 mm in diam., dark brown, rather densely short-hairy, glabrescent. Leaves 3—5-jugate, the axial parts rather densely fulvous-hairy; petiole terete, at base only slightly swollen, 11—14 cm long; petiolules slightly grooved above, 2-5 mm long. Leaflets oblong to elliptic, 18-26 by 7-13 cm, index 2.5—3, herbaceous, on both sides hairy on midrib and nerves, beneath moreover sparsely so all over the surface; base slightly oblique, rounded to subcordate, sometimes attenuate; apex tapering obtuse-acuminate; midrib prominent above, more strongly so and rounded beneath, nerves 1.5—3 cm apart, angle to midrib 50—55°, straight to slightly curved, the upper ones looped and joined near the margin, slightly sunk- en above, prominulous beneath, veins and veinlets finely reticulate, veins more or less scalariform, above hardly visible, prominulous beneath. Inflo- rescences axillary to pseudoterminal, thyrsoid, sometimes with a few short branches, 1.5—-5 cm long, dense, with sessile, c. 7-flowered cymes, densely puberulous; bracts deltoid, | mm; pedi- cels 1.5 mm. Flowers white. Sepals ciliate, inside glabrous, outer ovate, 2.5 by 1.8 mm, inner obo- vate to suborbicular, up to 3 by 2.5 mm, margin slightly petaloid. Petals 4, claw 1 mm, blade ellip- tic-ovate, 2 by 1.5 mm, outside slightly hairy just above the claw, woolly ciliate below insertion of scale, inside glabrous; scale folded, entire to emar- ginate, long-ciliate, doubled by the crest. Stamens: filaments 1.8 mm, rather thick, sparsely long-hairy in the upper half; anthers ellipsoid, | mm, connec- tive rather narrow, glabrous. Pistil with some scat- tered hairs; ovary obcordate, 3-lobed; style short, thick and terete; stigma cap-shaped; pistillode minute. Fruits not observed. Distribution — Malesia: New Guinea (Vogelkop Peninsula, three collections). Habitat & Ecology — Lowland forest. Fl. Aug. Notes — |. The position of the present species is not clear. Radlkofer originally described it in Eri- oglossum, but later referred it to Lepisanthes on the strength of anatomical characters. The ques- tion will be difficult to answer as long as the fruits are unknown; the obcordate pistil, however, looks more like that of Erioglossum or Aphania than of Lepisanthes s. str. 2. It is possible that the vegetative parts as de- scribed are not representative for the mature tree; Kostermans 2893 was collected from an un- branched tree only 2 m high, which does not ap- pear more ‘juvenile’ than the two collections made by Beccari; they all look like suckers. 13. Lepisanthes rubiginosa (Roxb.) Leenh., Blu- mea 17 (1969) 82. — Sapindus rubiginosa Adema, Leenhouts, Van Welzen — Sapindaceae Roxb., Pl. Corom. | (1796) 44, t. 62. — Moulin- sia rubiginosa (Roxb.) G. Don, Gen. Hist. 1 (1831) 667. — Erioglossum rubiginosum (Roxb.) Blume, Rumphia 3 (1847) 118; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 498; Craib, Fl. Siam. Enum. | (1926) 325; Radlk. in Engl., Pflanzenr. 98 (1932) 693; Kanehira & Hatusi- ma, Bot. Mag. Tokyo 57 (1943) 76; Steenis, FL. Sch. Indon. (1949) 253; Gagnep. in Fl. Indo- Chine, Suppl. 1 (1950) 933, f. 116, Backer & Bakh. f., Fl. Java 2 (1965) 134. — Type: Roxb. Pl. Corom. | (1796) t. 62. Sapindus edulis Blume, Cat. (1823) 64, nom. il- leg., non Aiton (1789). — Erioglossum edule (Blume) Blume, Bijdr. (1825) 229; Rumphia 3 (1847) 119, t. 166; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 154; Lecomte in FI. Indo- Chine | (1912) 1019; Koord. & Valeton, Atlas 1 (1913) f. 88; Merr., Sp. Blanc. (1918) 238; Ridley, Fl. Malay Penins. | (1922) 492, f. 49; Hend., Gard. Bull. Str. Settl. 4 (1928) 243. — Uitenia edulis (Blume) Steud., Nomencl. ed. 2, 2 (1841) 776, nom. inval. — Erioglossum edule (Blume) Blume var. genuina Blume ex Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 156, nom. illeg. — Type: Reinwardt 823 (L), Java (cultivated). Sapindus fraxinifolia DC., Prod. 1 (1824) 608. — Erioglossum edule (Blume) Blume var. frax- inifolia (DC.) Blume, Rumphia 3 (1847) 120. — Type: Riedlé s.n. (L, P), Timor. Erioglossum edule (Blume) Blume var. album Blume, Rumphia 3 (1847) 119. — Syntypes: Blume 120, s.n. (both L), W Java. Erioglossum edule (Blume) Blume var. subcorym- bosum Blume, Rumphia 3 (1847) 119. — Eri- oglossum edule (Blume) Blume var. corym- bosum Teijsm. & Binn., Cat. Hort. Bog. (1866) 215, in errore? — Syntype: Blume s.n. (L), W Java. Lepisanthes hirta Ridley, J. Fed. Mal. St. Mus. 10 (1920) 132; Radlk. in Engl., Pflanzenr. 98 (1932) 753. — Type: Ridley FMS 13161 (K, SING), Malay Peninsula. For complete synonymy see Leenhouts (1969). Shrub or small tree, up to 16 m high, dbh up to 28 cm (Forman 239 from N Celebes reported to be 30 m high with a diameter of 60 cm; from India also reported to be a fairly big tree). Indumentum ferrugineous to fulvous, sometimes and in some parts silvery-grey. Branchlets terete, grooved, c. 5(— 15) mm in diam., densely short-hairy when young. Leaves (2—)3—6(—9)-jugate, often with a pseudo- terminal leaflet, velvety when young; petiole about terete, 7.5—12(—20) cm long, densely short-hairy, 649 late glabrescent; petiolules up to 5(—10) mm long. Leaflets elliptic to lanceolate, (4.5—)6.5—18(—25) by (2—)3.5—8.5(—11) cm, stiff chartaceous, above greyish green to grey, beneath yellowish green to reddish brown, on both sides shortly and densely hairy on midrib and nerves, sparsely so all over the surface mainly beneath (velvety on the touch), more or less glabrescent; base rounded to broadly cuneate; apex obtuse to acute or acuminate, often mucronulate; midrib prominent above, rather strong and rounded beneath, nerves 8—12(—16) on either side, ascending to spreading, nearly straight to slightly curved, bent at the margin, not distinctly joined but for few upper ones, hardly prominent above, slightly so beneath, veins and veinlets in- conspicuous above, beneath densely reticulate-sca- lariform and prominulous. /nflorescences 25—35(— 50) cm long, densely ferrugineous tomentose; branches often long, ascending, spicate; cymes nearly sessile to short-stalked, glomerulous to dis- tinctly branched, few to several-flowered; bracts and bracteoles small, subulate; pedicels 1—2(—5) mm. Flowers sweet-scented. Sepals orbicular- ovate, slightly concave, green when fresh, margin sometimes petaloid, ciliate, inside glabrous or with inner three 1.8—2.8 by 2-3 mm, obtuse. Petals 4 (or 5), claw 0.5—1 mm, blade 24 by 1.5—2.2, crenu- late in upper half, white to yellowish when fresh, long ciliate in upper part of the claw and — sparse- ly — in the narrowed lower part of the blade, out- side with a few hairs at the base; scale c. 1.5-3 mm long, quadrangular to + bilobed, slightly nar- rowed from base to apex, bearded, the appendage deeply bilobed, the lobes sometimes also bilobed, hairy; the abaxial petals shorter and narrower than the adaxial ones, and with a shorter scale. Stamens longest abaxially; filaments flattened, long white hairy, in male flowers (1.5—)3—5 mm long, in fe- male 1.5 mm; anthers oblong-ovate, 0.8 mm, gla- brous. Ovary 3-lobed, 1.2—1.8 by 2—2.2 mm, dense- ly appressed-hairy; style cylindric, 2.2 mm long, bent near the obscurely 3-lobed apex, sparsely ap- pressed-hairy in the lower 2/3; pistilode in female flowers c. 0.8 mm high, long hairy. Fruits 1-, 2-, or 3-lobed, lobes spreading, 8—13 by 7-8 mm, faint- ly carinate, dark purple to nearly black when ripe, subglabrous; endocarp thin but hard and tough, gla- brous. Seeds oblong-ellipsoid, 9-11 by 4 by 4mm, hilum basal, small. — Fig. 52c. Distribution — Continental SE Asia from north- ern India to Indo-China and SE China (Kwangtung, Hainan), Malesia and NW Australia (York Sound, Brunswick Bay). Habitat & Ecology — Under seasonal as well as everwet conditions, preferably on periodically dry, 650 rarely on marshy, fertile as well as sterile soils, both on heavy clay, on sand, and on limestone. A sub- stage tree or shrub of more open vegetation: in de- ciduous forests (common in the teak forests of C and E Java), in young secondary forests, shrubland, etc., along forest edges, roadsides, river banks, along the inner side of the mangrove; up to 300(— 1200) m altitude. FIl., fr. Jan.—Dec. [in the Malay Peninsula twice a year after dry periods, accord- ing to Corner, Wayside Trees (1940) 587, t. 177]. Flowers much visited by Xylocarpa sp. For galls see Docters van Leeuwen, Zoocecidia (1926) 335, f. 604-608. Uses — In India, where the trees seem to be a better size, the timber is said to be valuable, but in Malesia it is used only for firewood and sometimes (Java) for rice-pounders and tool-handles. A de- coction of roots and leaves, sometimes also of fruits and seeds, is used medicinally against fever. The young leaves are eaten as a vegetable, and the as- tringent but sweet fruits are relished as a titbit, mainly by children. For further details see Burkill, Flora Malesiana ser. I, Vol. 11 (3) (1994) Dict. Econ. Prod. Malay Penins. (1935) 938; Heyne, Nutt. Pl. Indon. ed. 3 (1950) 989; Ochse & Bakh., Ind. Groenten (1931) 648, f. 396. Chromosomes — 2n = 26: Mehra et al., Silvae Gen. 21 (1972) 96-102. Notes — 1. The inflorescences in the upper leaf axils are sometimes paired in which case the up- per is the stronger one. 2. According to Corner (1940) the basal part of the inflorescence and of its branches bore female flowers which opened first, the more apical parts male flowers, which opened later. This is not con- firmed by the study of numerous herbarium speci- mens. The inflorescences seem to be nearly exclu- sively either female, or male; in old inflorescences either (nearly) all flowers have fallen (probably male inflorescences) or there are a number of young fruits all over the inflorescence (female inflores- cences; note that the infructescence, depicted by Corner, is also covered with fruits all over). Whether all inflorescences of a tree are the same, needs to be studied in the field. Subgenus Aphania Lepisanthes subg. Aphania (Blume) Leenh., Blumea 17 (1969) 60, 83 (see there for com- plete synonymy). — Aphania Blume. Leaves paripinnate, sometimes simple; petiole and/or rachis exceptionally winged, sometimes with pseudo-stipules. Leaflets mostly + opposite, nervation closed. Petals as long as the sepals, outside glabrous or partly sericeous, scale not crested. Disc mostly complete, glabrous. Stamens 5—7(—9); filaments as long as or longer than anthers. Ovary 2- (or 3-)celled, glabrous or sparsely pilose at the style base. Fruits lobed, glabrous, septa not interrupted. Distribution — As the genus; 3 species. 14. Lepisanthes dictyophylla (Radlk.) Leenh., Blumea 17 (1969) 83. — Cupaniopsis dictyo- phylla Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 359. — Aphania dictyophylla (Radlk.) Radlk., Bot. Jahrb. 56 (1920) 268; in Engl., Pflanzenr. 98 (1932) 713. — Type: Sayer s.n. (M), Papua New Guinea. Shrub or liana, glabrous except the fulvous hairy buds and bracts. Twigs terete, c. 2.5 mm in diam., light grey-brown to ash-grey, smooth or slightly pustular-lenticellate. Leaves 3—4-jugate; petiole terete, 3-5 cm long, slightly swollen at base; ra- chis terete to angular above; petiolules narrowly grooved above, 2—5 mm long. Leaflets + opposite, ovate-oblong, 6—-10.5 by 2.24.2 cm, stiff-charta- ceous, greyish green above, the same or dark brown beneath, minutely pellucid-dotted; base broadly cuneate and attenuate to rounded, equal-sided; apex obtuse to tapering acuminate, acumen short, broad, and obtuse; midrib prominulous and acute above, prominent and angular beneath, nerves 0.5—1 cm apart, angle to midrib almost 90°, almost straight, looped and joined at some distance from the mar- gin, equally prominulous on both surfaces, vena- tion finely reticulate, equally prominulous on both surfaces. Inflorescences solitary, simple, 3.5—-18 cm long; peduncle short, cymes distant, short-stalked to sessile, 1—5-flowered; pedicels up to 2 mm long. Sepals 5, sparsely ciliolate partly with glandular hairs, further glabrous or outside very sparsely puberulous, outer 2 distinctly smaller, ovate, 1.8 by 1.2 mm, inner broad-elliptic to orbicular, 2.5 Adema, Leenhouts, Van Welzen — Sapindaceae by 1.8 mm, thinner towards the crenulate margin. Petals 5, subsessile, oblong-elliptic, up to 3 by 2 mm, fairly densely ciliolate; scale small, biparted, villous. Disc of 5, + separate, erect, glabrous lobes. Stamens 5, filaments partly hairy; anthers ovate, | mm long, glabrous. Ovary 2-celled, glabrous; style short; stigma slightly bilobed. Fruits not observed. Distribution — Malesia: E New Guinea. Habitat & Ecology — Understorey of rain for- est; altitude 60 m. Fl. Aug.—Oct. Note — Closely related to L. senegalensis. 15. Lepisanthes mixta Leenh., Blumea 17 (1969) 83, f. 2. — Type: Docters van Leeuwen 11326 (K, L), Mamberano River, New Guinea. Shrub, glabrous except for the inflorescences. Twigs terete, 4-8 mm in diam., blackish, verrucu- lous by small, orbicular lenticels. Leaves sometimes with pseudoterminal leaflet, 3—6-jugate; petiole semiterete, 10-20 cm, swollen at base, narrowly winged at least in the upper part, wings if com- plete broadened at the base into a pair of up to 3 cm wide, semiorbicular pseudo-stipules, swollen at base; rachis flattened above, up to 5 mm wide winged. Leaflets + opposite, (sub)sessile with a swollen base, lanceolate, 17.5—30 by 3.5—5 cm, thin-chartaceous, yellowish to greyish green on either side; base equal-sided, cuneate to subcor- date; apex long-tapering, obtuse; midrib prominu- lous above, prominent and acute beneath, nerves 1—3.5 cm apart, angle to midrib 55—60°, straight to slightly curved, looped and joined at some distance from the margin, equally prominulous on both sur- faces, some intercalary veins strongly developed, venation rather coarsely reticulate, equally prominulous on both surfaces or more distinct be- neath. Inflorescences solitary, slender pyramidal thyrsoid, 25-30 cm long, minutely short fulvous hairy; peduncle up to 15 cm long, rather stout, branches patent, up to 6 cm, racemous, with many short-stalked, several-flowered cymes; pedicels 2— 3 mm long, slender. Flowers (female not observed) white. Sepals 5, outer 1 or 2 smaller, ovate, 1.2— 1.8 by 1—-1.5 mm, with some scattered glands along the margin, inner orbicular-obovate, 2—2.5 by 2— 2.2 mm, partly petaloid, ciliolate partly with glan- dular hairs. Petals (3—)5, short-clawed, oblong-el- liptic, 2 by | mm, sparsely ciliolate partly with glan- dular hairs; scale small, bilobed. Disc complete, flat, more or less distinctly lobed. Stamens 5, an- thers ovate, 1—1.5 mm, glabrous. Pistillode 2-mer- ous, glabrous. Fruits 2-lobed, not stipitate, red, parts globular, 7.5 mm in diam. Distribution — Malesia: W New Guinea (two collections seen). Habitat & Ecology — In forest up to 60 m alti- 651 tude. Fl. Aug., Nov.; fr. Aug. Note — This remarkable species has the fruits of subg. Aphania, but the habit of subg. Otophora because of its winged petiole and rachis. The lat- ter character is most conspicuous in the type spec- imen in which the broad wings are widened at base into a kind of pseudo-stipule; the other specimen (Aet 679, Babo on McCluer Bay) has very narrow wings and lacks the pseudo-stipules. In all other characters the two agree completely. 16. Lepisanthes senegalensis (Poir.) Leenh., Blu- mea 17 (1969) 85. — Sapindus senegalensis Juss. ex Poir., Encycl. 6 (1805) 666. — Apha- nia senegalensis (Poir.) Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 238; in Engl., Pflanzenr. 98 (1932) 703. — Type: Adanson & Geoffroi f. s.n. (P, Herb. Jussieu 11386), Senegal. Aphania montana Blume, Bijdr. (1825) 236; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 158: Atlas 1 (1913) t. 132; Radlk. in Engl., Pflan- zenr. 98 (1932) 711; Backer & Bakh. f., Fl. Java 2 (1965) 134. — Sapindus montana (Blume) Blume, Rumphia 3 (1847) 97. — Type: Anonymous s.n. (L, ?Blume), W Java. Euphoria verticillata Lindl., Bot. Reg. (1827) t. 1059. nom. illeg. — Nephelium verticillatum (Lindl.) G. Don, Gen. Hist. 1 (1831) 670. — Scytalia verticillata (Lindl1.) Roxb., Hort. Beng. (1814) 29; Fl. Ind. ed. Carey (1832) 273. — Didymococcus verticillatus (Lindl.) Blume, Rumphia 3 (1847) 103. — Sapindus verticil- latus (Lindl.) Kurz, Rep. Pegu, App. A (1875) 38. — Type: Roxburgh s.n., cult. Hort. Calcut- ta. Scytalia danura Roxb., [Hort. Beng. (1814) 29, nom. nud.] Fl. Ind. ed. Carey (1832) 274. — Aphania danura (Roxb.) Radlk., Sitzungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 8 (1878) 238; in Engl., Pflanzenr. 98 (1932) 716. — Type: Roxburgh s.n., India. Sapindus cuspidata Blume, Rumphia 3 (1847) 98. — Aphania cuspidata (Blume) Radlk., Sit- zungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 8 (1878) 238; in Engl., Pflan- zenr. 98 (1932) 706. — Type: Zippelius 210b (L), SW New Guinea. Otophora paucijuga Hiern in Hook. f., Fl. Br. India 1 (1875) 680. — Aphania paucijuga (Hiern) Radlk., Sitzungsber. Math.-Phys. Cl. KO6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 239; Hend., Gard. Bull. Str. Settl. 4 (1928) 243; Radlk. in Engl., Pflanzenr. 98 (1932) 708. — Type: Maingay 1529 (= KD 462) (K, L), Ma- lacca. 652 Aphania sphaerococca Radlk., [Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 238, nom. nud.] Sapind. Holl.-Ind. (1879) 7, 21; in Engl., Pflanzenr. 98 (1932) 705. — Syntypes: Beccari PP 503, PP 908 (both FI, M), New Guinea. Aphania longipes Radlk., [Sitzungsber. Math.- Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 8 (1878) 239, nom. nud.] Sapind. Holl.-Ind. (1879) 68; in Engl., Pflanzenr. 98 (1932) 705. — Type: Teijsmann HB 7872 (L, M), New Guinea. Aphania philippinensis Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 60; in Engl., Pflanzenr. 98 (1932) 709. — Syntypes: Ahern 216 p.p. (M), Luzon; Warburg 14597, Philippines, Sulu Arch. Aphania boerlagei Valeton, Ic. Bog. 2 (1906) 281, t. 185; Radlk. in Engl., Pflanzenr. 98 (1932) 706. — Syntypes: Hort. Bot. Bog. 43, 43a (= Teijsmann HB 12763) (BO, L,M), SW Celebes. Aphania angustifolia Radlk. in Elmer, Leafl. Philipp. Bot. 1 (1907) 209; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 499; Radlk. in Engl., Pflanzenr. 98 (1932) 710. — Type: Elmer 7330 (BO, K, L, M), Philippines, Leyte. Hydnocarpus tamiana Pulle, Nova Guinea 8 (1912) 671; cf. Sleumer in Fl. Males. I, 5 (1954) 33. — Type: Gjellerup 262 (L), NW New Guinea. Aphania loheri Radlk., Philipp. J. Sc., Bot. 8 (1914) 452; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 499; Radlk. in Engl., Pflanzenr. 98 (1932) 710. — Type: Loher 5874 (K, L, M), Philippines, Luzon. Aphania macrophylla Radlk. in Fedde, Rep. 18 (1922) 332; in Engl., Pflanzenr. 98 (1932) 713. — Type: Koorders 18015 (L, M), N Celebes. Aphania dasypetala Radlk. in Fedde, Rep. 18 (1922) 333; in Engl., Pflanzenr. 98 (1932) 714. — Type: Native collector 709 (BO, K, L, M), Sarawak. Aphania fascicularis Radlk. in Fedde, Rep. 18 (1922) 334; in Engl., Pflanzenr. 98 (1932) 714. — Type: Warburg 18168 (M), Moluccas, Ba- can. Aphania masakapu Melch., Notizbl. Berlin-Dahl. 10 (1928) 277; Radlk. in Engl., Pflanzenr. 98 (1934) 1491. — Type: Peekel 941, Bismarck Archipelago, New Ireland. Treelet or shrub, 0.5—24 m high, dbh up to 44 cm, exceptionally a liana (Peekel 1083, New Ire- land); young parts densely to sparsely, appressed- ly, shortly fulvous to ferrugineous hairy. Twigs terete, 2-6 mm in diam., greyish to brown or black, smooth to verruculose by many small, orbicular Flora Malesiana ser. I, Vol. 11 (3) (1994) lenticels. Leaves sometimes with a pseudotermi- nal leaflet, 1—4(—6)-jugate, or sometimes (some or all leaves, in the former case either scattered among pinnate leaves or only near the inflorescence) sim- ple; lower pair of leaflets sometimes attached near or at the base, smaller and more caducous than other ones; in vivo young leaves pinkish with reddish petioles; petiole flattened to terete, 0-14 cm long, often much swollen at base, sometimes lenticel- late like the twig; rachis terete to carinate above; petiolules terete, slightly flattened, or grooved, swollen, 2-15 cm long. Leaflets elliptic to lanceo- late, widest about or below the middle, 7—35 by 2— 15 cm (simple leaves up to 60 by 18 cm), charta- ceous to subcoriaceous, greyish green (rarely ol- ive or brown) above, greyish or yellowish green to light brown beneath, densely to sparsely minutely pellucid-dotted; base acute to obtuse, more or less attenuate (in sessile simple leaves subcordate, ob- lique or not); apex obtuse to acuminate, mucronu- late, acumen up to 5 cm long, obtuse to acute; mid- rib faint to prominulous above, beneath prominent, rounded, nerves 7—20 per side, 1—3 (in the large simple leaves up to 4) cm apart, angle to midrib 30-90°, straight to curved, looped and joined at some distance from the margin, prominulous on both surfaces, between every two nerves usually | or 2 intercalary veins stronger developed, nerves and veins rather densely reticulate, prominulous on both surfaces, especially beneath. Inflorescences hairy to subglabrous, few to several fascicled, sim- ple or sparsely, mostly slender branched, up to 40 cm long racemes or narrow thyrses, or sometimes a single, widely though sparsely branched, up to 60 cm long thyrse with a strong peduncle and ra- chis; cymes distant, short-stalked, 3-flowered; pedi- cels 1—-4(-8) mm long. Flowers slightly fragrant, female ones apparently developing first. Sepals 5 (rarely 4), in vivo mostly dark red, concave, outer 2 oblong-ovate to orbicular, rounded at apex, 1-3 by 0.8-1.2 mm, sometimes with a narrow, peta- loid, crenulate margin, sparsely ciliolate, inner 3 up to 4 by 3 mm, mostly with a broad, petaloid, crenulate to fimbriate, ciliolate margin. Petals 5 (rarely 4), in vivo creamish or greenish white, im- bricate, short-clawed to sessile, the blade elliptic to oblong, widest below to about the middle, 2.5— 4.5 by 1-2 mm, entire, rounded, membranous but often thickened towards the centre and the base, ciliate at least near the base, furthermore mostly glabrous, sometimes more or less sericeous out- side; scale minute to 2/5 of the length of the blade, simple to deeply bilobed (rarely divided into two minute auricles), sparsely ciliate to long-hairy (ex- ceptionally glabrous). Disc entire (+ interrupted if one of the petals is suppressed), annular to saucer- Adema, Leenhouts, Van Welzen — Sapindaceae shaped. Stamens 5—7(—9); filaments flattened or terete, narrowed from base to apex, 1—3.5 mm, in old flowers apparently longer than in mature ones, in vivo white, woolly in the lower 2/3, rarely sub- glabrous; anthers elliptic to ovate, 1—1.8 mm long, in vivo when young light yellow, old purplish brown, connective mostly apiculate. Ovary 2- (or 3-)lobed, 1.5—3 mm, hardly to up to 1.5 mm stipi- tate, mostly with some stiff hairs at the style base and in the basal part of the style; style cylindric, 1-2 mm long, straight to slightly curved; stigma sometimes decurrent halfway down the style, in vivo purple; pistillode often long pilose in the api- cal part. Fruits 2-lobed (often only 1 part devel- oped), sessile or up to 2 mm stipitate, lobes short- ellipsoid to globular, 8—15 by 5—15 mm, smooth, when ripe in vivo dark red to black; endocarp egg- shell-like, light brown. Seeds ovoid to globular, + attenuate towards the hilum, 7-8 by 6—7 mm, testa shining brown to black, hilum circular. — Fig. 52a. Distribution — Tropical Africa, Madagascar, Sri Lanka and the SW Deccan Peninsula, NE India, E Pakistan, Burma to Indo-China and Hainan, the Andamans and Nicobars, and Malesia: all regions except the Lesser Sunda Islands, in Borneo known only from Sarawak, Brunei, and Sabah. Habitat & Ecology — Under periodically dry as well as everwet conditions, in dry as well as marshy, fertile to rather sterile, well to badly aerated, acid to basic soils, clay or sand as well as marl and cor- al limestone rocks, mostly at low altitudes: 0-200(— 500, rarely up to 1400) m. An understorey tree of rather light forests — in Borneo mixed Dipterocarp forest, in Java teak forest — and secondary forests, of river banks and coastal rocks. Fl. mainly July— Nov.; fr. mainly Jan.—Apr. Uses — Wood hard, heavy, and durable, used in the Malay Peninsula for house-posts. The roots are used in medicine in Mindanao. From the leaves a shampoo is made (Mindanao). The fruits are edi- ble. See Burkill, Dict. Econ. Prod. Malay Penins. (1935) 190. Chromosomes — 2n = 28: Mangenot & Mang- enot, Rev. Cyt. Biol. Veg. 25 (1962) 411-447; 2n = 30: Saerkar et al. in Love, Taxon 31 (1982) 578. 653 Notes — The Malesian material of this widely distributed species 1s rather variable, as is clear from the above description. This variability is mainly gradual, and sometimes clinal. The main race (Su- matra, Java, Borneo, Philippines, S Celebes, Moluc- cas) is ‘montana’ with obtuse, rarely acuminate leaflets and short-stipitate fruits with short ellip- soid to subglobular lobes, grading to the east into ‘cuspidata’ with more acuminate leaflets and ses- sile fruits with more globular lobes. In the Philip- pines the leaflets tend to become narrower (the series philippinensis—angustifolia—loheri), or the leaves have more jugae, up to 3 pairs of leaflets being normal. Unifoliolate leaves characterize ‘macrophylla’ from N Celebes and ‘Hydnocarpus tamiana’ from NW New Guinea. These may not be real local races, but plants flowering when young, both being taken from shrubs only. The flowers of ‘montana’ are 5-merous, but some spec- imens from Sumatra have flowers with 5 and with 4 petals in the same inflorescence. These speci- mens have also some hairs on the outside of the petals, whereas the petals in ‘montana’ are gla- brous. In that way these Sumatran specimens form a transition to the races of continental SE Asia, as well as to the only other Malesian race which de- serves to be mentioned, which is made up of ‘pau- cijuga’ from the Malay Peninsula and ‘dasypeta- la’ from Borneo. This race differs from ‘montana’ by often having 4 petals that are slightly (‘pauci- juga’) to rather densely (‘dasypetala’) sericeous outside, by the longer, stipitate ovary, and by the inconspicuous venation, the nerves are more widely spreading and more distinctly looped and joined at a slightly greater distance from the margin. All specimens from the Malay Peninsula represent ‘paucijuga’, whereas both ‘dasypetala’ and ‘mon- tana’ are known from Borneo, there being appar- ently clearly distinct. It is not possible, however, to separate ‘montana’ and ‘paucijuga—dasypeta- la’ as subspecies or varieties, as they are connect- ed via Thailand and Indo-China by a chain of races. See for a more comprehensive discussion Leenhouts (1969: 55-59). LITCHI (P.W. Leenhouts) Litchi Sonn., Voy. Ind. Or. Chine 2 (1782) 230, t. 129; Radlk. in Engl., Pflanzenr. 98 (1932) 914; Leenh., Blumea 24 (1978) 398. — Scytalia Gaertn., Fruct. 1 (1788) 197, nom. illeg. — Euphoria Comm. ex Juss., Gen. (1789) 247, nom. illeg. — Type spe- cies: Litchi chinensis Sonn. 654 Flora Malesiana ser. I, Vol. 11 (3) (1994) Trees, monoecious. Indumentum of solitary, simple or 2-branched hairs; no glandular scales. Leaves spirally arranged or sometimes partly, nearly opposite, especially towards the inflorescence, paripinnate, 1—4- (?sometimes 5-)jugate; pseudo-stipules absent; peti- ole and rachis not winged. Leaflets mostly (sub)opposite, beneath dull, margin entire to undulate, mainly in the upper part. Inflorescences terminal and axillary, thyrsoid; bracts triangular, 0.5-2 mm long. Flowers actinomorphic, unisexual. Calyx 4- or 5-merous, outside and inside densely appressed short-hairy, the tube cupular, the lobes to 0.3-0.5 of the length of the tube, apert in bud, equal, not petaloid. Petals absent. Disc annular, small, without appendages. Stamens 6-11, far exserted in male flowers, short in female and in seemingly bisexual, but functionally male flowers; filaments variably hairy; anthers gla- brous. Pistil short-stalked; ovary 2- (?or 3-)celled; style terminal, shorter than the ovary; stigma consisting of 2 (or 3?) long, spreading to recoiled lobes. Ovules | per cell, basal. Fruits nearly always only 1 cell developed, hardly stipitate, ellipsoid, ovoid, or globular, not winged, indehiscent(?), glabrous, spiny to scaly; pericarp thin, coriaceous to rather hard, endocarp glabrous. Seeds glabrous, partly or completely enveloped by an entire, translucent, juicy arillode. — Fig. 57. Distribution — Monotypic. Note — The names Scyralia and Euphoria are illegitimate as in both cases Litchi was included. Dimocarpus, however, is, contrary to the opinion of most authors, not consid- ered synonymous, and hence not superfluous (see under that genus). Litchi chinensis Sonn., Voy. Ind. Or. Chine 2 (1782) 230, t. 129; Lecomte in Fl. Indo-Chine 1 (1912) 1047, f. 131; Groff, Lychee and Lon- gan (1921); Merr., Enum. Philipp. Flow. Pl. 2 (1923) 504; Radlk. in Engl., Pflanzenr. 98 (1932) 917, f. 21; Gagnep. in FI. Indo-Chine, Suppl. 1 (1950) 966, f. 121: 8-11; Backer & Bakh. f., Fl. Java 2 (1965) 137; Poilane, J. Agr. Trop. Bot. Appl. 12 (1967) 541; Leenh., Blu- mea 24 (1978) 398. — Sapindus edulis Aiton, Hort. Kew. 2 (1789) 36. — Euphoria lit-chi Des- font., Tableau (1804) 135, nom. illeg.; Blume, Bijdr. (1825) 233. — Nephelium lit-chi (Des- font.) Cambess., Mém. Mus. Nat. Hist. Nat. Paris 18 (1829) 30, nom. illeg.; Wight, Ic. 1 (1838) t. 43; Blume, Rumphia 3 (1847) 106; Hiern in Hook. f., Fl. Br. India 1 (1875) 687; Watt, Dict. 5 (1891) 346; Backer, Fl. Batavia | (1907) 348; Dunn & Tutcher, Kew Bull. add. ser. 10 (1912) 67; Ridley, Dispersal (1930) 487; Corner, Wayside Trees (1940) 592; K. Ramesh Rao, Ind. Woods 2 (1963) 225. — Type: Son- nerat 1062 (G, P), China. Euphoria lit-chi Desfont. var. undulata Blume, Bijdr. (1825) 233, nom. illeg. — Type: Blume s.n. (L), Java. Euphoria didyma Blanco, Fl. Filip. (1837) 288, nom. illeg.; ed. 2 (1845) 201; ed. 3, 2 (1878) 10. — Nephelium didyma (Blanco) Craib, FI. Siam. Enum. | (1926) 329, pro basionym, spec- imens excluded. — Neotype (Leenhouts 1978): BS 17429 (L, P), Philippines, Luzon. Litchi philippinensis Radlk. [ex Whitford, Philipp. J. Sc., Suppl. 1 (1906) 637, 645, 647, nom. nud. | Philipp. J. Sc., Bot. 8 (1914) 458; Schneider, Bull. For. Philipp. 14 (1916) 147; Groff, Ly- chee and Longan (1921) 34, f. 2; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 504; Radlk. in Engl., Pflanzenr. 98 (1932) 915. — Litchi philippin- ensis Radlk. f. genuina Radlk., Philipp. J. Sc., Bot. 8 (1914) 459, nom. illeg.; in Engl., Pflan- zenr. 98 (1932) 916. — Lectotype (Leenhouts 1978): FB 2995 (BO, L, M), Philippines, Lu- zon. Litchi philippinensis Radlk. f. mindanaensis Radlk., Philipp. J. Sc., Bot. 8 (1914) 459; in Engl., Pflanzenr. 98 (1932) 916. — Type: Elm- er 13270 (BO, FI, L, M, P, U), Philippines, Mindanao. Litchi chinensis Sonn. f. glomeriflora Radlk. in Engl., Pflanzenr. 98 (1932) 919. — Lectotype (Leenhouts 1978): Blume s.n. (L), Java. Branchlets terete, striate, smooth, or densely lenticellate. Petiole terete to semiterete, swollen and slightly hollowed at base, 1.2—7 cm long, early gla- brescent to glabrous, often pustular lenticellate; pet- iolules above deeply grooved, swollen toward the base. Leaflets smooth, above shining, glabrous to beneath fairly densely minutely appressed-hairy; Adema, Leenhouts, Van Welzen — Sapindaceae base equal-sided to sometimes oblique, acute, more or less attenuate; margin slightly recurved; apex mostly tapering (or more abruptly), short to long, obtuse- to acute-acuminate, sometimes rounded to slightly emarginate; midrib narrowly grooved above, nerves straight to curved, often slightly wavy or zigzagging, looped and joined near the margin or not, inconspicuous on both sides, veins coarse- ly reticulate, veinlets minutely tessellate; domatia absent. /nflorescences ferrugineous-strigose. Flow- ers greenish white or yellowish, fragrant. Stamens: filaments filiform; anthers elliptic, apiculate to emarginate at apex, c. | mm long. Pistil 1.5—1 mm stipitate; ovary with spreading lobes, densely warty; style terete. Seeds ellipsoid, c. 2 by 1.5 cm, testa shining (blackish) brown, hilum basal, circular, 6— 7 mm diam., arillode bluish white or light yellow to pinkish, up to 5 mm thick when fresh. Distribution — See subspecies. Notes — 1. Though being one of the Sapinda- ceous genera best known to the general public, to the taxonomist Litchi remains a genus full of mys- tery regarding its history, phytogeography, and tax- onomy. The written history of Litchi chinensis goes back to c. 100 B.C. when the emperor Wu Ti of the Han Dynasty tried in vain to introduce it from north- ern Indo-China to Central E China. About the 8th century many varieties were widely cultivated in the southeastern provinces of China, probably for at least three centuries. Culture concentrated until the end of the 18th century in the coastal districts of the provinces of Kwangtung and Fukien, appar- ently at some time also in some more southern prov- inces, especially Yunnan, and probably in north- ern Indo-China (for history see Groff 1921). It seems astonishing that the culture of this highly prized fruit tree remained restricted for such a long time to a relatively restricted region, although the Chinese were for centuries the most active colo- nists of the Far East — though mainly as traders, far less as farmers. One reason may be that the seeds of Litchi are viable for a few days only outside the fresh fruits; another, that the propagation and cul- ture of this species requires great skill, the species having very special demands as to climate and soil. As in so many cases of old economic plants it is difficult to tell whether wild-growing specimens are really indigenous or naturalized. However, the restricted region where it has been in cultivation for such a long time very probably includes its orig- inal area. Its climatologic requirements give fur- ther indications where to look for its original area. The culture of Litchi in SE China is restricted by frost and drought, so that its natural area was prob- ably a region with a somewhat warmer and wetter climate. On the other hand, subsp. chinensis re- 655 quires a drier and cooler season, otherwise it will hardly flower, so it is adapted to a region with a monsoon climate. All this agrees well with Poi- lane (1967) who reported Litchi chinensis as being fairly common growing in the wild in N Vietnam and Cambodia. 2. It is not clear whether the fruits of L. chinen- sis are finally dehiscent or not. They show a dis- tinct suture, at least on the inside. It seems proba- ble that at least the fruits of subsp. philippinensis are normally dehiscent, and possibly those of the other subspecies would behave in the same way when overripe. 3. Nephelium lit-chi is illegitimate as it refers back to Litchi chinensis via Sapindus edulis Aiton, and the epithet chinensis should have been used. KEY TO THE SUBSPECIES la. Twigs slender, to 3.5 mm in diam. Cymules up to, 5 mmulongs staked axe |. oe 2 b. Twigs thick, to 7 mm in diam. Cymules ses- WME GEWGS sass sene bn ac b. subsp. javensis 2a. Leaves 2—4-jugate. Stamens mostly 6(—10). Fruits nearly smooth or rarely with up to | mm high, acute, pyramidal warts a. subsp. chinensis b. Leaves 1- or 2- (or 3-)jugate. Stamens mostly 7 (or 6). Fruits with up to 3 mm high, acute, pyramidal warts... c¢. subsp. philippinensis a. subsp. chinensis — Litchi chinensis Sonn. — Euphoria lit-chi Desfont. — Nephelium lit-chi Cambess. Tree, up to 35 m high, dbh up to | m. Branch- lets 2.5-3.5 mm in diam., greyish brown, when young with appressed, short, 2-branched, brown hairs, early glabrescent. Axillary buds | per leaf axil. Leaves 2—4-, exceptionally 5-jugate; petiole 0.8—1 mm in diam.; rachis above flat or grooved and with a fine central rib (lower) to carinate (up- per interjugae), beneath rounded; petiolules 3-8 mm long. Leaflets elliptic, or (when relatively broad) obovate, (3—)8—11(—16) by 1.8—4 cm, in- dex 2-4, chartaceous to coriaceous, mostly rather stiff; base if oblique broadest at acroscopic side; midrib beneath prominent and rounded, nerves 0.5— 1 cm apart, angle to midrib 50—70°, veinlets clear. Inflorescences 15—30 cm long with few long erec- to-patent branches, these sparsely branched main- ly in their upper half; cymules erecto-patent, up to 5 mm long stalked, lax, S—12-flowered; bracts 0.5— 2 mm long; pedicels slender, 2—3(—4) mm long. Calyx 4- (or 5-)merous, c. 1.5 by 2 mm. Disc var- iably hairy to glabrous. Stamens 6(—10); filaments 2.5 mm long, rather densely hairy all over. Ovary 656 Flora Malesiana ser. I, Vol. 11 (3) (1994) JH von 73 Fig. 57. Litchi chinensis Sonn. Habit, flowers and fruits. — Subsp. philippinensis (Radlk.) Leenh. a. Habit: b. male flower; c. female flower; f. fruit. — Subsp. chinensis. d. Fruit; e. ibid., longitudinal section (a, b: FB 30836; c: BS 17429; d, e: L, alc. 8524). 2 by 5mm; style 1 mm. Fruits c. 3.5 by 3. cm, bright red to purplish when ripe, nearly smooth or scaly to densely set with up to | mm high, flat, pyrami- dal, acute warts. — Fig. 57d, e. Distribution — Probably originating from the northern part of the Indo-Chinese Peninsula or from SE China, now widely cultivated mainly in sub- tropical regions; in Malesia possibly indigenous in the Malay Peninsula, Borneo and the Philippines, rarely cultivated (Malay Peninsula, Philippines), exceptionally naturalized and apparently hardly ever fruiting. Habitat — Prefers a hot and wet climate with at least a short cool and dry season, but no frost. It Adema, Leenhouts, Van Welzen — Sapindaceae seems to favour compact, moist, and fertile soils, and is mostly grown at low altitudes on river banks, along dykes, between ditches or ponds, etc. The ‘mountain lychee’, which may represent the origi- nal form, however, is grown on dry land in the hills. Ecology — The fruits are eaten by bats and par- rots. Uses — Cultivated for its fruits (Litchi, Lychee), mainly in SE China, Florida, Hawaii, and South Africa, from where they are exported, fresh, dried, or canned, to other parts of the world. The wood is also highly prized, reported to be hard, durable, and taking a fine polish (for a description of the timber, see p. 428). See Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1546; Groff, Ding & Groff, Lingn. Agr. Rev. 2 (1924) 34; Menzel in Verheij & Coronel (eds.), Plant Res. SE Asia (PROSEA Handb. 2, Edible fruits and nuts) (1991) 146; Ochse, Indische Vruchten (1927) 246, f. 119. Chromosomes — 2n = 28: Chaudhuri, Curr. Sci. 9 (1940) 416; 2n = 30: Mehra et al., Silvae Gen. 21 (1972) 96-102. Note — The many races of cultivated lychee are arranged into two groups, the “water lychee’ and the ‘mountain lychee’. The former contains most (and the best) varieties, is cultivated in the low- lands, and has mostly nearly smooth fruits. The ‘mountain lychee’ is mainly used as a stock; it is grown in hilly regions, and has smaller and more prickly fruits. The latter may come nearer to the original wild form. b. subsp. javensis Leenh., Blumea 24 (1978 ) 401. — Litchi chinensis Sonn., auct. javan. — Eu- phoria lit-chi Desfont. var. undulata Blume — Litchi chinensis Sonn. f. glomeriflora Radlk. Branchlets up to 7 mm in diam., greyish brown, glabrous. Axillary buds often 2 per axil, the upper of which more developed. Leaves 2-4-jugate; pet- iole mostly c. 1.5 mm in diam.; rachis often flat- tened as well beneath as above; petiolules 3-8 mm long. Leaflets (narrowly) elliptic, 5.5S—10(—15) by 1.5—3.5(—5) cm, index 2.5—5, chartaceous to co- riaceous; base if oblique broadest at acroscopic side; margin tending towards being ‘shouldered’ towards the apex; midrib nearly always slender and angular beneath, nerves up to 1.8 cm apart, angle to midrib 50—70°, reticulations above always clear, beneath nearly always less conspicuous to some- times invisible. Inflorescences 12-20 cm long with few long erecto-patent, spicoid, thick; branches usually unbranched; flowers in sessile clusters; pedicels up to | mm long. Calyx 4- (rarely 5-)mer- ous, 0.8—1 by 2—2.5 mm. Disc sparsely hairy. Sta- mens 7—11; filaments 2 mm long, sparsely hairy in lower half only. Ovary 1 by 2 mm, according to 657 some authors exceptionally 3-celled; style 0.8 mm. Fruits as in subsp. chinensis. Distribution — Malesia: Java; known only cul- tivated from a few localities, apparently rare. Habitat & Ecology — Unlike subsp. chinensis this subspecies seems to thrive well under tropical everwet conditions. It is cultivated at altitudes up to c. 250 m. FI. Feb.; fr. Mar. Uses — The fruits are probably comparable in quality to those of subsp. chinensis. Notes — 1. Subsp. javensis was always consid- ered to be introduced from China, and that is why no special attention was paid to it. Only Radlkofer described it as a separate form but with the remark ‘status morbosus.’. It differs distinctly from subsp. chinensis, however, not only morphologically but also by being adapted to an everwet tropical cli- mate. To my knowledge it cannot be linked to any of the Chinese cultivars; the spicoid inflorescenc- es are especially distinctive. It is known only as a rarely cultivated fruit tree of West Java, at least from the 18th century onwards (J. Burman 1/9 in L), but it may be indigenous. 2. Litchi chinensis subsp. javensis Leenh. is based upon Litchi chinensis f. glomeriflora Radlk. c. subsp. philippinensis (Radlk.) Leenh., Blumea 19 (1971) 129. — Euphoria didyma Blanco — Litchi sp. nov. Merr., Philipp. J. Sc., Suppl. 1 (1906) 87. — Litchi philippinensis Radlk. — Nephelium didymum Craib. Tree, up to 35 m high, dbh up to 1.25 m. Branch- lets 1.5—3 mm in diam., (yellowish to silvery-)grey, glabrous. Axillary buds | per leaf axil. Leaves 1— 3-, mostly 2-jugate; petiole c. 1 mm in diam.; ra- chis terete or above slightly flattened and carinate; petiolules 4-10 mm long. Leaflets elliptic. (7—)8- 11(—14) by 2.8-4.5(—8) cm, index 2.5-3, stiff-co- riaceous; base if oblique broadest at basiscopic side; midrib beneath prominent and rounded, nerves 0.8— 1.5 cm apart, angle to midrib 60-85°, veinlets above invisible, beneath inconspicuous to clear (mainly as the surface over the veins and veinlets is not dull but sometimes even shining; see, however, f. mindanaensis). Inflorescences 6—14 cm long, of- ten tufted (terminal one with 2 slightly smaller ones at the same level from the upper leaf axils, axillary ones with up to 4 smaller ones or with 2 strong branches from the base); branches few, (erecto-)pat- ent, sparsely rebranched; cymules up to 5 mm long stalked, lax, 5—15-flowered; bracts 0.5—1 mm long; pedicels very slender, 2-5 mm long, in female flow- ers thickening shortly after fertilization. Calyx 4- or 5-merous, |.8—2 by 2-3 mm. Disc hairy. Sta- mens (6 or) 7; filaments 24 mm long, densely hairy all over. Ovary 1 by 3.5 mm; style 1 mm. Fruits 658 Flora Malesiana ser. I, Vol. 11 (3) (1994) 2.5—3 by 2—2.5 cm, densely set with acute, pyram- idal warts c. 4 mm diam. and up to 3 mm high. — Fig. 57a-c, f. Distribution — Malesia: Philippines (Luzon, Sibuyan, Samar, Mindanao), SE New Guinea (Northern Prov.: Hoogland 3684). Habitat & Ecology — Primary and secondary forests from sea level to 500 m altitude. Fl. Feb.— May; fr. Apr., July. Wood anatomy — See Desch, Mal. For. Rec. 15 (1954) 532. Uses — The wood is hard and durable and is used for house poles. Notes — 1. The above description has been based exclusively upon L. philippinensis f. genuina Radlk. The only collection known from Mindanao, Elmer 13270, was described by Radlkofer as a separate form, f. mindanaensis. It differs in a few characters, as follows: Branchlets 3—4 mm in diam. Leaves 1- or 2-jugate; petiole 1-2 mm in diam.; in- dex of leaflets 2—2.5, veins and veinlets beneath hardly visible, the whole lower surface being dull- papillose. As only this one specimen is known, and only 8 specimens of ‘f. genuina’, it is impossible to decide on the taxonomic status of this form. 2. Subsp. philippinensis is not cultivated, and this makes it more probable that the only speci- men known from New Guinea, collected from a 20 m high tree in the rain forest, was really wild. 3. Subsp. philippinensis is closely allied to sub- sp. chinensis; its small, prickly fruits are quite sim- ilar to those of the ‘mountain lychee’. Its arillode is said to be short, covering only part of the seed. It is not certain, however, whether the fruits from which this character was noted were fully mature, and in subsp. chinensis, too, the arillode develops only very late. 4. Contrary to the opinion of Merrill (1918) and of most subsequent authors but in accordance with Radlkofer (1932), I am of the opinion that Eupho- ria didyma Blanco is synonymous with Litchi chin- ensis subsp. philippinensis and not with Dimocar- pus longan var. malesianus. My arguments are: 1) in the description no mention is made of the corol- la which in Philippine Dimocarpus longan is very conspicuous, and 2) Dimocarpus longan var. male- sianus was already described by Blanco under the name of Euphoria lit-chi. MISCHOCARPUS (R.W.J.M. van der Ham) Mischocarpus Blume, Bijdr. (1825) 238, nom. cons.; Rumphia 3 (1849) 166; Radlk. in Engl., Pflanzenr. 98 (1933) 1288; R.W. Ham, Blumea 23 (1977) 251; S.T. Reynolds in Fl. Austral. (1985) 94; Yap in Tree Fl. Malaya 4 (1989) 449. — Cupania sect. Mischocarpus Migq., Fl. Ind. Bat. 1, 2 (1859) 566. — Type species: Mischocarpus sundaicus Blume. Pedicellia Lour., Fl. Coch. (1790) 655, nom. rejic. (see under dubious names in R.W. Ham 1977). — Type species: Pedicellia oppositifolia Lour. Mischocodon Radlk., Bot. Jahrb. 50 (1913) 79; in Engl., Pflanzenr. 98 (1933) 1327. — Type species: Mischocodon reticulatus Radlk. Shrubs or (large) trees (with a slender unbranched stem), monoecious (or dioecious in M. reticulatus?). Indumentum of mostly appressed, single hairs only. Leaves paripinnate, leaflets larger towards the top, 1-6-jugate, no pseudo-stipules or wings; petiole subterete (to dorsoventrally flattened); petiolules present. Leaflets alternate to subopposite; base symmetrical (to slightly oblique); margin entire, flat (to revolute), apex (emarginate to) rounded to acuminate, often mucronulate; lower surface not papillate, red glands absent, domatia often present in axils of nerves; the latter looped and joined in the upper part (to indistinctly so in the lower 1/2—3/4); intersecondary nerves present, sometimes indis- tinct; veins forming a very regular reticulate, dense pattern, + prominent on both surfac- es. Inflorescences pseudoterminal, axillary, and ramiflorous thyrses, nearly always branch- ing, branches erect to spreading; cymules 1—7(—10)-flowered; pedicels 1—3(—5) mm long; Adema, Leenhouts, Van Welzen — Sapindaceae 659 bracts triangular to lanceolate. Flowers unisexual. Calyx spreading or cup-shaped, 5- (or 6-)lobed, connate for up to 2/3, persistent in fruit; lobes subequal, (slightly imbricate at base). Petals 0-5, minute to slightly longer than the calyx, apert, clawed or not; blade basally usually with 2 auricles or scales without crest. Disc uninterrupted (or interrupt- ed), annular or cup-shaped. Stamens (5—)8(—9), exserted; filaments thread-like; anthers basifixed, dehiscence latrorse or latero-introrse, (connective with a lighter coloured wart at the top). Pistil (2- or) 3- (or 4-)locular; ovules one per locule; ovary stipitate or subses- sile; style apical, the upper part split into 3 + recurved stigmatic lobes. Fruits a loculicid- ally dehiscent capsule, nearly always stipitate (only up to | mm high in M. paradoxus), not lobed, not winged, the cells about equally developed but the ovules abortive in (1 or) 2 cells, red, smooth; stipe hollow; valves thin to almost woody, usually shrivelled after dehiscence; pericarp slightly fleshy; endocarp sclerenchymatic, complete and lining valves and distal parts of the septa, or incomplete, only along the sutures, glabrous or variably hairy. Seeds (hanging by the pseudo-funicle): testa shining, chestnut-brown, (nearly) com- pletely covered by a thin-fleshy, translucent arillode, with an abaxial pseudo-funicle de- scending into the stipe; cotyledons superposed. — Figs. 58, 59. Distribution — 15 species from SE Asia, throughout Malesia to Australia; in Malesia 9 species, 1 undescribed. Habitat & Ecology — Shrubs and understorey trees of primary and sometimes sec- ondary (rain) forest, from sea level (rocky sea coasts; saltwater creeks) up to 3000 m altitude. The fruits may be attractive to animals because of the showy, slightly fleshy arillode. KEY TO. THE SPECIES la. Midrib above either not visible, hidden in a narrow groove, or rarely visible as a Meny SiCAGC ial SHALY SIMIKEM AINE cs... mtn st « Fc Stans state Soa Mes as 2 b. Midrib above clearly visible, but sometimes covered with hairs.............. 4 2a. Nerves 12~30 per side. Calyx connate for 1/3. Petals not clawed. Disc annular or slightly cup-shaped. Stipe of fruit 0-6(-15) mm high...................... 3 b. Nerves 6-15 per side. Calyx connate for 1/2—2/3. Petals clawed. Disc cup-shaped. NepevG. fruits tO = 25, mihi hii eas. 2 MMe Sone alee ee 5. M. pyriformis 3a. Inflorescences axillary, pseudoterminal, and ramiflorous. Calyx connate for 1/4. Petals 0—2. Disc not interrupted, c. 1.5 mm in diam., puberulous. Stipe of fruit 1.5— 6(—15) mm high; endocarp pubescent to densely woolly....... 2. M. largifolius b. Inflorescences ramiflorous (rarely also axillary). Calyx connate for 1/3. Petals (3—)5. Disc often interrupted, 0.75—1 mm in diam., glabrous or locally puberulous. Fruit not (or hardly) stipitate; endocarp glabrous............. 3. M. paradoxus 4a. Midrib above more or less hairy in at least some leaflets ................ fa SD b. Midrib above always glabrous, rarely subglabrous (see M. reticulatus)........ 6 Sa. Petiolules 4—-18(—25) mm long. Cymules stalked, short sericeous. Calyx connate for 1/3—2/3. Petals (0) 1—S. Filaments hairy. Fruits outside glabrous 5. M. pyriformis b. Petiolules 2-5 mm long. Cymules almost sessile, long-hairy. Calyx connate for 1/3. Petals absent. Filaments glabrous. Fruit outside densely ferrugineous 1. M. lachnocarpus 660 Flora Malesiana ser. I, Vol. 11 (3) (1994) 10a. lla. 1. Mischocarpus a. Inflorescences ramiflorous (rarely also axillary) Inflorescences axillary and pseudoterminal ...........--.- +00 ++ ee esters 10 4. Inflorescence branches more than 11 cm long, branching. Fruit without a stipe or StiperalmiostySi mami lag ln Se Ae Peer ere eRe OS ete Pa Re 8 Inflorescence branches less than 11 cm long, branching or not. Fruit stipe at least 4 mm high a. Fruits not stipitate or stipe at most | mm high. Calyx outside sparsely pubescent, inside puberulous near the base, often as a row of short hairs. Disc often irregularly deeply lobed to interrupted. Arillode without pseudo-funicle ... 3. M. paradoxus Fruit stipe 1-3 mm high. Calyx outside glabrous, inside glabrous or with a few hairs. Disc slightly lobed. Arillode with short pseudo-funicle 9. Mischocarpus prob. spec. nov. _ Inflorescence axes up to 10 cm long, not or very sparsely branched. Calyx hardly connate. Petals (2—)5. Filaments sparsely hairy or glabrous 8. M. triqueter Inflorescence axes up to 5 cm long, branching. Calyx connate for 1/2-3/4. Petals absent. Filaments glabrous. ..........5.0222 220s ese see eee 6. M. reticulatus Midrib above prominent, angular to rounded. Calyx connate for 1/3 (if connate for more than 1/4, then petals absent and filaments glabrous). Petals well-developed but not clawed, or reduced to absent. Disc annular, 1-2(—3) mm in diam. Filaments hairy onelabrous tales. 2t0Neh). ier 2a aya re Pei ter eee ie lacs ieee 11 Midrib above a prominulous to flattened slender line only. Calyx connate for 1/3- 2/3. Petals 1-5, well developed, clawed. Disc cup-shaped, 0.5—1 mm in diam. Fila- ments + hairy 5. M. pyriformis Leaflets (1 or) 2-6 per side; midrib above angular (to locally to rarely completely rounded): reticulation usually coarse. Calyx outside + puberulous all over, hardly connate (rarely for 1/4). Petals 0-5. Filaments usually puberulous at least at the base (to glabrous); anthers glabrous or + hairy 4. M. pentapetalus _ Leaflets 1-3 (or 4) per side; midrib above rounded, locally (to completely) angular; reticulation usually fine and dense. Calyx outside + hairy in the lower half only, connate for 1/4—1/3. Petals 0 (rarely up to 3 in Peninsular Malaysia). Filaments glabrous; anthers glabrous .......--.--.. +--+ sseeeee Been: 7. M. sundaicus lachnocarpus (F. Muell.) Ham 1977): Dallachy s.n., 14 Apr. 1864 (MEL Radlk., Sapind. Holl.-Ind. (1879) 43; Sitzungs- ber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 536, 647; Domin, Bibl. Bot. 22 (1927) 910; C.T. White, Contr. Arnold Ar- bor. 4 (1933) 63; Radlk. in Engl., Pflanzenr. 98 (1933) 1304; Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 523; R.W. Ham, Blumea 23 (1977) 268; S.T. Reynolds in Fl. Austral. 25 (1985) 96, f. 21a, b, map 123. — Ratonia lach- nocarpa F. Muell., Fragm. Phyt. Austral. 4 (1864) 157; E.M. Bailey, Queensl. Fl. 1 (1899) 296; Queensl. Agr. J. 5 (1899) 396. — Cupania lachnocarpa F. Muell. [Fragm. Phyt. Austral. 4 (1864) 157, nom. inval.] Fragm. Phyt. Austral. 5 (1865) 6; 9 (1875) 91. — Lectotype (R.W. holo, sh. 56003), Australia. Shrub or tree, up to 20 m high; densely ferrugi- neous pubescent to puberulous when young (to sub- glabrous). Twigs 2-4 mm in diam. Leaves |- or 2- (or 3-)jugate; petiole 1—4.5(—7) cm long; petiolules 2-5 mm long. Leaflets elliptic to obovate, 4-14(— 17) by 1.8-6.5 cm, index 1.5—3.2, pergamentaceous to coriaceous, (bullate); base acute (to obtuse); above puberulous to pubescent on the midrib (and nerves to the entire upper side laxly pubescent), beneath pubescent mainly on the midrib and nerves and along the margin, domatia absent or inconspic- uously in the axils of the nerves; midrib above prominent, rounded; nerves 10—14 per side, above Adema, Leenhouts, Van Welzen — Sapindaceae 1cm 661 11cm Fig. 58. Mischocarpus Blume. Fruits and embryos. — M. lachnocarpus (FP. Muell.) Radlk. a. Fruit; b. embryo. — M. largifolius Radlk. c. Fruit. — M. paradoxus Radlk. d. Fruit. — M. pentapetalus (Roxb.) Radlk. e. Fruit. — M. pyriformis (F. Muell.) Radlk. subsp. papuanus (Radlk.) R.W. Ham. f. Fruit. — M. sundaicus Blume. g. Fruit. — M. triqueter Radlk. h. Fruit; i. embryo (a: MEL 56004; b: Dallachy s.n.; C: Schodde & Craven 3963; d: Forbes 309; e: PNH 34128, f: Chalmers s.n.; g: FB 24765; h: BS 42010; 1: McGregor 271). prominulous, beneath prominent; veins rather dense, above prominulous, beneath prominent, becoming more distinct towards the margin. /nflo- rescences axillary and pseudoterminal, up to 14cm long; axes branched; cymules almost sessile, dense- ly ferrugineous pubescent; bracts triangular, up to 2 by | mm, outside densely ferrugineous pubes- cent, inside nearly glabrous; pedicels up to | mm long. Calyx hardly connate; lobes triangular, c. 1.5 by 1 mm, subcoriaceous, outside pubescent to vil- lous, inside glabrous or with a few hairs near the base. Petals 0. Disc annular, up to 1.5 mm in diam., puberulous. Stamens 8, up to 2 mm long; filaments glabrous; anthers papillate or not, glabrous (or pu- bescent). Pisti/ rather densely velutinous. Fruits 12— 16(—20) mm high, densely ferrugineous-villous to velutinous, (partly glabrescent); stipe 3—7 by c. | mm, seed-bearing part globose to ellipsoid to obovoid, triangular in cross section, apiculate for 1—2 mm, inside laxly pubescent to villous, mainly along the sutures; endocarp completely sclerenchy- matic. Seeds globose, c. 7 mm long; cotyledons about equal, superposed. — Fig. 58a, b. Distribution — Australia (N and E Queensland, NE New South Wales) and Malesia: New Guinea (Aru Islands and S and E Papua New Guinea up to Normanby I.). Habitat & Ecology — Primary and secondary rain forest, also along streams in open savannah, up to 1300 m altitude. Fl. Dec.—July; fr. Mar.—Aug. 2. Mischocarpus largifolius Radlk., Bot. Jahrb. 56 (1920) 304, f. 4g—j; in Engl., Pflanzenr. 98 (1933) 1308, f. 38g—j; R.W. Ham, Blumea 23 (1977) 269. — Type: Ledermann 12693 (B+ holo; M), Papua New Guinea. Tree up to 33 m high, dbh up to | m; puberu- lous, early glabrescent; buttresses usually present, up to 1 m high, 2 m spreading, and 3 cm thick; bark grey to darkbrown (to white), smooth. 7wigs 3-10 mm in diam., sometimes grooved. Leaves |— 662 4-jugate; petiole 4-20 cm long; petiolules 4-15 mm long. Leaflets ovate to elliptic, 7-38 by 2-16 cm, index 2—3, pergamentaceous to coriaceous, (+ bul- late); base rounded or obtuse (to acute), domatia absent or inconspicuously in the axils of the nerves below; midrib not visible above and hidden in a narrow groove, (or a very slender and sharp, sunk- en line); nerves 12—30 per side, usually very regu- lar, moderately curved; veins usually very regular- ly reticulate, above prominent to indistinct, beneath prominent. Inflorescences axillary, pseudoterminal, and ramiflorous on branches up to 12 mm in diam., up to 32 cm long; axes branched; bracts up to 2 mm long. Calyx connate for up to 25%; lobes tri- angular, up to | by 1 mm, subcoriaceous to some- what fleshy, outside sparsely, inside more densely pubescent, near the base with a dense row of hairs. Petals 0-2, elliptic, up to 1 by 0.5 mm, not clawed, inside pubescent, with two hairy auricles at the base. Disc annular, c. 1.5 mm in diam., puberu- lous. Pistil densely short sericeous. Fruits 14—28(— 40) mm high, glossy, glabrous; stipe (0—)1.5—6(— 15) by c. 1.5 mm; seed-bearing part ellipsoid to obovoid, triangular to rounded-triangular in cross section, up to 1.5 cm wide; endocarp thin, com- pletely sclerenchymatic, pubescent to densely woolly; septa densely woolly to glabrous. Seeds ellipsoid, 10-14 mm long; cotyledons subcollater- al, lower cotyledon folded, usually smaller than the upper one, (upper one folded). — Fig. 58c. Distribution — Solomon Islands to Malesia: New Guinea (Irian Jaya: Vogelkop; Papua New Guinea: N coast up to Peninsula). Ecology — Primary and secondary forest, from sea level up to 1300(—2800) m altitude. Fl. Mar.— May; fr. May—Oct. Twigs and leaf rachises often show holes in the bark and cavities inside as if they were inhabited by ants. 3. Mischocarpus paradoxus Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1920) 306; Baker, J. Bot. 61, Suppl. (1923) 11; Radlk. in Fedde, Rep. 20 (1924) 40; in Engl., Pflanzenr. 98 (1933) 1310; R.W. Ham, Blumea 23 (1977) 270. — Type: Forbes 310 (M holo; BM, prob. L), New Gui- nea. Shrub or tree, up to 5(—20?) m high; puberu- lous, early glabrescent. Twigs 3-8 mm in diam. Leaves 3- or 4-jugate; petiole 5—11(—17) cm long; petiolules 4-11(—15) mm long. Leaflets ovate to elliptic, 12-27 by 4.5—11 cm, index 1.8—3.2, per- gamentaceous to subcoriaceous; base rounded (to obtuse or acute); domatia absent or inconspicuously in axils of nerves below; midrib above an angular Flora Malesiana ser. I, Vol. 11 (3) (1994) line, nearly always visible; nerves 12—24 per side, moderately curved; veins very regularly reticulate, prominent on both surfaces. Inflorescences rami- florous, on 7-13 mm thick branches (to rarely ax- illary); axes up to 25 cm long, sparsely to rather strongly branched; bracts triangular, up to | mm high. Calyx connate for 1/3; lobes triangular, usu- ally slightly acuminate, up to 1.2 by 0.5 mm, (un- equal), subcoriaceous, outside sparsely pubescent, inside puberulous near the base, often as a row of short hairs. Petals (3—)5(—6), + elliptic, up to 0.5 mm long, not clawed, glabrous or with a few hairs near the base, not auriculate. Disc annular or slight- ly cup-shaped, often irregularly, deeply lobed to interrupted, 0.75-1 mm in diam., glabrous or lo- cally sparsely puberulous. Stamens 8, up to 2mm long; filaments pubescent; anthers smooth or lo- cally inconspicuously papillate, glabrous. Pistil not or shortly stipitate, pubescent. Fruits globose, up to 1 cmin diam., not stipitate or stipe c. 1 mm long in unripe fruits, glabrous, with many light brown dots and faint remnants of a waxy covering, very shortly apiculate, thin-walled; endocarp incom- pletely sclerenchymatic, consisting of a c. 2 mm wide tapering plate at either side of the suture, gla- brous; septa glabrous. Seeds up to 9 mm long, glo- bose to ellipsoid, arillode without appendix; coty- ledons subcollateral, lower cotyledon folded, usu- ally smaller than the upper one. — Fig. 58d. Distribution — Malesia: Papua New Guinea (Central Prov.). Ecology — Primary and secondary forest; 400— 1400 m altitude. Fl. June—Sept.; fr. Oct.—Nov. 4. Mischocarpus pentapetalus (Roxb.) Radlk., Sapind. Holl.-Ind. (1879) 43; Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 646, 648; Lecomte in FI. Indo-Chine. 1 (1912) 1028; Radlk. in Engl., Pflanzenr. 98 (1933) 1293; Gagnep. in FI. Indo- Chine, Suppl. 1 (1950) 984; R.W. Ham, Blu- mea 23 (1977) 271; Yap in Tree Fl. Malaya 4 (1989) 449; Jansen et al. in Verheij & Coronel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991) 347. — Schlei- chera pentapetala Roxb., [Hort. Beng. (1814) 29, nom. nud.] Fl. Ind. ed. Carey (1832) 275. — Schmidelia pentapetala Wight, Ic. 2 (1840) t. 402, nom. illeg. — [Cupania pentaphylla Wight, Ic. 2 (1840) t. 402, nom. inval. — Schleichera pentaphylla Roxb. in Wight, Ic. 2 (1840) t. 402, nom. inval. —] Cupania roxburghii Wight, Ic. 2 (1840) t. 402, nom. il- leg. — Cupania pentaphylla Hiern in Hook. f., FI. Br. India 1 (1875) 678. — Pedicellia penta- petala Pierre, Fl. Coch. (1895) text with t. 324a. Adema, Leenhouts, Van Welzen — Sapindaceae — Type: M.R. Smith (30417), 1811 (BM holo), India. Mischocarpus sumatranus Blume, Rumphia 3 (1847) 168; Radlk., Sapind. Holl.-Ind. (1879) 12, 44; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 646, 648; King, J. As. Soc. Beng. 65, IT (1896) 448; Vale- ton, Bull. Inst. Bot. Buitenzorg 15 (1902) 10; Ridley, Fl. Malay Penins. 1 (1922) 508; Merr., Pl. Elmer. Born. (1929) 176; Radlk. in Engl., Pflanzenr. 98 (1933) 1298; Burk., Dict. Econ. Prod. Malay Penins. (1935) 1480; Corner, Way- side Trees (1940) 589; Adelb., Blumea 6 (1984) 234. — Cupania sumatrana Miq., Fl. Ind. Bat. 1, 2 (1859) 566; Hiern in Hook. f., Fl. Br. India 1 (1875) 678; Kurz, Rep. Pegu (1875) 32; Fl. Burma | (1877) 285; Kanjilal & Das, Fl. As- sam | (1936) 317. — Ratonia sumatrana Kurz, Rep. Pegu (1875) App. A38, B40. — Pedicel- lia sumatrana Pierre, Fl. Coch. (1895) text with t. 323b. — Lectotype (R.W. Ham 1977): Anon- ymous s.n. (L holo, sh. 908.269-1030), S Su- matra. Mischocarpus fuscescens Blume, Rumphia 3 (1847) 169; Radlk., Sapind. Holl.-Ind. (1879) 12, 43; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 646, 648; King, J. As. Soc. Beng. 65, II (1896) 447; Vale- ton, Bull. Inst. Bot. Buitenzorg 15 (1902) 10; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 221; Lecomte in Fl. Indo-Chine 1 (1912) 1028, excl. var. bonii H. Lecomte; Ridley, Fl. Malay Penins. | (1922) 508; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513; Radlk. in Engl., Pflan- zenr. 98 (1933) 1294; Ramesh Rao, Ind. Woods 2 (1963) 223, f. 325; Backer & Bakh. f., Fl. Java 2 (1965) 141. — Cupania fuscescens Migq,., FI. Ind. Bat. 1, 2 (1859) 567. — Pedicellia fusces- cens Pierre, Fl. Coch. (1895) text with t. 323b.; Hu, Bull. Fan Mem. Inst. Biol. 1 (1929) 31. — Lectotype (R.W. Ham 1977): Anonymous s-.n. (L holo, sh. 908.269-555), Java. Schleichera subundulata Turcz., Bull. Soc. Nat. Moscou 21 (1848) 574. — Cupania subundu- lata Rolfe, J. Bot. 23 (1885) 211; Vidal, Phan- er. Cuming. (1885) 6, 105. — Type: Cuming 507 (KW holo, n.v.; A, FI, K, L, M, P, SING; KW, n.v.), Philippines. Mischocarpus salicifolius Radlk. in Perkins, Fragm. Fl. Philipp. 1 (1904) 64; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513; Radlk. in Engl., Pflan- zenr. 98 (1933) 1297. — Type: Merrill 1842 (B¥ holo; M, US), Philippines. Mischocarpus ellipticus Radlk. in Elmer, Leafl. Philipp. Bot. | (1907) 210; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513; Radlk. in Engl., 663 Pflanzenr. 98 (1933) 1294. — Type: Elmer 7272 (M holo; A, BO, K), Philippines. Mischocarpus endotrichus Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1615; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513; Radlk. in Engl., Pflanzenr. 98 (1933) 1303. — Type: Elmer 12977 (M holo; A, BM, BO, FI, K, L, P, U, US), Philippines. Mischocarpus brachyphyllus Radlk., Philipp. J. Sc., Bot. 8 (1914) 472; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 512; Radlk. in Engl., Pflanzenr. 98 (1933) 1304. — Lectotype (R.W. Ham 1977): Vanoverbergh 1175 (M holo; A, L, P, U, WRLS), Philippines. Mischocarpus sublaevis Radlk., Philipp. J. Sc. 20 (1922) 662; in Engl., Pflanzenr. 98 (1933) 1303. — Type: Wenzel 660 (error 600 in Radlk.) (M holo; A, BM), Philippines. Nephelium hosei Ridley, Kew Bull. (1933) 191, type only; Harms in Engl., Pflanzenr. 98 (1934) 1500. — Type: Hose 124 (BM? holo, n.v.; CGE, L), Sarawak. Shrub or tree up to 15(—25) m high; puberulous on all young parts and especially in the inflores- cences, brownish ferrugineous, partly glabrescent. Twigs (2—)3—9(—12) mm in diam., brownish to red- dish brown, (becoming greyish brown). Leaves (1—)2—5(-6)-jugate; petiole 3-25 cm long; peti- olules 3-12 mm long. Leaflets ovate to elliptic, (S—) 7-20(-40) by (1—-)2-6(-15) cm, index 2-5(-8), pergamentaceous to coriaceous; base rounded or angular; domatia usually present in the axils of the nerves below; midrib above prominent, angular (to rounded in the basal part or completely so), rather flat and broad in the basal part in big leaflets; nerves (6—)9—20(—25) per side, at least prominent on the lower surface, (hardly distinct from veins); veins usually coarsely reticulate, above hardly or not prominent, beneath prominent. /nflorescences ax- illary and pseudoterminal; main axis up to 40 cm long; secondary axes 1—20 cm long; cymules up to 11 mm long; bracts triangular, up to 6 by 1.5 mm (to 8 by 4 mm), (those of the secondary axes cadu- cous). Calyx hardly connate, rarely so for 1/4; lobes triangular to ovate, 0.75—2 by 1—2 mm, subcoria- ceous to slightly fleshy, sometimes 1(—3) nerves visible, outside puberulous to pubescent (to very sparsely so in the upper part), inside puberulous, at least at the base as a more or less dense row. Petals 0-S, usually unequal, ovate to elliptic, minute to 2 by 1 mm, glabrous or pubescent (usually at the base), not clawed, usually auriculate. Disc an- nular, (1—)1.5—2(—3) mm in diam., sometimes ir- regularly lobed, rarely with lobes protruding be- tween the stamens, puberulous or glabrous. Sta- 664 Flora Malesiana ser. I, Vol. 11 (3) (1994) mens 7 or 8, 3-4 mm long; filaments puberulous (to glabrous); anthers papillate, sparsely pubescent or glabrous. Pistil puberulous. Fruits (0.8—)1—2(-3) cm long, glabrous; stipe 2-11 by 1-2 mm; seed- bearing part globose to ellipsoid to obovoid, trian- gular (or triquetrous) in cross section, 0.4—1.4 cm in diam., inside glabrous (or completely, rather densely hairy); endocarp incomplete (2 mm thick at either side of the suture) to complete, usually less so in the fertile than in the sterile cells. Seeds up to 8(-12) mm long, ellipsoid to globose; coty- ledons subcollateral, both or only the lower one folded, lower one usually smaller. — Fig. 58e. Distribution — From India, Burma, SW China, throughout SE Asia, to W Malesia up to the line Philippines—Borneo—Java. Habitat & Ecology — Primary and secondary forest; (300—)800—2000 m altitude. Fl. and fr. throughout the year, but the main periods different in the various regions, e.g. Java: fl. Nov., fr. Jan.— Mar.; Philippines: fl. Mar.—May, fr. Mar.—June. Notes — 1. Many species formerly kept sepa- rate are included in M. pentapetalus. Two of these, ‘M. endotrichus’ and ‘M. brachyphyllus’, were dis- tinguished by the presence of a hairy endocarp. Fur- ther species were differentiated on the variation in the petals. However, the whole range of variation, from 5 well-developed auriculate petals via 5 or less, reduced, non-auriculate ones, to petals absent, occurred in each of these species. Variation in the leaflets led to the description of several more spe- cies; ‘M. sumatranus’ has leaflets with many pro- nounced nerves and is common in SE Asia, Su- matra, Borneo, and Luzon; ‘M. fuscescens’ has leaf- lets with few, usually less pronounced nerves; it occurs throughout western Malaysia. The variation is extreme in N Borneo and the Philippines; leaflet length ; breadth ratio varies here from 2 to 8 (‘M. ellipticus’ and ‘M. salicifolius’ resp.), and leaflet sizes range from 5 by 2 cm up to 38 by 15 cm. Collections of the ‘salicifolia’ type are widespread in the Philippines, while the ‘edlipticus’ type comes usually from Samar, Leyte, Negros, and northern Mindanao, outside the Philippines being represent- ed also in Indo-China (‘M. pentapetalus var. cam- bodianus’). However, in spite of their variation in flowers and fruits, they are not different from ma- terial from India, China, Java, or elsewhere. Not- withstanding all this variation M. pentapetalus is a well-defined species. 2. See also note 2 under M. sundaicus. 5. Mischocarpus pyriformis (F. Muell.) RadIk., Sapind. Holl.-Ind. (1879) 43; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 536, 647; Maiden & Betche, Cens. N.S.W. Pl. (1916) 129; Domin, Bibl. Bot. 22 (1927) 910; Francis, Austral. Rain-For. Trees (1929) 235; ed. 2 (1951) 261; Radlk. in Engl., Pflanzenr. 98 (1933) 1305; Merr. & L.M. Per- ry, J. Arnold Arbor. 21 (1940) 523: R.W. Ham, Blumea 23 (1977) 275; S.T. Reynolds in FI. Austral. 25 (1985) 97. — Schmidelia pyriformis F. Muell., Fragm. Phyt. Austral. 1 (1858) 2, comb. illeg. — Cupania pyriformis F. Muell., Fragm. Phyt. Austral. 2 (1860) 76; 5 (1865) 147; 9 (1875) 90; Ch. Moore, Fl. N.S.W. (1893) 91. — Ratonia pyriformis Benth., Fl. Austral. 1 (1863) 461. — Lectotype (R.W. Ham 1977): Hill & Mueller s.n. (MEL holo, sh. 56010; K, M, MEL), Queensland. Mischocarpus. papuanus Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1924) 39; in Engl., Pflanzenr. 98 (1933) 1307; P. Royen, Man. For. Trees Papua & New Guinea 2 (1964) f. 14. — Type: Chalm- ers s.n., 1885 (M holo), New Guinea. Mischocarpus retusus Radlk., Bot. Jahrb. 56 (1920) 304, f. 4a-f; in Engl., Pflanzenr. 98 (1933) 1308, f. 38a-f. — Lectotype (R.W. Ham 1977): Le- dermann 11231 (B¥ holo; M), New Guinea. Shrub or tree up to 30 m high; puberulous, gla- brescent. Twigs 1.5—6 mm in diam., Leaves 1—4- jugate; petiole 1-8 cm long; petiolules 4-18(—25) mm long. Leaflets ovate to elliptic, 3.5—18 by 1.2— 7 cm, index 2—3.5, pergamentaceous to coriaceous; base acute to obtuse (or rounded); margin flat to revolute; domatia absent or present, (conspicuous), usually restricted to the axils of the basal nerves below; midrib above not visible and hidden in a narrow groove, or (locally) visible as a slender, sunken to prominulous line, beneath prominent till the apex, (puberulous on both sides); nerves 6-15 per side, moderately (to irregularly) curved, above prominulous or slightly sunken (to hardly distinct from venation), beneath prominent to prominulous; veins rather densely reticulate, above prominulous to sunken, beneath prominent to prominulous. /n- florescences axillary and pseudoterminal, up to 25 cm long, axes branched; cymules often crowded to the end of the axes; bracts triangular, up to 1.5 mm long. Calyx cup-shaped, up to 2 mm high, con- nate for 1/2—2/3; lobes triangular, 0.3—0.8 mm long, subcoriaceous, outside densely appressedly short- hairy especially in the upper half, inside glabrous or puberulous. Petals 0-5, up to 1.5 mm long, clawed for 1/3—3/4, variably hairy, ciliate, mainly on the claw, scales, and auricles; claw usually clear- ly tubular, split inside; blade varying from trian- gular to ovate, usually with 2 scales or auricles at the base, usually with 2 or 3 rounded or acute lobes Adema, Leenhouts, Van Welzen — Sapindaceae at the apex. Disc annular to cup-shaped, uninter- rupted, lobed, or interrupted once to several times, 0.5S—1 mm in diam., glabrous. Stamens (7) 8, up to 3 mm long; filaments pubescent; anthers papillate, pubescent to puberulous or glabrous. Pistil stipi- tate; stipe usually grooved; ovary not grooved, short sericeous mainly in the upper half. Fruits up to 3.5 cm high, glabrous; stipe up to 2.5 cm long by 1.5 mm in diam. at the base; seed-bearing part glo- bose to ellipsoid, rounded to triangular in cross section, up to 1 cm in diam., apiculate; endocarp partly or completely sclerenchymatic, glabrous (or locally hairy). Seeds ellipsoid, up to 7 mm long; cotyledons subcollateral, lower folded, usually smaller than the upper one. Distribution — Australia (NE and E Queensland, NE New South Wales) and Malesia: New Guinea. Habitat & Ecology — Subsp. retusus is found in the high mountains, subsp. papuanus at lower alti- tudes. Note — Three subspecies can be recognized, of which one, subsp. pyriformis, is found only in Aus- tralia. The remaining two subspecies are readily separable in Papua New Guinea, but only with dif- ficulty so in the Vogelkop in Irian Jaya. KEY TO THE SUBSPECIES la. Leaflets not or very shortly acuminate; doma- tia | or 2(—7), usually conspicuous, very rare- ly absent. Disc c. | mm in diam. Anthers hairy b. subsp. retusus b. Leaflets with long and slender acumen, or acu- men rarely short and broad; domatia | or 2 small ones or absent. Disc 0.5—1 mm in diam. Anthers glabrous (or with few hairs) a. subsp. papuanus a. subsp. papuanus (Radlk.) R.W. Ham, Blumea 23 (1977) 277. — Mischocarpus papuanus Radlk. Twigs 1.5—4 mm in diam. Leaves (1-) 2- or 3- (4-)jugate; petiole 2-8 cm long; petiolules 4—20 mm long. Leaflets 5-16 by 2—6 cm, index 2-3, pergamentaceous to subcoriaceous; apex with a long and slender (or short and broad) acumen; domatia absent or | or 2 small ones in the axils of the basal nerves; nerves 7-15 per side. /nflores- cences up to 20 cm long. Calyx lobes up to 0.5 mm long. Petals 1-5, up to | mm long. Disc 0.5—1 mm in diam. Stamens 7 or 8; anthers glabrous or with a few hairs. Fruits up to 3.5 cm high; stipe (10—)15— 25 mm long, up to | mm in diam. at the base; seed- bearing part up to | cm in diam.; endocarp gla- brous. — Fig. 58f. Distribution — Malesia: Irian Jaya (N coast from 665 Vogelkop to Jayapura) and Papua New Guinea (Western, Morobe, and Gulf Prov.). Habitat & Ecology — Primary rain forest; sea level up to 1200(—1800) m altitude. Fl. Mar.—July; fr. Mar., Apr., Aug. b. subsp. retusus (Radlk.) R.W. Ham, Blumea 23 (1977) 277; S.T/. Reynolds in Fl. Austral. 25 (1985) 98, map 127. — Mischocarpus retusus Radlk. Twigs 2-3 mm in diam. Leaves 1- or 2- (3-)ju- gate; petiole 1-5 cm long; petiolules 4-18 mm long. Leajlets 3.5—13 by 1.2-4.5 cm, index 2—3.5, coria- ceous; apex not or shortly acuminate; domatia (rare- ly absent to) usually conspicuous, | or 2(—7), usu- ally in the axils of the basal nerves; nerves 6—12 per side. Inflorescences up to 15 cm long. Calyx lobes up to 0.8 mm long. Petals 4 or 5, up to 1.5 mm long. Disc c. | mm in diam. Stamens 8; an- thers puberulous. Fruits up to 2.5 cm high; stipe 10—17(—20) mm long, up to 1.5 mm in diam. at the base; seed-bearing part up to 8 mm in diam.; en- docarp sometimes with a few hairs. Distribution — Australia (NE Queensland) and Malesia: Irian Jaya (Jayapura and Snow Mountains Prov.) and Papua New Guinea (E Sepik, W High- lands, S Highlands, Morobe, and Milne Bay Prov.). Habitat & Ecology — Primary forest at (1000—) 1300-3000 m altitude. Fl. and fr. the whole year round. 6. Mischocarpus reticulatus (Radlk.) R.W. Ham, Blumea 23 (1977) 280. — Mischocodon retic- ulatus Radlk., Bot. Jahrb. 50 (1913) 80; 56 (1920) 308; in Engl., Pflanzenr. 98 (1933) 1328; Merr. & L.M. Perry, J. Arnold Arbor. 21 (1940) 524; P. Royen, Man. For. Trees Papua & New Guinea 2 (1964) 33; Hartley et al., Lloydia 36 (1973) 270. — Lectotype (R.W. Ham 1977): Schlechter 17683 (B¥ holo; M), New Guinea. Tree up to 15 m high, dbh up to 0.75 m; puber- ulous, early glabrescent. Twigs 4-7 mm in diam., slightly grooved. Leaves 2—-4-jugate; petiole 7-20 cm long; petiolules 7-14 mm long. Leaflets ovate to elliptic, 7-28 by 3.4—10 cm, index 2—3.2, perga- mentaceous to subcoriaceous, slightly bullate; base acute to obtuse or rounded; domatia small, in the axils of the nerves (and intersecondary nerves) below; midrib prominent above, angular or locally obtuse, (sub)glabrous, (beneath puberulous); nerves 10-14 per side, moderately curved, prominent on both surfaces; veins prominent above and beneath. Inflorescences ramiflorous, on branches of at least 6—10 mm in diam., (to axillary), probably also cau- liflorous; axes up to 5 cm long, branched, ferrugi- Flora Malesiana ser. I, Vol. 11 (3) (1994) \ \ Si A Fig. 59. Mischocarpus reticulatus (Radlk.) R.W. Ham. a. Part of twig, top leaflet from above; b. male inflorescence; d. female flower; e. ibid., calyx removed; f. infructescence (a: Hartley 12423; b, c: Hartley 12534; d, e: Carr 14994; f: Clemens 8619). c. male flower, calyx removed; ’ Adema, Leenhouts, Van Welzen — Sapindaceae 667 neous puberulous; flowers solitary or rarely in pairs; pedicels up to 5 mm long; bracts lanceolate to ovate, up to 3 mm long. Calyx cup-shaped, 3—S mm high, connate for ]/2—3/4, membranous, greenish white; lobes triangular, up to 2 by 2 mm, with 3 or 5 nerves, outside sparsely pubescent, inside glabrous or with a few hairs near the base. Petals absent. Disc an- nular to pentagonal, broad, nearly completely cov- ering the receptacle, surrounding the base of the stamens, confluent with the base of the pistil, 2-3 mm in diam., glabrous or laterally puberulous. Sra- mens 5—7; filaments up to 11 mm long, filiform, glabrous; anthers smooth (or papillate at the apex), glabrous. Pistil glabrous or sparsely hairy, distinctly angular in cross section; style with 1—-1.5 mm long, recurved, twisted stigmatic lobes. Fruits up to 2.5 cm high, glabrous; stipe 5—14 by c. 1.5 mm; seed- bearing part globose, up to | cm in diam., apicu- late for 1-2 mm, inside glabrous; endocarp scler- enchymatic for 2/3 of the width of a valve. Seeds not known ripe. — Fig. 59. Distribution — Malesia: Papua New Guinea (Madang, E Highlands, Morobe, and Central Prov.). Habitat & Ecology — Mountain rain forest from 600-1650 m altitude. Fl. Mar., May, Sept.—Dec.: fr. July, Aug., Oct. 7. Mischocarpus sundaicus Blume, Bijdr. (1825) 238; Rumphia 3 (1847) 167; Radlk., Sapind. Holl.-Ind. (1879) 12, 91; Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 646; King, J. As. Soc. Beng. 65, II (1896) 447; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 223; Lecomte in Fl. Indo-Chine 1 (1912) 1029, f. 128; Radlk., Bot. Jahrb. 56 (1920) 303; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513; Craib, Fl. Siam. Enum. 1 (1926) 333; Radlk. in Engl., Pflanzenr. 98 (1933) 1299; Burk., Dict. Econ. Prod. Malay Penins. (1935) 1479; Corner, Wayside Trees (1940) 589, f. 211, 212; Adelb., Blumea 6 (1948) 324; Gagnep. in Fl. Indo-Chine, Suppl. 1 (1950) 986; Backer & Bakh. f., Fl. Java 2 (1965) 141; R.W. Ham, Blu- mea 23 (1977) 281; Yap in Tree Fl. Malaya 4 (1989) 449. — Cupania lessertiana Cambess., Mém. Mus. Hist. Nat. Paris 18 (1829) 28, 46, t. 3, comb. illeg.; Hiern in Hook. f., Fl. Br. India 1 (1875) 678. — Cupania mischocarpus Steud., Nomencl. ed. 2, 1 (1840) 454, comb. illeg. — Pedicellia sundaica Pierre, Fl. Coch. (1895) t. 323b. — Lectotype (R.W. Ham 1977): Anony- mous S.n. (L holo, sh. 908.269-749), Java. Schleichera revoluta Turcz., Bull. Soc. Nat. Mos- cou 21 (1848) 574. — Cupania revoluta Rolfe, J. Bot. 23 (1885) 211; Vidal, Phan. Cuming. (1885) 49, 105, non Radlk., comb. illeg. (ICBN 1972, art. 64). — Type: Cuming 1387 (KW holo, n.v.; BM, K, L, P), Philippines. Cupania erythrorhachis Miq., Sumatra (1861) 199, 509. — Type: Teijsmann 4557 (U holo; BO, K, L, M), Sumatra. Mischocarpus lessertianus Ridley, Fl. Malay Penins. | (1922) 508; Burk., Dict. Econ. Prod. Malay Penins. (1935) 1479. — Lectotype (R.W. Ham 1977): Ridley 1908 (SING holo), Singa- pore. |Mischocarpus vulcanicus Elmer ex Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513, nom. nud.; Elmer, Leafl. Philipp. Bot. 10 (1939) 3809, nom. nud. — Based on Elmer 17092, 17116 (A, BM; BO) FE Kol, NY, BU; UE SUSPENG 7), Philippines. ] Mischocarpus pyriformis auct. non Radlk.: T.C. White, Contr. Arnold Arbor. 4 (1933) 63. Shrub or tree, up to 10(—30) m high; puberu- lous in the young parts and inflorescences. Twigs 1.5—5 mm in diam., usually reddish brown. Leaves 1—3- (or 4-)jugate (or simple); petiole 1-11 cm long; petiolules 3-8 mm long. Leaflets ovate to elliptic, 4-17(—26) by 1.5—7(—10) cm, index 24.5, pergamentaceous to subcoriaceous; base rounded or angular; (margin slightly revolute); domatia usu- ally present in the axils of the nerves below; mid- rib prominent above, broad to rather narrow, usu- ally rounded, often locally angular (to completely so); nerves 8—15 per side, moderately curved, be- neath prominent; veins densely reticulate, prominu- lous to smooth above, beneath more prominent to the margin. /nflorescences axillary and pseudoter- minal, 1.5—25 cm long, branched; cymules shortly stalked; pedicels 1-3 mm long; bracts triangular, up to 1.5 mm long (to up to 6 by 2 mm), usually caducous. Calyx connate for 1/4—1/3; lobes trian- gular, 0.65—2 by 0.5-1 mm, subcoriaceous, out- side puberulous to pubescent in the lower half (to rarely so apically), inside glabrous (to a few hairs). Petals 0 (up to 3 in Peninsular Malaysia), up to 1.5 mm long, elliptic to oblong, glabrous or with a few hairs near the base, inconspicuously auriculate. Disc annular, 1-2 mm in diam., glabrous or sparsely puberulous. Stamens (6—)8(—9), up to 4 mm long: filaments glabrous; anthers not papillate, glabrous. Pistil puberulous. Fruit 7—17(—20) mm high, gla- brous or very sparsely puberulous; stipe 2—14 by 1—1.5 mm; seed-bearing part globose to ellipsoid, round to faintly triangular in cross section, apicu- lus 1-2 by 0.5—1(—2) mm, inside glabrous (to pu- bescent along the sutures); endocarp completely sclerenchymatic. Seeds globose to ellipsoid, up to 7 mm long; cotyledons superposed, about equal. — Fig. 58g. 668 Flora Malesiana ser. I, Vol. 11 (3) (1994) Distribution — From India and S China through SE Asia and throughout Malesia. Absent from Australia, see note 3. Habitat & Ecology — Primary and secondary forest in lowland and coastal regions; sea level up to 800(—1600) m altitude. Fl. mostly Jan.—Aug.; fr. mostly May—Dec. Notes — 1. In a few collections, restricted to Peninsular Malaysia, petals occur. The petals strongly resemble those of M. pentapetalus and other species in the ‘pentapetalus-group’ (M. pen- tapetalus, M. triqueter, M. largifolius, M. paradox- us, M. reticulatus, and M. grandissimus). 2. The delimitation between M. sundaicus and M. pentapetalus is not sharp for some characters and that is why Gagnepain (Not. Syst. 13, 1947, 34, 35) suggested hybridization between these two species. Detailed studies on this problem are needed (see also below, the note under Mischocarpus prob. spec. nov.). 3. §.T. Reynolds (in Fl. Austral. 25, 1985, 101) reports that M. sundaicus is absent from Australia. The specimens attributed to M. sundaicus by R.W. Ham (1977) were described by her as three new species: M. australis, M. macrocarpus, and M. stip- itatis. They differ from M. sundaicus in the much- branched panicle with cymules on longer pedicels, the hairy inner surface of the fruit, the persistent broad style base, the larger bracts and dorsiven- trally flattened petioles. As Australia is outside the scope of Flora Malesiana, no decision about the status of these new species has been made. 8. Mischocarpus triqueter Radlk. in Perkins, Fragm. Fl. Philipp. | (1904) 65; Merr., Enum. Philipp. Flow. Pl. 2 (1925) 514; Radlk. in Engl., Pflanzenr. 98 (1933) 1297. — Mischocarpus fuscescens auct. non Blume: Vidal. Revis. (1886) 95, no. 720, 1039; Cat. Herb. (1892) 83, no. 720, 1039. — Lectotype (R.W. Ham 1977): Ahern 262 (B+ holo; BO, US), Philippines. Mischocarpus cauliflorus Radlk., Philipp. J. Sc., Bot. 8 (1914) 471; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 513; Radlk. in Engl., Pflanzenr. 98 (1933) 1298. — Type: McGregor 271 (M holo; K, US), Philippines. Small tree, up to 6 m high, dbh up to 12 cm, not or hardly branching; puberulous, early glabrescent. Twigs 4-9 mm in diam., usually grooved. Leaves (2- or) 3—5-jugate; petiole 7-17 cm long; petiolules 8-15 mm long. Leaflets ovate to obovate, 5—26 by 2-10 cm, index 2—4.5, pergamentaceous to subco- riaceous; base acute (to obtuse); domatia absent or inconspicuously in the axils of the nerves below; midrib above prominent, angular; nerves 7—13 per side, moderately curved; nerves and veins promi- nent on both surfaces. Inflorescences ramiflorous on up to 15 mm thick branches consisting of 1—7, up to 10 cm long axes, simple (or sparsely branched at the base); bracts triangular, up to | mm long. Calyx lobes hardly connate, triangular to ovate, 1.5— 2.5 by 1-2 mm, membranous with | or 3 visible nerves, outside sparsely pubescent, inside glabrous or with a few hairs near the base. Petals (2—)5, el- liptic, c. 1.5 by | mm, sparsely accrescent to 3 by 2 mm in young fruits, not clawed, glabrous; auri- cles usually present, sparsely pubescent. Disc an- nular, usually with lobes protruding between the stamens, + confluent with the base of the fruit, 1.5— 2 mm in diam., glabrous. Stamens 8, 3-4 mm long; filaments glabrous to sparsely puberulous; anthers papillate, glabrous. Pistil hardly stipitate, distinct- ly triangular, hairy. Fruits up to 3 cm high, gla- brous; stipe 4—12 mm high, triangular in cross sec- tion; seed-bearing part ellipsoid to obovoid, trian- gular (or rounded-triangular) in cross section, up to 1.2 cm in diam., apiculate for 1-3 mm, inside glabrous; endocarp thin, (nearly) completely scle- renchymatic. Seeds ellipsoid, up to 1.2 cm long; cotyledons subcollateral, both folded, the lower usually smaller than the upper. — Fig. 58h, i. Distribution — Malesia: Philippines (Luzon, Mindoro, Guimaras, Negros). Habitat & Ecology — Primary forest up to 500 m altitude. Fl. Oct., Jan.; fr. Feb.—July. 9. Mischocarpus prob. spec. nov.: R.W. Ham, Blu- mea 23 (1977) 286. — Based on UPNG (Mil- lar) 1042 (K), Papua New Guinea. Small tree, c. 6 m high; puberulous in infruc- tescences, glabrescent. Twigs 6-8 mm in diam. Leaves c. 3-jugate; petiole c. 17 cm long; petiolules c. 1 cm long. Leaflets elliptic, 12-16 by 4-5 cm, index c. 3, subcoriaceous; base acute; domatia very inconspicuous, in the axils of the nerves below; midrib above prominent, angular to locally round- ed: nerves c. 15 per side, moderately curved; veins regularly reticulate, prominent on both surfaces. Inflorescences and flowers unknown. Infructes- cences ramiflorous, 13-16 cm long, branched. Calyx in fruit connate for 1/4, lobes slightly un- equal, triangular to ovate, c. 1.5 by | mm, outside glabrous, inside glabrous or with a few hairs, sub- coriaceous at the base, towards the top somewhat membranous with visible nerves, not accrescent. Petals: 1 was found together with | scar of a petal: elliptic, c. 0.75 by 0.5 mm, not clawed, without scales or auricles, glabrous. Disc in fruit annular, 1.5—2 mm in diam., slightly lobed, glabrous with lenticel-like dots. Fruits immature, green, proba- bly all sterile, 8-10 mm high, glabrous; stipe 1-3 mm high, slightly lobed at the base, grading into Adema, Leenhouts, Van Welzen — Sapindaceae the seed-bearing part; the latter about rounded-tri- angular in cross section, 5—7 mm in diam., inside pubescent on axial and sutural parts, sclerenchy- ma just reaching into the septa. Seeds hardly de- veloped (although fruit seemingly mature); arillode with a slightly lobed pseudo-funicle + as long as the stalk. Distribution — Malesia: Papua New Guinea (Central Prov.). 669 Ecology — In rain forest. Fruiting at least in March. Note — This is doubtlessly a Mischocarpus in the ‘pentapetalus-group’, probably closest related to M. largifolius and M. paradoxus. It may be a new species, but the specimen may also be a hy- brid, because of the sterile fruits, either between M. largifolius and M. paradoxus, or between M. sundaicus and one of the latter two species. NEPHELIUM (P.W. Leenhouts) Nephelium L., Mant. Pl. 1 (1767) 18; Radlk. in Engl., Pflanzenr. 98 (1932) 950; Leenh., Blumea 31 (1986) 373 [see there for more synonyms on the (sub)sectional level]. — Type species: Nephelium lappaceum L. Medium-sized to tall trees or rarely shrubs, probably dioecious, sometimes monoe- cious. Jndumentum of solitary simple hairs, glandular scales absent. Leaves spirally ar- ranged, paripinnate, (1-foliolate or) 1—5(—18)-jugate, without pseudo-stipules; neither petiole nor rachis winged. Leaflets alternate to more rarely opposite, beneath mostly dis- tinctly glaucous, finely papillate, domatia often present, margin entire or rarely repand, nervation open (except in N. subfalcatum). Inflorescences in some species all axillary, in most at least partly pseudoterminal, in some truly terminal, in N. cuspidatum also rami- and cauliflorous. Flowers actinomorphic. Sepals (4 or) 5 (or 6), free to more than half- way up connate, valvate or sometimes (when the sepals are nearly free) slightly imbri- cate, all equal, not petaloid, entire, outside and inside hairy, outside more sparsely and with shorter hairs than inside, not ciliate, without glands. Petals shorter than the calyx, 5 (or 6), clawed, well developed, with a bilobed scale without appendages, hairy on both sides, entire, or 1-4 reduced, or often none. Disc entire, often slightly lobed, without appendage. Stamens 4—10, exserted in male flowers; filaments rather densely long-hairy at least in the basal part, anthers nearly always with at least a few hairs. Pistil densely hairy; ovary short-stalked, (1- or) 2(—4)-celled, mostly warty; style usually well devel- oped; stigmas fairly long, spreading to finally recurved. Ovules | per cell, half enveloped by an outgrowth of the placenta. Fruits 1-, exceptionally 2-lobed; stipe short to incon- spicuous; body ellipsoid to subglobular, the surface warty to spiny, exceptionally nearly smooth; pericarp thin- to thick-coriaceous or exceptionally corky to nearly woody, inside glabrous, apparently often at least in the apical part finally loculicidal, either rather irreg- ular or with two about equal valves. Seeds: hilum nearly basal; micropylar wart apical or mostly subapical; sarcotesta covering the whole seed, with the exception of the micro- pylar region, or at least perforated in front of the micropyle, sometimes with a collar- like outgrowth around the hilum; endotesta tough or exceptionally rather hard. — Figs. 60-63. Distribution — 22 species; SE Asia from Yunnan and Assam to Hainan and Malesia: W Malesia, Philippines, Celebes, and Moluccas. Habitat & Ecology — Middle storey of rain forests at low to medium altitudes, some- 670 Flora Malesiana ser. I, Vol. 11 (3) (1994) times in deciduous or savannah forest. The fruits are mainly eaten by monkeys and fruit bats: the fruits of some species seem to be dispersed by water. Uses — The sarcotesta of a few species is eaten, but only N. appaceum (Rambutan) is commonly cultivated as a fruit tree. The timber is hardly of any importance. Note — Identification is hardly possible without + mature fruits. Therefore, under many species notes have been given regarding differences with other comparable species from the same area. KEY TO THE SPECIES la. Pistil 2(—4)-merous. (Exceptionally a few flowers with a |-merous ovary) ..... 2 By aia desnitecoll ents eas meen meen cin eG gle 60 oGiaig ooo Oba rom 13. N. maingayi 2a. Fruit wall coriaceous or eggshell-like, rarely more than 1(—7) mm thick....... 3 b. Fruit wall hard, corky, 3-6 mm thick, the saccate base and the carina of the fruit 6 hs Eamets great eat ner Re era be Raat eM neetete SEER 2. N. compressum 3a. Fruits warty, the warts without a distinct apical appendage. (In a few cases this character may be difficult. Try both ways) ..........--. 6. see ee eee e eee 4 b. Fruits not warty, the appendages usually distinctly divided into a broader or thicker basal and a. narrower apical part 2.22000... 6 a2 sti ets > hd oie ee 9 4a. At least the upper half of the fruit appendages densely puberulous .........-. 5 by Enuitsiglabrousisyadee a: Paste remot eR GE fe Stott atin ae ee 6 5a. Leaf axes densely tomentellous, sometimes glabrescent. Leaflets hairy all over above, soon glabrescent, the midrib sometimes excepted, tomentose on midrib and nerves beneath, densely sericeous in between. Calyx 3 mm high....... 5. N. daedalum b. Leaf axes sparsely puberulous to glabrous. Leaflets glabrous above, sparsely seri- ceous all over beneath, sometimes glabrescent. Calyx up to 1.75 mm high 15. N. melanomiscum 6a. Leaflets mostly widest at about 1/3 above the base; reticulation on upper side dense. Liruitsaculeatess rena Saeniecs wee ee fed ie ee. SSeS rake 7 b. Leaflets widest at or sometimes slightly below the middle; reticulation on upper sidesdlax. Pruitsuberculates. .. 2.4. ayeeia-rl acikis Ae Seite SRI Se eee 8 7a. Leaflets obtuse to acute; veins reticulate; domatia usually present. Burma, Thai- land Alma oehinmalyey. eons eeveriness 2 issih: NS Sethe Bid « Beslet < 8. N. hypoleucum b. Leaflets tapering into a long and broad acumen; veins especially on the lower side rather distinctly scalariform; domatia absent. Borneo ......... 1. N. aculeatum 8a. Leaves up to 7-jugate. Leaflets usually almost parallel-sided, abruptly rounded at the base, flat when dried. Fruits coarsely warty, the warts + arranged into longitudi- nal rows. Malay Peninsula, Sumatra, Java .............- 9. N. juglandifolium b. Leaves 2- or 3-jugate. Leaflets with strongly curved sides, tapering to the cuneate base, often curled or rolled up when dried. Fruits densely knobby like a small Pan- danus fruit head, the knobs not confluent. Borneo An unusual form of 18. N. ramboutan-ake 9a. Fruit appendages distinctly puberulous all over........--..--+2+ +e esses 10 b. Fruits (sub)glabrous or exceptionally with a hair tuft on the tops of the append- AGES ars cepstral ces veers SP RASS ERD. AE er aS carat iy ernst of RPI Soa heuer V5) Adema, Leenhouts, Van Welzen — Sapindaceae 671 I4a. 18a. 19a. 20a: . Fruit appendages mostly filiform. Leaflets mostly flat when dried........... 22 7 beniolnes above,erooved: Leafletsisymmetricals js): 2enian aagbulien fui... 1] Petiolules above not grooved. Leaflets slightly falcate ... 12. N. macrophyllum mmertiiets pemeath clabrousonmearhyyse ip .2sne Assess enw bs vad adie heb. 12 Leaflets beneath densely tomentose on and along midrib and nerves, distinctly to moderately densely sericeous in between... . 2s. os oe ea 6. N. hamulatum . Veins and veinlets densely reticulate, above not very conspicuous, beneath incon- Ppmeriienass Meal lets: Gc ying DROW ac52y8 A + A Reet Bera et eto ns ea a bet te a 3) b. Petiole of leaves absent. Flowering twigs 10-15 mm thick. Leaflets + coriaceous 3. R. congesta Adema, Leenhouts, Van Welzen — Sapindaceae 705 3a. 6a. 7a. 4a. Rpecinict MOWw rere’. 6s (Nt ee. ce TE Pe AI Kiera dee alien ee 4 2 ETE SITE SIG EEL gins ea a atic: A, cam ibe i A hi USGA GH SNEIC Sa ee ere Al gee a a A ee ne oe 5 Botmbscales foldedimarams OF petals: . os.) 62 2s sc ne os aye ek ew cael ee sap 8 . Flowers 4-5 mm diam. Petals pilose on both sides towards the base; margins glabrous or pilose. Lateral nerves of leaflets 0.5—1.8 cm apart. Petiole 1.3-6 cm OMI Tee et Ee Baths Salah latent ke juinys.» = S/T TEE ema Le ee ON EM footy ei diye 6 . Flowers 6—8 mm diam. Petals outside pilose, inside glabrous or very laxly hairy: margins pilose. Lateral nerves of leaflets 0.5—3 cm apart. Petiole 2-17 cm ME te EN ano hag tee’ <4 «duals «fie Aes RaRAd hee peewee) BAS ia 7 Inflorescences paniculate. Leaflets opposite to alternate, margin (slightly) recurved. Petiolule a pulvinus only, rarely up to 4 mm long. Margin of petals pilose 10. R. robertsonii Inflorescences thyrsoid, branching only at the base, giving the impression of a tuft of inflorescences. Leaflets opposite, sometimes subopposite, margin flattened to slightly recurved. Petiolule 5—10 mm long. Margins of petals glabrous 7. R. multiscapa Leaflets ovate to elliptic, pergamentaceous to coriaceous, apex usually acuminate, sometimes obtuse, acute or cuspidate. Petiolule 7-22 mm long, if shorter, then al- Mau mLOreseencesmamillOrousieser. 36. ok Ss see sae ae Gee 9. R. ramiflora Leaflets obovate to elliptic, coriaceous, apex obtuse to distinctly acuminate, then always abruptly narrowing. Petiolule a pulvinus only or up to 10 mm long 6. R. koordersii . Leaflets ovate, index 3.2—4.5. Petals pilose outside, glabrous inside 4. R. elongata Leaflets elliptic, index 2.1—2.8. Petals pilose on both sides .... 1. R. applanata BeeRL SHCULES SIADTOUS. « ..0,o¢.,decad aspen ares SIS A CARTR OPES CRUE St rea 10 Prose sutures densely Setose 2s. L IHRE. osc. sans ee see ee 4. R. elongata meape Olinnt less than jn OMS 5...) opm nk Su oe i Oe, Sank 1] Sipe Of fruit longer fhan3 Mm: fo... cela ss aes BAR ee en ee 14 . Stipe of fruit present. Apex of leaflets acute to caudate; midrib prominent above 12 Stipe of fruit absent. Apex of leaflets obtuse with a slightly emarginate tip; midrib flamened to slishtly sunken above! . / <> Ahab ohne 8. R. obtusa a. Petiolule usually a pulvinus only, up to 13 mm long. Apex of leaflets acute to cuspi- aber, Pb irs Qa iat, Martens Rae tL aoe ee desc Fe ES. Petiolule 9-18 mm long. Apex of leaflets caudate............. 5. R. gracilipes a. Fruit reniform to depressed globose, 1.7—2.5 by 2—2.5 cm. Arillode present. Pulvi- HHS jobleatlets 3-2 mim lon ge ivetnncs. - . . «\ncesenie were ee eee 10. R. robertsonii Fruit obcordate, 1.2—1.5 by 1.2—1.7 cm. Sarcotesta present. Pulvinus of leaflets 5— 1 Seal LOWE ae ee re ere eee ere ne ee LE A Ree a 1. R. applanata Leaflets ovate to elliptic, pergamentaceous to coriaceous, apex usually acuminate, sometimes obtuse, acute or cuspidate. Petiolule 7-22 mm long, if shorter, then al- WaYS- INEOLESCEUGES FANATIGEOMS ts a acs koe cu ees na wie sn ee 9. R. ramiflora Leaflets obovate to elliptic, coriaceous, apex obtuse to acuminate. Petiolule usually a palyinus Onlyoup PorkO mmdOne yn. . 8 ie hk Oh ee hays 7. R. Kkoordersii 706 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1. Rhysotoechia applanata Etman, Blumea 39 (1994) in press. — Type: Floyd & Hoogland 3834 (A, CANB, L, LAE, US), Papua New Guinea. Tree, 15-20 m high, dbh c. 45 cm, outer bark dark brown, inner bark orange brown, heartwood pinkish brown. Branchlets slightly rough, reddish brown with green dots to greyish black; flowering twigs 5—6.5 mm thick. Leaves 1—3-jugate, some- times with a terminal leaflet; petiole 2.5—7 cm long, terete to flattened above, sometimes slightly winged below the lowermost pair of leaflets, slightly ribbed, glabrous; rachis 3.5—7 cm long, terete to angular, slightly winged below the jugae, slightly ribbed, glabrous; petiolule a pulvinus only, 5-13 mm long, grooved, glabrous. Leaflets opposite to suboppo- site, elliptic, 9.5-20 by 4-8.5 cm, index 2.1—2.8, pergamentaceous, both sides glabrous; base sym- metric, usually acute, sometimes slightly attenu- ate; margin (slightly) recurved; apex acuminate; midrib prominent, slightly ribbed to smooth, lat- eral nerves 1-3.2 cm apart, sometimes slightly sunken above, basally indistinctly, apically distinct- ly looped, intercalated veins curved towards the base, veins laxly reticulate, on upper surface flat- tened to slightly raised. Inflorescences axillary, thyrsoid, branching at the base, to 3 cm long, den- sely puberulous; cymules 1-flowered; bracts and bracteoles to 1 mm long; pedicels 1—1.5 mm long. Flowers only seen in bud. Sepals glabrous on both sides, margin glabrous to very laxly ciliate, outer ones ovate, inner ones orbicular. Petals broadly el- liptic, outside pilose only at the base, inside pilose towards the base; claw very short, glabrous; mar- gins coarsely lobed, pilose towards the base; apex rounded; scales as folded margins of petal, thick- ened, velutinous. Disc glabrous. Stamens 8; fila- ments velutinous; anthers laxly puberulous. Pis- til: ovary sericeous; style and stigma very imma- ture. Fruits obcordate, 1.2—1.5 by 1.2—1.7 cm, out- side rugose, very wrinkled, glabrous, inside densely papillose, stipe 2-3 mm high, lobes 1—3, well de- veloped, 1.2—1.4 by 0.6—0.7 cm; style 0.5—1 mm long. Seeds ellipsoid to obovoid, 1—-1.3 by 0.7-0.8 cm, sarcotesta covering the lower part of the seed; hilum 0.5—1 mm diam.; pseudohilum 1—1.2 cm diam. Distribution — Malesia: Papua New Guinea (Owen Stanley Range). Habitat & Ecology — Coastal rain forest or hill forest; altitude from sea level up to 400 m. FI., fr. Sept. 2. Rhysotoechia bilocularis Etman, Blumea 39 (1994) in press. — Type: van Royen 4710 (A, CANB, L), Irian Jaya. Small tree, c. 4m high, dbh c. 10 cm. Branch- lets slightly rough, (reddish) brown; flowering twigs 4-5 mm thick. Leaves 4-jugate, sometimes with a terminal leaflet; petiole 6-8 cm long, terete to flattened above, smooth, glabrous; rachis 13— 19 cm long, terete to flattened above, very slightly winged below the jugae, smooth, glabrous; peti- olules 0.5—1.2 cm long, flattened, glabrous; pulvi- nus 2-4 mm long, grooved, glabrous. Leaflets sub- opposite to alternate, ovate to elliptic, 7-19.5 by 3-6.5 cm, index 2.3—3.1, (thin) pergamentaceous, both sides glabrous; base usually symmetric to slightly oblique, attenuate; margin (slightly) re- curved; apex not abruptly narrowing, acuminate, sometimes acute; midrib slightly prominent, an- gular, lateral nerves 0.5—1.5 apart, almost perpen- dicular to the midrib towards the base, marginally distinctly looped, intercalated veins not distinctly curved towards the base, veins very densely retic- ulate, on upper surface slightly raised. Inflorescenc- es axillary, thyrsoid, branching at the base, to 7 cm long, laxly puberulous; copiously flowering; bracts and bracteoles 0.5—1 mm long; pedicels 1— 2 mm long. Flowers only seen in bud. Sepals: both sides glabrous, margin laxly ciliate, outer ones broadly elliptic, inner ones orbicular. Petals broadly cate, outside glabrous, inside pilose; claw distinct; m irgins coarsely lobed, thickened and recurved towards the base, pilose; apex rounded; scales ab- sent. Disc glabrous. Stamens 7 or 8; filaments ve- lutinous; anthers puberulous. Ovary 2-locular, seri- ceous; style and stigma very immature. Fruits not observed. Distribution — Malesia: Irian Jaya (Merauke Dist.). Habitat & Ecology — River bank in primary rain fores; altitude 70 m. FI. Aug. 3. Rhysotoechia congesta Etman, Blumea 39 (1994) in press. — Type: NGF 29309 (A, CANB, L, LAE), Papua New Guinea. Tree, 6-21 m high, dbh 42 cm, not buttressed, with somewhat gnarled divided trunk, narrow crown of erect branches, outer bark dark brown with minute fissures, inner bark light brown. Branch- lets rough, greenish brown to greenish black; flow- ering twigs 10-15 mm thick. Leaves 1—3-jugate; petiole absent; rachis to 20 cm long, terete to flat- tened below the jugae, sometimes slightly winged below the jugae, ribbed, glabrous; petiolule a pul- vinus only, 5—8 mm long, flattened above, glabrous. Leaflets opposite, elliptic to obovate, 12-30 by 6— 14cm, index 1.6—2.8, + coriaceous, both sides gla- brous; base distinctly oblique, obtuse to rounded; margin (strongly) recurved; apex acute to obtuse; midrib flattened, slightly sunken, ribbed, lateral Adema, Leenhouts, Van Welzen — Sapindaceae 707 f a it ij Wessensorp ‘9 Fig. 68. Rhysotoechia congesta Etman. Habit (NGF 29309). nerves 2—5 cm apart, usually slightly sunken, ba- sally indistinctly, apically distinctly looped, inter- calated veins curved towards the base, veins very laxly reticulate, on upper surface slightly raised. Inflorescences axillary or subterminal or rami- florous, paniculate, branching at the base, very laxly puberulous to glabrous when old; bracts, bracte- oles, pedicels and flowers not observed. Fruits glob- ular with an emarginate apex, attenuate at the base, 2-2.5 by 1.8—2.3 cm, outside slightly rugose, gla- 708 Flora Malesiana ser. I, Vol. 11 (3) (1994) nnn nnn yaEE a anna brous, inside papillose; stipe 3-5 mm high; lobes 1-3, well developed, 1.5—1.7 by 1—1.1 cm; style c. 1 mm long. Seeds ovoid, c. 12 by 10 mm; hilum c. 4 mm diam.; pseudohilum c. 6 mm diam. — Fig. 68. Distribution — Malesia: Papua New Guinea (E Highlands and Morobe Prov.). Habitat & Ecology — Forest; altitude 210-1260 m. Fl. Jan.; fr. Nov. 4. Rhysotoechia elongata Radlk., Bot. Jahrb. 56 (1920) 291; in Engl., Pflanzenr. 98 (1933) 1211; Etman, Blumea 39 (1994) in press. — Lecto- type (Etman 1994): Fitzgerald s.n., New Gui- nea. Tree or shrub, up to 2 m high. Branchlets smooth, greyish brown to greyish black; flowering twigs 2-6 mm thick. Leaves 2—4-jugate; petiole 4— 12 cm long, terete to flattened above, sometimes ribbed, glabrous; rachis 4-34 cm long, angular, slightly winged below the jugae, ribbed, glabrous; petiolules 1-10 mm long, (slightly) grooved, gla- brous, pulvinus 1-3 mm long, grooved, glabrous. Leaflets opposite to subopposite, ovate, 10-27 by 3-7 cm, index 3.24.5, papery to (thin) pergamen- taceous, both sides glabrous; base symmetric or slightly oblique, acute to attenuate; margin slight- ly recurved; apex usually abruptly narrowing but not very distinctly so, acute to cuspidate; midrib prominent, angular, lateral nerves 0.6—2 cm apart, almost perpendicular to the midrib towards the base, marginally distinctly looped, intercalated veins not distinctly curved towards the base, veins very densely reticulate, on the upper surface slightly raised. Inflorescences axillary, thyrsoid, not branch- ing at the base, to 2.5 cm long, puberulous; bracts and bracteoles 0.5—1 mm long; pedicels 1-3 mm long. Flowers c. 5 mm in diam. Sepals: both sides glabrous, margin very laxly ciliate; outer ones broadly ovate, c. 2 by 1.5 mm; inner ones elliptic, c. 3 by 2 mm. Petals obovate, 2-3 by 1-2 mm, outside velutinous, proximally pubescent, inside glabrous; claw c. 1 mm high, both sides densely velutinous; margins lobed, slightly recurved to- wards the base, glabrous; apex obtuse; scales as folded margins of petal, lobed, 0.3-0.6 mm high, glabrous. Disc glabrous. Stamens 8; filaments c. 2 mm long, velutinous; anthers c. 1 mm long, laxly puberulous. Pistil immature; ovary sericeous; style and stigma not observed. Fruits obcordate, 1—1.5 by 1—1.2 cm, outside rugose, laxly puberulous, in- side smooth, along the sutures densely setose; stipe 1-3 mm high; lobes 3, well developed, c. 6 by 5 mm; style not observed. Seeds not observed. Distribution — Malesia: Papua New Guinea (Owen Stanley Range). Habitat & Ecology — (Poor) lowland forest; al- titude 100-360 m. FI. June—Sept.; fr. July. 5. Rhysotoechia gracilipes Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 347; Bot. Jahrb. 56 (1920) 291; in Engl., Pflanzenr. 98 (1933) 1215; Etman, Blumea 39 (1994) in press. —Type: MacGregor s.n. (M, MEL), Papua New Guinea. Branchlets smooth to slightly rough, reddish black to brownish black; flowering twigs c. 4 mm thick. Leaves 1—4-jugate; petiole 6.5—8 cm long, terete, very slender, ribbed, glabrous; rachis 12— 16 cm long, terete, very slender, ribbed, glabrous; petiolules 11.8 cm long, flattened, glabrous, pul- vinus 3-8 mm long, distinctly grooved, glabrous. Leaflets opposite, usually ovate, sometimes ellip- tic, 10-15 by 4.5—6.8 cm, index 2.1—3.5, papery to thin pergamentaceous, both sides glabrous; base symmetric, attenuate; margin not recurved; apex abruptly narrowing, caudate; midrib prominent, distally flattened, slightly ribbed to smooth, later- al nerves 0.9-1.8 cm apart, marginally looped, in- tercalated veins curved towards the base, veins densely reticulate, on upper surface slightly raised. Inflorescences subterminal, not branching at the base; bracts, bracteoles, pedicels and flowers not observed. Fruits obcordate, c. 1.5 by c. 1.2 cm, outside rugose, laxly puberulous, inside densely papillose; stipe c. 3 mm high; lobes 3, well devel- oped, 0.8-1.5 by 0.50.8 cm; style c. 1 mm long. Seeds c. 1.2 cm long; hilum c. 2 mm diam; pseu- dohilum c. 3 mm diam. Distribution — Malesia: Papua New Guinea (Central Prov.). Ecology — Fr. Dec. Note — All vegetative parts look rather vulnera- ble and slender. The rachis, petiole and petiolule are, in some specimens, only 1 mm thick. 6. Rhysotoechia koordersii Radlk. in Engl. & Prantl, Nat. Pflanzenfam., Nachtr. 3 (1907) 206; in Fedde, Rep. 18 (1922) 343; in Engl., Pflan- zenr. 98 (1933) 1213; Etmam, Blumea 39 (1994) in press. — Type: Koorders 18844 (BO, K, L, M), Menado. Rhysotoechia mortoniana auct. non (F. Muell.) Radlk.: Koord., Minah. 19 (1898) 407. Tree, up to 19 m high, dbh 2-25 cm, outer bark smooth, dark green to blackish brown, inner bark yellowish to ochre, thin. Branchlets rough to smooth, greenish brown to greyish black to red- dish black; flowering twigs 3-6 mm thick. Leaves |—-4-jugate; petiole 2-11 cm long, slightly angular to terete, sometimes slightly winged below the low- Adema, Leenhouts, Van Welzen — Sapindaceae ermost pair of leaflets, (slightly) ribbed, glabrous; rachis 2.5—20 cm long, angular, slightly winged below the jugae, ribbed, glabrous; petiolule usual- ly a pulvinus only, up to 10 mm long, slightly grooved, glabrous; pulvinus 2—5 mm long, grooved, glabrous. Leaflets usually subopposite, sometimes opposite, obovate to elliptic, 6-23 by 1-11 cm, index 2—3.7, coriaceous, both sides glabrous; base sometimes very slightly oblique but never distinctly so, acute to attenuate; margin (not) slightly re- curved; apex obtuse to distinctly acuminate, then always abruptly narrowing; midrib prominent, smooth to angular, lateral nerves 0.5—3 cm apart, basally indistinctly, apically distinctly looped, in- tercalated veins curved towards the base, veins laxly reticulate, on the upper side slightly raised. /nflo- rescences axillary, thyrsoid, branching at the base, to 19 cm long, laxly puberulous; cymules 1|-flow- ered; bracts and bracteoles 0.5—1 mm long; pedi- cels 4-9 mm long. Flowers c. 7 mm in diam. Se- pals: both sides glabrous, margin ciliate; outer ones broadly ovate, 3—2.5 by 2.5 mm; inner ones orbic- ular, 3.5—3 by 3 mm. Petals broadly obovate, c. 2.5 by 2 mm; outside pilose, inside laxly pilose; claw c. 0.8 mm high, both sides pilose; margins coarsely lobed, slightly thickened towards the base, densely pilose; apex rounded; scales absent. Disc glabrous. Stamens 8; filaments c. 3 mm long, ve- lutinous towards the base; anthers c. 0.5 mm long, puberulous. Ovary c. | mm high, densely sericeous; style c. 2 mm high, laxly puberulous. Fruits ob- cordate, attenuate at the base, 1.5—1.7 by 1.3-1.5 cm, outside rugose, very laxly puberulous, inside smooth; stipe slender, 5—7 mm high; lobes 1—3, well developed, 0.8—1 by 0.6-0.7 cm; style c. 0.5 mm long. Seeds not observed. Distribution — Malesia: Borneo (Kalimantan: Kota Belud and Beluran District), Celebes (Mali- li, Minahassa). Habitat & Ecology — Forest; altitude from sea level up to 700 m. FI. Apr.; fr. May. Note — Rhysotoechia koordersii shows an over- all similarity with R. ramiflora, although there are a few differences. These are best observed in the specimens of both species from Celebes. The over- lap in the third character in the table below is caused by the R. koordersii specimens from Kalimantan. The differences are, in my opinion, large enough for RK. koordersii and R. acuminata to remain sep- arate species, although they must be closely relat- ed. leaflet form R. ramiflora R. koordersti ovate to elliptic obovate to elliptic 709 7. Rhysotoechia multiscapa Etman, Blumea 39 (1994) in press. — Type: Carr 14999 (CANB, L), Papua New Guinea. Tree, up to 16.5 m high. Branchlets rough, grey- ish black; flowering twigs 4—7 mm thick. Leaves 2—4-jugate; petiole 2-6 cm long; rachis 2—8.5 cm long; petiole and rachis terete to flattened above, ribbed, glabrous; petiolules 5-10 mm long, flat- tened to grooved towards the pulvinus, glabrous, pulvinus 3—5 mm long, grooved, glabrous. Leaf- lets opposite to subopposite, usually elliptic, some- times obovate or ovate, 8—14 by 2.5—5 cm, index 2.3—3.6, coriaceous, both sides glabrous; base sym- metric, not abruptly narrowing, acute to attenuate; margin not to slightly recurved; apex acute to acu- minate, then slightly abruptly narrowing; midrib prominent, smooth to angular, very laxly puberu- lous, lateral nerves 0.5—1.8 cm apart, basally in- distinctly, apically distinctly looped, intercalated veins curved towards the base, veins densely retic- ulate, on upper surface slightly raised. /nflorescenc- es axillary or subterminal, thyrsoid, branching at the base, to 9 cm long, very laxly puberulous, co- piously flowered; cymules 1- or 2-flowered; bracts and bracteoles c. | mm long; pedicels c. 2 mm long. Flowers c. 4 mm in diam. Sepals: both sides gla- brous, margin very laxly ciliate, outer ones broad- ly ovate, 1.5—1.7 by 1.8—2 mm, inner ones orbicu- lar, 2.2-2.5 by 2 mm. Petals broadly obovate, | .8— 2 by 2—2.4 mm, both sides pilose towards the base; claw 0.5—0.6 mm high, inside glabrous, outside densely pilose; margins coarsely lobed, thickened towards the base, sometimes slightly recurved to- wards the base, glabrous; apex rounded; scales absent. Disc glabrous. Stamens 8; filaments 2.2— 2.5 mm long, velutinous; anthers c. 0.5 mm long, glabrous to very laxly puberulous. Pistillode seri- ceous. Fruits not observed. — Fig. 69. Distribution — Malesia: Papua New Guinea (Owen Stanley Range). Habitat & Ecology — Forest; altitude 1280 m. Fl. Nov. 8. Rhysotoechia obtusa Etman, Blumea 39 (1994) in press. — Type: NGF 27986 (A, CANB, L, LAE), Papua New Guinea. Tree or shrub, 4-6 m high, outer bark grey to dark grey to light brown, middle bark light green, inner bark cream, wood of medium hardness, straw- leaflet apex petiolule 7-22 mm 0-10 mm acute to cuspidate obtuse to acuminate 710 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1mm Fig. 69. Rhysotoechia multiscapa Etman. a. Male flower; b. ibid., one sepal removed; c. ibid., one petal left; d. sepal; e. petal (a—e: Carr 14999). coloured. Branchlets rough, greenish black to grey- ish black to reddish black; flowering twigs 4-7 mm thick. Leaves 1—4-jugate, often with a terminal leaf- let; petiole 2.5—6 cm long, terete to angular, usual- ly slightly winged below the lowermost pair of leaf- lets, ribbed, glabrous; rachis 2.5—10.5 cm long, angular, slightly winged below the jugae, ribbed, glabrous; petiolules 6-11 mm long, grooved, gla- brous; pulvinus 2-5 mm long, grooved, glabrous. Leaflets opposite to subopposite, ovate to elliptic, rarely obovate, 7-16 by 3-6 cm, index 2.6-3.1, (thick) pergamentaceous, both sides glabrous; base sometimes very slightly oblique, usually abruptly narrowing, attenuate; margin (slightly) recurved; apex obtuse to slightly emarginate; midrib flattened to slightly sunken, angular, lateral nerves 0.7—2.5 apart, basally indistinctly, apically distinctly looped, intercalated veins curved towards the base, veins laxly reticulate, on upper surface slightly raised. Inflorescences axillary or subterminal, pan- iculate, branching at the base, to 4 cm long, puber- ulous; bracts and bracteoles to 1 mm long; pedi- cels 1.5-2 mm long. Flowers not observed. Fruits 1.5-1.8 by 2-2.5 cm, outside smooth to slightly rugose, laxly strigose, laxly papillose, inside dense- ly papillose; stipe absent; lobes 3, well developed, 0.9-1.4 by 1.1-1.3 cm; style not observed. Seeds immature or only partly present, ellipsoid; hilum 0.5 mm diam.; pseudohilum 5 mm diam. Distribution — Malesia: Papua New Guinea (Central Prov.). Habitat & Ecology — Low lying country in or at the edge of mixed secondary forest; often subject to flooding. Altitude about sea level. Fl. Sept.; fr. Dec.—Jan. 9. Rhysotoechia ramiflora Radlk., Sapind. Holl.- Ind. (1879) 19, 62; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 542, 657; in Engl., Pflanzenr. 98 (1933) 1214; Etman, Blumea 39 (1994) in press. — Type: Beccari it. sec. 10 (M), Celebes. Rhysotoechia acuminata Radlk., Philipp. J. Sc., Bot. 8 (1913) 465, 466; Merr., Enum. Philipp. Adema, Leenhouts, Van Welzen — Sapindaceae Flow. Pl. 2 (1923) 510; Radlk. in Engl., Pflan- zenr. 98 (1933) 1213. — Type: Loher 5882 (K, M), Luzon. Rhysotoechia striata Radlk., Philipp. J. Sc., Bot. 8 (1913) 466; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 510; Radlk. in Engl., Pflanzenr. 98 (1933) 1213. — Lectotype (Etman 1994): Clemens 1067 (M), Mindanao. Paratypes: Cle- mens ‘K’, 778, 978. Tree or shrub, up to c. 8 m, dbh 3-30 cm, sparse- ly branched, bark smooth, grey brown, thin. Branchlets smooth to slightly rough, greyish black to reddish black to light brown; flowering twigs 4-13 mm thick. Leaves 1—4-jugate; petiole 3-17 cm long, terete to flattened above, sometimes slight- ly winged below the first jugum, (slightly) ribbed, glabrous; rachis 4.5—22 cm long, terete to flattened or slightly angular above, usually slightly winged below the jugae, (slightly) ribbed, glabrous; peti- Olules 0.7—2.2 cm long, flattened to slightly grooved, glabrous, pulvinus 3-10 mm _ long, grooved, glabrous. Leaflets opposite to suboppo- site, rarely alternate, ovate to elliptic, rarely obo- vate, 9-31 by 4-12 cm, index 1.8—3.3, pergamen- taceous to coriaceous, both sides glabrous; base symmetric (or slightly oblique), attenuate, rarely acute; margin not to slightly recurved; apex acu- minate, sometimes obtuse, acute or cuspidate, of- ten abruptly narrowing; midrib prominent, angu- lar to ribbed, lateral nerves 1—3 cm apart, basally indistinctly, apically distinctly looped, intercalat- ed veins curved towards the base, veins laxly re- ticulate, on upper surface slightly raised. Inflores- cences axillary, subterminal or ramiflorous, thyr- soid, branching at the base or not, to 13 cm long, very laxly puberulous to glabrous; cymules 1-flow- ered; bracts and bracteoles 0.5—1 mm long; pedi- cels 2-4 mm long. Flowers 6-8 mm in diam. Se- pals glabrous except for (laxly) ciliate margins, outer ones broadly elliptic, c. 2.5 by 2 mm, inner ones broadly ovate, c. 3.5 by 2.5 mm. Petals broadly obovate, 1.6—2 by 1.8—2 mm, outside (laxly) pi- lose, inside glabrous, claw 0.2-0.4 mm high, mar- gins coarsely lobed, apex rounded; scales absent. Disc glabrous. Stamens 8; filaments 3—5 mm long, velutinous towards the base; anthers c. | mm long, puberulous. Ovary c. | mm long, densely sericeous; Style and stigma c. 2 mm long, the first laxly pu- berulous. Fruits obcordate, 1.8—2.5 by 1.6—2.1 cm, outside rugose to ribbed, laxly puberulous, inside (densely) papillose; stipe 0.5—1 cm long; lobes 1- 3, well developed, 1—1.5 by 0.7—0.9 cm; style 0.5- 1 mm long. Seeds ellipsoid or ovate, 1—1.7 cm by 6—10 mm; hilum 2—3 mm diam.; pseudohilum 0.4— 1.1 cm diam. Distribution — Malesia: Borneo (E Kalimantan), 711 Philippines (Luzon, Samar, Mindanao, Sulu), Celebes. Habitat & Ecology — Mixed dipterocarp forest, in forest margins, river banks or forest slopes or limestone hills; altitude 30-600 m. Fl. May—June and Oct.—Nov.; fr. Feb.—May. Note — This species is very variable in a number of characters, for example: form and thickness of the leaflets, smoothness of petiole and rachis, way in which the inflorescences are borne on the branch- es, number of cells developed in the fruits, and length of the stipe. 10. Rhysotoechia robertsonii (F. Muell.) Radlk., Sitzungsber. Mat.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 542, 657; in Engl., Pflanzenr. 98 (1933) 1214; S.T. Rey- nolds, Austrobaileya 2 (1984) 42; in Fl. Aus- tral. 25 (1985) 65. — Cupania robertsonii F. Muell., Fragm. Phyt. Austral. 5 (1866) 146; Etman, Blumea 39 (1994) in press. — Type: Dallachy s.n. (L, M, MEL), Rockingham Bay, Australia. Rhysotoechia contermina Domin, Bibl. Bot. 89 (1927) 905. — Type: Domin s.n., p.p., Harvey’s Creek, Australia. Tree, 6-25 m high, dbh 10-75 cm, low but- tressed, outer bark patchy grey to dark grey to brown to green, (moderately) smooth except for the fine cracking and stippling, scrape red-brown, inner bark fibrous, cream. Branchlets rough to smooth, greenish brown to greyish brown; flower- ing twigs 4-6 mm thick. Leaves 1—4-jugate; peti- ole 1.3—6 cm long, terete to flattened above, slightly winged below the lowermost pair of leaflets, slight- ly ribbed, very laxly puberulous to glabrous; ra- chis 2.5—8 cm long, angular, slightly winged be- low the jugae, slightly ribbed, laxly puberulous to glabrous; petiolule usually a pulvinus only, up to 4 mm long, flattened above, very laxly puberulous; pulvinus |—2 mm long, slightly grooved, very lax- ly puberulous. Leaflets opposite to alternate, ovate to elliptic, 6-20 by 2-7 cm, index 2.2-4.8, perga- mentaceous, both sides very laxly puberulous; base sometimes slightly oblique, acute to attenuate; margin (slightly) recurved; apex sometimes abrupt- ly narrowing, acute to cuspidate; midrib prominent, angular, lateral nerves 0.5—1.5 apart, basally indis- tinctly, apically distinctly looped, intercalated veins not very distinct, veins densely reticulate, on up- per surface slightly raised. /nflorescences axillary, paniculate, 10.5-17.5 cm long, laxly puberulous; bracts and bracteoles 0.5—0.7 mm long; pedicels 1-2 mm long. Flowers 4-5 mm in diam. Sepals: both sides glabrous, margin ciliate; outer ones broadly ovate, 1.5—2 by 1—1.2 mm; inner ones or- Tae bicular to elliptic, 2-2.5 by 1.5—2 mm. Petals obo- vate, 1.2-1.5 by 1—-1.5 mm, outside pilose only at the base, inside pilose towards the base; claw c. 0.2 mm high, glabrous; margins lobed, thickened towards the base, pilose; apex rounded; scales ab- sent. Disc glabrous. Stamens 8; filaments c. 3 mm long, velutinous towards the base; anthers c. 0.8 mm long, sometimes with a few hairs. Ovary c. 2 mm high, laxly sericeous towards the style; style c. | mm high, laxly sericeous; stigma 0.5—0.7 mm high, laxly strigose. Fruits reniform to depressed globose, 1.7—2.5 by 2—2.5 cm; outside rugose, gla- brous, inside densely papillose; stipe 2-3 mm high; lobes 3, well developed, 1—1.2 by 1.2—2.2 cm; style c. 1 mm long. Seeds obovoid, 1—1.3 by 0.8-1 mm; hilum c. 0.5 mm diam.; pseudohilum c. 2 mm diam. Distribution — Malesia: Papua New Guinea (Western Prov.); Australia (Queensland). Habitat & Ecology — Found in rain forests from low hills to riverine, and on and near creek banks. Soil: alluvial, dark clay loam; altitude 20-450 m. Vegetation with Eugenia, and Callistemon as dom- inant species. Fl. July—Oct.; fr. Oct.—Dec. IMPERFECTLY KNOWN SPECIES Rhysotoechia grandifolia Radlk., Sapind. Holl.- Ind. (1879) 89, 132, 249; Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 542, 657; in Engl., Pflanzenr. 98 (1933) 1212; Etman, Blumea 39 (1994) in press. — Lectotype (Etman 1994): Korthals 1] (L), Bor- neo. Paratype: Teijsmann 7488, Moluccas, Ge- ben. Rhysotoechia grandiflora Radlk. ex Merr., J. Str. Br. Roy. As. Soc., special issue (1921) 361, mis- spelled name. Shrub, up to 2 mhigh. Branchlets slightly rough, reddish black to brownish black; flowering twigs c. 6 mm thick. Leaves 2—4-jugate, rarely with one subterminal leaflet; petiole 4-10 cm long; rachis 3-20 cm long; petiole and rachis terete to flattened above, slightly ribbed, glabrous; petiolules 1.5—2 cm long, flattened to slightly grooved, glabrous; pulvinus 5~7 mm long, grooved, bulbously thick- ened, glabrous. Leaflets opposite, rarely suboppo- site, ovate to elliptic, 13-24 by 6.5—9.5 cm, index 2.13, + coriaceous, both sides glabrous; base sym- metric, sometimes slightly oblique, usually abruptly narrowing, attenuate; margin not to slightly re- curved; apex acuminate; midrib prominent, ribbed, lateral nerves 1.5—3 cm apart, basally indistinctly, apically distinctly looped, intercalated veins curved towards the base, veins laxly reticulate. /nflores- cences, flowers and fruits not observed. Flora Malesiana ser. I, Vol. 11 (3) (1994) Distribution — Malesia: Borneo (Kalimantan), Moluccas (Geben, Obi Island, W Part, Jikodolong). Habitat & Ecology — Rather open coastal for- est, transition zone between coral beach sand and red porous nickel soil; altitude 30 m. Fr. Nov. Note — None of the three collections studied has inflorescences or fruits. The foliar characters of the collection made by Korthals are remarkably simi- lar to R. acuminata. The leaves (including the pul- vini) in the other two collections (made by De Vo- gel and Teijsmann) of R. grandifolia are much thicker than in R. acuminata. According to Radl- kofer, R. grandifolia has sessile fruits, in contrast with R. acuminata whose fruits are distinctly stipi- tate. Also, the pulvini of R. grandifolia are “bul- bously thickened’ and not normally thickened as in R. acuminata. This character cannot be observed in the Korthals-collection. Although I doubt whether R. grandifolia can be separated from R. acuminata, the absence of flowers and fruits in the specimens studied keeps me from concluding that R. grandifolia and R. acu- minata are conspecific. Rhysotoechia longipaniculata Kanehira & Hatusi- ma, Bot. Mag. Tokyo 57 (1943) 79, f. 13. — Type: Kanehira & Hatusima 11534, New Gui- nea, Nabire (n.v.). Rhysotoechia momiensis Kanehira & Hatusima, Bot. Mag. Tokyo 57 (1943) 81, f. 13. — Type: Kanehira & Hatusima 14148, New Guinea, Momii (n.v.). Rhysotoechia spec. Tree, up to 1.8 m high. Branchlets smooth to slightly rough, brownish black to greyish black; flowering twigs c. 5 mm thick. Leaves 3—5-jugate; petiole 5~10 cm long, terete, smooth, glabrous; rachis 8-17 cm long, terete, sometimes slightly winged below the jugae, smooth, glabrous; peti- olules 5—15 mm long, slightly grooved, glabrous; pulvinus 1-4 mm long, grooved, sometimes wrin- kled, glabrous. Leaflets opposite to subopposite, elliptic, 10-22 by 4-7 cm, index 2.3—3.1, (thick) pergamentaceous, both sides glabrous; base sym- metric, attenuate; margin not to slightly recurved; apex abruptly narrowing, acuminate to cuspidate; midrib flattened to prominent, smooth to ribbed, lateral nerves 0.5—2.5 cm apart, basally open, api- cally looped, intercalated veins curved towards the base; veins densely reticulate, on upper surface slightly raised. Inflorescences axillary, thyrsoid, branching at the base, to 8 cm long, very laxly pu- berulous; cymules 1-flowered; bracts and bracte- oles 0.1-0.3 mm long; pedicels 1—1.5 mm long. Adema, Leenhouts, Van Welzen — Sapindaceae Wis Flowers only seen in bud. Sepals: both sides gla- brous, margin laxly ciliate, outer ones broadly ovate, inner ones orbicular. Petals orbicular, out- side velutinous towards the base, inside pilose to- wards the base; claw absent to very short; margins lobed, glabrous; apex rounded; scales free, not very distinct, pilose. Disc glabrous. Stamens 8; filaments densely velutinous towards the base; anthers gla- brous. Ovary 3-locular, glabrous; style and stigma very immature, glabrous. Fruits not observed. Distribution — Malesia: Irian Jaya (Waigeo Is- land). Habitat & Ecology — Upper stretches of creek in primary forest; altitude c. 30 m. Fl. Feb. Note — This may be a new species, but the ma- terial is too scanty to be certain of this. SAPINDUS (P.W. Leenhouts) Sapindus L., Gen. Pl. ed. 5 (1754) 171; Sp. Pl. (1753) 367; Radlk. in Engl., Pflanzenr. 98 (1932) 630-668. — Type species: Sapindus saponaria L. Electra Norofia, Verh. Bat. Gen. K. W. 5 (1791), nom. nud. — Dittelasma Hook. f. in Benth. & Hook. f., Gen. Pl. 1 (1862) 395. — Type species: Dittelasma rarak (DC.) Hook. f. [= Sapindus rarak DC.}. Trees, monoecious. /ndumentum of solitary, simple hairs. Leaves paripinnate (in young specimens sometimes imparipinnate), in Malesian species up to 13-jugate; no pseudo- stipules; petiole and rachis marginated to winged or not. Leaflets often more or less ob- lique to falcate, entire. Inflorescences terminal, thyrsoid, widely branched. Flowers reg- ular or zygomorphic. Calyx 5-merous, sepals free, imbricate, outer 2 smaller, (hardly) petaloid. Petals 4 or 5, imbricate, equal, as long as to slightly longer than the calyx, short-clawed, with a scale which may be nearly as long as the petal itself, reduced to a transverse ridge, or represented by a pair of auricles. Disc annular or semi-annular, gla- brous or hairy. Stamens 8, free, all about equal, not or hardly exserted; filaments hairy at least at the base; anthers ellipsoid, cleft at base. Ovary 3-lobed, 3-celled; style terminal, simple, slender, about as long as the ovary, straight or bent, with 3 stigmatic lines; pistil- lode in male flowers minute. Ovules | per cell, sessile on a thickened angular placenta. Fruits 3-parted, often 1 or 2 parts abortive, breaking up into globular to obovoid drupes, pulp containing much saponin, endocarp pergamentaceous, hairy around the placenta. Seeds without arillode or sarcotesta, globular or ellipsoid, smooth, black, testa bony, hilum linear, slightly impressed. Distribution — 10 species, 2 of which restricted to Central and southern North Amer- ica, a third one throughout tropical America and (autochthonous?) in the Tropics (and Subtropics) of Africa, Asia, and the Pacific; 6 species in continental S and SE Asia, one of these also in western Malesia, one species endemic in Hawaii. — Fig. 70. Habitat & Ecology — Tropical and subtropical regions, as well under everwet as under seasonal conditions; at low to medium altitudes; in forests and thickets. Uses — The fruits of some species are widely used as a substitute for soap (‘Soap nuts’). Notes — 1. Apparently closest related to the African genera Deinbollia Schum. & Thonn. and Hornea Baker. 2. Radlkofer subdivided the genus into four sections. Of these, the following three are represented in Malesia: sect. Sapindus (S. saponaria L.), sect. Dasysapindus Radlk. (S. trifoliatus L., sometimes planted) and sect. Dittelasma (Hook. f.) Radlk. (S. rarak DC.). 714 Flora Malesiana ser. I, Vol. 11 (3) (1994) a ge a a nn Pe a a a KEY TO THE SPECIES la. Leaves 1—5-jugate. Petals 5, the scale represented by a hairy ridge or a pair of auri- cles Discvannular s2.<.cc.%, cee: . 5 08 SP SL CR 2 b. Leaves (7-)9-13-jugate. Petals 4, with a scale of the same shape and size. Disc Ser ATNU A snr, ote hee eee ees ga eke ee 1. S. rarak 2a. Calyx outside with some hairs at the base only. Disc, pistil, and fruit glabrous 2. S. saponaria b. Calyx outside hairy all over. Disc, pistil, and fruit hairy. India and Ceylon. In Malesia Sometimes: planted (Pigs (06) oo icra cyte 2-0) ea inten soe S. trifoliatus L. 1. Sapindus rarak DC., Prod. | (1824) 608; Blume, Rumphia 3 (1847) 91, t. 167; Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 150; At- las 1 (1913) f. 89; Radlk. in Engl., Pflanzenr. 98 (1932) 663; Steenis, Fl. Sch. Indon. (1949) 252; Backer & Bakh. f., Fl. Java 2 (1965) 133. — [Saponaria Rarak Rumph., Herb. Amb. 2 (1741) 134.] — Dittelasma rarak (DC.) Hook. f. in Benth. & Hook. f., Gen. Pl. 1 (1862) 395. — Type: Lahaye s.n. in Herb. Prod., Granata. Sapindus angustifolius Blume, Rumphia 3 (1847) 99. — Type: Van Royen s.n. (L). Tree, up to 42 m high, dbh up to | m. Branch- lets terete, up to 1 cm thick, inconspicuously lenti- cellate, brownish to blackish, fulvous tomentellous, glabrescent. Leaves (7—)9-13-jugate, up to 50 cm long, glabrous; petiole + terete, somewhat flattened and thickened at base, up to 9 cm long; rachis not winged; petiolules 2-5 mm long. Leaflets subop- posite to alternate, lanceolate-ovate, mostly oblique and slightly falcate, 7-16 by 2—3.5 cm, chartaceous; base very oblique, lower half acute, upper half rounded-attenuate to cuneate; apex obtuse to ta- pering acute-acuminate, mucronate; midrib slightly raised above, more strongly so beneath, nerves many, rather dense, oblique, curved, not joined, rather well developed, prominulous on both sur- faces, like the intermediate veins and rather densely reticulate veinlets. Inflorescences up to c. 35 cm long, densely fulvous- to ferrugineous-tomentel- lous. Flowers zygomorphic, white. Sepals flat, hardly petaloid, outside densely appressed fulvous- hairy, outer broad-ovate to suborbicular, 2-3 by 1.5-2 mm, inner obovate, 3-4 by 1.8—2 mm. Pet- als 4, lanceolate-ovate to elliptic, 3 by 1-2 mm, short-clawed, outside densely appressed long ful- vous-hairy, woolly along the margin; scale 1/3 shorter than the petal, of about the same shape, trun- cate and slightly incurved at apex, along the mar- gin and especially at the apex densely woolly. Disc semi-annular, glabrous. Stamens: filaments densely long-hairy outside and along the margin, the api- cal part excepted, in male flowers 2.5 mm, in fe- male ones 1.2—1.5 mm; anthers 0.3—0.5 mm, in male flowers glabrous, in female ones sparsely hairy. Pistil glabrous, 3.5—4 mm. Fruits: parts subglobu- lar, 2 by 1.8 cm, carinate, red, glabrous. Seeds sub- globular, 1.2—-1.5 cm diam. — Fig. 70d. Distribution — Assam, Burma, Thailand, Indo- China, Taiwan; in Malesia: Sumatra (known from a few localities only), Bangka, Malay Peninsula, Java, Madura, Lombok, Sumbawa; widely culti- vated, also in other parts of Malesia. It is difficult to distinguish between wild and naturalized speci- mens. Habitat & Ecology — Forests, mainly under sea- sonal conditions; c. 200-1600 m altitude. Fl. Jan.— Feb., June, Sept.—Nov.; fr. Jan., Feb., Apr.—Aug. Uses — Wood hard, but not durable. Fruits and seeds used like those of S. saponaria. See Heyne, Nutt. Pl. Indon. ed. 3 (1950) 988; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1958. Note — Sapindus pinnatus Mill., Gard. Dict. ed. 8 (1768) has since De Candolle often been cited as a possible synonym. From Miller’s short descrip- tion — based upon seedlings cultivated in England — it is clear, however, that S. pinnatus cannot be identified with S. rarak. He clearly mentions the leaves as being winged; moreover, he says that the species is more hardy than S. saponaria, which, on comparing the areas of distribution of the two species, is hardly to be expected of S. rarak. Ap- parently, his name refers either to seedlings of a race of S. saponaria with 8 or 10 pairs of leaflets, more than usual, or to another genus with seeds like those of Sapindus. 2. Sapindus saponaria L., Sp. Pl. (1753) 36072 Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 364; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 498; Radlk. in Engl., Pflanzenr. 98 (1932) 639, f. 14; P. Royen, Man. For. Trees Papua & New Gui- nea 2 (1964) f. 19. — Lectotype (Pennington 1992): Plukenet, Phytographia (1692) t. 217, mee Adema, Leenhouts, Van Welzen — Sapindaceae 715 Imm lem lcm lcm Fig. 70. Sapindus saponaria L. a. Habit; b. petal; c. fruit. — S. rarak DC. d. Fruit. — S. trifoliatus L. e. Fruit (a, b: Jboet 255; c: Schodde 2779; d: Boerlage s.n.; e: Kostermans 25836). Sapindus mukorossi Gaertn., Fruct. 1 (1788) 342, t. 70, f. 3g, h; Radlk. in Engl., Pflanzenr. 98 (1932) 651; Corner, Wayside Trees (1940) 595, f. 212. — Type unknown. Sapindus forsythii DC., Prod. 1 (1824) 607. — Type: Forsyth s.n. in Herb. Prod., Granata. Quassia tricarpa Blanco, FI. Filip. (1837) 351, p.p.; ed. 3 (1878) t. 388; Radlk., Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 16 (1887) 399. — Neotype: Merrill Sp. Blanc. 183 (BO, L), Luzon. Sapindus vitiensis A. Gray, U. S. Expl. Exp. Bot. 1 (1854) 251; Radlk. in Engl., Pflanzenr. 98 (1932) 655. — Type: U. S. Expl. Exp. s.n. (US), Fiji. Sapindus microcarpus Jardin, Mém. Soc. Sc. Nat. Math. Cherb. 5 (1857) 295, 307; non Ruiz & Pavon (1804). — Sapindus saponaria L. f. mi- crocarpa (Jardin) Radlk. in Martius, Fl. Bras. 13, 3 (1900) 517. — Sapindus saponaria L. var. jJardiniana F.B.H. Brown, Bull. Bish. Mus. 130 (1935) 160. — Type: Jardin s.n., Marquesas, c. 1845. Sapindus balicus Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 396; Sapind. Holl.-Ind. (1879) 7, 20, 67; 716 Flora Malesiana ser. I, Vol. 11 (3) (1994) a —— eee in Engl., Pflanzenr. 98 (1932) 655. — Type: Teijsmann s.n., Bali. ?Sapindus oocarpus Radlk., Not. Syst. 1 (1910) 302; in Engl., Pflanzenr. 98 (1932) 654; Gag- nep. in Fl. Indo-Chine, Suppl. 1 (1950) 939. — Type: Balansa 3430 (P), Tonkin. Tree, up to 25 m high, dbh up to 56 cm. Branch- lets terete, up to 7 mm thick, often conspicuously lenticellate, ashy grey to brown, glabrous to rather densely fulvous-hairy and glabrescent. Leaves |— 5-jugate, up to 40 cm long; petiole terete to 3-an- gular in cross section, marginated to winged to- wards the lower pair of leaflets or not, 1.5—5.5 cm long, hardly to distinctly swollen at base; rachis marginated to winged beneath every pair of leaf- lets or not. Leaflets (sub)opposite, elliptic to lan- ceolate, widest about or below the middle, mostly slightly oblique and falcate, 6-16 by 3—6 cm, chart- aceous, glabrous; base more or less oblique, cuneate; apex emarginate or obtuse to acute; mid- rib flat to prominulous above, prominent beneath, nerves 10-15 pairs, slightly curved to nearly straight, upper ones looped and joined, prominu- lous on both surface as are the often strongly de- veloped intermediate veins and the densely reticu- late veinlets. Inflorescences up to 25 cm long, densely shortly fulvous-tomentose. Flowers regu- lar, cream. Sepals orbicular to broad-ovate, con- cave, mostly with a broad petaloid margin, ciliola- te and with some appressed hairs near the base, outer 1—-1.2 mm in diam., inner 2 by 1.5—2 mm. Petals 5, oblong-ovate to ovate, 1.5—2.5 by 1—1.2 mm, short-clawed, woolly-ciliate and outside at least at the base long hairy, inside above the claw either with a hairy ridge or with two involute, hairy auricles. Disc annular, glabrous. Stamens: filaments 0.5—1 mm, variably hairy; anthers 0.5—1 mm, gla- brous. Pistil glabrous, 2 mm. Fruits: parts subglob- ular, 0.8—1.2 cm in diam., not carinate, glabrous. Seeds subglobular, 0.8—1 cm diam. — Fig. 70a—c. Distribution — Apparently originating from trop- ical and subtropical America (from Florida to N Argentine), widely cultivated and naturalized in the Tropics and Subtropics; in Asia distributed from Central Japan, Quelpaert I. near Korea, and Cen- tral China to India, partly at least under cultiva- tion; in Malesia known from Philippines, Lesser Sunda Islands, and Papua New Guinea (Central Prov., around Rigo only), possibly always intro- duced by man. Habitat & Ecology —In more or less open, most- ly secondary forests at up to 500 m altitude. In Less- er Sunda Islands and New Guinea: fl. Mar—June; fr. May, Aug.; in Philippines: fl. Oct., Nov.; fr. Jan., Mar., Apr. For dispersal (over short distances pos- sibly by sea-currents, but mainly by man) see Rid- ley, Dispersal (1930) 268. Uses — Ornamental tree. Fibres of inner bark used for ropes. Roots, bark, leaves, but especially fruits used as a substitute for soap because of the high amount of saponin; for the same reason the fruits are used as a fish poison. Seeds formerly used as buttons and beads. Several parts used in medi- cine. See Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1959; Brown, Useful Pl. Philipp. 2 (1950) 369; Quisumbing, Philipp. J. Sc. 77 (1948) 162; Walker, Imp. Trees Ryukyu (1954) 197, f. 119. Chromosomes — 2n = 22: Ono, Mem. Nat. Sci. Mus. Tokyo 10 (1977) 63-76; 2n = 30: Mehra et al., Silvae Gen. 21 (1972) 96-102; 2n = 36: Sarkar et al. in Love, Taxon 26 (1976) 636, 649. Note — The delimitation of Sapindus saponaria as given here is wider than Radlkofer’s by the in- clusion of S. balicus, mukorossi, oocarpus, and vitiensis. The distinction between these ‘species’ was very vague. The continental Asian forms — S. mukorossi and allies — are closely related to the American ones. The Malesian and Pacific forms were distinguished by Radlkofer as f. microcar- pus, characterized only by the slightly smaller fruits. There are three distinguishable Malesian races, each rather uniform. In ‘S. balicus’ the pet- iole and rachis are never winged; the leaflets are rather thin, obtuse and often emarginate at the apex, with 10-12 pairs of nerves, the upper ones joined at some distance from the margin, and intercalary veins not strongly developed; the flowers are small with petaloid sepals and only partly hairy petals. In the Philippine race the petiole and rachis are often winged; the leaflets are thicker, often acute to acuminate at apex, with 12-15 pairs of nerves, the upper ones of which are joined quite near the margin, and with often 3 intermediate veins strongly developed between every pair; the flowers are slightly bigger, with petaloid sepals and nearly gla- brous petals. The New Guinea race is character- ized mainly by elliptic, hardly oblique and not fal- cate leaflets, by hardly petaloid sepals, and by pet- als which are outside entirely and rather densely hairy. Adema, Leenhouts, Van Welzen — Sapindaceae PLT SARCOPTERYX (P.C. van Welzen) Sarcopteryx Radlk., Sapind. Holl.-Ind. (1879) 19, 57, 97, 98; Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 486-490, 500, 544, 658, 659; in Engl., Pflanzenr. 98 (1933) 1232-1238, f. 36; S.T. Reynolds, Austrobaileya 2 (1984) 53-57, f. 5; in Fl. Austral. 25 (1985) 72-77, f. 18a—f, map 92-96; Welzen, Blumea 36 (1991) 87-103, figs., maps. — Lectotype species (S.T. Reynolds 1984): Sarcopteryx squamosa (Roxb.) Radlk. Shrub to tree. /ndumentum of simple solitary hairs and red glandular hairs only. Branch- lets smooth to slightly grooved. Leaves paripinnate, 1|—7-jugate, pseudo-stipules absent, not winged; petiolules usually present. Leaflets: base attenuate; margin entire; both sur- faces smooth; on lower surface domatia at most shallow pockets in axils of nerves, small red glands present; venation raised; nerves especially towards the apex marginally looped: veins reticulate. /nflorescences usually in the upper leaf axils and pseudoterminal, usual- ly only with a few branches; cymules cincinnate (to dichasial). Flowers apparently bisex- ual but presumably functionally male or female. Sepals 5, equal in size, basally connate. Petals 5, clawed, margin sometimes auriculate near base, outside and inside sericeous at base; scales 2, densely pilose along outer margin, apex, and inner surface; crest a small enation or well developed, clavate and glabrous. Disc uninterrupted. Stamens 8; fila- ments especially basally pilose; anthers dehiscence latrorse, usually papillate, usually slightly pilose, connective usually appendaged. Pistil: ovary 3- (or 4-)locular, densely hirsute; style and stigma triangular, not lobed, grooved, elongating in fruit. Fruits an obcordate loculicidally dehiscent capsule, smooth, somewhat lobed, slightly winged, wings narrow at most, stipe usually present; inside glabrous except for a hair tuft below the placenta and hidden by the arilloid appendage (absent in S. caudata); mesocarp woody in stipe and lower part of seed chambers. Arillode completely covering seed, usually with a straight, sometimes a curled appendage (S. caudata). Seeds obovoid; hilum triangular, relatively large. — Figs. 71-73. Distribution — 12 species; in Malesia 7 species, one widespread over Moluccas and New Guinea, the other 6 endemic in New Guinea. The remaining 5 species endemic in the rain forests of E Australia. Habitat & Ecology — Mainly found in primary, secondary, and montane (moss) for- ests, also along roads, rivers, and edges of forests, sometimes in regularly burned vegeta- tion. Soils: clay, sand, sandstone, volcanic soil, peat, occasionally ultrabasic; sometimes marshy; sea level up to 2800 m altitude. Sometimes branches myrmecophilous. KEY TO THE SPECIES la. Leaflets below glabrous or sericeous on basal part of midrib ................ 2 b. Leaflets below hirsute or villose on basal part of midrib...............0..... 3 2a. Leaflets with very distinct venation, apex cuspidate to caudate. Sepals |.3—2.3 by 718 Flora Malesiana ser. I, Vol. 11 (3) (1994) Sa. 6a. Leaflets with distinct venation, apex acuminate to caudate. Sepals 0.7—1.7 by 0.7— 1.8 mm. Petals 1.3—3.5 by 1.1—3 mm, apex truncate. Arillode with a straight appen- dix sblamssbelow, placemtdiac me tergcu)=) iain «a eaeraeeear - 7. S. squamosa a. Leaves 1—5-jugate; petiolule up to 11 mm long. Leaflets ovate to elliptic, 3-18.3 by e2=7iem,tslightly todenselytpilosc?.men Arie. seine tele ete cise) eh -ntg eeee eS 4 Leaves (1—)2-7-jugate; petiolule a pulvinus only. Leaflets ovate, 0.9-4.2 by 0.3- leSicmirdenselyapiloseyipe pase ot tee aes oes eee ee 3. S. coriacea . Leaflets punctate or not, with or without domatia; nerves at base marginally looped or not. Petals 1.1—2.5 by 0.8—2 mm, claw 0.1—0.5(—1.3) mm high; crest on scales absent (Fig. 71c) to well-developed (Fig. 71a). Fruit with a 3.5-4.5 mm high SUS A hegre ahi aie A. Papert sen eNthe loonie 6 sd caioaevaecNed Pato Shoes ops eee 5 Leaflets punctate, usually with shallow domatia; nerves marginally looped. Petals 2.6-2.8 by 2.4—2.8 mm, claw c. 0.7 mm high; crest on scales well-developed (Fig. (lant, withsaio—o Mn MieWEStIPS 2... pecs atin oie ele 1. S. brachyphylla Crest on petal scales absent (Fig. 71c) or slightly developed (Fig. 71a). Leaflets Olten with, shallow domatia 2: = 00s eae oes tes ae oer ae eee een 6 Crest on petal scales large (Fig. 71a). Leaflets seldom with domatia 4. S. crispata Leaflets ovate to elliptic, 3.7-17.2 by 1.5-5.5 cm, with shallow domatia below; apex acuminate to cuspidate (to caudate). Sepals 1.5-3 by 1—-1.8 mm. Petals 1.3— 2.5 by 1-2 mm, margin auriculate near claw (Fig. 71a)............ 5.S. rigida Leaflets ovate (to elliptic), 3-8 by 1.8-4 cm, domatia absent; apex acuminate. Se- pals 1.2-1.8 by 0.6-0.8 mm. Petals 1.1-1.4 by 0.8—-1 mm, margin not auriculate MEA Claw) (Hie w/ lib —G)) p-pmesiteit. e ncsys io aes chee Sueieaes cl teu ot 6. S. rubiginosa 1. Sarcopteryx brachyphylla Radlk., Sitzungs- Tree, 3-25 m high; dbh 10-40 cm; outer bark ber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. | brown to patchy grey and white, minutely tuber- Miinchen 20 (1890) 265; Bot. Jahrb. 56 (1920) culate, inner bark fawn with pale pink spots; sap- 297; in Engl., Pflanzenr. 98 (1933) 1237; wood cream with black streaks; indumentum ru- Welzen, Blumea 36 (1991) 92, f. la, 3, map 1. — fous. Branchlets crispy hirsute especially when — Type: W. Sayer s.n., 1887 (M holo), Papua —_—- young; flowering twigs 2—4.5 mm thick. Leaves New Guinea. 1-3(—4)-jugate; petiolule up to 7(-11) mm long. Fig. 71. Sarcopteryx Radlk. Petals from inside. — a. S. brachyphylla Radlk., margin with auricles (arrow). — b. S. squamosa (Roxb.) Radlk., margin without auricles (arrow). — c. S. rigida Radlk., margin without auricles. — d. S. caudata Welzen, margin without auricles (a: Fisher 62; b: Atje 283; c: van Valkenburg 33; d: Schodde & Craven 4884). Adema, Leenhouts, Van Welzen — Sapindaceae 719 1cm Fig. 72. Sarcopteryx brachyphylla Radlk. Fruit, typical are the stipe (arrow) and the winged edges (Pullen 7969). Leaflets ovate to elliptic, 3.3—13.5 by 1.2-5.8 cm, asymmetrical, coriaceous, punctate; base asymmet- rical; margin flat (to slightly recurved); apex (acute to) acuminate to cuspidate, usually mucronulate; upper surface mainly crispy hirsute on midrib, of- ten with wax; lower surface hirsute, domatia usu- ally as shallow pockets; nerves marginally looped; veins distinct. /nflorescences in upper leaf axils and pseudoterminal, rather slender, hirsute, up to 12.5 cm long, usually with a few branches; cymules cincinnate (to partly dichasial), 2—4-flowered; bracts and bracteoles triangular, sericeous; bracts up to 3 mm long; bracteoles up to 1.2 mm long. Flowers c.6 mm in diam. Sepals ovate, 0.7—1.6 by 1—1.5 mm, sericeous, green. Petals obovate, white, blade elliptic, 2.6—2.8 by 2.4-2.8 mm, claw up to 0.7 mm high, broadly cuneate, margin auriculate near claw, apex rounded; scales 1.3—1.8 mm long, yellow; crest large, clavate, usually glabrous. Sta- mens: filaments 2.6—3.2 mm long; anthers 0.8—0.9 by 0.5—0.7 mm. Pistil: male ovary c. 0.9 mm high, style and stigma c. 0.2 mm long; female ovary c. 2.8 mm high, green, style and stigma c. 2.9 mm long, yellow to reddish. Fruits 1.8—2.2 by 1.6-2.1 cm, red, stipe 5~9 mm high, broadly triangular, lobes with 2-4 mm broad wings; hair tuft below placenta. Arillode yellow to orange, with straight appendage. Seeds obovoid, 7-12.5 by 5—6.7 mm. — Figs. 71a, 72. Distribution — Malesia: Papua New Guinea (Northern, Central, and Milne Bay Prov.). Habitat & Ecology — Found in primary hilly to montane rain forest, Castanopsis forest, moss for- est, edge of forest; soil marshy; 1500-2065 m alti- tude. Fl. June—Aug.; Fr. June—Dec. Note — See note under Sarcopteryx rigida. 2. Sarcopteryx caudata Welzen, Blumea 36 (1991) 95, f. ld, 4, map 1. — Type: Hoogland & Pullen 5437 (L holo; BM, BRI, K, US), Pa- pua New Guinea. Shrub to tree, up to 23 m high, dbh up to 45 cm; outer bark dark red-brown to grey-brown, smooth to with many pustular lenticels, with fine reticulate cracks and with fairly numerous horizon- tal ridges, inner bark (straw-)brown; wood straw, heartwood pale brown to brown. Branchlets seri- ceous (to hirsute) when young; flowering twigs 1- 5 mm thick. Leaves 2—5-jugate; petiolule up to 7 mm long. Leaflets elliptic, 3.5—11.1 by 1.4-3.9 cm, almost symmetrical to asymmetrical, coriaceous, not (to slightly) punctate; base somewhat asym- metrical; margin flat; apex cuspidate to caudate, mucronulate; upper surface (slightly) sericeous on midrib; lower surface (slightly) sericeous; doma- tia absent; nerves marginally looped; veins highly distinct. /nflorescences in upper leaf axils and pseu- doterminal, rather sturdy, sericeous-hirsute, up to 12.3 cm long, with a few up to 3.7 cm long branch- es; cymules dichasial to cincinnate, 2—4-flowered: bracts and bracteoles triangular, sericeous; bracts up to 2.3 mm long; bracteoles up to 1.9 mm long. Flowers 4.2—-7 mm in diam. Sepals ovate, 1.3—2.3 by 1.2—2 mm, hirsute, green. Petals obovate, white, apex acute; crest on scales absent to minute. Male flowers: petals 3.8—4.3 by 2.2—2.5 mm, claw 0.2- 0.3 mm high, scales 1.8—2.8 mm long; stamens: filaments 3.2—9.6 mm long, pale green, anthers 0.8— 1.1 by 0.4—0.7 mm, dark red; pistil: ovary 0.4—0.7 mm high, style and stigma 0.1—0.3 mm long. Fe- male flowers: petals 2.8—3.9 by 2.2—2.3 mm, claw c. 0.2 mm high, scales 1.3—2.3 mm long; stamens: filaments 1.3—2.2 mm long, pale green, anthers 0.8— 1.1 by 0.4—0.7 mm; pistil pale green, ovary 1.8— 2.8 mm high, style and stigma 1.44.2 mm long. Fruits c. 1.4 by 1.5 cm, smooth, glabrescent, red- brown, stipe low, c. 3.5 mm high, lobes with c. | mm broad wings; no tuft of hairs below placenta. Arillode with a curled appendage. Seeds obovoid, c. 6 by 4.5 mm. — Figs. 71d, 73. Distribution — Malesia: Irian Jaya (Wissel Lakes, Baliem Valley, Valentijn Mts); Papua New Guinea (E Highlands and Morobe Prov.). Habitat & Ecology — Found in primary (Noth- ofagus), secondary, and montane forest. Soil: clay, 720 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 73. Sarcopteryx caudata Welzen. Habit (Schodde & Craven 4884). peat, white sandstone. Rather scarce to locallycom- 3. Sarcopteryx coriacea Radlk., Sapind. Holl.- mon; 1750-2500 m alt. Fl. Apr.—June; fr. Aug. Ind. (1879) 98; Sitzungsber. Math.-Phys. Cl. (based on | specimen). Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) Note — Specimens from western New Guinea 544, 659 (typification); in Engl., Pflanzenr. 98 have leaflets with a somewhat shorter tip and less (1933) 1235; Welzen, Blumea 36 (1991) 96, mucronulate than the other specimens of this spe- map 1. — Type: D’Urville 29118 (P holo; M), cies. Irian Jaya. Adema, Leenhouts, Van Welzen — Sapindaceae 721 -_ CO EEE Shrub, 1-3 m high. Branchlets hirsute when young; flowering twigs 1.5—2.5 mm thick. Leaves (1—)2-7-jugate; petiolule a pulvinus only. Leaflets ovate, 0.9-4.2 by 0.3-1.8 cm, (slightly) asymmet- rical, coriaceous, not punctate; base sometimes asymmetrical; margin flat; apex acuminate, some- times asymmetrical, mucronulate; upper surface hirsute, glabrescent; lower surface densely hirsute; domatia absent; nerves marginally looped; veins very distinct. /nflorescences axillary (to pseudo- terminal), slender, hirsute, up to 6.2 cm long, not branching or with a few up to 5 cm long branches; cymules cincinnate, 2- or 3-flowered:; bracts and bracteoles triangular, sericeous; bracts up to 2 mm long; bracteoles up to 1.1 mm long. Flowers less than 2 mm in diam., young? Sepals deltate, 0.8— 1.4 by 0.6—1.1 mm, hirsute, light green to yellow- ish brown. Petals orbicular, white (to yellow). Male flowers: petals 1.3-1.8 by 1.1—1.6 mm, claw c. 0.3 mm, scale c. 0.8 mm high with a small glabrous crest; stamens: filaments c. 2.7 mm long, anthers 0.4—0.6 mm long, yellow to orange to brownish pink; pistil: ovary c. 0.6 mm high, style and stig- ma 0.1—0.3 mm long. Female flowers: petals c. 1 by | mm, claw c. 0.2 mm, scales c. 0.4 mm high, without crest; stamens: filaments c. 0.7 mm long, anthers 0.4—0.6 mm long; pistil: c. 1.2 mm high, brownish green; style and stigma 0.1—0.3 mm long. Fruits 1-1.4 by 0.9-1.2 cm, smooth, glabrous, red to purplish, stipe absent, lobes with less than 0.5 mm wide wings; tuft of hairs below placenta. Arillode with a straight appendage. Seeds obovoid, c.9 by 6mm. Distribution — Malesia: Irian Jaya (Vogelkop). Habitat & Ecology — Found in primary and sec- ondary forest, edge of mossy forest, open heath, fire vegetation. Soil: coarse sand, grey clay. Rath- er scarce to locally common; 800-2300 m altitude. Fl. Jan.—Apr.; fr. Oct.—Mar. 4. Sarcopteryx crispata Welzen, Blumea 36 (1991) 96. — Type: Clemens 709 (L), Papua New Guinea. (Shrub to) tree, up to 33 m high,dbh up to 60 cm; no buttresses; outer bark brownish to black with numerous small pustular lenticels which sometimes form longitudinal lines, inner bark brown or red- brown streaks on paler background; sapwood pale to brown, heartwood yellow-brown. Branchlets crispy hirsute especially when young; flowering twigs 2.5-6.5 mm thick. Leaves (1—)2—5-jugate; petiolule up to 7 mm long. Leaflets ovate to ellip- tic, 3.7-18.3 by 1.8-7 cm, symmetrical to asym- metrical, thin to coriaceous, punctate or not; base (sometimes) asymmetrical; margin flat to slightly recurved; apex acute to acuminate (to cuspidate), usually mucronulate; upper surface mainly crispy hirsute on midrib; lower surface hirsute; domatia absent (or few shallow pockets); nerves mainly upwards marginally looped; veins distinct. /nflo- rescences in upper leaf axils and pseudoterminal, often rather sturdy, hirsute, up to 23.6 cm long, usually with a few up to 7.6 cm long branches; cymules cincinnate (to partly dichasial), up to 5- flowered; bracts and bracteoles triangular, seri- ceous; bracts up to 2 mm long; bracteoles up to 0.9 mm long. Flowers 34.8 mm in diam.; buds brown pilose. Sepals ovate, 1.1—1.6 by 0.6—-1.4 mm, sericeous, green. Petals obovate, blade rhombic, 1.5—2.2 by 1.2-2 mm, white, claw 0.3—-0.5 mm high, rather slender, margin not auriculate, apex truncate to rounded, outside and inside basally sericeous; scales 0.6—1.7 mm long, yellow; crest large, broad to high, clavate, usually glabrous. Disc yellow. Male flowers: stamens: filaments 34.4 mm long, white, anthers 0.7—1 by 0.6-0.8 mm, pinkish to red; ova- ry 0.6—0.7 mm high, style and stigma 0.2-0.4 mm long. Female flowers: stamens: filaments 1.4—2.5 mm long, anthers 0.6—0.9 by 0.4-0.7 mm; pistil: ovary 1.4—2.2 mm high, style and stigma 1.8—3.4 mm long. Fruits 1.9-2 by 1.5—1.7 cm, smooth, gla- brescent, red, stipe 3.5-4.5 mm high, broadly tri- angular, lobes with 24 mm broad wings; axis with hairs below placenta. Arillode yellow, with straight appendage. Seeds obovoid, 7.5-8.5 by 4.5-6 mm. Distribution — Malesia: Irian Jaya (S of the Vogelkop, Biak I., Noemfoor I., and along the N coast near the border of Papua New Guinea); Pa- pua New Guinea (W Highlands, Morobe, Western, and Central Proy.). A somewhat disjunct distribu- tion which is probably due to insufficient collect- ing. Habitat & Ecology — Found in understorey of primary and secondary rain forest, in oak and No- thofagus forest, and along the road. Soil: well- drained volcanic soil, sand, stone, clay; 10-2000 m altitude. Fl. June—Nov.; young fr. Nov.—Mar. Stems sometimes myrmecophilous. 5. Sarcopteryx rigida Radlk, Bot. Jahrb. 56 (1920) 296, f. 2; in Engl., Pflanzenr. 98 (1933) 1236, f. 36; Welzen, Blumea 36 (1991) 98, f. Ic, map 2. — Lectotype (Van Welzen 1991): Ledermann 11500 (M), Papua New Guinea. Arytera sordida Radlk., Bot. Jahrb. 56 (1920) 301; in Engl., Pflanzenr. 98 (1933) 1279. — Sarco- pteryx sordida (Radlk.) R.W. Ham, Blumea 23 (1977) 290. — Type: Ledermann 12492 (Bt holo; M), E Papua New Guinea. Tree, 3-15 m high, dbh 3—12 cm; outer bark brown to light cream-grey, smooth, inner bark or- ange; wood white to light straw, hard. Branchlets 722 crispy hirsute especially when young; flowering twigs 2-4 mm thick. Leaves 1—3(—4)-jugate; peti- olule up to 9 mm long. Leaflets ovate to elliptic, 3.7-17.2 by 1.5—-5.5 cm, subsymmetrical to asym- metrical, coriaceous, usually not punctate; base slightly asymmetrical; margin flat to slightly re- curved; apex acuminate to cuspidate (to caudate), not mucronulate; upper surface mainly crispy hir- sute on midrib, often with wax; lower surface ru- fous-hirsute; domatia present as shallow pockets; nerves usually marginally looped; veins distinct. Inflorescences in upper leaf axils and pseudoter- minal, rather slender, hirsute, up to 16.2 cm long, sometimes with a few up to 2.8 cm long branches; cymules cincinnate, 1—3-flowered; bracts and bracteoles triangular, sericeous; bracts up to 2.7 mm long; bracteoles up to 1.1 mm long. Flowers 2.7—-4 mm in diam.; buds brown pilose. Sepals del- tate, 1.5—3 by 1—-1.8 mm, sericeous, green. Petals obovate, white, blade rhombic, 1.3—2.5 by 1-2 mm, claw 0.5—1.3 mm high, rather slender, margin au- riculate between claw and blade, apex rounded, outside and inside basally sericeous; scales 0.5— 1.2 mm long, yellow; crest absent or very small and then clavate and usually glabrous. Stamens: female filaments 1.8—2.9 mm long, white; anthers 0.6-1 by 0.4-0.7 mm, red. Pistil: female ovary 0.8— 1 mm high, light green, style and stigma 0.20.6 mm long. Fruits seen immature, red, winged, gla- brescent; stipe low; axis below placenta with few hairs. Arillode with straight appendage. Seeds seen immature, obovoid. — Fig. 71c. Distribution — Malesia: Papua New Guinea (W Sepik, W Highlands, S Highlands, E Highlands, and Morobe Prov.). Habitat & Ecology — Found in understorey of Nothofagus-Castanopsis forest, lower montane rain forest; 1600-2800 m altitude. Fl. Jan., Feb.; young fr. May—Feb. (or Oct.—May). Note — Sarcopteryx rigida differs from S. brach- yphylla in having petal scales without or with a small crest only and a smaller fruit with a short stipe. 6. Sarcopteryx rubiginosa Welzen, Blumea 36 (1991) 98, map 2. — Type: Vinas 18 (L holo; LAE; A, BFC, CBG, K, UPNG, n.v.), Papua New Guinea. Small tree, 7-10 m high, dbh up to 16 cm; out- er bark (reddish) brown, rough, inner salmon to brown; wood white, hard. Branchlets hirsute when young; flowering twigs 2—3.5 mm thick. Leaves 2- or 3-jugate; petiolule up to 6 mm long. Leaflets ovate (to elliptic), 3-8 by 1.8-4 cm, somewhat asymmetrical, coriaceous, not punctate; base some- what asymmetrical; margin flat; apex acuminate, Flora Malesiana ser. I, Vol. 11 (3) (1994) not mucronulate; upper surface pilose on midrib; lower surface hirsute; domatia absent; nerves mar- ginally looped; veins distinct. Inflorescences in upper leaf axils to pseudoterminal, slender, hirsute, up to 8 cm long, not branching or with a few up to 2.4 cm long branches; cymules cincinnate, 1- or 2- flowered; bracts and bracteoles triangular, seri- ceous; bracts up to 2 mm long; bracteoles up to 1.1 mm long. Flowers c. 3 mm in diam. Sepals ovate, 1.2—1.8 by 0.6-0.8 mm, hirsute. Petals or- bicular to elliptic, 1.1—1.4 by 0.8—1 mm, white, claw 0.1-0.2 mm high, margin not auriculate near claw, apex obtuse; scales 0.4—0.7 mm high, without crest. Stamens: filaments 0.7—1 mm long; anthers 0.5— 0.7 mm long. Pistil: ovary 0.6-1 mm high; style and stigma 0.1—0.2 mm long. Fruits unknown. Distribution — Malesia: Papua New Guinea (Morobe Prov.). Habitat & Ecology — Found in montane (faga- ceous) moss forest; 1800-2300 m altitude. Fl. Nov., Dec. 7. Sarcopteryx squamosa (Roxb.) Radlk., Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 303, nom. illeg. (ge- nus not yet described); Sapind. Holl.-Ind. (1879) 19, 57, 97; Sitzungsber. Math.-Phys. Cl. Konig]. Bayer. Akad. Wiss. Miinchen 9 (1879) 544, 659; in Engl., Pflanzenr. 98 (1933) 1234; Welzen, Blumea 36 (1991) 101, f. 1b, 5, map 2. — Sa- pindus squamosus Roxb., [Hort. Beng. (1814) 88, nom. nud.] Fl. Ind. 2 (1832) 282. — Type: Roxburgh s.n. (BR holo), Irian Jaya. Sarcopteryx melanophloea Radlk., Sapind. Holl.- Ind. (1879) 19, 57; Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 544, 659; in Engl., Pflanzenr. 98 (1933) 1234; Streimann, Pl. Upper Watut Watershed (1983) 170. — Type: Beccari PP 15 (Fl holo; M), New Guinea. Sarcopteryx holconeura Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 266; in Engl., Pflanzenr. 98 (1933) 1233. — Type: MacGregor s.n., 1890 (M holo; MEL, sh. 31988), E New Guinea. (Shrub to) tree, 2-25 m high, dbh up to 30 cm; (low buttresses); outer bark rather smooth, thin flaky to finely fissured, grey-green to dark brown, inner bark pinkish straw to purplish to red-brown, hard, non-fibrous; wood straw to orange-pink, sur- face corrugated; indumentum brown. Branchlets sericeous when young; flowering twigs 2—6.5 mm thick. Leaves 1—4(-5)-jugate; petiolule up to 5(— 7) mm long. Leaflets ovate to elliptic, 4.6-23.5 by 1.3-8.4 cm, usually asymmetrical, rather thin to coriaceous, usually punctate; base usually asym- Adema, Leenhouts, Van Welzen — Sapindaceae 723 oe O——————————————eeeeeeeeeee—e—e—e——eeoOoooo metrical; margin flat (to somewhat revolute); apex acuminate to caudate, mucronulate; upper surface at most sericeous on midrib, usually with wax; low- er surface not to (slightly) sericeous on midrib: domatia absent (to few pockets apically); nerves mainly apically marginally looped; veins distinct. Inflorescences in upper leaf axils and pseudoter- minal, rather slender, 0.8—2 mm thick, sericeous, up to 25.5 cm long, usually with a few up to 11.4 cm long branches; cymules cincinnate (to partly dichasial), 2- or 3(—6-)flowered; bracts and bracte- oles triangular, sericeous; bracts up to 3 mm long; bracteoles up to 1.3 mm long. Flowers 2.5—5.2 mm in diam., fragrant. Sepals deltate to ovate, 0.7—1.7 by 0.7—1.8 mm, sericeous, pale green to yellowish brown. Petals obovate, apex truncate, frayed, white: scales yellow; crest small to large, often flat and linear, apically glabrous. Male flowers: petals 1.7— 3.5 by 1.8-3 mm, claw 0.2-0.7 mm high, scales I-1.6 mm long; stamens: filaments 2.8-6.6 mm long, white, anthers 0.5—0.8 by 0.3-0.7 mm, or- ange to red; pistil: ovary 0.3—1 mm high, style and stigma 0.1—1 mm long. Female flowers: petals 1.3- 1.8 by 1.1-1.7 mm, claw 0.1—0.3 mm high, scales 0.5—0.9 mm high; stamens: filaments 0.8—3.3 mm long, anthers 0.4—0.8 by 0.3-0.5 mm; pistil yel- lowish green, ovary 0.8—1.5 mm high, style and stigma 0.3—2.5 mm long. Disc yellowish green to red. Fruits 1.2—2.1 cm high by 1-2.2 cm broad, smooth, glabrescent, purplish red, stipe low, 3-6.5 mm high, lobes with 1-3 mm broad wings; tuft of hairs below placenta. Arillode yellow to red, with a straight appendage. Seeds obovoid, 5.5—10 by 3.7-6 mm, brown. — Fig. 71b. Distribution — Malesia: Moluccas, N Irian Jaya, Papua New Guinea(mainly in thecoastal provinces). Habitat & Ecology — Found in primary and mainly secondary forest, especially along edges: roads, mangrove, savannah, rivers. Often found on sometimes inundated land. Soil: often (silt) clay, also ultrabasic; sea level up to 1575 m altitude. Usually common. Branches sometimes hollow and filled with ants. Fl. Jan.—Apr., June—Sept.; fr. Mar.— July, Sept.—Dec. SARCOTOECHIA (P.W. Leenhouts) Sarcotoechia Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 501; in Engl., Pflanzenr. 98 (1933) 1256; S.T. Reynolds, Austrobaileya 2 (1985) 181; in Fl. Austral. 25 (1985) 82; Leenh., Blumea 33 (1988) 198. — Lectotype species (S.T. Reynolds 1985): Sarcotoechia cuneata Radlk. Tree, monoecious. /ndumentum of solitary, simple, densely appressed, short brown hairs only. Twigs glabrous except for the tip. Leaves paripinnate, unifoliolate or 1- or 2- jugate (Malesian species); no pseudo-stipules; neither petiole nor rachis winged; peti- olules strongly swollen. Leaflets opposite (Malesian species), not papillose, (sub)glabrous, without domatia or red glands; base symmetrical, acute; margin entire (except for 2 Aus- tralian species); midrib above prominulous, beneath prominent; nerves widely spread- ing. Inflorescences axillary and sometimes ramiflorous, racemose, simple or with | or 2 basal branches. Flowers unisexual, actinomorphic. Sepals 5, free, slightly imbricate, equal, not petaloid, outside sparsely, inside densely hairy. Petals 5 or 0, shorter than the sepals, outside near the base sparsely hairy, inside long-hairy in the lower half, with woolly auricles, no crest. Disc complete, densely hairy (New Guinea) or glabrous (Australia). Stamens (5—)7 (or 8), hardly exserted; filaments hairy; anthers glabrous. Pistil hairy; ovary 2- or 3-locular with | ovule per locule; smooth, not lobed: style apical, about as long as the ovary; stigma slightly lobed. Fruits capsular, dehiscence loculicidal, com- pletely 2- or 3-celled, slightly 2- or 3-lobed, not winged, shortly stipitate, outside + smooth, very sparsely shortly sericeous, pericarp + fleshy, completely 2- or 3-celled, inside densely woolly. Seeds ellipsoid, glossy blackish brown; sarcotesta annular around the hilum to cupular and covering up to about a third part of the seed. — Figs. 74, 75. 724 Flora Malesiana ser. I, Vol. 11 (3) (1994) SS eee Distribution — 10 or 11 species; 6 of these, occurring in northern Queensland, belong to subgenus Sarcotoechia, characterized by a glabrous disc; 4 or possibly 5 species occur in Papua New Guinea and belong to subgenus Pilosodiscus Leenh., characterized by a hairy disc. KEY TO THE SPECIES la. Petals present (scars visible under the fruit). ............ esses e eee e cerns ps b. Petals absent or only | or 2 present... 2a. Ovary 3-locular. Twigs fairly persistently angular. Leaflets bullate, the apex emar- ginate, rounded or obtuse........... Ps, Sa A Me NEE OS 1. S. angulata b. Ovary 2-locular. Twigs terete. Leaflets flat, the apex acutely acuminate 3. S. bilocularis 3a. Leaflets 7-20 by 3-6 cm, flat, the apex tapering into a broad rounded acumen. Bowland’ cc wae eee 4. S. planitiei b. Leaflets 4.5-8 by 1.5—-3.5 cm, + bullate, the apex emarginate, rounded, or obtuse. IMontanes et le eee ees 1. Sarcotoechia angulata Leenh., Blumea 33 (1988) 200. — Type: LAE (Stevens) 51061 (L holo), New Guinea. Tree up to 24 m high, dbh up to 35-40 cm. Young twigs characteristically angular when dried, grad- ually becoming terete, c. 5 mm diam., pustular len- ticellate, light greyish brown. Leaves |-jugate; pet- iole semiterete, c. 1.5 cm by 1.5 mm; leaf axes sparsely hairy to subglabrous. Leaflets elliptic, 5.5— 7 by 3-3.5 cm, index c. 2, stiff-pergamentaceous, + bullate, glabrous; base attenuate; margin strong- ly recurved; apex emarginate, rounded, or obtuse; nerves c. | cm apart, slightly curved, ending free, about equally prominulous on both sides; veins and veinlets very laxly reticulate, prominulous on both sides. Inflorescences up to 7 cm long, densely hairy, glabrescent; pedicels c. 2 mm long. Sepals ovate, c. 1.5 by 1.1 mm. Petals 5, elliptic, short-clawed, c. 1.2 by 1 mm. Stamens 7; anthers c. 0.8 mm long. Pistillode 3-locular. Fruits and seeds unknown. — Fig. 75a. Distribution — Malesia: Papua New Guinea (1 collection from the Eastern Highlands Prov.). Habitat & Ecology — In mixed forest at 2550 m altitude. Fl. Nov. Note — The species resembles vegetatively S. apetala with twigs also + angular at first. 2. Sarcotoechia apetala Leenh., Blumea 33 (1988) 202. — Type: Hartley 12505 (L holo; K), New Guinea. Cf. Mischocarpus: Hartley et al., Lloydia 36 (1973) 270. 2. S. apetala Tree, 10-20 m high, dbh up to 30 cm. Twigs at first angular, becoming terete, 2.5—5 mm diam., ca- naliculate, pustular lenticellate, light grey to light greyish brown. Leaves |-jugate; petiole semiterete to sometimes narrowly and deeply grooved above, 0.5-1.5 cm by 1—2.5 mm; petiolules almost absent; leaf axes glabrous. Leaflets elliptic to obovate, 4.5— 8 by 1.53.5 cm, index 2-3, stiff-pergamentaceous, often bullate, glabrous; base attenuate; margin strongly recurved; apex emarginate, rounded, or ob- tuse; nerves 0.5—1 cm apart, slightly curved, ner- vation dense, distinctly looped and joined near the margin, or nerves more distant and ending free; veins laxly reticulate, inconspicuous or distinct above if nerves are more distant. Inflorescences \— 7 cm long, densely hairy, glabrescent; pedicels c. 2 mm long. Sepals ovate, 1-1.3 by 1-1.6 mm. Pet- als absent. Stamens 5 or 7; anthers 0.6—0.8 mm long. Pistil 3-locular. Fruits transversely triangu- lar ellipsoid, c. 8 by 8 mm, at base narrowed into a short stipe. Seeds partly covered by a sarcotesta with a narrow free margin. — Fig. 74. Distribution — Malesia: Papua New Guinea (Morobe, Northern, and Eastern Highlands Prov.). Habitat & Ecology — Subcanopy of montane rain forest at 1800-2200 m altitude. Fl. Aug.; fr. Sept— Oct. 3. Sarcotoechia bilocularis Leenh., Blumea 33 (1988) 200. — Type: NGF (Sayers) 21569 (L holo), New Guinea. Treelet up to 4.5 m high, dbh at least 7.5 cm. Twigs very early terete, 2.5-3.5 mm diam., rather Adema, Leenhouts, Van Welzen — Sapindaceae 75 Fig. 74. Sarcotoechia apetala Leenh. a. Habit; b. detail of lower surface of leaflet; c. fruit (a-c: Brass 31795). 1om Flora Malesiana ser. I, Vol. 11 (3) (1994) tem Fig. 75. Sarcotoechia Radlk. Lower surfaces of leaflets in detail and fruits. — S. angulata Leenh. a. Detail of lower surface of leaflet. — S. bilocularis Leenh. b. Detail of lower surface of leaflet; c. fruit. — S. planitiei Leenh. d. Detail of lower surface of leaflet; e. fruit (a: LAE 51061; b, c: NGF 21569; d, e: Pullen 8401). smooth, light greyish brown. Leaves |-jugate; pet- iole semiterete, 1-2 cm long by c. 1.5 mm broad; petiolules 3-5 mm long; leaf axes glabrous. Leaf- lets elliptic, 9.5-12 by 3.25-4.5 cm, index c. 3, flat, stiff pergamentaceous, glabrous; base attenu- ate; margin slightly recurved; apex mostly taper- ing into a short acute acumen; nerves I1-1.75 cm apart, strongly curved, the upper ones looped and joined at some distance from the margin, above prominulous, beneath prominent; veins laxly re- ticulate, equally prominulous on both sides. Infruct- escences up to 12 cm long, thinly puberulous; pedi- cels slender, up to 4 mm long. Flowers described from remains under fruits. Petals: scars present. Stamens: 7 or 8 scars present. Pistil 2-locular. Fruits transversely ellipsoid to subglobular, c. 8 by 10 mm, at base narrowed into a short stipe. Seeds with a small sarcotesta around the hilum. — Fig. 75b, c. Distribution — Malesia: Papua New Guinea (1 collection from Wagau, Morobe Prov.). Habitat & Ecology — Forest edge on ridge or steep slope; 1500 m altitude. Fr. Jan. Note — This species vegetatively resembles S. planitiet. 4. Sarcotoechia planitiei Leenh., Blumea 33 (1988) 201. — Type: Pullen 8401 (L holo), New Guinea. Trees up to 25 m high, dbh up to 40 cm. Twigs terete, finely grooved, 2-4 mm diam., densely pus- tular lenticellate, yellowish to greyish brown. Leaves unifoliolate or 1- or 2-jugate; petiole terete or above flattened with a narrow groove, (0.7—)1.5— 3 cm by 1-2 mm; petiolules 2-3 mm long; leaf axes densely puberulous, glabrescent. Leaflets el- liptic to obovate, 7-20 by 3-7 cm, index 2.5-3.5, thin-pergamentaceous, glabrous; base sometimes slightly attenuate; margin flat; apex obtuse or + tapering into a broad rounded acumen; nerves (0.5—)1—2.5 cm apart, strongly curved to nearly straight, only the few uppermost ones looped, above prominulous, beneath prominent; veins and vein- lets laxly reticulate, prominulous on both sides. Inflorescences up to 15 cm long, puberulous. Bracts and bracteoles + deltoid, 0.2—0.5 by 0.2-0.5 mm, outside and inside appressed-hairy. Pedicels c. 2.5 mm long, hairy. Male flowers somewhat smaller than the female ones. Sepals deltoid, 0.7—1.1 by (0.51.1 mm, inside at the base also with a row of longer hairs. Petals absent or | or 2 reduced ones present, + spathulate, 0.6—1.1 by 0.2-0.6 mm. Sta- Adema, Leenhouts, Van Welzen — Sapindaceae 727 ——————ee——————————————————————— — — — | >>> >_> Se mens 7 or 8; in male flowers: filaments 2—2.9 mm long, anthers c. 0.5 mm long; in female flowers: filaments 0.7—1 mm long, glabrous or with | or 2 hairs, anthers 0.4-0.5 mm long. Pistil 3-locular; style c. 0.9 mm long; stigma c. 0.4 mm long; pis- tillode c. 0.6 by 0.7 mm. Fruits immature, 3-celled, + globular. Seeds with at least the lower third cov- ered by a cupular sarcotesta with a narrow, lobed, free margin. — Fig. 75d, e. Distribution — Malesia: Moluccas (Ambon, Ceram), SE New Guinea (Milne Bay Prov., near Mayu R.). Habitat & Ecology — Rain forest on plain; soil terete, 1-1.75 cm by c. | mm; petiolules c. 3 mm long; leaf axes slightly puberulous, glabrescent. Leaflets obovate, 7.5—9.5 by 2-3 cm, index 3-3.75, pergamentaceous, above slightly puberulous on the base of the midrib, furthermore glabrous; apex mostly tapering into a short, broad, rounded acu- men, sometimes narrowly rounded; nerves 0.75—1 cm apart, usually strongly curved, only the few uppermost ones looped and joined, above prominu- lous, beneath slightly more so; veins and veinlets above laxly, beneath rather densely reticulate, prominulous. Young inflorescences like erect cat- kins, densely minutely hairy. Flowers and fruits unknown. Distribution — Malesia: SE New Guinea (North- ern Prov., Hydrographers Range). Habitat — Rain forest at 1100 m altitude. Note — As the flowers are too young for analy- sis it is unknown whether the disc is hairy or gla- brous. When hairy the specimen, Pullen 5452, re- sembles S. planitiei, but, when glabrous, it may be one of the Australian species, probably S. cuneata Radlk. or possibly S. lanceolata (C.T. White) S.T. Reynolds. well-drained gravel; 150-350 m altitude. Fr. July. Note — The description of the flowers is based on material from the Moluccas. 5. Sarcotoechia sp.: Leenh., Blumea 33 (1988) 202. — Based on Pullen 5452 (L), SE New Guinea. Tree up to 28 m high. Twigs terete, finely grooved, c. 3 mm diam., greyish brown, densely pustular-lenticellate. Leaves 1-jugate; petiole semi- SCHLEICHERA (P.W. Leenhouts) Schleichera Willd., Sp. Pl. 4, 2 (1806) 1096, nom. cons.; Radlk. in Engl., Pflanzenr. 98 (1933) 872. — Cussambium Lamk, Encycl. 2 (1786) 230. — Type species: Schleichera trijuga Willd. [= Schleichera oleosa (Lour.) Oken]. Trees, dioecious. Indumentum of solitary, simple hairs; young parts with glandular hairs and more or less sticky resinous. Leaves paripinnate, 2—4-jugate; rachis not winged; no pseudo-stipules. Leaflets subopposite (to alternate); margin entire; domatia absent. Inflorescences in the defoliated part of the branchlets above the leaf-scars, sometimes axillary, fascicles of a few simpe (female) or sparsely branching (male) thyrses, the basal part with scattered, many-flowered fascicles, the upper part spicate. Flowers: sometimes male and female flowers present in the same inflorescence, then one of the two kinds predominating, and the other may be sterile. Sepals 4 or 5 (or 6), connate in lower fourth, slightly valvate but soon apert, the lobes all equal, not petaloid. Petals absent. Disc unin- terrupted, more or less patelliform, sinuate, glabrous or sparsely hairy. Stamens (5 or) 6— 8 (or 9), long exserted in male flowers; filaments filiform; anthers basifixed in a cleft, dehiscence introrse, glabrous. Pistil (2- or) 3- (or 4-)locular; ovary sparsely to densely pilose and glandular; style subglabrous, with recurved stigmatic lobes: ovules | per loc- ule. Fruit a hard-crustaceous, dry berry, 1- (or 2-)celled, not winged, either smooth or with patent, simple or branched, strong thorns, glabrescent; inside glabrous. Seeds com- pletely enveloped by an arillode, which is thin-papery and often adhering to the endocarp when dried. — Fig. 76. Distribution — Monotypic. 728 Flora Malesiana ser. I, Vol. 11 (3) (1994) eS Schleichera oleosa (Lour.) Oken, Allg. Naturgesch. Bot. 2 (1841) 1341; Merr., Int. Rumph. (1917) 337; Coster, Ann. Jard. Bot. Buitenzorg 33 (1923) 138; Ochse & Bakh., Ind. Groent. (1931) 650, f. 397; Merr., Comm. Lour. (1935) 247; Japing & Oey Djoen Seng, Tectona 29 (1936) 552, f. 35; Steup, Trop. Natuur 27 (1938) 140; Nath, Fam. Burm. Flow. Pl. 1 (1963) 176; Backer & Bakh. f., Fl. Java 2 (1965) 136. — Cussambium Rumph., Herb. Amb. 1 (1741) 154, t. 57, nom. inval. — Pistacia oleosa Lour., Fl. Coch. (1790) 615. — Schleichera trijuga Willd., Sp. Pl. 4, 2 (1806) 1096; Blume, Rumphia 3 (1847) 147; Bedd., Fl. Sylv. (1871) t. 119; Bran- dis, For. Fl. (1874) 105, t. 20; Hiern in Hook. f., Fl. Br. India 1 (1875) 681; Koord. & Vale- ton, Bijdr. Booms. Java 9 (1903) 177; Atlas (1913) t. 86; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 516; Craib, Fl. Siam. Enum. | (1926) 328; Dammerman, Nat. Tijd. Ned.-Indié 86 (1926) 42; Radlk. in Engl., Pflanzenr. 98 (1932) 874; Kanjilal & Das, Fl. Assam | (1936) 320; Gagnep. in FI. Indo-Chine, Suppl. 1 (1950) 988, f. 126. — Melicocca trijuga Juss., Mém. Mus. Hist. Nat. Paris 3 (1817) 187, t. 8. — Stadman- nia trijuga Spreng., Syst. 2 (1825) 243. — Cus- sambium glabrum Ham., Mem. Wern. Nat. Hist. Soc. 5 (1826) 356. — Cussambium spinosum Ham., Mem. Wern. Nat. Hist. Soc. 5 (1826) 356. — Schleichera aculeata Kostel., Allg. Med. Pharm. FI. 5 (1836) 1829, nom. illeg. — Stad- mannia sideroxylon (non DC.) Hassk., Tijd. Nat. Gesch. Phys. 10 (1843) 130. — Cussambium oleosum O. Kuntze, Rev. Gen. Pl. 1 (1891) 143; Pierre, Fl. Coch. (1895) t. 328, f. b. — Type: Rumphius (1741) t. 57. Tree, up to 40 m high, dbh up to 2 m, but usual- ly much less; (with slight buttresses), the bole usu- ally crooked. Branches terete, striate, 2—5(—8) mm diam., black when young, later yellowish brown to ashy; young parts sparsely, shortly fulvous-seri- ceous and with sessile glands. Leaves (2- or) 3- (or 4-)jugate; axial parts usually early glabrescent; young leaves deep purple; petiole terete to more or less flattened or slightly grooved above, 2—6(— 8) cm long, pulvinate; rachis terete to 3-angular; petiolules swollen, slightly grooved above, 1-3 mm long. Leaflets elliptic to obovate, 4.5—18.5(—25) by 2.5—9 cm, chartaceous to coriaceous, dark brown or greyish green above, medium brown to green- ish beneath, (sub)glabrous; base subacute to cu- neate, often oblique; margin entire to repandous; apex obtuse or emarginate (to shortly acuminate); nerves 12-15 or more per side, straight to slightly curved, looped and joined near the margin with the exception of the lower ones; intersecondary nerves often more or less strongly developed; reticulation fine, dense, prominulous on both surfaces. Inflo- rescences 6-15 cm long, sparsely hairy. Flowers pale yellow or pale green. Sepal lobes ovate to del- toid, c. 1.5 mm high, obtuse to acute, thin-hairy on both sides, the margin ciliate (and glandular), de- ciduous in fruit. Stamens: filaments c. 2 mm long, sparsely hairy; anthers broad-elliptic, c. 0.75 mm long, slightly emarginate at apex. Pistil strongly reduced in male flowers; ovary ovoid, slightly 3- angular and indistinctly 3-sulcate, c. 1.25 mm long; style rather thick, 1.25—1.5 mm long. Fruits broad- ovoid to subglobular, c. 15 by 13 mm when 1-seed- ed, or transversely ellipsoid, slightly flattened, somewhat bilobed, 17—20 by c. 18 by 14 mm when 2-seeded, narrowed at base, pointed at apex, gran- ular, yellow. Seeds subglobular, c. 12 by 10 by 8 mm; hilum orbicular; testa dull medium-brown, smooth and glabrous; arillode yellow and subacid. — Fig. 76. Distribution — Tropical SE Asia from Sri Lan- ka and the western Deccan to Indo-China; in Malesia: Java, Lesser Sunda Islands, Central and SW Celebes, the islands of Saleyer, Kabaéna, and Muna, Moluccas (Ambon, Banda, cited also from Ceram and the Kai Islands). Although cited by Miquel (Sumatra, 1861, 199) from Sumatra, I saw only cultivated specimens from there, and none of them seen by him. As to the probably erroneous, old records for the Philippines, see Radlkofer (in Engl., Pflanzenr. 98, 1932, 877, footnote), and Merrill (Enum. Philipp. Flow. Pl. 2, 1923, 516). I am uncertain about the status of this tree in Malesia. It is possible that it is not autochthonous, but in- troduced by man. It has several uses and is often cultivated. IThe species thrives only in the drier parts, however, hence the distribution gap in ever- wet W Malesia. Tradesmen may have brought the seeds to several parts of the Archipelago, and it is interesting that the common Hindu name, kusam, is very like the most common name in Malesia, kusambi. See Carthaus (Tectona 2, 1909, 318). Habitat & Ecology — Typical in regions subject to a dry period, e.g., in Java the parts with natural teak-forest. On dry to periodically rather swampy soils of several types; in monsoon forest as well as parkland or savannahs with only scattered trees. The plants are fire-resistant (cf. Meijer Drees, Comm. For. Res. Inst. 33, 1951, 110). Usually at low altitudes, up to 900(—1200) m. Seedlings are light-demanding. Fl. in the beginning of the dry season; fr. about half a year later. Deciduous, but completely leafless during a few days only. The seeds are eaten by mammals (probably mainly Viverridae) and birds; furthermore, dispersal would Adema, Leenhouts, Van Welzen — Sapindaceae 729 Fig. 76. Schleichera oleosa (Lour.) Oken. a. Habit. b. fruit with spines (a: Buwalda 3177; b: Colfs 290). be by termites (cf. Ridley, Dispersal, 1930, 167, SoU): Uses — Wood for timber and especially for an excellent charcoal; bark used for dyeing and in native medicine. For a description of the timber, see p. 427. Young leaves eaten as a vegetable; fruits (arillode) eaten as a titbit; from the seed an oil is pressed which is used for several purposes, among others in native medicine and as a constituent of the true Makassar oil. See Heyne (Nutt. Pl. Indon. ed. 3, 1950, 990-996). 730 Flora Malesiana ser. I, Vol. 11 (3) (1994) SYNIMA (P.W. Leenhouts & F. Adema) Synima Radlk., Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 501; in Engl., Pflanzenr. 98 (1933) 1254-1255; Reynolds, Austrobaileya 2 (1985) 186: in Fl. Austral. 25 (1985) 82, 202. — Type species: Synima cordieorum CE Muell.) Radlk. Trees, monoecious. Indumentum of solitary simple hairs; no glandular scales. Twigs terete. Leaves spirally arranged, paripinnate, 2—7-jugate; no pseudo-stipules; neither pet- iole nor rachis winged. Leaflets opposite to alternate, beneath smooth, without domatia or glands, margin entire or crenate to serrate. Inflorescences axillary, together sometimes pseudoterminal, thyrsoid. Flowers unisexual, regular. Sepals 5, free, equal, not or slight- ly imbricate, not petaloid, on both sides hairy. Petals 5, longer than the sepals, ciliate but otherwise glabrous, inside with 2 recurved, woolly, not to distinctly crested scales nearly as long as the petal. Disc entire, glabrous. Stamens 8, exserted, filaments and sometimes anthers hairy. Ovary 3-celled, densely hairy; style apical, about as long as the ovary, densely hairy; stigma slightly lobed. Ovules 1 per cell. Fruits 3-celled, faintly 3-lobed, loculicidal, valves in lower part remaining fused, marginated at the corners, stipe short, pericarp fleshy. Seeds covered with a sarcotesta except for an adaxial strip; cotyledons unequal, collateral. — Fig. 77. Distribution — Australia (N Queensland) and SE New Guinea; two species. KEY TO THE SPECIES la. Inflorescences with patent branches ..............+--+5-- 1. S. cordierorum be intlorescences unbranched ee. ere. ey. ete en oe 2. S. macrophylla oblique, acute (or rounded), attenuate; margin en- tire, undulate, or crenulate; apex rounded to acute or slightly acuminate; midrib above slightly raised, beneath prominent and rounded, nerves c. | cm apart, widely spreading, strongly curved, ending free or sometimes + looped and joined, above hard- ly raised, beneath strongly raised, veins and vein- 1. Synima cordierorum (F. Muell.) Radlk., Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 9 (1879) 513, 546 (‘cordierii’); in Engl., Pflanzenr. 98 (1933) 1255; Hender- son & S.T. Reynolds, Austr. Syst. Bot. Newsl. 43 (1985) 21; S.T. Reynolds, Austrobaileya 2 (1985) 186, f. 6G—I; in Fl. Austral. 25 (1985) 82. — Cupania cordierii F. Muell., Fragm. 9 _ lets moderately coarsely reticulate, beneath some- (1875) 93. — Type: Dallachy s.n. (L, MEL), — what more prominent than above. Inflorescences Australia. up to 30 cm long, densely hairy, with few erecto- patent branches, the flowers in short, dense, many- flowered cymules. Flowers white. Sepals 1-1.1 mm high, inside densely sericeous. Petals 1.2-1.4 by 1.1-1.5 mm, short-clawed, scales not crested. Sta- Tree, up to 27 m high, dbh up to 40 cm. /ndu- mentum densely fulvous tomentellous. Twigs 4—6 mm in diam. Leaves 3—5-jugate; petiole flattened above, rounded below, 4-6 cm long; axes hairy, especially so at base, upwards glabrescent; peti- olules 2-10 mm long, grooved above, strongly swollen at base. Leaflets elliptic (or ovate), 6.25— 13 by 2.25—4.5 cm, index (2—)3, stiff-pergamenta- ceous, very sparsely hairy below towards the base on midrib and nerves, otherwise glabrous; base in the lower leaflets symmetric, in the upper ones mens: filaments 2.5 mm long; anthers 0.6 mm long. Fruits 20 by 18 mm, bright red, outside sparsely puberulous, inside densely woolly. Seeds oblique- ly obovoid-ellipsoid, 13 by 6 mm, testa shining black, sarcotesta yellow. — Fig. 77a-f. Distribution—Australia(Queensland: Cook Dist.) and Malesia: Ceram, New Guinea (Irian Jaya: Mer- auke Dist.; Papua New Guinea: Fergusson Island). Adema, Leenhouts, Van Welzen — Sapindaceae 731 a a a a Fig. 77. Synima Radlk. Habit, fruits, seeds and embryos. — S. cordierorum (F. Muell.) Radlk. a. Habit; b. fruit; c. seed ventral view; d. seed, dorsal view; e. embryo, lateral view; f. embryo, ventral view. — S. macrophylla Reynolds. g. Fruit; h. seed, ventral view; i. seed, dorsal view (a: Pullen 7445; b-f: Gray 765; g-i: Brass 26088). 1e2 Habitat & Ecology — Lowland rain forest; alti- tude 700 m. Fl. Oct.; fr. Dec. 2. Synima macrophylla S.T. Reynolds in Fl. Aus- tral. 25 (1985) 82, 202; Austrobaileya 2 (1985) 187. — Type: Webb & Tracey 8223 (BRI, L), Queensland. Small tree, 1.5—3 m high (in Australia up to 11 m). Indumentum short-tomentose, brownish. Twigs 4-10 mm in diam., grooved. Leaves 2—6-jugate; petiole 4.5—18 cm, slightly to strongly pulvinate, rachis 11-28 cm, both terete or flattened above, rounded below; petiolule S—15 mm, grooved above. Leaflets elliptic to narrowly ovate, 8-23 by 3.5-11 Flora Malesiana ser. I, Vol. 11 (3) (1994) midrib with few short hairs; base oblique, cuneate to rounded; apex acute to obtuse or acuminate; midrib slightly prominent above, nerves 5—10 per side, 9-20 mm apart, angle to midrib 60—90°. In- florescences 5.5-28 cm, not branched. Flowers white. Fruits obpyramidal, rounded deltoid in cross section, red, 17—21 by 15-20 mm, stipe c. 6 mm, outside thinly appressed-hairy, inside villous or more appressed-hairy. Seeds narrowly obovoid, | 1— 13 by 5-6 mm, testa shiny black, sarcotesta yel- low; hypocotyl with very short hairs. — Fig. 77g-1. Distribution — Australia (N Queensland) and Malesia: Papua New Guinea (Western and Milne Bay Prov.). Habitat & Ecology — Lower montane or mossy cm, index 2—3.6, dark green above, mid green be- oak forest; altitude 150-900 m. Fr. June, Dec. low, chartaceous, almost glabrous on both surfaces, TOECHIMA (P.W. Leenhouts) Toechima Radlk., Sapind. Holl.-Ind. (1879) 60; in Engl., Pflanzenr. 98 (1933) 1249; S.T. Reynolds, Austrobaileya 2 (1985) 176; in FI. Austral. 25 (1985) 77; Leenh., Blumea 33 (1988) 203. — Lectotype species (S.T. Reynolds, Austrobaileya 2): Toechima ery- throcarpum (F. Muell.) Radlk. Trees, monoecious. Indumentum of solitary simple hairs only. Twigs terete. Leaves paripinnate, 1—6-jugate; neither pseudo-stipules, nor wings; petiolules above slightly grooved, pulvinate. Leaflets opposite to alternate, densely minutely warty beneath, small red glands absent; margin entire (Malesian species). Inflorescences axillary, usually branch- ing, together sometimes pseudoterminal, thyrsoid. Flowers unisexual. Sepals 5, nearly free, valvate to narrowly imbricate in bud, all equal, not petaloid. Petals 5, as long as to distinctly longer than the calyx, hardly to longly clawed; blade with 2 erect or recurved, densely woolly, distinctly crested scales nearly as long as the petal. Disc uninterrupted, erect, thick, + 5-lobed, glabrous. Stamens 8, in male flowers long exserted; filaments filiform; anthers glabrous, dehiscence latero-introrse. Pistil sessile, densely hairy; ovary 3-locular (Malesian species) with 1 ovule per locule; septa complete; style apical, longer than the ovary; stigma grooved or slightly lobed. Fruits capsular, dehiscence loculicidal, 3-angular-globular, sessile, not winged; outside + smooth, glabrous or tomentose, inside densely woolly but for a fleshy cupular part around the placenta; pericarp thick, hard- fleshy. Seeds ellipsoid; testa smooth, shiny brownish black; a small sarcotesta with a narrow, free arillode margin at both sides of the nearly basal hilum. — Fig. 78. Distribution — 6 species, in Australia (Queensland and northern New South Wales to the Wilson R.) and Malesia (New Guinea, | species). 19a—d, map 100; Leenh., Blumea 33 (1988) 204. — Cupania erythrocarpa F. Muell., Fragm. Phyt. Austral. 5 (1985) 7. — Type: Dallachy s.n. (iso in NSW, sh. 166318, 166319), Australia. Toechima erythrocarpum (F. Muell.) Radlk., Sa- pind. Holl.-Ind. (1879) 60; in Engl., Pflanzenr. 98 (1933) 1252; S.T. Reynolds, Austrobaileya 2 (1985) 178; in Fl. Austral. 25 (1985) 79, f. Adema, Leenhouts, Van Welzen — Sapindaceae — —— J WH VTE WY i fi (a8 \ a 733 Ae Fig. 78. Toechima erythrocarpum (F. Muell.) Radlk. subsp. papuanum Leenh. a. Habit; b. seed, ventral view; c. seed, lateral view (a: Schodde 2866; b, c: NGF 8568). subsp. papuanum Leenh., Blumea 33 (1988) 204. — Toechima livescens Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 266; in Engl., Pflanzenr. 98 (1933) 1253; Hartley et al., Lloydia 36 (1973) 270. — Syntypes: Forbes 374 (FI, L, M, P), 637, 761, 804 (FI, L,M), New Guinea. Toechima subteretes Radlk., Sapind. Holl.-Ind. (1879) 19, 60; in Engl., Pflanzenr. 98 (1933) 1254. — Type: Beccari it. sec. 17 (Fl holo), New Guinea. Toechima hirsutum Radlk. in K. Schum. & Hollr., Fl. Kais. Wilh. Land (1889) 67; Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 266; in Engl., Pflanzenr. 98 (1933) 1253. — Type: Hollrung 820 (M holo), New Guinea. Toechima sp.: Hartley et al., Lloydia 36 (1973) 270. — Based on Hartley 10724. Not cf. Toechima: Hartley et al., Lloydia 36 (1973) 270 (Hartley 10716) (= Alectryon affinis Radlk.). Treelet or tree, up to 30 m high by 60 cm dbh; (with low buttresses). Twigs appressedly minutely brown-hairy and early glabrescent (to persistently loosely appressedly to + patently brown hirsute to tomentose). Leaves (1—)3—6-jugate; petiole sub- terete (to terete), 4-9 cm long, 1.5—3.5 mm thick; 734 Flora Malesiana ser. I, Vol. 11 (3) (1994) ON petiolules 1-10 mm long; axes (sub)glabrous (to hairy). Leaflets (sub)elliptic, 4-21 by 2—6.5 cm, index 2-3, thin- to stiff-pergamentaceous, green- ish to bluish grey above, dark brown beneath when dry, glabrous to on the midrib and beneath on the nerves sparsely hairy, (with some small domatia or axillary hair tufts beneath); base slightly oblique to symmetrical; apex acute to acuminate; midrib above prominulous, nerves less than 10 per side, up to 3 cm apart, usually erecto-patent and strong- ly curved, ending free, above hardly prominent; intersecondary nerves variably developed; reticu- lation lax. Inflorescences up to 30 cm long with few to several long patent to rather erect branches, the flowers in sessile, dense, many-flowered tufts; densely puberulous. Sepal lobes triangular, 1.25— 1.8 by 1.25—2 mm, on both sides variably hairy, + ciliate, no glands, margin entire. Petals shortly and broadly to long and narrowly clawed; blade ellip- tic to transversely elliptic, 1.2-2 by 1.5—2 mm, gla- brous or outside slightly hairy near the base, in- side densely appressed-hairy, entire or slightly toothed to emarginate at apex. Stamens: filament 3.54.5 mm long, densely (to sparsely) woolly usu- ally with the exception of the uppermost part, an- ther suborbicular, c. 0.4 mm long. Pistil: ovary c. 1.8 mm long; style slender, c. 2.5 mm high. Fruits 2-2.5 cm diam., glabrous (to tomentellous); wall 5~7 mm thick. Seeds 14-18 by 8-10 mm, hilum and sarcotesta together c. 10 by 3 mm. — Fig. 78. Distribution — Malesia: New Guinea. A second subspecies, subsp. erythrocarpum, in northern Queensland. Habitat & Ecology — An understorey tree of primary and secondary rain forest on well-drained but mostly alluvial soils, sometimes in mangrove or on river banks; up to 900 m altitude. Fl. Jan.— Oct., mainly June—Sept.; fr. Jan.—Mar. and June— Oct. EXCLUDED Toechima plurinerve Radlk. in Fedde, Rep. 20 (1924) 36; in Engl., Pflanzenr. 98 (1933) 1252. Based upon material cultivated in the Bogor Botanic Garden of unknown origin, this seems iden- tical with the Australian species 7. daemelianum Radlk. TRIGONACHRAS (P.W. Leenhouts) Trigonachras Radlk., [Sitzungsber. Math.-Phys. Cl. K6ningl. Bayer. Akad. Wiss. Miinchen 8 (1878) 299, 304, nom. inval.] Sapind. Holl.-Ind. (1879) 46; in Engl., Pflanzenr. 98 (1933) 1243: Leenh., Blumea 33 (1988) 204; Yap in Tree FI. Malaya 4 (1989) 459. — Lectotype species (Leenhouts 1988): Trigonachras acuta Radlk. Trees, monoecious. Indumentum of solitary simple hairs only. Leaves paripinnate, 1— 9-jugate; no pseudo-stipules; neither petiole nor rachis winged. Leaflets opposite to al- ternate, pergamentaceous; margin entire; upper surface without wax, below sometimes with brownish glands in axils of nerves, without papillae; midrib above prominulous; nerves prominulous on both sides. /nflorescences axillary, together usually pseudotermi- nal, thyrsoid, hairy, + glabrescent; cymes lax, |- or few-flowered. Flowers regular, uni- sexual. Sepals 5, free, slightly imbricate, all about equal or sometimes the outer two slightly smaller, sometimes thinned out to the margin, no glands, margin entire. Perals 5, longer than the calyx, distinctly clawed; claw densely ciliate; blade entire, inside with (1 or) 2 erect, densely woolly, not crested scales nearly as long as the blade. Disc uninter- rupted, annular, + thick, glabrous. Stamens (7) 8 (9), in male flowers long exserted; fila- ments filiform, broadened to the base; anthers obovoid, dehiscence latero-introrse to la- trorse. Pistil sessile, densely hairy; ovary 3-locular with | ovule per locule; septa com- plete; style apical, about as long as the ovary; stigma grooved or slightly lobed. Fruits distinctly stipitate, capsular, loculicidal, dehiscing into 3 equal valves, not winged, smooth, outside minutely hairy or glabrous, inside usually densely and woolly hairy; pericarp Adema, Leenhouts, Van Welzen — Sapindaceae 735 hard-fleshy. Seeds ellipsoid, testa smooth, shining, black to brown; hilum nearly basal, elliptic, fairly big, but covering less than one third of seed; no arillode. — Figs. 79, 80. Distribution — 8 species in Malesia: Malay Peninsula, Sumatra, Borneo, Philippines, Celebes, and New Guinea. Habitat & Ecology — Medium-sized trees from lowland forest, primary as well as secondary, often in marshes. KEY TO THE SPECIES 2a. Fruits 2—3 by 1.5—2 cm. Philippines A ey ee Ee Ce ee re ee 3. T. cultrata Beets. 4. bY 2.25.Cm. onmores NotLehilippines .4.'°. 5. . 2). 2220.) Mae. 3 3a. Fruits up to 3 cm wide, the wall c. 1 mm thick. West Malesia ....... b. Fruits c. 4 cm wide, the wall c. 10 mm thick. New Guinea...... 4a. Leaflets tapering into an acute acumen Bb Leatlets not acuminate . 0... 2s... 5,.:. 5a. Leaves 7-jugate; petiolules 3-5 mm long. Borneo............. 1. T. acuta 7. T. nov. sp. B b. Leaves 1—4-jugate; petiolules 7-10 mm long. Philippines. ...... 4. T. cuspidata 6a. Leaves 3—5-jugate; petiole terete. Borneo, Philippines ..................... 7 b. Leaves 1- or 2-jugate; petiole flat above. Celebes ............. 7a. Petiolules 3—9 mm long; leaflets S—10 by 1.754 cm. Borneo ... 2. T. celebensis 6. T. nov. sp. A b. Petiolules 12-15 mm long; leaflets 11-15 by 5-6 mm. Philippines 1. Trigonachras acuta (Hiern) Radlk., Sapind. Holl.-Ind. (1879) 46; in Engl., Pflanzenr. 98 (1933) 1245; Corner, Wayside Trees (1940) 596, f. 212; Gard. Bull. Sing., Suppl. 1 (1978) 153; H. Keng, Gard. Bull. Sing. 35 (1982) 91; Leenh., Blumea 33 (1988) 206; Yap in Tree Fl. Malaya 4 (1989) 459. — Sapindacea Wall., Cat. (1848) nr. 9036. — Cupania acuta Hiern in Hook. f., Fl. Br. India 1 (1875) 677. — Syntypes: Main- gay KD 445 (CGE, K, L), Malay Peninsula; Wallich 9036 (K), Singapore. Tree up to 25 m high, dbh up to 50 cm; with buttresses. Twigs subterete, when young + angu- lar, 3-6 mm diam., grey to black, brown tomentel- lous, glabrescent. Leaves 3—-8-jugate; petiole sub- terete to terete, 1.5—6 cm long by 1-2 mm broad; petiolules subterete to grooved above, 3-6 mm long. Leaflets opposite to alternate, ovate to elliptic, 4.5— 12 by 1.54 cm, index 2.25-4.5, slightly falcate, thin- to rather stiff-pergamentaceous, glabrous or with some appressed hairs on the lower side of the midrib with small glands in the nerve axils beneath; base symmetrical to oblique, narrower at the basi- scopic side, obtuse to acute (or rounded), slightly 8. T. nov. sp. C attenuate; apex usually tapering into an acute acu- men (or acute to obtuse); nerves 0.5—1.5 cm apart, spreading, strongly curved, ending free; reticula- tion fairly coarse, equally raised on both sides. Jn- florescences pseudoterminal (and terminal), 10—20 cm long, without or with a few long branches, usu- ally few-flowered, fulvous-puberulous. Sepals broad-ovate, 1.3—1.75 by 1.25-1.75 mm, fairly densely short sericeous on both sides to rather Sparse inside, margin ciliate. Petals 2-3 by 1.6— 2.5 mm; (claw with a few hairs outside); blade or- bicular, glabrous, inside with 2 scales somewhat more than half as long as the petal. Stamens 7 or 8; filaments up to 5.5 mm long, fairly densely pat- ently long hairy except for the upper fourth; an- thers c. 1 mm long, glabrous. Pisti/ 3- (or 4-)locu- lar, shortly brown-hairy; ovary slightly trigonous, tapering into the at least equally long style; stigma grooved. Fruits + triangular-clavate, apiculate, 4.5— 5 by 2.25-3 cm, brown tomentellous, red when fresh; wall c. 1 mm thick, hard, thick and fleshy when fresh, endocarp wrinkled, sparsely to dense- ly woolly. Seeds c. 1.5 by 1 cm, black and shiny; hilum basal, transversely elliptic, c. 7 by 5 mm. — Fig. 79. 736 Flora Malesiana ser. I, Vol. 11 (3) (1994) Oe ee ee ee eS EEE EE SS See eee Fig. 79. Trigonachras acuta (Hiern) Radlk. a. Habit; b. fruit (a, b: SAN 40845). Adema, Leenhouts, Van Welzen — Sapindaceae aa7 Oc ee Distribution — Malesia: Sumatra, Malay Penin- sula, and Borneo (Sabah). Habitat & Ecology — In primary and secondary forests, often in swamps, along river banks, along roads, on slopes and ridges, often on sandy soils; sea level up to 225 m altitude. Fl. Apr—June; fr. Feb., July—Sep., Nov. 2. Trigonachras celebensis Leenh., Blumea 33 (1988) 211. — Type: Prawiroatmodjo & Soe- woko 1747 (L holo), SE Celebes. Tree up to 25 m high, dbh up to 35 cm; with buttresses. Twigs terete, 3-4 mm diam., dark red- brown to blackish, glabrous. Leaves |- or 2-jugate; petiole subterete, 3—9 cm long by c. 1.5 mm broad; petiolules above flattened, 5-12 mm long. Leaf- lets opposite to alternate, ovate to elliptic, 6-15 by 2—6 cm, index 2—3.5, straight to subfalcate, varia- bly pergamentaceous, glabrous, with some small glands along the basal part of the midrib beneath, not glaucous below; base hardly to distinctly ob- lique, acute or rounded, attenuate; apex rounded to acute; nerves 1—2 cm apart, spreading, rather strongly curved or at first nearly straight, ending free; intersecondary nerves + strongly developed; reticulation rather coarse, prominulous on both sides. Inflorescences pseudoterminal, 6-15 cm long, sparsely branched, rather densely appress- edly shortly yellowish brown hairy. Sepals broad- ovate, 1.3—2.25 by 1.2-1.6 mm, outside sparsely appressed-hairy, inside the same to glabrous. Pet- als long-clawed, 1.6—2.5 by 1—-1.5 mm; claw and base of plate outside sparsely appressed-hairy to glabrous; blade elliptic, inside glabrous, with 2 scales or sometimes | entire scale. Stamens 8 or 9; filaments c. 6 mm long, woolly in the lower 2/5; anthers 1.1—1.4 mm long, subglabrous to fairly densely appressed-hairy. Fruits triangular-ellipsoid, 2.75—3 by c. 2 cm, abruptly narrowed into a 0.5— 0.75 cm long stipe, + apiculate, glabrous, orange; wall c. 3 mm thick, fleshy; endocarp densely woolly. Seeds unknown. — Fig. 80a, b. Distribution — Malesia: Celebes (Central, South- east). Habitat & Ecology — Primary and secondary dryland forest on limestone or clay; up to c. 250 m altitude. Fl. Nov.—Dec.; fr. Nov. Uses — The timber is worthless because it soon decays. 3. Trigonachras cultrata (Turcz.) Radlk., [Sit- zungsber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 299, nom. inval.] Sa- pind. Holl.-Ind. (1879) 46; in Engl., Pflanzenr. 98 (1933) 1246; Leenh., Blumea 33 (1988) 207. — Sapindus cultratus Turez., Bull. Soc. Imp. Nat. Moscou 31 (1858) 403. — Type: Cuming 1304 (FI, L, M, P, SING), Philippines. Trigonachras brachycarpa Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1614; in Engl., Pflanzenr. 98 (1933) 1248. — Type: Elmer 10949 (M holo; FI, K, L, NY, U), Philippines. Trigonachras obliqua Radlk., Philipp. J. Sc., Bot. 8 (1914) 467; in Engl., Pflanzenr. 98 (1933) 1247, p.p. — Type: FB (Bernardo) 13108 (M), Philippines (Luzon, Cagayan Prov.). Trigonachras rigida Radlk., Philipp. J. Sc., Bot. 8 (1914) 467, p.p.; in Engl., Pflanzenr. 98 (1933) 1247, p.p. — Lectotype (Leenhouts 1988): Mer- rill 2967 (M holo; BM, K, NY), Philippines. Trigonachras membranacea Radlk., Philipp. J. Sc., Bot. 8 (1914) 468; in Engl., Pflanzenr. 98 (1933) 1248. — Syntypes: FB (Clark) 1073 (M, NY); Vidal 2488, 2500 (K); all Philippines. Tree up to 22 m high, dbh up to 50 cm. Twigs terete, when young angular, 3.5—9 mm diam., light to purplish grey, early to late glabrescent. Leaves 5—9-jugate; petiole (sub)terete, mostly flattened at the base, 3-13.5 cm long by 1-4 mm thick; peti- olules slightly grooved above, 2.5-8 mm long. Leaflets opposite to alternate, ovate (to elliptic), 3.5—18 by 1.5—7 cm, index 2-4, straight to falcate, (thin-)pergamentaceous, glabrous to sometimes sparsely hairy on the basal half of the midrib, usu- ally small glands in some to most of the nerve ax- ils beneath; base oblique to sometimes symmetri- cal, rounded to acute, attenuate; apex acute (to rounded); nerves 1-3 cm apart, widely to steeply spreading, variably curved, ending free; reticula- tion rather coarse (or dense beneath), beneath most- ly more prominent than above. /nflorescences to- gether pseudoterminal, up to 40 cm long, with few spreading branches, rather densely ferrugineous puberulous, + glabrescent. Sepals ovate, 2—2.2 by 1.6-2 mm, outside densely, inside densely to sparsely hairy, not ciliate. Petals: claw c. 1 mm long, blade transverse-elliptic, in total 3—3.5 by 2- 3 mm, claw and basal half of blade outside sparse- ly woolly, inside glabrous, inside with 2 scales. Sta- mens 8; filaments 6.5—7.5 mm long, in the lower half variably woolly; anthers 1.4—-1.5 mm long, glabrous. Fruits globular to triangular-obovoid, 2— 3 by 1.5—2 cm, stipe up to 1 cm long, usually api- culate, densely ferrugineous or fulvous puberulous to tomentose; wall 1—3 mm thick, fleshy, endocarp finely transversely wrinkled, sparsely (to densely woolly). Seeds 11—12.5 by 7-8 mm, black; hilum suborbicular to transverse-elliptic, 2-6 by 2—6 mm. — Fig. 80c. Distribution — Malesia: Philippines. Habitat & Ecology — In primary forest, on ridg- es, along rivers, and near the sea shore; sea level 738 Flora Malesiana ser. I, Vol. 11 (3) (1994) icm Fig. 80. Trigonachras Radlk. Leaflets and fruits. — T. celebensis Leenh. a. Lower surface of leaflet; b. fruit. — T cultrata (Turcz.) Radlk. c. Fruit. — 7. cuspidata Radlk. d. Leaflet lower surface. — T. papuensis Leenh. e. Fruit (a, b: Prawiroatmodjo & Soewoko 1747; c: PNH 37151; d: BS 20467, e: NGF 42923). up to 500 m alt. Fl. Apr. and May; fr. June—Aug. Note — Most of the collections are nearly gla- brous, but some from Luzon have densely tomen- tose twigs, leaf axes, and inflorescences. 4. Trigonachras cuspidata Radlk., Philipp. J. Sc., Bot. 6 (1911) 182; in Engl., Pflanzenr. 98 (1933) 1246; Leenh., Blumea 33 (1988) 210. — Type: FB (Hagger) 1411 (M), Philippines. Trigonachras falcatocuspidata Radlk., Philipp. J. Sc. 20 (1922) 661; in Engl., Pflanzenr. 98 (1933) 1247. — Lectotype (Leenhouts 1988): BS (Ra- mos) 14920 (M holo; BM), Philippines. Trigonachras rigida Radlk., Philipp. J. Sc., Bot. 8 (1914) 467, p.p., excl. type. Tree up to 15 m high, dbh up to 40 cm. Twigs terete, 4-5 mm diam., blackish, glabrous. Leaves 1-4-jugate; petiole (sub)terete, 3-6.5 cm long by 1-2 mm broad; petiolules above slightly grooved, 7-15 mm long. Leaflets opposite, elliptic, 5—14 by 2-4.75 cm, index 2-3, slightly falcate, stiff-perga- mentaceous, glabrous, without glands; base ob- lique, acute, attenuate; apex with a + tapering long acute acuminate; nerves 1-2 cm apart, spreading, strongly curved, ending free; reticulation coarse, prominulous on both sides. Inflorescences seem- ingly terminal as the vegetative terminal bud is mostly shifted aside and suppressed, c. 12 cm long, sparsely branched, densely appressedly shortly brown hairy. Sepals broad-ovate, c. 1.5 by 2 mm, outside and inside sparsely hairy. Petals: blade broad-ovate, 1.5—3 by c. 1.3 mm, glabrous, inside with 2 short scales. Stamens 8; filaments c. 5 mm long, woolly in lower two thirds; anthers c. 1.5 mm long, sparsely ciliate. Pistil 3-locular. Fruits trian- gular-ellipsoid, 4-5 by 2-3 cm, stipitate and apicu- late or not, glabrous; wall fleshy, c. 5 mm thick, endocarp densely brown woolly. Seeds c. 1.5 by 0.75 cm, brown; hilum basal, suborbicular. — Fig. 80d. Distribution — Malesia: Philippines (Luzon, Polillo, possibly also Mindanao). Habitat & Ecology — Behind mangrove and in lowland forests. Fl. Feb.—Mar.; fr. June-July. Adema, Leenhouts, Van Welzen — Sapindaceae 739 ————— a 5. Trigonachras papuensis Leenh., Blumea 33 (1988) 212. — Type: NGF (Henty) 20513 (L holo; K), New Guinea. Tree up to 30 m high, dbh up to 100 cm; with buttresses. Twigs terete, when young + angular, 4— 7 mm diam., grey to blackish, brown tomentellous, early glabrescent. Leaves 3—S-jugate; petiole terete, flattened to the base, 5—11 cm long by 1.5—2.5 mm broad; petiolules above slightly grooved, 5—15 mm long. Leaflets opposite to alternate, ovate, 5.5—14 by 2-5.25 cm, index 2—S, falcate, thin- to stiff-per- gamentaceous, glabrous, often with glands in the nerve axils (and in the branching of the veins) be- neath; base oblique (or symmetrical), acute (to obtuse), attenuate; apex acute (to tapering into an acute acumen); nerves 0.75—1.5 cm apart, widely spreading, usually strongly curved, ending free (or + looped and joined); reticulation coarse, above more prominent than beneath. /nflorescences pseu- doterminal, up to 12 cm long, sparsely branched, fulvous puberulous. Sepals broad-ovate, outside and inside sparsely sericeous. Petals glabrous, in- side with 2 scales. Stamens 8; anthers ciliate. Pis- til 3-locular, densely brownish hairy. Fruits ellip- soid to obovoid, not or shortly stipitate, not apicu- late, 4.5—6 by c. 4 cm, light brown tomentellous, + glabrescent; wall c. 1 cm thick, + fleshy, endocarp wrinkled, ferrugineous tomentose. Seeds unknown. ~ Fig. 80e. Distribution — Malesia: Papua New Guinea (Central and Milne Bay Provy., Fergusson Is.). Habitat & Ecology — Rain forest or secondary vegetation on hills, flood plains, and river banks; up to 200 m altitude. Fl. Dec.; fr. Mar., June, Nov. Uses — Used as a fish poison. 6. Trigonachras nov. sp. A: Leenh., Blumea 33 (1988) 207. — Based on SAN 75360 (L, SAN), Sabah. Tree, 30 m high, dbh up to 35 cm. Twigs terete, c.5 mm diam., grey, glabrous. Leaves 3—S-jugate; petiole terete, 3.5—4 cm long by 1.5—2 mm thick: petiolules above slightly grooved, 3-9 mm long. Leaflets opposite, elliptic, 5-10 by 1.75—4 cm, in- dex 2.5—3, slightly falcate, stiff-pergamentaceous, glabrous, without glands; base hardly to distinctly oblique, acute, strongly attenuate; apex acute; nerves 1—1.5 cm apart, spreading, straight, ending free; reticulation coarse, on both sides prominu- lous. Inflorescences pseudoterminal, 12-15 cm long, in fruit sparsely branched, fairly densely pu- berulous. Flowers only known from remains un- der the fruit. Sepals outside sparsely, inside densely hairy. Fruits (probably young) 3-angular-obovoid, c. 2.75 by 1.5 cm, narrowed to the base, apiculate, glabrous; wall somewhat fleshy, at least | mm thick, endocarp densely woolly. Distribution — Malesia: Borneo (Sabah, | col- lection). Habitat & Ecology — Hill side at 180 m alti- tude. Young fruits in July. Note — Presumably allied with T. acuta. 7. Trigonachras nov. sp. B: Leenh., Blumea 33 (1988) 209. — Based on S 28037 (L), Sarawak. Tree, 25 m high, dbh up to 65 cm; with but- tresses. Twigs terete, c. 4 mm diam., brownish black, subtomentellous, early glabrescent. Leaves 7-jugate; petiole subterete, c. 5 cm long by 2 mm thick; petiolules above slightly grooved, 3—5 mm long. Leaflets (sub)opposite, ovate, 7.5—11.5 by 34 cm, index 2.5—3, slightly faleate, membranous (young?), above glabrous, beneath tomentellous on the midrib, small glands in the lower nerve axils; base oblique, basiscopic side acute and strongly attenuate, ascroscopic side rounded and hardly attenuate; apex tapering into acute acumen; nerves c. | cm apart, straight to slightly curved, ending free; reticulation above dense, beneath coarse, on both sides prominulous. /nflorescences and flowers unknown. Fruits triangular ellipsoid, c. 3 by 2 cm, stipitate, apiculate, wrinkled, gla- brous. Distribution — Malesia: Borneo (Sarawak, 1 collection). Habitat & Ecology — On limestone slope, grey sandy soil; 250 m altitude. Fr. Sept. Note — Possibly related to T. nov. sp. C from Mindanao. 8. Trigonachras nov. sp. C: Leenh., Blumea 33 (1988) 210. — Based on BS 83920 (NY), Phil- ippines. Tree, up to 20 m high, dbh up to 40 cm. Twigs terete, c. 8 mm diam., dark grey, glabrous. Leaves 5-jugate; petiole terete, c. 9.5 cm long by 2.5 mm thick; petiolules above broadly shallowly grooved, 12-15 mm long. Leaflets (sub)opposite, elliptic, 11-15 by 5-6 cm, index 2-3, straight to subfal- cate, pergamentaceous, glabrous, some glands in the lower nerve axils below; base hardly oblique, acute, attenuate; apex acute or the very apex round- ed; nerves 1.75—2.25 cm apart, spreading, moder- ately curved, ending free; reticulation rather coarse, beneath more prominent than above. /nflorescenc- es and flowers unknown. /nfructescences pseudo- terminal, c. 30 cm long, hardly branched, puberu- lous, glabrescent. Fruits globular, 3—3.25 by 2.25-— 2.5 cm, abruptly narrowed into ac. 5 mm long stipe, apiculate, slightly wrinkled, glabrous; wall fleshy, 740 Flora Malesiana ser. I, Vol. 11 (3) (1994) 1-2 mm thick; endocarp densely woolly. Seeds Habitat & Ecology — In dry forest at low alti- unknown. tude. Distribution — Malesia: Philippines (Mindanao, Note — ResemblesT7. nov. sp. B and T. celeben- 1 collection). sis in the alliance of 7. cuspidata. TRISTIRA (P.W. Leenhouts) Tristira Radlk., Sapind. Holl.-Ind. (1879) 63, 98; in Engl., Pflanzenr. 98 (1932) 867. — Lectotype species (here proposed): Tristira harpullioides Radlk. |= Tristira triptera (Blanco) Radlk.]. Tree, monoecious. Indumentum of solitary, simple hairs only. Leaves paripinnate, 3— 7-jugate, without pseudo-stipules, neither petiole nor rachis winged. Leaflets opposite to alternate, not papillate beneath, velutinous beneath and on midrib and nerves above, gla- brescent above; base oblique, the acroscopic side broadest. Inflorescences terminal and in the upper leaf-axils, thyrsoid; bracts conspicuous, up to c. 6 mm long; cymules up to c. 7-flowered. Flowers actinomorphic, unisexual, male and female in the same inflores- cences. Sepals 5, free, imbricate, outer 2 slightly smaller, the inner more or less petaloid towards the margin, outside and inside densely appressedly short-hairy. Petals absent. Disc uninterrupted, woolly to glabrous, without appendages. Stamens 8 or 9, exserted in male flowers; filaments densely patent hairy at base; anthers glabrous (in female flowers ciliate), dehiscing latero-introrse. Ovary short-stipitate, smooth, densely fulvous-pilose, 3-locular, tapering into the terminal style; latter slightly longer than the ovary and sparse- ly appressed-hairy, bearing in the upper part 3 stigmatic grooves; ovules | per locule. Fruit indehiscent, sessile or stipitate, trigonous-ellipsoid to -globular, slightly grooved, winged; latter higher than broad; exocarp coriaceous (thin-fleshy when fresh), mesocarp fibrous-woody, endocarp hard-woody, inside rather densely white woolly. Seeds glabrous, without arillode, not winged. — Fig. 81. Distribution — Monotypic. Tristira triptera (Blanco) Radlk., Sapind. Holl.- Ind. (1879) 62, 63; Merr., Sp. Blanc. (1918) 239; Enum. Philipp. Flow. Pl. 2 (1923) 502; Radlk. in Engl., Pflanzenr. 98 (1932) 868; Kraemer, Trees West. Pacific Reg. (1951) 224, f. 80, excl. seed. — Melicocca triptera Blanco, Fl. Filip. ed. 2 (1845) 203; ed. 3, 2 (1878) 16. — Zollin- geria triptera Rolfe, J. Linn. Soc. Bot. 21 (1884) 309. — Type: Blanco s.n. (G-DC, acc. to Ra- dik.), Philippines. Tristira harpullioides Rad\k., Sapind. Holl.-Ind. (1879) 19, 62, 63; in Engl., Pflanzenr. 98 (1932) 869. — Type: Beccari hb. 2838 (FI holo), Moluccas. Guioa sp.: Vidal, Rev. Pl. Vasc. Filip. (1886) 95, pro no. 220; Ceron, Cat. Pl. Herb. Manila (1892) 53, pro Vidal 220. Tristira celebica Boerl. & Koord. in Koord., Mi- nah. (1898) 407; Koord., Suppl. Cel. 2 (1922) t. 54; Radlk. in Engl., Pflanzenr. 98 (1932) 869. — Type: Koorders 18840 (BO holo; M), N Celebes. Tristira pubescens Merr., Publ. Gov. Lab. Philipp. 6 (1904) 12; Radlk. in Engl., Pflanzenr. 98 (1932) 869. — Tristira pubescens Merr. f. gen- uina Radlk., Philipp. J. Sc., Bot. 8 (1914) 456, nom. illeg. — Type: Merrill 2842 (PNH# holo; BM, K, M), Philippines. Tristira pubescens Merr. f. hemidasya Radlk., Philipp. J. Sc., Bot. 8 (1914) 456. — Type: Ahern FB 421 (M holo; BO, K, SING), Philip- pines. Zollingeria macrocarpa auct. non Kurz: Fern.-Vill., Nov. App. (1880) 53; Vidal, Sinopsis (1883) t. 35) it, Gz Tree, 6—12(—20) m high, dbh up to 15 cm, bark white. Branchlets terete, 3—6 mm diam., ferrugin- Adema, Leenhouts, Van Welzen — Sapindaceae 741 icm lcm Fig. 81. Tristira triptera Radlk. a. Habit fruiting branch; b. cross section of fruit; c. female flower, stami- nodes already dropped; d. male flower; e. stamen (a: Clemens 17902: b: Santos 5153; c—e: PNH 7442). 742 eous-velutinous, early to late glabrescent, shiny greyish to purple-brown or black, scattered (often very sparsely) minutely pustular-lenticellate. Leaves (3—)5(—7)-jugate (those near the inflores- cences |- or 2-jugate), axial parts sparsely hairy, + glabrescent, to tardily ferrugineous-velutinous; petiole + flattened to grooved, at least at base, 3.5— 8 cm long, base somewhat pulvinate; rachis above flat with a narrow keel; petiolules above flat to deeply grooved, 2—4(—10) mm long. Leaflets ovate (lower leaflets) to elliptic (upper leaflets), rarely obovate, (3—)5—10(—16) by (1.2—)1.5—4(—6.2) cm, index 2—3.5, thin-chartaceous, above shiny; base acute to rounded, distinctly attenuate; margin en- tire [to (partly) crenate]; apex obtuse to emargin- ate, mucronulate or not; midrib above slender, prominulous to slightly sunken, with a fine keel, beneath prominent and rounded; nerves 0.8—1 cm apart, slightly curved, only the upper ones more or less looped and joined towards the margin, prominulous on both sides; intersecondary nerves few; reticulations fine, prominulous on both sur- faces. Inflorescences 7-30 cm long, with few erec- to-patent branches; latter sparsely rebranched, sparsely to densely fulvous- to ferrugineous-veluti- nous, cymules up to 7 mm long, with short stalk; pedicels slender, 2-3 mm long, hairy; bracts per- sistent, patent, linear, up to c. 6 by | mm, with a short petiole, chartaceous, margin recurved, apex acute, midrib distinct. Flowers creamy. Sepals: margin densely ciliate; outer broad-obovate to el- liptic, 2.2-2.8 by 1.8-2.5 mm, thick, base broad, margin subentire; inner orbicular to elliptic, 2.2—3 by 1.8—2.8 mm, thinner, base narrow, margin crenu- late to entire. Disc broad and flat. Stamens: fila- ments thread-like, tapering from base to apex, 2.2— Flora Malesiana ser. I, Vol. 11 (3) (1994) 3.5 mm long; anthers basifixed, oblong, 1.2—2.3 mm long, at base cleft over c. 40%, at the apex with a rounded (glandular?) appendage. Ovary sharply trigonous-ellipsoid, c. 2.5 mm high; style slender, c. 3 mm long, stigmatic part curved in old flowers. Infructescences sometimes still velutinous; sepals persistent and slightly accrescent. Fruits 2.5— 3.5 by 1.5—2.5 cm; wings coriaceous, up to 1 cm broad, centrally widest or broadening from base to apex, apically connate, either emarginate or taper- ing into the 3-4 mm long style rest; pericarp c. 1 mm thick. Seeds with basal, orbicular, small hilum; testa shining blackish-brown. — Fig. 81. Distribution — Malesia: E Philippines (from Luzon to Mindanao), Celebes, Moluccas (Sula Is- lands, Ceram). Habitat & Ecology — In forest on coral lime- stone, on cliffs along the beach, in hill forest, from sea level to c. 200 m altitude. Fl. Oct.—Feb., fr. Aug.—Mar. Note — The infraspecific variation is mainly in the degree of hairiness, the shape of the fruit, and the crenation of the leaflet-margins. The first char- acter, primarily used by Merrill and Radlkofer to delimit some species, grades throughout the range of the species. The shape of the fruit also grades from globular with the wings only touching at the apex (emarginate) to ellipsoid with the wings unit- ed and tapering into the stylar apiculum. The laxly crenate leaf-margin, used by Koorders and Radl- kofer to characterize T. celebica from Celebes, is also and even better shown by FB (Miranda) 20542 from Mindanao, while other specimens show a ten- dency towards it. None of these characters are cor- related. TRISTIROPSIS (P.W. Leenhouts) Tristiropsis Radlk. in Dur., Index Gen. Phan. (1888) 76; in Engl., Pflanzenr. 98 (1932) 861; S.T. Reynolds in Fl. Austral. 25 (1985) 11. — Lectotype species (here proposed): Tristiropsis acutangula Radlk. Palaoea Kanehira, Bot. Mag. Tokyo 49 (1935) 271. — Type species: Palaoea falcata Kanehira [= Tristiropsis acutangula Radlk.]. Trees, monoecious. Indumentum of solitary, simple hairs only. Branches (sub)terete, densely minutely tomentose, glabrescent. Leaves bipinnate or the upper ones at least forked, with several leaflets per branch; neither pseudo-stipules, nor winged; petiole pulvinate, + flattened above; pinnae alternate, rarely opposite, unequally pinnulate. Leaf- lets alternate to subopposite, chartaceous, glabrous or variably hairy; base oblique; mar- gin entire; midrib prominent beneath; nerves dense, spreading, reticulations elongated Adema, Leenhouts, Van Welzen — Sapindaceae 743 LS ee a parallel with the nerves, rather lax, prominulous on both surfaces. Inflorescences thyrses, axillary towards the end of the branches; peduncle pulvinate; branches few, spreading, bearing few to many, short- to long-stalked, few- to many-flowered (in the upper parts reduced to | flower) cymes; pedicels articulated at base. Flowers fairly big. Sepals 5, free, imbricate, concave, the two outer ones slightly or hardly smaller, fleshy, margin petaloid or not, outside appressedly, densely, shortly tomentose, margin ciliate. Petals 0 or 5, slightly shorter than the sepals, clawed or not, outside mainly towards the base sericeous, margin ciliate; scale single, (somewhat) bifid, hairy, not crested. Disc uninter- rupted. Stamens 8(—13), hardly or not exserted; filaments filiform; anthers basifixed, ovate- oblong, cleft at base for a third at most, the apex of the connective usually rounded- apiculate by a gland which is dark when dry, dehiscence latrorse. Pistil densely, appress- edly fulvous-hairy; ovary conical-ovoid, faintly 3(—5)-gonous, 3(—5)-locular, tapering into a short style with a 3-grooved stigma. Ovules | per locule. Fruits subdrupaceous, (1—)3(— 5)-celled, shortly attenuate to short-stipitate at base, shortly apiculate at apex, densely minutely tomentose, subglabrescent; wall thin, pericarp fibrous, endocarp woody; inside variably hairy. Seeds brown, hilum basal, orbicular: no arillode. — Fig. 82. Distribution — 3 species from the Marianas to Christmas Island (Pacific Ocean), Queensland, and the Solomon Islands; in Malesia: from Philippines and Borneo towards the east. See Leenh. & Balgooy, Blumea, Suppl. 5 (1966) 198, map 109. Habitat — Canopy trees of primary and secondary forests at low to medium altitudes. KEY TO THE SPECIES Peeeseiicmentn youOwiSi DROWN <'s 5... \. lush otae .chlon vest ue. 2 Bomanmenrum reddish browic.. ws... sk..e es. dak ode ee 3. T. ferruginea 2a. Sepals with a few hairs only inside. Petals present. Ovary 3(—5)-locular 1. T. acutangula b. Sepals densely hairy inside. Petals absent. Ovary 2-locular ....... 2. T. apetala 1. Tristiropsis acutangula Radlk. [in Dur., Index Type: Weinland 251 (B+ holo; K), New Gui- Gen. Phan. (1888) 76, nom. nud.] Sitzungsber. nea. Math.-Phys. Cl. K6énigl. Bayer. Akad. Wiss. Tristiropsis canarioides Boerl. ex Valeton, Ic. Bog. Miinchen 20 (1890) 248; in Engl., Pflanzenr. 98 (1932) 863; Whitmore, Gard. Bull. Sing. 22 (1967) 17; Foreman, Check List Bougainville (1971) 176, fig. — Type: Guppy 272 (K holo), Solomon Islands. Burseraceae? Hemsl., J. Linn. Soc. Bot. 25 (1890) 353. Tristiropsis obtusangula Radlk. {in Dur., Index Gen. Phan. (1888) 76, nom. nud.] Sitzungsber. Math.- Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 248; in Engl., Pflanzenr. 98 (1932) 863; Hosokawa, Trans. Nat. Hist. Soc. Form. 25 (1935) 30. — Type: Gaudichaud s.n. (P holo), Mariana Islands. 2 (Feb. 1906) 285, t. 186 & 187; Radlk. in Engl., Pflanzenr. 98 (1932) 865; Meijer Drees, Comm. For. Res. Inst. 33 (1951) 110; P. Royen, Man. For. Trees Papua & New Guinea 2 (1964) 49: Backer & Bakh. f., Fl. Java 2 (1965) 136; S.T. Reynolds in Fl. Austral. 25 (1985) 12, map 8. — Type: Teijsmann s.n. (BO holo), W New Guinea. Tristiropsis nativitatis Hemsl. ex Ridley, J. Str. Br. Roy. Asiat. Soc. 45 (June 1906) 182; Guillau- min, Ann. Jard. Bot. Buitenzorg 26 (1912) 214; Radlk. in Engl., Pflanzenr. 98 (1932) 866. — Tristiropsis ridleyi Hemsl., Hooker's Icon. PI. (Dec. 1906) t. 2812, nom. illeg. — Type: Rid- ley 67 (K, P), Christmas Island. Tristiropsis ovata Radlk. in Elmer, Leafl. Philipp. Bot. 5 (1913) 1605; in Engl., Pflanzenr. 98 Tristiropsis subangula K. Schum. in K. Schum. & Laut., Nachtr. Fl. Schutzgeb. Siidsee (1905) 310; Radlk. in Engl., Pflanzenr. 98 (1932) 864. — 744 Flora Malesiana ser. I, Vol. 11 (3) (1994) Fig. 82. Tristiropsis Radlk. Habit, sepals and ovaries. — T. ferruginea Leenh. a. Habit; b. petal from inside; c. ovary. — T. acutangula Radlk. d. Petal from inside; e. ovary. — T. apetala Leenh. f. Ovary (a: Kostermans 21330; b, c: Anderson 4626; d, e: LAE 4092; f: LAE 62105). Adema, Leenhouts, Van Welzen — Sapindaceae 745 (1932) 865. — Type: Elmer 11909 (M holo; BM, K), Philippines. Tristiropsis dentata Radlk., Denkschr. Kais. Akad. Wiss. Math.-Naturwiss. KI. 89 (1913) 572; in Engl., Pflanzenr. 98 (1932) 866. — Syntypes: Rechinger 4916 & 4620 (W, n.v.), Bougainville. Tristiropsis subfalcata Radlk., Philipp. J. Sc., Bot. 8 (1914) 455; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 502; Radlk. in Engl., Pflanzenr. 98 (1932) 864. — Type: Hallier 4584 (PNH? holo; L), Philippines. Tristiropsis oblonga Radlk., Philipp. J. Sc., Bot. 8 (1914) 456; in Engi., Pflanzenr. 98 (1932) 864. — Type: FB (Merritt) 4063 (PNH# holo; K, M), Philippines. Palaoea falcata Kanehira, Bot. Mag. Tokyo 49 (1935) 271, f. 25. — Tristiropsis falcata Kane- hira & Hatusima, Bot. Mag. Tokyo 57 (1943) 83 (‘Tristylopsis’). — Syntypes: Kanehira 2426 (K, P), Caroline Islands, Palau, Peliliu; Kane- hira 2427 (n.v.), Caroline Islands. Tristiropsis novoguineensis Kanehira & Hatusima, Bot. Mag. Tokyo 57 (1943) 83, f. 15 (‘Tristy- lopsis’). — Type: Kanehira & Hatusima 14249 (n.v.), W New Guinea. Tree, up to 35(—53) m, dbh up to at least 6 cm, often with buttresses up to 3 m high, 4 m wide, and 6 cm thick. Branchlets 4-10 mm thick, ful- vous-tomentose, glabrescent and then shiny pur- ple-brown, older parts more or less pustular lenti- cellate. Leaves up to 2 m long; petiole up to 20 cm long or more and up to 1.5 cm thick, rachis terete, the ultimate parts flattened to flat above; petiolules 1-8 mm long, above with a broad and flat groove, slightly pulvinate. Leaflets ovate to elliptic, 5-18 by 1.5—8 cm, index 2-4, glabrous (or bearded in the nerve-axils beneath); base acute to obtuse (or truncate); margin entire to subundulate; apex (emar- ginate to) obtuse (to acute), often mucronulate; midrib above slightly sunken, broad, nerves 0.75— 1.5 cm apart, nearly straight to slightly curved, all looped and joined near the margin, faint; intersec- ondary nerves well developed, often joined with the marginal arches. Thyrses up to 40 cm long, densely, appressedly, shortly, brownish hairy; pe- duncle up to 15 cm long; stalks to cymes up to 8 mm long; pedicels slender, 3-10 mm long. Sepals cream to greenish, persistent and black in fruit, outer 2 elliptic-ovate to orbicular, c. 3 by 2.5 mm; inner + obovate, c. 4 by 3 mm, margin petaloid, crenulate at apex, with a few hairs inside. Petals cuneate at base, broad-elliptic to broad-ovate, 2.5— the insertion of the scale long-ciliate, furthermore sparsely ciliate, apex crenulate, inside glabrous; scale about 0.5—0.6th the length of the petal, di- vided till the base, woolly on the inner surface. Disc divided into a thick, spreading outer and a thin, erect inner wall, black, glabrous or the latter cili- ate. Stamens: filaments c. 3 mm long, patent-hairy at least in the upper 0.6th part; anther c. 1.5 mm long, densely hairy on the dorsal side of the con- nective, sometimes also on the ventral side. Pistil: ovary c. 3 mm long; style slightly curved, c. 2mm long. Fruits ellipsoid to subglobular, widest about or above the middle, narrowed to short-stipitate at base, 3—4-angular to 3—4-ribbed in cross section, 20-30 by 14—25 mm, yellowish green to dark-yel- low when ripe, patently short-hairy inside, often sterile but well developed. — Fig. 82d, e. Distribution — Marianas (Guam), Carolines (Pa- lau Is.), Solomon Islands, Queensland, and Male- sia: Christmas Island, Kangean Archipelago, Ba- wean, Madura (acc. to Meijer Drees), Flores, Timor, SE Borneo, Philippines (Mindoro, Guima- ras I., Basilan, Mindanao), P. Muna near SE Celebes, Moluccas (Halmahera, Sula and Kai Islands), New Guinea, and the Admiralty Islands. Habitat & Ecology — A canopy tree of primary or secondary forests on soils with permanently high groundwater; mainly of low altitudes, up to about 500(—850) m altitude. Fl. Jan. -Aug., Oct.—Nov.; fr. Jan._Nov. Flowers with a sweet smell. Uses — The wood is said to be hard, heavy, and durable, and is used as a timber. For a description of the timber, see p. 427. Note — Tristiropsis obtusangula Radlk., the other syntype of the genus, is almost certainly conspe- cific with T. acutangula. The type specimen of the former (Gaudichaud s.n. in P) differs only by the distinctly stipitate, flattened-ellipsoid (not at all angular) fruits. It is from the Marianas; just like the other specimen available (Moran 4680, Guam), which is in flower and shows no differences with T. acutangula. 2. Tristiropsis apetala Leenh., Blumea 24 (1878) 509. — Type: LAE (Katik) 62105 (L holo), New Guinea. Tree, 8 m high, dbh up to 26 cm. Branchlets unknown. Leaves at least c. 1.5 m long, glabrous; petiole terete, canaliculate, c. 26 cm long by 1.25 cm thick; rachis terete, on the upper side with a slender rib; petiolules |—4 mm long, above with a broad and flat groove, pulvinate. Leaflets alternate, ovate, falcate, c. 18 by 7.5 cm, index c. 2.5, perga- mentaceous, glabrous; base acute at the narrow basiscopic side, rounded at the broad acroscopic side, attenuate; apex tapering to a short, cuneate, acute acumen; midrib above prominulous, slender; nerves 1—1.75 cm apart, nearly straight to slightly curved, ending free, faint at both sides; intersec- 746 Flora Malesiana ser. I, Vol. 11 (3) (1994) ondary nerves common but not reaching more than halfway the margin. Thyrsus c. 30 cm long, thinly minutely hairy, only the ultimate branches more densely so; peduncle hardly or not pulvinate, c. 3 cm long; cymules sessile (or with up to 2 mm long stalks); pedicels slender, 0.75—2.5 mm long. Se- pals yellow, broad-ovate, outer c. 3 by 3.5 mm, inner c. 3.5 by 4 mm, margin thin, slightly crenu- late, outside and inside densely appressedly short- hairy. Petals absent. Disc broad, flat, adnate to the base of the sepals, glabrous. Stamens 8, slightly exserted; filaments attached in the basal part, c. 3.5 mm long, sparsely woolly all over; anthers c. 1.75 mm long, the apex without a distinct gland, connective as well as thecae fairly densely woolly. Pistillode 2-locular. Fruits unknown. — Fig. 82f. Distribution — Malesia: Papua New Guinea (Madang Prov.); known from the type only. Habitat & Ecology — Rain forest at 90 m alti- tude. Note — The present species is unique in the ge- nus because it lacks petals. Moreover, it differs from T. acutangula, the species it resembles most, in the sepals being as densely hairy inside as outside. The apetalous flowers with a broad and flat disc are also found in other genera like Glenniea Hook. f. and Mischocarpus Blume. 3. Tristiropsis ferruginea Leenh., Blumea 13 (1966) 395. — Type: SF (Carr) 27276 (SING holo), Borneo. Tree, up to 30 m, dbh up to 40 cm. Branchlets c. | cm thick, when young densely orange-brown to ferrugineous tomentellous, glabrescent. Leaves: petioles 6-10 cm long; ultimate parts of the rachis above flattened to keeled; petiolules up to 2 mm long, above keeled, pulvinate. Leaflets alternate, ovate, 5—10 by 1.8—3 cm, index c. 3, beneath beard- ed in the nerve-axils, furthermore glabrous or thinly tomentose on the nerves beneath and on the mid- rib; base acute to obtuse, decurrent; apex tapering acuminate, acumen acute; midrib above slender and prominulous, nerves 0.5—1 cm apart, curved to straight, only the upper ones looped and joined near the margin, prominulous on both sides, but more strongly so beneath; intersecondary nerves more or less well developed, parallel to the nerves. Thyrses up to 17 cm long, densely tomentellous; peduncle 2-5 cm long; cymes with 1-2 mm long stalks; pedicels 1-2 mm long. Sepals dark purple, hardly or not persistent in fruit, outer and inner hardly different (outer 2 broad-ovate, inner broad- obovate), c. 2.5 by 2—2.2 mm, inside rather dense- ly appressed-hairy, margin entire, apparently not petaloid. Petals with a c. 1.25 mm long claw, the blade transversely half-elliptic, c. | by 1.5 mm, margin below the insertion of the scale densely woolly, upper part densely ciliate, apically crenu- late, inside woolly, the claw more densely so than the blade; scale rearching about halfway the blade, slightly bilobed, completely woolly. Disc 5-lobed, the lobes in the centre deeply hollowed, hence each + annular, the part towards the centre of the flower densely hairy, furthermore glabrous. Stamens 8; filaments 1-2 mm long, rather densely woolly in the lower 0.7th part; anthers c. 0.8 mm long, gla- brous. Fruits subglobular, c. 22 by 17 mm, at base contracted into ac. | mm long stipe, densely fer- rugineous tomentellous, (1-) 2- (or 3-)celled, in- side densely ferrugineous velvety. — Fig. 82a-c. Distribution — Malesia: Borneo (Sarawak, Ba- ram Dist., G. Api; Sabah, Mt Kinabalu; E Kutei, Sangkulirang Dist.). Habitat & Ecology — Primary forest on lime- stone; up to 500 m altitude. Fl. May, July; fr. Aug., Oct. Uses — For a description of the timber, see p. 427. XEROSPERMUM (P.W. Leenhouts) Xerospermum Blume, Rumphia 3 (1847) 99; Radlk. in Engl., Pflanzenr. 98 (1932) 936; Leenh., Blumea 28 (1983) 389; Yap in Tree Fl. Malaya 4 (1989) 460. — Xerosper- mum sect. Tetrasepalum Radlk. [in Dur., Index Gen. Phan. (1887) 76, nom. nud.] in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 331, nom. illeg. — Type species: Xero- spermum noronhianum Blume. Xerospermum sect. Pentasepalum Radlk. [in Dur., Index Gen. Phan. (1887) 76, nom. nud.] in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 331. — Syntype species: Xero- spermum acuminatum Radlk. (= X. laevigatum Radlk.), X. laevigatum Radlk. Adema, Leenhouts, Van Welzen — Sapindaceae 747 a 8 Medium-sized trees (or shrubs), dioecious. Indumentum of solitary simple hairs only. Branchlets terete, lenticels many, scattered, pustular but usually inconspicuous; wood reddish brown. Leaves paripinnate, (unifoliolate to) 1- or 2- (or 3-)jugate, without pseu- do-stipules; petiole subterete to terete, at the base slightly swollen and hollowed above: neither petiole nor rachis winged. Leaflets opposite, neither papillate nor glaucous be- neath, glabrous or puberulous on the nerves below and on the midrib. glabrescent, usual- ly on the lower side with few to several orbicular flat glands in the nerve axils and/or scattered all over the leaf surface, mainly in the basal half: no domatia: base (sub)symmetrical; margin entire. /nflorescences usually solitary in the lower leaf axils and tufted in the upper leaf axils; these tufts consist of a central axis and two to several branches in the axils of bud scales; the more branches to a cluster the shorter and the more equal they are; the central axis of such a cluster may end in a vegetative terminal bud and develop later in a vegetative branch: bracts usually caducous. Flowers actin- omorphic, 4- or 5-merous, unisexual. Sepals free or slightly connate, either all about equal or the outer two slightly smaller, the outer usually with a narrow, the inner with a broad membranous margin to nearly completely membranous. Petals about equal to or slightly shorter than the sepals, sessile to variably clawed, without a scale, woolly ciliate. Disc uninterrupted or interrupted, not lobed, in female flowers inconspicuous. Stamens (7) 8 (9), hardly to distinctly exserted in male fowers: filaments at least partly woolly; anthers attached dorsally at the base, dehiscence latrorse, glabrous or with a few hairs, sometimes ciliate. Pistil 2- (or 3-)locular; ovary deeply lobed, warty with on top of each wart a stiff brown hair; style apical, columnar, broadened to the apex, relatively short, with a few hairs or glabrous; stigma arched, elliptic with a longitudinal groove, (deeply cleft in fruit); ovules 1 per locule, nearly basally attached. Fruits with 1 (or 2) lobes developed, in the latter case lobes widely spreading, not winged, the lobes ellipsoid to subglobular, capsular, probably finally loculicidally dehiscent; the fruit wall outside spiny, warty, or colliculate to granular, soon glabrous, spines broader than high; inside smooth or slightly colliculate, glabrous. Seeds ellipsoid to subglobular, completely covered by a thin sarcotesta except for the basal hilum; testa inside with a pocket in which the radicle fits. — Figs. 83, 84. Distribution — Bangla Desh, Assam, Burma, the Indochinese Peninsula, and Malesia: Malay Peninsula, Sumatra, Java, Borneo. Two species. Habitat & Ecology — Often common in the midle and lower stories of the lowland and lower montane rain forests. The fruits are said to be eaten by birds, monkeys, and bats. Uses — The thin yellow to orange sarcotesta is eaten, but is not of importance. The opinions on the timber quality are rather divergent: fire wood, inferior timber, or a good, tough, and durable timber. See Heyne, Nutt. Pl. Indon. ed. 3 (1950) 997: Burkill, Dict. Econ. Prod. Malay Penins. ed. 2, 2 (1966) 2313. Note — In the older literature there was some confusion regarding the sarcotesta. With Hiern in Hook. f., Fl. Br. India 1 (1875) 686, the description reads: “testa ... fleshy, pilose, simulating an aril.” This became with Kurz, For. Fl. Burma | (1877) 295: “Seeds without arillus, the testa pilose, fleshy outside and arillus-like” (see also Kanjilal, Das & Purkay., Fl. Assam 1, 1936, 232). The mention of the sarcotesta as being pilose is incom- prehensible. 748 Flora Malesiana ser. I, Vol. 11 (3) (1994) nnn nnn EEE anna KEY TO THE SPECIES la. Flowers 5-merous. Pedicels in fruit not much swollen, 1.5—2.5(—3.5) mm thick 1. X. laevigatum b. Flowers 4-merous. Pedicels in fruit distinctly swollen, (3.5—)5—7 mm thick 2. X. noronhianum 1. Xerospermum laevigatum Radlk., [Sitzungs- 98 (1932) 949; Burk., Dict. Econ. Prod. Malay ber. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Penins. (1935) 2272; Wyatt-Smith & Kochum- Miinchen 8 (1878) 305, nom. nud.] Sapind men, Mal. For. Rec. 17, rev. ed. (1965) 364; Holl.-Ind. (1879) 23, 25; in Engl., Pflanzenr. Leenh., Blumea 28 (1983) 391; Yap in Tree Fl. 4 \Wessendor p 94 Fig. 83. Xerospermum laevigatum Radlk. subsp. laevigatum. Habit (de Wilde & de Wilde-Duyfjes 18907). Adema, Leenhouts, Van Welzen — Sapindaceae 749 Malaya 4 (1989) 461. — Type: Griffith KD 1006/1 (K holo), Burma. Xerospermum acuminatum Radlk., Sapind. Holl.- Ind. (1879) 7, 25; in Engl., Pflanzenr. 98 (1932) 948. — Type: Beccari PB 3468 (not 3408 as given by Radlk., 1879) (FI holo; K, P), W Bor- neo. Xerospermum unijugum Radlk., Rec. Bot. Serv. India 3 (1907) 351; in Engl., Pflanzenr. 98 (1932) 948. — Type: F. Kehding 090 (FI holo), Malay Peninsula. Xerospermum muricatum auct. non Radlk.: J.A.R. Anderson, Gard. Bull. Sing. 20 (1963) 169. (Shrub to) tree, up to 36 m high, dbh up to | m. Twigs 1-5 mm thick, glabrous. Leaves: petiole 0.4-7 cm long; petiolules 0.3—1.5 cm long; leaf axes glabrous. Leaflets elliptic, 4.5-18 by 1.75— 10 cm, index 1.75—3.75, coriaceous, glabrous, (without or) with few to several glands; base acute to rounded, decurrent; apex (rounded to acute or) variably acuminate; the upper (to most) nerves + distinctly joined; intersecondary veins variable; veinlets rather finely reticulate, prominulous at both sides, sometimes beneath more so than above. Inflorescences up to 20 cm long if solitary, no more than 5 cm if tufted; well developed axes with scat- tered side-branches, these patent and short or erec- to-patent and long; branches as well as the main axis (in its upper part) bearing several lax and of- ten several-flowered cymes, the lower long-stalked, consisting of a central flower and two sometimes long and many-flowered monochasial branches, the upper sessile and monochasial, often, if the axis is short, together forming a dense cluster of branch- es and flowers; bracts deltoid to lanceolate, up to 1.5 mm long, sparsely hairy; pedicels 1.5—5 mm long. Flowers 5-merous. Sepals free, all about equal or the outer two distinctly smaller, ovate to obovate, especially the outer two concave, 1.6—2.8 by 1.4—2.5 mm, ciliate (to woolly-ciliate), glabrous or nearly so (in Sumatra exceptionally outside and sometimes also inside thinly puberu- lous). Petals obovate to spathulate, 1—3 by 0.5—1.2 mm, bigger in male than in female flowers, whitish, variably woolly. Disc uninterrupted or interrupted, yellow. Stamens (7 or) 8; filaments 2—5 mm long, woolly in the lower half to all over but for the apex; anthers 0.6-0.9 mm long, gla- brous. Pedicels in fruit only slightly swollen, 1.5— 2.5(-3.5) mm thick. Fruit lobe(s) globular to oblong-ellipsoid, 2.5—3.75 by 1.5—2.5 cm, densely aculeate, orange to pinkish; wall coriaceous, c. 0.5 mm thick. Distribution — Burma (Mergui Archipelago) and Malesia: Malay Peninsula, Sumatra, Borneo. KEY TO THE SUBSPECIES la. Disc uninterrupted. Leaflets usually oblong to elliptic, not or slightly acuminate a. subsp. laevigatum b. Disc in female flowers nearly always, in male flowers sometimes interrupted. Leaflets broad- elliptic, distinctly acuminate b. subsp. acuminatum a. subsp. laevigatum: Leenh., Blumea 28 (1983) 392, f. la; Yap in Tree Fl. Malaya 4 (1989) 461. For synonyms, type, and more literature see X. /ae- vigatum Radlk. and X. unijugum Radlk. under the species. (Shrub to) tree, up to 36 m high, dbh up to | m; buttresses up to 2.5 m high, up to 2.3 m spreading, thick, (branching). Twigs rather slender, up to 4 mm thick. Leaves: petiole up to 4.5 cm long; ra- chis 2—3 cm long; leaflets usually oblong to ellip- tic, flat to slightly dorsiventrally curved, apex usu- ally not or shortly acuminate. /nflorescences up to 12 cm long, thin-puberulous, glabrescent. Sepals all nearly equal, usually thin and petal-like, not always glabrous. Petals up to 2 by 0.8 mm. Disc uninterrupted. — Fig. 83. Distribution — As the species. Habitat & Ecology — Primary and sometimes secondary forests, on dry land, crests, etc., on sandy clay or sandstone; up to 700 m altitude. Fl. Feb.— Mar., Sept., Dec.; fr. July, Aug., Dec. b. subsp. acuminatum (Radlk.) Leenh., Blumea 28 (1983) 393, f. 1b, d, e; 2d. See for synonyms, type, and literature X. acumi- natum Radlk. and X. muricatum auct. under the species. (Shrub to) tree, up to 20 m high, dbh up to 24 cm; (with slight buttresses or) with stiltroots. Twigs 1.5—5 mm thick. Leaves: petiole up to 7 cm long; rachis also up to 7 cm long; leaflets usually broad- elliptic, fairly strongly dorsiventrally curved, apex usually distinctly acuminate, acumen usually cuneate and acute, up to 3 cm long. /nflorescences if axillary 2 cm long or more, glabrous or rarely sparsely hairy. Sepals: outer ones distinctly small- er than inner ones, all fleshy or inner partly peta- loid, glabrous but for the margin. Petals up to 3 by 1.2 mm. Disc in female flowers nearly always in- terrupted in front of the inner sepal which lies in between the two outer ones (in S 9007, however, at the same place as in male flowers), in male flow- ers either uninterrupted or interrupted in front of the outermost one of the three inner sepals. — Fig. 84a. 750 Flora Malesiana ser. I, Vol. 11 (3) (1994) Distribution — Malesia: Borneo (Sabah, Bru- nei, Sarawak, West Kalimantan). Habitat & Ecology — Peat swamp forest; up to 30 m altitude. Fl. apparently throughout the year; ripe fr. known from Mar. and Apr. 2. Xerospermum noronhianum (Blume) Blume, Rumphia 3 (1847) 100; Miq., Fl. Ind. Bat. 1, 2 (1859) 552: Hiern in Hook. f., Fl. Br. India | (1875) 686, p.p.; Kurz, For. Fl. Burma | (1877) 295; Ridley, Trans. Linn. Soc. London, Bot. 3 (1893) 289: Koord. & Valeton, Bijdr. Booms. Java 9 (1903) 182; Atlas 4 (1918) pl. 797; Radlk. in Engl., Pflanzenr. 98 (1932) 946; Kan- jilal, Das & Purkay., Fl. Assam 1 (1936) 322; Adelb., Blumea 6 (1948) 324; Backer & Bakh. f., Fl. Java 2 (1965) 137; Leenh., Blumea 28 (1983) 394, f. 1c, 2a—c, e; Yap in Tree Fl. Ma- laya 4 (1989) 461; Jansen et al. in Verheij & Coronel (eds.), Pl. Res. SE Asia (PROSEA Handb.) 2, Edible fruits and nuts (1991) 365. — Euphoria noronhiana Blume, Bijdr. (1825) 234, comb. illeg. — Nephelium noronhianum Cambess., Mém. Mus. Hist. Nat. Paris 18 (1829) 30. — Lectotype (Leenhouts 1983): Blume s.n. (L sh. 908.272-748), Java. Euphoria xerocarpa Blume, Bijdr. (1825) 234, comb. illeg. (for lectotypification see Blume, Rumphia 3, 1847, 100). — Nephelium xerocar- pum Cambess., Mém. Mus. Hist. Nat. Paris 18 (1829) 30. — Arytera xerocarpa Adelb., Blu- mea 6 (1948) 324, nom. superfl. — Type: un- known (described from Nusa Kembangan near Java). [Sapindus glabratus Wall., Cat. (1847) no. 8095, nom. nud.] Cupania glabrata Kurz, J. As. Soc. Beng. 41, I (1872) 303; not Fern.-Vill., Nov. App. (1883) 349 [= Guioa koelreuteria (Blanco) Merr.|. — Xerospermum glabratum Radlk., Sitzungsber. Math.-Phys. Cl. Kénigl. Bayer. Akad. Wiss. Miinchen 8 (1878) 300; Sapind. Holl.-Ind. (1879) 23; Craib, Fl. Siam. Enum. | (1926) 329; Radlk. in Engl., Pflanzenr. 98 (1932) 946; Gag- nep. in Fl. Indo-Chine, Suppl. 1 (1950) 957. — Type: Kurz 2058 (K, M), Burma. Xerospermum lanceolatum Radlk., Sapind. Holl.- Ind. (1879) 7, 25; in Engl., Pflanzenr. 98 (1932) 943. — Type: Beccari PB 1031 (Fi holo; K, M, P), Sarawak. [Nephelium muricatum Griff., Cat. (1865) no. 1004, nom. nud. —] Xerospermum muricatum Radlk. [Sapind. Holl.-Ind. (1879) 23, 69, nom. inval.] in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1895) 331, f. 168; in Engl., Pflanzenr. 98 (1932) 940, f. 23; Corner, Wayside Trees (1940) 596, f. 215; non J.A.R. Anderson, Gard. Bull. Sing. 20 (1963) 169 (= Xerospermum laevigatum subsp. acuminatum). — Type: Griffith KD 1004 (M holo; K, L, P), Malay Peninsula. Xerospermum wallichii King, J. As. Soc. Beng. 65, II (1896) 432: Ridley, Fl. Malay Penins. 1 (1922) 498; Radlk. in Engl., Pflanzenr. 98 (1932) 943; M.R. Henderson, J. Mal. Br. As. Soc. 17 (1939) 42; Chin, Limest. Fl. Malaya (1973) 483. — Lectotype (Leenhouts 1983): King’s coll. 8725 (CAL holo?, n.v.; K), Malay Peninsula. Xerospermum brachyphyllum Radlk., Rec. Bot. Surv. India 3 (1907) 348; in Engl., Pflanzenr. 98 (1932) 942. — Type: Forbes 452 (G holo, n.v.; BO, L), W Java. Xerospermum cylindrocarpum Radlk., Rec. Bot. Surv. India 3 (1907) 348; in Engl., Pflanzenr. 98 (1932) 942. — Type: Forbes 2715 (B¥ holo; FI, L, P, SING), Sumatra. Xerospermum echinulatum Radlk., Rec. Bot. Surv. India 3 (1907) 350; Ridley, Fl. Malay Penins. 1 (1922) 497; Radlk. in Engl., Pflanzenr. 98 (1932) 944. — Syntypes: King’s coll. 8637 (K, L), Malay Peninsula, Perak, near Ulu Kerling; Scortechini 2104 (K, L, P), Malay Peninsula. Xerospermum intermedium Radlk., Rec. Bot. Surv. India 3 (1907) 348; Ridley, Fl. Malay Penins. | (1922) 497; Craib, Fl. Siam. Enum. | (1926) 329; Radlk. in Engl., Pflanzenr. 98 (1932) 944; Gagnep. in FI. Indo-Chine, Suppl. 1 (1950) 958. — Syntypes: Curtis 3436 (K, SING), Malay Pe- ninsula, Dindings, Lumot; Helfer KD 1005 (FI, K, L, M, P), Burma, King’s Island; Helfer 143 = KD 1006 (FI, K, L, M), Tenasserim and An- damans; Kehding 90 (FI), Malay Peninsula. Xerospermum fallax Radlk. in Fedde, Rep. 18 (1922) 340; in Engl., Pflanzenr. 98 (1932) 942. — Type: Blume in Hb. Martens (M holo), Java. Xerospermum testudineum Radlk. in Fedde, Rep. 18 (1922) 340; in Engl., Pflanzenr. 98 (1932) 941. — Syntypes: Bogor Bot. Gard. III.H.17 = Sutrisno 1 = Teijsmann 6676 (BO, L, M, P), Sumatra, Lampongs; Bogor Bot. Gard. IIL.1.28 = Diepenhorst s.n. (BO, L, M), Sumatra W Coast, Priaman; Forbes 1216a (B, n.v.), Java; Jelinek (W, n.v.), Java. Xerospermum xanthophyllum Radlk., Flora 118— 119 (1925) 400; in Engl., Pflanzenr. 98 (1932) 941. — Type: Bogor Bot. Gard. III.1.50a (M holo; BO, L, P). Xerospermum spec.: Corner, Wayside Trees (1940) SOT eZ: Nephelium maingayi auct. non Hiern: Hiern in Hook. f., Fl. Br. India 1 (1875) 688, p.p., ex- cluding the type (cf. Radlk. in Engl., Pflanzenr. 98, 1933, 964). Adema, Leenhouts, Van Welzen — Sapindaceae YW Tree, up to 25(—30) m high, dbh up to 30(—75) cm, (with buttresses). Twigs 1.5—5.5 mm thick, var- iably densely short hairy, early glabrescent. Leaves: petiole 1—7 cm long; petiolules 1-12 mm long: leaf axes glabrous or thinly to densely brownish or ful- vous puberulous, glabrescent. Leaflets + elliptic, up to 50 by 30 cm, index 1.5—2.5(—6.5), pergamen- taceous to coriaceous, glands few to many; base obtuse to acute; apex rounded to variably acumi- nate; nerves ending free but for the few uppermost ones; intersecondary nerves not conspicuous; veins and veinlets finely to laxly reticulate, about equal- ly raised on both sides to smooth (or slightly sunk- en) on the upper side. /nflorescences up to 25 cm long if solitary, much shorter if tufted; axes simple or with some short patent branches in the lower part, all branches bearing few to many sessile or subsessile, very condensed, few- to several-flow- ered cymes, towards the apex reduced to solitary flowers; bracts deltoid to lanceolate, up to 1.5 mm long, sparsely hairy; pedicels c. 2 mm long. Flow- ers 4-merous. Sepals free or slightly connate, the outer two usually slightly smaller than the inner ones, ovate to obovate, 1—2(—3) by 1—2.4 mm, out- side and inside glabrous or hairy (nearly always inside at the base), ciliate. Petals obovate to broadly Fig. 84. Xerospermum Blume. Detail of leaflet, fruits. — X. laevigatum Radlk. subsp. acuminatum (Radlk.) Leenh. a. Fruit. — X. noronhianum (Blume) Blume. b. Detail lower surface of leaflet: c—e. fruits showing differences in sculpture (a: SAN 27301; b: FRI Ja 6185: c: KEP/FRI 10748; d: Stone & Sidek 12523; e: KEP 29472). 752 Flora Malesiana ser. I, Vol. 11 (3) (1994) spathulate, 1-2.8 by 0.5—1.7 mm, short- to long- clawed with an ovate to transversely elliptic blade, variably woolly, nearly always with the exception of the base outside, inside often sparsely hairy to glabrous. Disc uninterrupted. Stamens 8 (9); fila- ments 1.5—2.5 mm long, woolly with the excep- tion of base and apex to woolly in the upper half (to glabrous); anthers 0.3—0.8 mm long, glabrous or with a few hairs, (ciliate). Pedicels in fruit strong- ly swollen, (3.5—)5—7 mm thick. Fruit lobes ellip- soid to subglobular (or obovoid), 1.75—5 by 1.25— 5 cm, aculeate or tuberculate (and/or colliculate to granulate), red or darkbrown; wall coriaceous, corky, or woody, 0.65—2.5 mm thick. — Fig. 84 b-e. Distribution — As the genus. Habitat & Ecology — Primary and secondary rain forest, sometimes in peat forest, heath forest, or bamboo forest, on plains as well as on slopes and crests, in dry places but also on river banks, along marshes, or in periodically flooded localities, on different kinds of soil (sand, sandy clay, fertile volcanic loam, peat, subsoil granite, sandstone, or limestone); sea level up to 300(—1500) m altitude. Especially in continental Asia (Thailand, Indo- China) often common. FI. throughout the year but mainly Jan.—Apr. and also rather frequently Aug.— Dec.; fr. mainly Jan.—Aug. The fruits are eaten by birds and monkeys. Notes — 1. The variation in leaves, flowers, and especially the fruits, is extensive but continuous in this species. At first sight it is difficult to imagine that a rough fruit without any appendages could belong to the same species as a fairly densely spiny fruit. However, the rough fruit shows a basic pat- tern corresponding with bundles of spines in the aculeate fruits, moreover, fruits with short spines are still rough in the grooves between the bundles of spines. As to the flowers, the greatest variation is in the hairiness of the sepals and petals. The only entity that is reasonably well characterized is ‘Xerospermum wallichii’. \t has relatively broad leaflets with a laxly reticulate venation; the sepals are glabrous on both sides, the petals are glabrous inside, and the fruits are rough and without spines. The only one of these characters that seems to be exclusive is that of the petals; all other characters are extremes in a continuous series. Typical ‘X. wal- lichii’ is only known from the Malay Peninsula where it appears to be rather common. It is unde- sirable to segregate such a local, morphologically rather extreme but hardly sharply delimited form from the rest of the species. 2. The two combinations under Euphoria, viz. E. noronhiana and E. xerocarpa, are illegitimate as Euphoria is illegitimate; see Leenh., Blumea 19 (1971) 116. 3. Blume’s Euphoria xerocarpa was based upon both flowering and fruiting material. Both the im- portance Blume attached to the fruit characters in the Sapindaceae in general and the specific epi- thet point to the fruiting part as the most logical choice for a lectotype. This is fully in accordance with Blume’s own choice: in 1847 (Rumphia 3: 100) he recognized the fruiting part as belonging to his new genus Xerospermum, where he placed it in the synonymy of X. noronhianum. The flower- ing part became the basis of a new genus and spe- cies, Arytera litoralis. A century later, Adelbert (Blumea 6, 1948, 324) made a different choice. The fruiting part remained under X. noronhianum, and the flowering part became the lectotype of Eupho- ria xerocarpa. Accordingly, Euphoria xerocarpa became synonymous with Arytera litoralis, and as the epithet xerocarpa was older than litoralis, a new combination had to be made, Arytera xero- carpa. As Blume’s lectotypification was much ear- lier and more in accordance with all evidence avail- able, the combination Aryfera xerocarpa was su- perfluous and hence illegitimate. 4. The 1879 publication of Xerospermum muri- catum was invalid as it was actually a nomen nu- dum. The few characters mentioned were only dif- ferences between Nephelium and Xerospermum, and lead to the conclusion that the species belongs to Xerospermum. The references to older publica- tions are either to nomina nuda or to mixtures. See Radlk., Sapind. Holl.-Ind. (1879) 69-70. ZOLLINGERIA (F. Adema) Zollingeria Kurz, J. As. Soc. Beng. 41, I (1872) 303, nom. cons.; Ra. Math.-Phys. Cl. K6nigl. Bayer. Akad. Wiss. Miinchen 20 (1890) 293; in Engl., Pflanzenr. 98 (1933) 724: Adema, Blumea 37 (1992) 73. — Type species: Zollingeria macrocarpa Kurz. Adema, Leenhouts, Van Welzen — Sapindaceae 753 1cm 1cm Barend Fig. 85. Zollingeria borneensis Adema. a. Habit; b. male flower; c. female flower; d. ovary, longitudinal section; e. ovary, cross section; f. fruit; g. fruit, bottom view; h. seed; i. seed, bottom view (a-e: SAN 90057; f-i: Elmer 20888). 754 Flora Malesiana ser. I, Vol. 11 (3) (1994) Trees. monoecious. Indumentum of solitary simple hairs, no glands. Twigs terete. Leaves spirally arranged, paripinnate, 1—8-jugate; pseudo-stipules absent; neither petiole nor rachis winged. Leaflets opposite to alternate, margin entire. Inflorescences axillary, thyrsoid, branched. Flowers unisexual, regular (or zygomorphic). Sepals 5, free, slightly to dis- tinctly unequal, (not) imbricate, not petaloid, outside hairy (or glabrous). Petals 5 (or 4), shorter (or longer) than the sepals, with or without scales or auricles. Disc entire (or interrupted), glabrous. Stamens 8, (slightly) exserted in male flowers, filaments hairy, anthers glabrous. Ovary at the base 3-celled, higher up 1-celled; style 3-lined or 3-lobed. Ovules | per cell. Fruit 1-celled, 3-winged, indehiscent. Seeds | per fruit, without arillode or sarcotesta. — Fig. 85. Distribution — 2 or 3 species in Burma, Laos and Thailand; | in Malesia: Borneo (Sabah). Zollingeria borneensis Adema, Blumea 37 (1992) 73-76. — Type: Elmer 20888 (A, F, L, U), Sabah. Trees, 60 m high, clear bole 3 m high, dbh 70 cm. Bark whitish, inner bark pale yellow, 0.7 cm; sapwood white. Indumentum ferrugineous tomen- tellous. Twigs 2-3 mm in diam., striate, silvery- grey, with many small lenticels, soon glabrous. Leaves (1-) 2- or 3-jugate; petiole 1-6.5 cm long, pulvinate; rachis 1-10.5 cm long; petiole and ra- chis semiterete, striate, short patently hairy; peti- olules reduced to pulvinus, 1-4 mm long, semi- terete, short-hairy. Leaflets + elliptic, 9.5—20 by 2-6 cm, index 2.9-3.6, chartaceous, above and below glabrous, midrib and nerves sparsely to densely short-hairy; base slightly oblique, (broad- ly) cuneate; apex acuminate, very apex rounded; midrib prominent above, nerves 9-12 per side, 8— 28 mm apart, angle to midrib 60-65°. Inflorescenc- es together pseudoterminal, 6—14 cm long; cymes several-flowered; bracts and bracteoles acicular, 0.6-12 by 0.1-0.4 mm, outside and inside hairy; pedicels 1.5—2.5 mm long, articulated near the base, angular, hairy. Flower buds about globular, 1.7 by 1.9 mm. Flowers yellowish to green; male ones somewhat smaller than the female ones. Sepals tri- angular to spathulate, not imbricate, thinned to- wards the margin especially apically, slightly un- equal, 1.7—2.6 by 1.2-1.6 mm, apex rounded to retuse, outside and inside appressed-hairy, ciliate. Petals cuneate to spathulate, distinctly clawed, 1.4— 2.4 by 1.2-1.6 mm, ciliate, claw 0.4-0.7 mm long, hairy on both surfaces, auricles ciliate. Disc en- tire, annular or saucer-shaped. Filaments of sta- mens 1.5 mm long, anthers 0.4 mm long; in fe- male flowers filaments 1.2-1.6 mm long, anthers 0.5—0.6 mm long. Ovary outside hairy, inside gla- brous; style | mm long, stigma + capitate, 3-lobed; pistillode 3 by 1.9 mm. Fruits prismatoid, sharply deltoid in cross section, 6-7 by 2.5—5 cm; wail thin, coriaceous, outside striate, densely velutinous, in- side glabrous. Seeds prismatoid, 33 by 14 mm. — Fig. 85. Distribution — Malesia: Borneo (Sabah). Habitat & Ecology — Primary forest, altitude Sim. EL Feb: CULTIVATED, EXOTIC SAPINDACEAE Filicium decipiens (Wight & Arn.) Thw., Enum. Pl. Zeyl. (1864) 58, sub Pteridophyllum decip- iens, 408; Radlk. in Engl., Pflanzenr. 98 (1933) 1427; Backer & Bakh. f., Fl. Java 2 (1965) 142. Trees. Indumentum sparse, sericeous, most parts with scale hairs. Leaves alternate, paripinnate, 5—many-jugate, rachis (broadly) winged. Leaflets narrow, long, shiny; nerves many, straight, scalar- iform; domatia absent. Inflorescences axillary to pseudoterminal panicles. Flowers unisexual. Sepals 5, equal, ovate, ciliate, with scale hairs outside. Petals 5, elliptic, about as large as sepals, ciliate; scales and auricles absent. Disc flat, complete, 5-lobed, densely pilose. Stamens 5, glabrous, exserted in male flowers, anthers not opening in female flowers. Ovary compressed- globose, finely hairy at base, 2-locular, rudimen- tary in male flowers; style short; stigma 2-lobed; one ovule per locule. Fruit a drupe, smooth, glabrous, thin-walled. Seeds without fleshy struc- ture. — Fig. 86. Adema, Leenhouts, Van Welzen — Sapindaceae 755 & ey ———s , icm ‘Q a Fig. 86. Filicium decipiens (Wight & Arn.) Thw. a. Habit; b. fruit (a: SAN 7/299: b: SAN 7358/). Distribution — Indigenous in S§ India and Sri Koelreuteria paniculata Laxm., Nov. Comm. Lanka, but now a pantropical ornamental. In Acad. Petrop. 16 (1772) 561, t. 18; P. Royen, Malesia locally cultivated as an ornamental (e.g. Manual For. Trees Papua New Guinea 2, Sa- Street tree) in Java and the Lesser Sunda Islands. pindaceae (1964) 28. 756 Flora Malesiana ser. I, Vol. 11 (3) (1994) Deciduous trees. Young parts puberulous. Leaves alternate, imparipinnate or bipinnate. Leaf- lets dentate to divided. Inflorescences large termi- nal panicles. Flowers unisexual, zygomorphic. Se- pals 5, connate at base, slightly unequal. Petals 4, longer than sepals, yellow, reddish at base, turned upwards; scales 2. Disc undulate, lowest or inter- rupted at the place of the missing petal. Stamens 8, exserted in male flowers. Fruits bladder-like, in- flated, loculicidal capsules. Seeds globose, with- out fleshy structures. Distribution — Indigenous in Korea and China; widely cultivated elsewhere. In Malesia according to Van Royen, I.c., cultivated in New Guinea as an ornamental. No herbarium material seen. Several indigenous species are also found cultivated as fruit trees: Dimocarpus longan, Litchi sinensis, Nephelium species, especially N. lappaceum. They have been treated under the respective genera. Index to scientific plant names 757 Se nee e cena ee INDEX TO SCIENTIFIC PLANT NAMES Suprageneric epithets have been entered under the family name to which they belong preceded by the indication of their rank (subfamily, tribe, etc.). Infrageneric epithets have been entered immedi- ately under the generic name to which they belong, preceded by the indication of their rank (subgenus, 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. Aceraceae 425, 431, 433, 434 (Alectryon) (Alectryon cobbe) Affonsea pteropoda Kosterm. 613 Aglaia chartacea Kosterm. 632 Alectryon Gaertn. 422, 430, 446*, 449, 450 subg. Alectryon 438, 444, 447, 449, 451, 454 subg. Synalectryon (Radlk.) Leenh. 439, 447, 449, 451, 452 sect. Eualectryon Radlk. 454 sect. Spanoghea (Blume) Radlk. 454 sect. Synalectryon Radlk. 452 affinis Radlk. 451, 452, 733 cardiocarpus Leenh. 438, 452, 453*, 455 celebicus Radlk. 456 connatus Radlk. 446*, 451, 453*, 454 coriaceus Radlk. 454 excisus Radlk. 456 ferrugineus (Blume) Radlk. 446*, 452, 453* 455 ferrugineus auct. 457 fuscus Radlk. 451, 456 glaber (Blume) Radlk. 446*, 452, 453*, 456 inaequilaterus Radlk. 456 kangeanensis Leenh. 451, 454 macrophyllus Kaneh. & Hatus. 455 mollis Radlk. 455 myrmecophilus Leenh. 451, 453*, 457 ochraceus Radlk. 456 repandodentatus Radlk. 422, 436, 443*, 451, 453*, 457 reticulatus Radlk. 451, 453*, 458 reticulatus auct. 455 serratus Radlk. 456 sp., Leenh. 544 sphaerococcus Radlk. 456 strigosus Radlk. 455 subdentatus (Benth.) Radlk. forma pseudostipularis Radlk. 458 Allophylus L. 421, 425, 430, 459 amboinensis Blume 461 apiocarpus Radlk. 461 bartlettii Merr. 462 betongensis Craib 462 blancoi Blume 460 brevipetiolatus Radlk. 462 cambessedei Blume 461 celebicus Blume 461 chlorocarpus Radlk. 462 cobbe (L.) Raeuschel 424, 433, 436, 441, 459, 464* forma blancoi Fern.-Vill. 461 forma fulvinervis (Blume) Backer 460 subforma bantamensis (Blume) Backer 460 forma /ittoralis (Blume) Backer 460 forma villosus (Roxb.) Hiern 460 var. blancoi (Fern.-Vill.) Vidal 461 var. glaber Corner 462, 465 var. glabra (Roxb.) Prain 460 var. grossedentata (Turez.) Vidal 461 var. limosus Corner 462 var. marinus Corner 462 var. racemosus (L.) Ridley 460 var. velutinus Corner 462 var. villosus Corner 462, 465 dasythyrsus Radlk. 462 dimorphus Radlk. 461 edulis 429, 433 var. edulis 432 filiger Radlk. 461 fulvinervis (Blume) Blume 460 var. bantamensis (Blume) Blume 460 var. bullatus Blume 460 var. burmannianus Blume 460 var. fuscus Blume 460 var. hircinus Blume 460 var. montanus Blume 460 var. reclinatus Blume 460 var. rufescens (Blume) Mig. 461 var. sericeus Blume 460 var. velutinus Blume 460 glaber Boerl. 460 granulatus Radlk. 462 grossedentatus (Turcz.) Fern.-Vill. 461 hymenocalyx Radlk. 462 insignis Radlk. 462 integrifolius Blume 460 Javensis (Blume) Blume 460, 465 var. cuspidata 460 var. genuinus Radlk. 460 var. racemosa 460 var. rigida 460 var. robusta 460 var. striata Blume 460 laetevirens Ridley 462 758 (Alectryon) largifolius Radlk. 462 leptocladus Radlk. 462 leptococcus Radlk. 461 leptostachyus (Blume) Boerl. 461 leucochrous Radlk. 462 leucocladus Radlk. 462 ligustrinus Blume 461 littoralis (Blume) Blume 460 var. repanda Blume 460 var. serrulata Blume 460 lopezii Merr. 462 macrostachys Radlk. 461 malvaceus Radlk. 462 micrococcus Radlk. 461, 465 mutabilis (Blume) Boerl. 461 obliquus Radlk. 462 peduncularis Radlk. 462 quinatus Radlk. 461 racemosus (L.) Boerl. 460, 465 repandodentatus Radlk. 462 rufescens Blume 461 var. cuspidatus Blume 461 var. heterodon Blume 461 rugosus Blume 461 samarensis Merr. 462 scandens Ridley 462 sessilis Blume 461 setulosus Radlk. 461 simplex Quis. 462 simplicifolius Radlk. 462 stenophyllus Merr. 462 subincisodentatus Radlk. 462 sumatranus Blume 461, 465 var. crenulatus Blume 461 var. elongatus Blume 461 var. glabriusculus Blume 461 var. politus Blume 461 sundanus Migq. 461 ternatus Lour. 460 ternatus (Forst. & Forst.) Radlk. 460, 465 timoriensis (DC.) Blume 460, 465 tomentosus Boerl. 461 triphyllus (Burm.) Mert. 460 unifoliolatus Radlk. 461 Flora Malesiana ser. I, Vol. 11 (3) (1994) (Alectryon) villosus (Roxb.) Blume 460 zeylanicus L. 460 Amesiodendron Hu 465 chinense (Merr.) Hu 439, 449, 466*, 467 integrifoliolatum Lo 467 tienlinensis Lo 467 Ampacus litorea (angustifolia) minor Rumph. 459 Ampacus litorea prima Rumph. 459 Anomosanthes longifolia (Hiern) Pierre 631 Aphania Blume 627, 650 sect. Euaphania Radlk. 627 angustifolia Radlk. 652 boerlagei Valeton 652 cuspidata (Blume) Radlk. 651 danura (Roxb.) Radlk. 651 dasypetala Radlk. 652 dictyophylla (Radlk.) Radlk. 650 fascicularis Radlk. 652 loheri Radlk. 652 longipes Radlk. 652 macrophylla Radlk. 652 masakapu Melch. 652 montana Blume 651 paucijuga (Hiern) Radlk. 651 philippinensis Radlk. 652 senegalensis (Poir.) Radlk. 651 sphaerococca Radlk. 652 Aphanococcus celebicus Radlk. 634 Apiocarpos Montrouz. 598, 603 moguinii Montrouz. 599, 603 Aporetica Forst. & Forst. 459 gemella DC. 460 penicellata Blanco 460 ternata Forst. & Forst. 460 Aporrhiza 430 Arytera Blume 440, 447, 467 angustifolia Radlk. 473 bifoliolata S.T. Reynolds 470 brachyphylla Radlk. 449, 470, 471 bullata H. Turner 449, 469, 471, 472* (Arytera) densiflora Radlk. 469, 472 divaricata auct. 478 foveolata auct. 479 geminata Radlk. 474 gigantosperma Radlk. 473 karang Miq. 570 lineosquamulata H. Turner 469, 473 litoralis auct. 476, 601 litoralis Blume 427, 469, 470, 472*, 473, 752 forma angustifolia Radlk. 473 forma genuina Radlk. 473 forma major Radlk. 474 forma rufescens Radlk. 473 var. major King 474 macrobotrys (Merr. & Perry) R.W. Ham 449, 469, 474 miniata H. Turner 470, 475 montana Blume 623 montana auct. 571 morobeana H. Turner 469, 476 multijuga H. Turner 469, 476, 477* musca H. Turner 470, 478 novaebrittanniae H. Turner 469, 478 ochracea Blume ex Koord. 473 pseudofoveolata H. Turner 469, 479 rufescens Radlk. 473 silaka Miq. 591 sordida Radlk. 721 sp., S.T. Reynolds 479 xerocarpa (Blume) Adelb. 473, 750, 752 xerocarpa auct. 597 Atalaya Blume 421, 437, 444, 479 sect. Atalaya 480 bijuga Span. 482 papuana (Radlk.) Leenh. 480, 481* salicifolia (DC.) Blume 443*, 480, 482* Ay-Assa Rumph. 606 Blancoa Blume 598, 601 arborea Blume 601 Blighia 430 sapida 432, 433 Index to scientific plant names 759 Boa massy Rumph. 491 (Cupania) Cupanieidites 425 Boswellia obliqua Blanco sect. Guioa G. Don 548 Cupaniopsis Radlk. 422, 437, 540 sect. Mischocarpus Miq. 449, 450, 493 Burseraceae? 743 Capura alata (Blume) Teijsm. & Binn. 646 fruticosa (Roxb.) Vidal 643 multijuga Hook. f. ex Radlk. 642 spectabilis (Blume) Teijsm. & Binn. 636 Cardiospermum L. 421—423, 425, 430, 432, 435, 440, 441, 483 corindum L. 484, 486 grandiflorum Swartz 432, 483, 484 halicacabum L. 484, 485* var. /uridum (Blume) Adelb. 484 var. microcarpum (H.B.K.) Blume 484 lyridum Blume 484 microcarpum H.B.K. 484 Caryophyllaster litoreus Rumph. 523, 526, 527 Chonopetalum Radlk. 541 Chytranthus 430 Cnemidiscus thorelii Pierre 544 Cnesmocarpon Adema 440, 441, 486 dasyantha (Radlk.) Adema 487, 488* dentata Adema 440, 444, 487, 489* discoloroides Adema 487, 489* 490 montana Adema 487, 489*, 490 Connarus jackianus Wall. 623 Crossonephelis Baill. 540 palawanicus Leenh. 544 penangensis Leenh. 542 philippinensis Leenh. 542 thorelii Leenh. 544 Cubilia Blume 422, 423, 490 blancoi Blume 491 blancoi auct. 690 cubili (Blanco) Adelb. 436, 438, 445, 491, 492* 493* rumphii Blume 491 Cupania 430 sect. Dimereza G. Don 549 sect. Elattostachys Blume 527 658 acuminata (Blume) Miq. 508 acuta Hiern 735 anacardioides A. Rich. 497 blumei Steud. 606 brachyphylla F. Muell. 471 cordierii F. Muell. 730 cunninghamii Hook. 520 diphyllostegia F. Muell. 520 diplopetala Hassk. 570 distachya Blume 533 erythrocarpa F. Muell. 732 erythrorhachis Miq. 667 fuscescens Miq. 663 fuscidula Kurz 570 glabrata Kurz 750 glabrata auct. 571 griffithiana Kurz 585 griffithiana auct. 571 Jackiana Hiern 623 lachnocarpa F. Muell. 660 lessertiana Cambess. 667 minjalilen Blume 570 mischocarpus Steud. 667 mutabilis Miq. var. coriacea Mig. 533 obtusa (Blume) Mig. 510 pallidula Hiern 591 pentaphylla Hiern 662 pentaphylla Wight 662 perrottetii Cambess. 624 pleuropteris Blume 585 forma apiculata Radlk. 585 var. apiculata Hiern 585 var. bijuga Hiern 567 pseudorhus A. Rich. 617 pyriformis F. Muell. 664 regularis Blume 570 regularis auct. 576 revoluta Rolfe 667 richii A. Gray 624 robertsonii F. Muell. 711 roxburghii Wight 662 rupestris Cambess. 606 salicifolia (DC.) Decaisne 482 semiglauca var. acutifolia F. Muell. 564 subundulata Rolfe 663 sumatrana Miq. 663 zippeliana Blume 535 sect. Elattopetalum Radlk. 494 sect. Macropetalum Radlk. 494 acuticarpa Adema 496* anacardioides (A. Rich.) Radlk. 495, 497 angustifolia Radlk. 499 bilocularis Adema 437, 495, 496*, 497 brachythyrsa Radlk. 501 bullata Adema 495, 498 celebica Adema 437, 495, 496*, 499 curvidens Radlk. 496*, 499 denticulata Radlk. 499 dictyophylla Radlk. 650 euneura Adema 496*, 500 flaccida Radlk. 499 gigantophylla Radlk. 499 grosseserrata Radlk. 501 insularis Radlk. 499 longifoliata Kaneh. & Hatus. 499 macropetala Radlk. 495, 496*, 501 multidens Radlk. 499 multijuga Merr. & Perry 499 napaensis Adema 495, 502*, 502 oxypetala Radlk. 504 papuana Radlk. 499 patentivalvis Radlk. 546 phanerophlebia Merr. & Perry 495, 503 platycarpa Radlk. 437, 447, 495, 502*, 503 remotidens Merr. & Perry 499 reticulata Merr. & Perry 499 rhytidocarpa Adema 495, 496*, 504 serrata (F. Muell.) Radlk. forma vestita Radlk. 499 stenopetala Radlk. 496, 504, 505* forma genuina Radlk. 504 strigosa Adema 495, 506 subdentata Radlk. 499 subserrata Radlk. 499 760 Cussambium Rumph. 728 glabrum Ham. 728 oleosum O. Kuntze 728 spinosum Ham. 728 Dabanus Rumph. ex Kuntze 698 acuminatus (Hook. f.) Kuntze 702 Deinbollia Schum. & Thonn. 430, 713 Dictyoneura Blume 435, 507 acuminata Blume 447, 507, 508 subsp. acuminata 446*, 508, 509* subsp. microcarpa J. Dijk 510 bamleri K. Schum. & Laut. 510 bamleri auct. 508, 510 integerrima Radlk. 511, 539 microcarpa Radlk. 510 obtusa Blume 449, 507, 509*, 510 philippinensis Radlk. 508 rhomboidea Radlk. 508 sp., J. Dijk 507, 511 sp., Streimann 510 sphaerocarpa Radlk. 508 sphaerocarpa auct. 510 subhirsuta Radlk. 508 Didymococcus verticillatus (Lindl) Blume 651 Dimereza Labill. 549 Dimocarpus Lour. 422, 430, 436, 445, 446, 511 dentatus Meijer ex Leenh. 513, 514* foveolatus (Radlk.) Leenh. 513 fumatus (Blume) Leenh. 512. 513, 515, 516* subsp. fumatus 515 subsp. javensis (Radlk.) Leenh. 515 subsp. philippinensis Leenh. 515, 516* longan Lour. 424, 427, A34, 512,517,518, 756 subsp. longan 518 var. longan 442*, 516%, 518 Flora Malesiana ser. I, Vol. 11 (3) (1994) Ne (Dimocarpus longan) subsp. malesianus Leenh. 518 var. echinatus Leenh. AAD*-SI6*, 518, 519 var. malesianus 442%*, 512, 516*, 518, 519, 658 Diploglottis Hook. f. 520 australis (G. Don) Radlk. 429, 440, 449, 520, 521%, 538 cunninghamii Hook. f. 520 diphyllostegia F. Muell. ex F.M. Bailey 520 Diplopeltis 424 Diplopetalon Spreng. 549 Distichostemon 424 Dittelasma Hook. f. 713 rarak (DC.) Hook. f. 714 Dodonaea Miller 420, 421, 424, 430, 431, 435, 441, 522 angustifolia L.f. 523, 525* angustifolia auct. 526 burmanniana DC. 523, 526 candolleana Blume 526 var. minor Blume 526 dioica auct. 526 elaeagnoides 523 eriocarpa Smith var. minor Sherff 523 var. waitziana Sherff 523 ferrea Jungh. 523 lamponga Mig. 527 littoralis Jungh. 526 madagascariensis 523 montana Jungh. 523 polyandra Merr. & Perry 523,524. 525* repanda Schumach. & Thonn. 526 schiedeana Schlechtendal 523 spec. nov., Turcez. 523 triquetra auct. 523 viscosa Jacq. 427, 443%, 52352575526 forma angustifolia Sherff 523 forma burmanniana West 523 forma minor Backer 526 forma repanda Radlk. 526 (Dodonaea viscosa) forma schiedeana Radlk. 523 forma viridula Kuntze 526 subforma excisa Radlk. 523 subforma waitziana Radlk. 523 var. angustifolia Benth. 523 var. candolleana Backer 526 var. vulgaris Benth. 526 var. waitziana Kuntze 523 viscosa auct. 524 waitziana Blume 523 zollingeri Turcz. 523 Donatophorus Zipp. eX Macklot 598, 603 erythrospermus Zipp. ex Macklot 607 Dysoxylum euneuron Miq. 537); Dysoxylum sp. 540 Eccremanthus Thwaites 698 Elattostachys (Blume) Radlk. 439, 444, 447, 527 aiyurensis Adema 528, 529*, 530* angulosa Adema 528, 529*, 530* duplicato-serrata Radlk. 535 erythrocarpum Adema 528, 529*, 530*,,53h globosa Adema 528, 529%, 53075551 goropuensis Adema 528, 529*, 530*, 531 obliquinervis Radlk. 528, 529*, 530*, 532 rubrofructus Adema 528, 529% 35307, 532 tetraporandra Radlk. 528, 529*, 530*, 533 verrucosa (Blume) Radlk. 528, 533, 534* zippeliana (Blume) Radlk. 528, 530*, 535 Electra Noronia 713 Erioglossum Blume 627, 648 edule (Blume) Blume 649 var. album Blume 649 var. corymbosum Teijsm. & Binn. 649 Index to scientific plant names 761 i (Erioglossum edule) Ganophyllum Blume 431,538 = (Guioa) var. fraxinifolia (DC.) Blume 649 var. genuina Blume ex Koord. & Valeton 649 var. subcorymbosum Blume 649 edule auct. 503 membranifolium Radlk. 648 rubiginosum (Roxb.) Blume 649 Euatalaya Radlk. 480 Euphoria Comm. ex Juss. 653 cinerea (Turcz.) Radlk. 517, 519 cubilii Blanco 491 didyma Blanco 654, 657 foveolatus Radlk. 513 glabra Blume 681 gracilis Radlk. 517, 519 lit-chi Desfont. 654, 655, 658 var. undulata Blume 654, 657 lit-chi auct. 517 longan (Lour.) Steud. 517 longana Lam. 517, 518 longana auct. 690 malaiensis (Griff.) Radlk. 3175519 forma genuina Radlk. 517 microcarpa Radlk. 517, 519 nephelium DC. 680 nephelium Poir. 680 noronhiana Blume 750 ramb-outan Labill. 681 ramboutan-ake Labill. 689 spec. nov., Meijer 513 spec. nov., Merr. 517 verruculosa Salisb. 517 verticillata Lindl. 651 xerocarpa Blume 473, 750 Euphoria auct. 511, 512 Euphorianthus Radlk. 536 euneurus (Miq.) Leenh. 440, 447, 522, 536, 3372 longifolius Radlk. 537 obtusatus Radlk. 537 pallidus Radlk. 537 Euphoriopsis Radlk. 536 longifolia Radlk. 537 Exothea copalillo 433 Filicium Thw. 421, 430, 431 decipiens (W. & A.) Thw. 427, 435, 445, 754, 755* falcatum Blume 427, 436, 445, 511, 538, 539* giganteum Hauman 540 obliquum Merr. 427, 540 Garuga javanica Blume 615 Gemella Lour. 459 trifolia Lour. 460 Gleditschia sp., Kaneh. & Hatus. 510 Glenniea Hook. f. 422, 436, 446, 540 penangensis (Ridley) Leenh. 541, 542, 543* philippinensis (Radlk.) Leenh. 541, 542 thorelii (Pierre) Leenh. 436, 443, 541, 543*, 544 Gloeocarpus Radlk. 450, 544 patentivalvis (Radlk.) Radlk. 437, 545*, 546 crenatus Radlk. 546 philippinensis Elmer 546 Gongrospermum Radlk. 548 philippinense Radlk. 438, 445, 547*, 548 Guioa Cav. 421—423, 437, 450, 548, 551*, 554*, 595" sect. Euguioa Radlk. 548 sect. Hemigyrosa Radlk. 549 acuminata Radlk. 557, 558, 563 acutifolia Radlk. 553, 560, 562, 564 amabilis Kaneh. & Hatus. 550, 561, 565, 598 aptera Radlk. 585 aryterifolia Radlk. 556, 563, 565 aryterifolia auct. 569 asquamosa Welzen 556, 560, 566 bicolor Merr. 555, 560, 566 bijuga (Hiern) Radlk. 550, 952, 555; 557/o56; 560, 567 bullata Radlk. 571 comesperma Radlk. 550, 555, 556, 561, 562, 568 comesperma auct. 569 contracta Radlk. 444, 450, 557, 563, 569 crenifoliola Merr. & Perry 590 curvidens Radlk. ex Dur. & Jacks. 499 dasyantha Radlk. 487 dasyantha auct. 594 diplopetala (Hassk.) Radlk. 448*, 553, 556, 558, 560, 570, 572*, 573* forma dentata Radlk. 591 forma microcarpa Radlk. 571 forma borneensis Radlk. 570 forma genuina Radlk. 570 var. borneensis Radlk. 570 diplopetala auct. 576 discolor Radlk. 553, 559, 574 elegans Radlk. 598 eriantha Merr. & Perry 480 falcata Radlk. 580 ferruginea Merr. ex Salvosa 580 forbesii Bak. f. 585 fuscidula (Kurz) Radlk. 570 var. glabrescens King 571 geminata Laut. & K. Schum. 473 glauca auct. 583 grandifoliola Welzen 556, 563, 574 hirsuta Welzen 552, 560, 575 hospita Radlk. 552, 562, 575 koelreuteria (Blanco) Merr. 552, 555-560, 576, 750 lasiothyrsa Radlk. 585 forma elmeri Radlk. 585 lentiscifolia Cav. 555* leptoneura Radlk. 571 macropetala Radlk. ex Dur. & Jacks. 501 malukuensis Welzen 552, 560, 577 melanopoda Merr. & Perry 550, 561, 578 membranifolia Radlk. 557, 560, 563, 578 membranifolia auct. 596 microphylla Radlk. 571 mindorensis Merr. 576 minjalilen Radik. 570 misimaensis Welzen 555, 562, 579 762 Flora Malesiana ser. I, Vol. 11 (3) (1994) (Guioa) molliuscula Radlk. 556, 561, 579 molliuscula auct. 594 multijuga Welzen 552, 561, 562, 580 multipunctata Radlk. 583 myriadenia Radlk. 555, 559, 580 normanbiensis Welzen 557, 563, 581 novobritannica Welzen 553, 562, 582 obtusa Merr. 580 oligotricha Merr. & Perry 552, 556, 561, 582 palawanica Welzen 553, 559, 583 parvifoliola Merr. 553, 555, 559, 583 patentinervis Radlk. 556, 560, 583 pauciflora Radlk. 550, 561, (Guioa) sp., Verheijen 566 sp., Vidal 576, 740 Spl esp:-2:0Mspy3. Lap, 571 squamosa Radlk. 570 forma genuina Radlk. 570 forma lineolata-punctata Radlk. 570 subapiculata Radlk. 585 subsericea Radlk. 552, 561, 562, 594, 595* subsericea auct. 568 sulphurea Radlk. 592 truncata Radlk. 552, 558, 596 unguiculata Welzen 556, 557, 563, 596 venusta Radlk. 550, 562, 597 venusta auct. 588 waigeoensis Welzen 550, 561, 597 (Harpullia) crustacea Radlk. 600, 606 crustacea auct. 605 cupanioides Roxb. 600, 604*, 606 var. latifolia Miq. 607 cupanioides auct. 601, 606, 610, 611 fraxinifolia Blume 607 fruticosa Blume 607 giganteacapsula Vente 600, 604*, 608 glanduligera Radlk. 601 hirsuta Radlk. 600, 608 hirsuta auct. 610 imbricata Thwaites 601 juglandifolia Blume 607 var. multiflora Blume 607 leptococca Radlk. 600, 604*, 608 longipetala Leenh. 600, 609 longithyrsifera Kaneh. & Hatus. 607 macrocalyx Radlk. 607 myrmecophila Merr. & Perry 600, 609 obscura Radlk. 607 584 Halicacabus peregrinus perrottetii Radlk. 576 Rumph. 484 perrottetii auct. 563, 591 Handeliodendron 425 platycarpa Radlk. 503 Harpullia Roxb. 422, 424, 430, pleuropteris (Blume) Radlk. 550=593;,9997,557— 559, 584, 586*,587* var. bijuga King 567 pleuropteris auct. 590 plurinervis Radlk. 553, 562, 588 pseudoamabilis Welzen 550, 562, 588, 589* pteropoda Radlk. 444, 550, 560, 561, 590 pterorhachis Welzen 550, 557, 590 pubescens (Zoll. & Mor.) Radlk. 553, 558, 559, 591 regularis Radlk. 570 reticulata Radlk. 553, 559, 592 rigidiuscula Radlk. 557, 563, 593 rigidiuscula auct. 568, 576 rubrofusca Radlk. ex Merr. 567 salicifolia Radlk. 576 scalariformis Welzen 556, 563, 593 sp., Hartley 593 sp., Merr. 583 sp., Streimann 568 436, 446, 598, 603 subg. Euharpullia Radlk. 603 subg. Harpullia 603 subg. Otonychium (Blume) Radlk. 601 sect. Euotonychium Radlk. 601 sect. Otonychidium Radlk. 601 sect. Thanatophorus Radlk. 603 aeruginosa Radlk. 611 angustifolia Radlk. 611 arborea (Blanco) Radlk. 599, 601, 602*, 604* var. megalocarpa Merr. 601 austrocaledonica Baillon 599, 603 blancoi Fern.-Vill. 601 camptoneura Radlk. 600, 605 camptoneura auct. 608, 611 carrii Leenh. 600, 605 cauliflora K.Schum. & Laut. 600, 604*, 605 cauliflora auct. 611 confusa Blume 607 odorata 427 oococca Radlk. 600, 610 pedicellaris Radlk. 601 peekeliana Melch. 600, 610 peekeliana auct. 605 pendula Planch. ex F. Muell. 431, 601 petiolaris Radlk. 600, 610 subsp. moluccana Leenh. 611 var. decidens Vente 611 var. moluccana 611 subsp. petiolaris 611 ramiflora Radlk. 600, 604*, 611 reticulata Radlk. 611 reticulata auct. 607 rhachiptera Radlk. 599, 612*, 613 rupestris Blume 606 solomonensis Vente 600, 613 sp., Ceron 601, 607 sphaeroloboa Radlk. 601 thanatophora Blume 607 tomentosa Ridley 601 vaga Merr. & Perry 600, 614 weinlandii K. Schum. 611 Index to scientific plant names 763 eee, Hebecoccus Radlk. 627, 633 falcatus Radlk. 633 ferrugineus Radlk. 634 inaequalis Radlk. 633 Hedyachras Radlk. 540 philippinensis Radlk. 542 Hemigyrosa Blume 549 longifolia Hiern 631 perrottetii Blume 576 Heterodendrum Desf. 451 oleaefolium 432 Hippocastanaceae 425, 431, 433, 434 Hornea Baker 713 Houssayanthus 430 Hydnocarpus tamiana Pulle 652 [rina |Norona] Blume 698 alnifolia Blume 701 glabra Blume 701 integerrima Blume 704 tomentosa Blume 701 var. alnifolia (Blume) Mig. 701 var. cuspidata Blume 701 Jagera Blume 436, 438, 441, 614 dasyantha (Radlk.) Reynolds 487 discolor L.S. Smith 487 glabra Hassk. 533 javanica (Blume) Blume ex Kalkman 615, 616* subsp. javanica 442*, 616* latifolia Radlk. 455 macrophylla Radlk. 615 pseudorhus (Rich.) Radlk. 615, 616*, 617 var. integerrima Reynolds 617 var. pseudorhus 617 forma pilosiuscula Radlk. 617 forma pseudorhus 617 roxburghii Blume 615 serrata (Roxb.) Radlk. 615 forma fulvinervis Radlk. 615 forma genuina Radlk. 615 speciosa Blume 615 Koelreuteria Laxm. 421, 430 arborea Blanco 576 edulis Blanco 643 paniculata Laxm. 755 Koordersiodendron pinnatum Merr. 540 Lachnopetalum Turcz. 620 glabrum Turcz. 624 Lepiderema Radlk. 435, 450, 618 melanorrhachis Merr. & Perry 618, 619* papuana Radlk. 618 Lepidopetalum Blume 439, 444, 447, 620 fructoglabrum Welzen 449, 621, 622 hebecladum Radlk. 623, 626 hebecladum auct. 623 Jackianum Radlk. 623 micans Laut. & K. Schum. 621, 622*, 623 montanum (Blume) Radlk. 621, 622*, 623 perrottetii (Cambess.) Blume 621, 622*, 624 subdichotomum Radlk. 621, 622*, 624, 625* subdichotomum auct. 622 xylocarpum Radlk. 621, 622*, 626 Lepisanthes Blume 422, 430, 437, 445, 627 subg. Aphania (Blume) Leenh. 437, 627, 629, 650 subg. Erioglossum (Blume) Leenh. 627, 629, 648 subg. Lepisanthes 627, 628, 630 sect. Hebecoccus (Radlk.) Leenh. 627, 628, 633 sect. Lepisanthes 627, 628, 628 subg. Otophora (Blume) Leenh. 422, 435, 436, 443, 627, 629, 635 sect. Anomotophora (Radlk.) Leenh. 628, 629, 646 sect. Otophora (Blume) Leenh. 627, 629, 635 sect. Pseudotophora (Radlk.) Leenh. 627, 629, 642 sect. Eulepisanthes Radlk. 627, 630 acutissima Radlk. 632 alata (Blume) Leenh. 629, 631*, 646 (Lepisanthes) amoena (Hassk.) Leenh. 630, 636, 637* amplifolia (Pierre) Leenh. 648 andamanica King 633 angustifolia Blume 631 aphanococca Leenh. 634 appendiculata (Hook. f.) Sym. 632 bengalan Leenh. 630, 643, 644* blumeana Koord. & Valeton 632 borneensis Leenh. 634 cuneata Hiern 631 dictyophylla (Radlk.) Leenh. 629, 650 divaricata (Radlk.) Leenh. 630, 638 forma divaricata 639, 640* forma lunduensis (RadIk.) Leenh. 639 eriolepis Radlk. 631 falcata (Radlk.) Leenh. 629, 633 subsp. borneensis (Leenh.) Leenh. 631*, 634 subsp. celebica (Radlk.) Leenh. 634 subsp. falcata 634 ferruginea (Radlk.) Leenh. 629, 631*, 634 fruticosa (Roxb.) Leenh. 435, 630, 631*, 643 hirta Ridley 649 kinabaluensis Leenh. 630, 639, 641* Kunstleri King 632 longifolia (Hiern) Radlk. 631 macrocarpa Radlk. 632 membranifolia (Radlk.) Radlk. 629, 648 mixta Leenh. 629, 651 montana Blume 630 multijuga (Hook. f.) Leenh. 448*, 630, 631*, 642 palawanica Radlk. 544 ramiflora (Radlk.) Leenh. 629, 647 rubiginosa (Roxb.) Leenh. 629, 631*, 648 schizolepis Radlk. 632 forma genuina Radlk. 632 764 (Lepisanthes) scortechinii King 632 senegalensis (Poir.) Leenh. 436, 629, 630, 631*, 651 sessiliflora Blume 631 tetraphylla (Vahl) Radlk. 437, 445, 629, 630, 631* viridis Radlk. 632 Linkeng Rumph. 517 Litchi Sonn. 422, 430, 653 chinensis Sonn. 428, 433— 436, 438, 442*, 445, 654, 655, 656*, 657 forma glomeriflora Radlk. 654, 657 subsp. chinensis 655, 656* subsp. javensis Leenh. 655, 657 subsp. philippinensis (Radlk.) Leenh. 655, 656*, 657 philippinensis Radlk. 654, 657 forma genuina Radlk. 654 forma mindanaensis Radlk. 654 ramboutan Labill. 681, 682 ramboutan-ake Labill. 689 sinensis 756 sp. nov., Merr. 657 Litsea ramboutan 681 Llagunoa 430, 431 Magonia 423, 430 Majidea 430 Melicocca L. 430 amoena Hassk. 636 javanica Hassk. 533 trijuga Juss. 728 triptera Blanco 740 Meliosma sumatrana (Jack) Walp. 704 Mildea Migq. 693 gibbosa Miq. 697 xystophyllum Mig. 696 Mischocarpus Blume 421, 423, 438, 440, 447, 449, 658, 724 australis S.T. Reynolds 668 brachyphyllus Radlk. 663 cauliflorus Radlk. 668 ellipticus Radlk. 663 endotrichus Radlk. 663 fuscescens Blume 663 fuscescens auct. 668 lachnocarpus (F. Muell.) Radlk. 659, 660, 661* Flora Malesiana ser. I, Vol. 11 (3) (1994) eee. ——E (Mischocarpus) largifolius Radlk. 448*, 659, 661* lessertianus Ridley 667 macrobotrys Merr. & Perry 474 macrocarpusReynolds 668 papuanus Radlk. 664, 665 paradoxus Radlk. 438, 448%, 659, 660, 661*, 662 pentapetalus (Roxb.) Radlk. 439, 660, 661*, 662, 688 var. cambodianus 664 prob. spec. nov. 660, 668 pyriformis (F. Muell.) Radlk. 659, 660, 664 subsp. papuanus (Radlk. ) R.W. Ham. 448*, 661%, 665 subsp. pyriformis 665 subsp. retusus (Radlk.) R.W. Ham 665 pyriformis auct. 667 reticulatus (Radlk.)R.W. Ham 659, 660, 665, 666* retusus Radlk. 664, 665 salicifolius Radlk. 663 stipitatis Reynolds 668 sublaevis Radlk. 663 sumatranus Blume 663 sundaicus Blume 660, 661*, 667 triqueter Radlk. 439, 448*, 660, 661*, 668 vulcanicus Elmer ex Merr. 667 Mischocodon Radlk. 658 reticulatus Radlk. 665 More Gaertn. 511 Moulinsia rubiginosa (Roxb.) G. Don 649 Nephelium L. 430, 438, 439, 445, 669 aculeatum Leenh. 670, 672 acuminatum Hook. f. 702 altissimum Teijsm. & Binn. 679 bassacense Pierre 674, 676 beccarianum Radlk. 674, 677 caudifolium Ridley 684 chryseum Blume 681, 683 chryseum auct. 674, 690 compressum Radlk. 439, 670, 672, 673* (Nephelium) costatum Hiern 671, 673 cuspidatum Blume 672, 673*, 674, 675 var. bassacense (Pierre) Leenh. 675, 676 var. cuspidatum 676 subvar. cuspidatum 675, 676 subvar. dasyneurum (Radlk.) Leenh. 675, 676 var. eriopetalum (Miq.) Leenh. 675, 676 var. multinerve (Radlk.) Leenh. 675, 676 var. ophiodes (Radlk.) Leenh. 677 subvar. beccarianum (Radlk.) Leenh. 675, 677 subvar. ophiodes 675, 677 var. robustum (Radlk.) Leenh. 675, 677 daedaleum Radlk. 670, 673*, 677 dasyneurum Radlk. 674, 676 didyma (Blanco) Craib 654, 657 eriopetalum Migq. 674, 676 ferrugineum F. Muell. 455 forbesii Baker 697 fumatum Blume 515 glabrum Cambess. 681 glabrum Norofia 680, 682 var. album Hassk. 690 var. nigrum Hassk. 690 var. rubrum Hassk. 690 var. sufferrugineum (Radlk.) Ridley 681 glabrum Reinw. ex Blume 681, 682 glabrum auct. 685, 690 hamulatum Radlk. 671, 678, 688 havilandii Leenh. 671, 678 herveyi Ridley 678, 684 hosei Ridley 663 hypoleucum Kurz 670, 673%, 678, 680*, 688 intermedium Radlk. 690 intermedium auct. 515, 516 juglandifolium Blume 442*, 670, 673*, 679 Index to scientific plant names 765 ee (Nephelium) (Nephelium) (Otolepis) lappaceum L. 434, 671, 672, 680, 682, 756 var. glabrum Blume 681 var. glabrum auct. 685 var. lappaceum 682 var. maingayi (Hiern) Valeton 685 var. pallens (Hiern) Leenh. 683 var. xanthioides (Radlk.) Leenh. 683 lappaceum auct. 683, 685 laurinum Blume 671, 673*. 684, 688 lit-chi (Desfont.) Cambess. 654, 655 longan (Lour.) Hook. 517, 518 longana (Lam.) Cambess. SLTVOLS var. hypoleuca (Kurz) King 674, 678 longana auct. 519 long-yan Blume 517, 518 macrophyllum Radlk. 671, 680*, 684 maculatum Radlk. 681, 682 maingayi Hiern 449, 670, 685, 686* maingayi auct. 750 malaiensis Griff. 517, 519 meduseum Leenh. 442*, 671, 673*, 686 melanomiscum Radlk. 670, 687 melliferum Gagnep. 671, 674, 688 multijuga Hook. f. 642 multinerve Radlk. 674, 676 muricatum Griff. 750 mutabile Blume 689 var. pallens Hiern 681, 683 var. rigida Blume 690 var. trigyna Blume 690 mutabile auct. 474 noronhianum Cambess. 750 nov. spec., Leenh. 693 obliquinervis Radlk. 674, 676 obovatum Ridley 681, 682 ophiodes Radlk. 674, 677 pallens (Hiern) Radlk. 681 papillatum Leenh. 672, 688, 689* parviflorum Gagnep. 688 philippinense Monsalud et al. 690 ramboutan-ake (Labill.) Leenh. 670-672, 673*, 689 rambutan Schnizl. 681 reticulatum Radlk. 671, 672, 691 robustum Radlk. 674, 677 rubescens Hiern 684 semiglaucum var. acutifolium F. Muell. ex Bailey & White 564 sp., Ceron 690 sp., Merr. 674 subfalcatum Radlk. 672, 680*, 692 sufferrugineum Radlk. 681, 682 tuberculatum Radlk. 679 uncinatum Radlk. ex Leenh. 442*, 671, 673*, 680*, 692 verticillatum (Lindl.)G. Don 651 winterianum F.M. Bailey 568 xanthoides Radlk. 681, 683 xerocarpum Cambess. 473, 750 xerospermoides Radlk. 687 Ornitrophe Comm. ex Juss. 459 glabra Roxb. 460 integrifolia Willd. 460 repanda Roxb. 460 schmiedelia Pers. 460 villosa Roxb. 460 Ostodes appendiculata Hook. f. 632 Otolepis Turcz. 627 sect. Anomotophora (Radlk.) Kuntze 628 sect. Otophora (Blume) Kuntze 627 sect. Pseudotophora (Radlk.) Kuntze 627 alata (Blume) Kuntze 646 amoena (Hassk.) Kuntze 636 cordigera Kuntze 636 erythrocalyx (Hiern) Kuntze 643 fruticosa (Roxb.) Kuntze 643 imbricata (Blume) Kuntze 636 nigrescens Turcz. 627 pubescens (Blume) Kuntze 636 sessilis King 643 spectabilis (Blume) Kuntze 636 Otonephelium 512 Otonychium Blume 598, 601 imbricatum Blume 601 retusum Miq. 614 Otophora Blume 627, 635 subg. Pseudophora Blume 627, 642 sect. Anomotophora Radlk. 628, 646 sect. Euotophora Radlk. 627, 635 sect. Pseudophora Blume 627, 642 sect. Pseudotophora Radlk. 627, 642 acuminata Radlk. 643 alata Blume 646 amoena (Hassk.) Blume 636 blancoi Blume 643 confinis Blume 636 cordigera Radlk. 636 divaricata Radlk. 638, 639 edulis Fischer 646 erythrocalyx Hiern 643 fruticosa (Roxb.) Blume 643 glandulosa Radlk. ex Ridley 643 imbricata Blume 636 imbricata auct. 642 javanica (Hassk.) Miq. 533 latifolia Ridley 643 lunduensis Radlk. 638, 639 macrocarpa Ridley 638, 639 multijuga (Hook. f.) Merr. 642 paucijuga Hiern 651 pubescens Blume 636 pyramidalis Radlk. 638 ramiflora Radlk. 647 resecta Radlk. 643 spectabilis Blume 636 var. pubicosta Blume 636 styligera Radlk. 636 tricocca Radlk. 642 766 Palaoea Kaneh. 742 falcata Kaneh. 745 Pancovia 430 Papaja litorea boeronensis Attehu dicta Rumph. 615 Paranephelium Miq. 435, 439, 443-445, 693 acanthocarpum Radlk. ex Koord. 697 chinense Merr. 467 gibbosum Teijsm. & Binn. 697 joannis Davids 695* macrophyllum King 695, 696 nitidum King 697 spirei Lecomte 695*, 696 xestophyllum Mig. 695*, 696, 697* Paullinia 430 cupana 433 seriana auct. 460 sorbilis 433 triantennata 433 yoco 433 Pedicellia Lour. 658 fuscescens Pierre 663 oppositifolia Lour. 658 pentapetala Pierre 662 sumatrana Pierre 663 sundaica Pierre 667 Physalis L. 484 alkekengi L. 484 Physalis Norona 484 halicacabum Crantz 484 Pistacia oleosa Lour. 728 Placodiscus Radlk. 541 Pometia Forst. & Forst. 422, 425, 436, 437, 441, 446%, 698 acuminata (Hook. f.) Radlk. 702 alnifolia (Blume) King 701 coriacea Radlk. 702 curtisii King 517, 519 forbesii Baker 698 glabra (Blume) Teijsm. & Binn. 701 macrocarpa Kurz 702 pinnata Forst. & Forst. 426, 427, 448*, 700*, 701, 103% forma acuminata (Hk. f.) Jacobs 702 forma alnifolia (Blume) Jacobs 701 Flora Malesiana ser. I, Vol. 11 (3) (1994) a 55058585 ——050— (Pometia pinnata) forma cuspidata (Blume) Jacobs 702 forma glabra (Blume) Jacobs 701 forma macrocarpa (Kurz) Jacobs 702 forma repanda Jacobs 702 forma tomentosa (Blume) Jacobs 701 var. javanica Koord. & Valeton 701 ridley King ex Radlk. 701, 703* tomentosa (Blume) Teijsm. & Binn. 701 var. cuspidata (Blume) J. Britten 701 Pseudonephelium Radlk. 511, a2 fumatum (Blume) Radlk. 5155516 javanicum Radlk. 515 Ptelea arborea Blanco 601 viscosa L. 526 Pteleocarpa lamponga (Miq.) Bakh. ex Heyne 527 Pteridophyllum decipiens 754 Quassia simaruba auct. 576 tricarpa Blanco 715 Ratonia lachnocarpa F. Muell. 660 lachnopetala Turcz. 624 litoralis Teijsm. & Binn. 473 montana Fern.-Vill. 623 pyriformis Benth. 664 rufescens Fern.-Vill. 473 sumatrana Kurz 663 zygolepis Turcz. 473 Rhus cobbe L. 459 taitensis Guillemin 614 Rhysotoechia Radlk. 437, 449, 704 acuminata Radlk. 710 applanata Etman 449, 705, 706 bilocularis Etman 704, 706 congesta Etman 704. 706, Oe contermina Domin 711 elongata Radlk. 705, 708 gracilipes Radlk. 705, 708 grandiflora Radlk. ex Merr. IA (Rhysotoechia) grandifolia Radlk. 712 koordersii Radlk. 705, 708, 709 longipaniculata Kaneh. & Hatus. 712 momiensis Kaneh. & Hatus. Te mortoniana auct. 708 multiscapa Etman 705, 709, 710* obtusa Etman 705, 709 ramiflora Radlk. 705, 709, 710 robertsonti (F. Muell.) Radlk. 705, 711 sp. 712 striata Radlk. 711 Sapindacea Wall. 735 Sapindaceae 419-768 subfam. Dodonaeoideae 423, 424, 430, 434 tribe Cossinieae 430, 434 tribe Dodonaeeae 426, 430, 434 tribe Doratoxyleae 424, 426, 430, 434 tribe Harpullieae 425, 426, 430, 434 tribe Koelreuterieae 430, 434 subfam. Dyssapindaceae 434 subfam. Eusapindaceae 434 subfam. Sapindoideae 423, 424, 434 tribe Aphanieae 434 tribe Cupanieae 421, 423 426, 428, 430, 434 tribe Lepisantheae 424, 430, 434 tribe Melicocceae 424, 426, 430, 434 tribe Nephelieae 421, 424, 426, 430, 435 tribe Paullinieae 423, 424, 429, 430, 435 tribe Sapindeae 426, 430, 434 tribe Schleichereae 424, 430, 435 tribe Thouinieae 424, 429, 430, 435 Sapindaceae sp. A, Koord.- Schum. 697 Index to scientific plant names 767 ————— SSE eee Sapindus L. 421, 430, 437, 445, 18 sect. Dasysapindus Radlk. 713 sect. Dittelasma (Hook. f.) Radlk. 713 sect. Sapindus 713 angustifolius Blume 714 baccata Blanco 643 balicus Radlk. 715 cinereus Turcz. 517 cultratus Turcz. 737 cuspidata Blume 651 edulis Aiton 654 edulis Blume 649 forsythii DC. 715 fraxinifolia DC. 649 fruticosa Roxb. 643 glabratus Wall. 750 guisian Blanco 588 koelreuteria Blanco 576 laurifolia auct. 635 longifolius auct. 537 microcarpua Jardin 715 montana (Blume) Blume 651 mukorossi Gaertn. 715 oocarpus Radlk. 716 papuana Radlk. 480 pinnatus Mill. 714 pubescens auct. 576 pubescens Zoll. & Mor. 591 rarak DC. 437, 714, 715* rubiginosa Roxb. 649 salicifolius DC. 482 saponaria L. 433, 714, 715* forma microcarpa (Jardin) Radlk. 715 var. jardiniana F.B.H. Brown 715 senegalensis Juss. ex Poir. 651 serrata Roxb. 615 sp., Ceron 542 squamosa Wall. 570 squamosus Roxb. 722 tetraphylla Vahl 630 trifoliatus L. 714, 715* verticillatus (Lindl.) Kurz 651 vitiensis A. Gray 715 Saponaria rarak Rumph. 714 Sarcopteryx Radlk. 421—423, 439, 444, 450, 717 brachyphylla Radlk. 443*, 118*,. 719* (Sarcopteryx) caudata Welzen 450, 717, T18*, 719, 720* coriacea Radlk. 718, 720 crispata Welzen 718, 721 holconeura Radlk. 722 melanophloea Radlk. 722 rigida Radlk. 718*, 721 rubiginosa Welzen 718, 722 sordida (Radlk.) W. R. Ham 721 squamosa (Roxb.) Radlk. 718*, 722 Sarcotoechia Radlk. 438, 439, 447, 723 subg. Pilosodiscus Leenh. 724 subg. Sarcotoechia 724 angulata Leenh. 724, 726* apetala Leenh. 724, 725* bilocularis Leenh. 724, 726* cuneata Radlk. 727 lanceolata (C. T. White) S.T. Reynolds 727 planitiei Leenh. 436, 724, 726* sp., Leenh. 727 Schleichera Willd. 430, 727 aculeata Kostel. 728 amoena (Hassk.) Walp. 636 oleosa (Lour.) Oken 427, 431, 436, 444, 449, 728, 729* pentapetala Roxb. 662 pentaphylla Roxb. 662 revoluta Turcz. 667 subundulata Turcz. 663 trijuga Willd. 728 Schmidelia L. 459 bantamensis Blume 460 cobbe (L.) DC. 459 fulvinervis Blume 460 var. macrophylla Teijsm. & Binn. 461 glabra (Roxb.) Steud. 460 grossedentata Turcz. 461 Javensis Blume 460 leptostachya Blume 461 ligustrina Blume 461 littoralis Blume 460 macrophylla Zipp. ex Span. 460 mutabilis Blume 461 var. subglabra Miq. 461 obovata A. Gray 461 parviflora Zipp. ex Span. 460 (Schmidelia) pentapetala Wight 662 pyriformis F. Muell. 664 racemosa L. 460 ternata (Forst. & Forst.) Cambess. 460 timoriensis DC. 460 tomentosa Hook. f. 461 Scyphopetalum Hiern 693 Scytalia Gaertn. 653 danura Roxb. 651 verticillata (Lindl.) Roxb. 651 Seringia lanceolata Blanco 601 Serjania 424, 430 piscatoria 433 serrata 433 Smelophyllum Radlk. 451 Spanoghea Blume 450 connata F. Muell. 454 ferruginea Blume 455 glabra Blume 456 Stadmania Lam. 451 australis G. Don 520 sideroxylon (non DC.) Hassk. 728 trijuga Spreng. 728 Streptostigma Thwaites 598, 601 Synima Radlk. 444, 730 cordierorum (F. Muell.) Radlk. 440, 730, 731* macrophylla S.T. Reynolds 439; 730: 7317732 Thanatophorus Zipp. ex Walp. 598 Thouinia Sm. 459 Thouinidium Sm. 480 Tina rupestris Blume 606 Toechima Radlk. 422, 446, 732 daemelianum Radlk. 734 erythrocarpum (F. Muell.) Radlk. 732 subsp. erythrocarpum 734 subsp. papuanum Leenh. 439, 447, 598, 732, 1337 hirsutum Radlk. 733 livescens Radlk. 733 plurinerve Radlk. 734 sp., Hartley et al. 733 subteretes Radlk. 733 Trigonachras Radlk. 436, 440, 445, 734 acuta (Hiern) Radlk. 735, 736* 768 Flora Malesiana ser. I, Vol. 11 (3) (1994) a TT (Trigonachras) (Tristiropsis) (Xerospermum) brachycarpa Radlk. 737 celebensis Leenh. 439, 735, 137, 1380 cultrata (Turcz.) Radlk. 735, TT. 138% cuspidata Radlk. 735, 738* falcatocuspidata Radlk. 738 membranacea Radlk. 737 nov. sp. A 735, 739 nov. sp. B 735, 739 nov. sp. C 735, 739 obliqua Radlk. 737 papuensis Leenh. 735, 738%, 739 rigida Radlk. 737, 738 Tristira Radlk. 438, 444, 740 celebica Boerl. & Koord. 740 harpullioidea Radlk. 740 penangensis Ridley 542 pubescens Merr. 740 forma genuina Radlk. 740 forma hemidasya Radlk. 740 triptera (Blanco) Radlk. 438, 443*, 444, 740, 741* Tristiropsis Radlk. 435, 440, 441, 742 acutangula Radlk. 427, 743, 744* apetala Leenh. 743, 744%, 745 canarioidea Boerl. ex Valet. 743 dentata Radlk. 745 falcata Kaneh. 745 ferruginea Leenh. 427, 743, 744*, 746 nativitatis Hemsl. ex Ridley 743 novoguineensis Kaneh. & Hatus. 745 oblonga Radlk. 745 obtusangula Radlk. 743 ovata Radlk. 743 ridleyi Hemsl. 743 subangula K. Schum. 743 subfalcata Radlk. 745 Uitenia edulis (Blume) Steud. 649 Ungnadia 430 Urvillea Kunth 424, 430, 484 uniloba var. uniloba 429 Usubis Burm. f. 459 triphylla Burm. f. 460 Walsura villosa auct. 607 Xanthoceras 430 Xerospermum Blume 430, 436, 438, 445, 746 sect. Pentasepalum Radlk. 746 sect. Tetrasepalum Radlk. 746 acuminatum Radlk. 749 brachyphyllum Radlk. 750 cylindrocarpum Radlk. 750 echinulatum Radlk. 750 fallax Radlk. 750 glabratum Radlk. 750 intermedium Radlk. 750 laevigatum Radlk. 748* subsp. acuminatum Leenh. 749, 751* subsp. laevigatum 749 lanceolatum Radlk. 750 muricatum Radlk. 750 muricatum auct. 749 noronhianum (Blume) Blume 442*, 748, 750, 751* sp., Corner 750 sp., Merr. 685 testudineum Radlk. 750 thorelii Pierre 544 unijugum Radlk. 749 wallichii King 750 xanthophyllum Radlk. 750 Zollingeria Kurz 439, 444, 752 borneensis Adema 439, 444, 753*, 754 macrocarpa auct. 740 triptera Rolfe 740 macrocarpa Kurz 752 Zygolepis Turcz. 468 rufescens Turcz. 473 WTA AT 3 5185 00203 | (continued from inside front cover) 1431 Punicaceae 4; 226 Sparganiaceae 4: 233;10: 718 Trimeniaceae 10: 327 Restionaceae 5: 416 Sphenocleaceae 4: 27 Triuridaceae 10: 109 Rhizophoraceae Sphenostemonaceae 10: 145 Turneraceae 4: 235 5: 429; 6: 965; 8: 550 Stackhousiaceae 4: 35 Typhaceae 4: 242: 6: 982 Rosaceae 11: 227 Staphyleaceae 6: 49 Ulmaceae 8: 31 Sabiaceae 10: 351 Stemonaceae 11: 399 Umbelliferae 4: 113, 595 Salicaceae 5: 107 Stylidiaceae 4:529; 6: 976 5: 555; 6: 983 Salvadoraceae 4: 224 Styracaceae 4: 49; 9: 568 7: 830; 9: 569 Sapindaceae 11: 419 Symplocaceae Valerianaceae 4: 253 Sarcosperma(ta)ceae 4: 32 8: 205; 9: 569; 10: 718 Violaceae Saururaceae 4: 47 Taccaceae 7: 806 7: 179, 831; 10: 720 Scyphostegiaceae Taxaceae 10: 347 Xyridaceae 5: 297; 6: 967 Thymelaeaceae 4: 366, 598; 9: 571 Simaroubaceae _ 6: 193, 968 4: 349; 6: 1,976; 7: 830 Zygophyllaceae 4: 64 Sonneratiaceae Trapaceae 4; 43 4: 280, 513; 6: 973 Trigoniaceae 4: 59 Index to revised families in Series II (Pteridophyta) Cyatheaceae 1; 65 Lindsaea group 1: 177 Tectaria group Fy Gleicheniaceae 1: 1 Lomariopsis group 1: 255 Thelypteridaceae 1: 331 Isoetaceae 1: 62 Schizaeaceae j Fey 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 J. Burley, Cambridge, USA K. Iwatsuki, Tokyo K. Larsen, Aarhus A. Latiff Mohamad, Bangi G.LI. Lucas, Kew ISBN 90-71236-21-8 FLORA MALESIANA Compiled and published under the auspices of Foundation Flora Malesiana D.A. Madulid, Manila Ph. Morat, Paris F.S.P. Ng, Rome D.H. Nicolson, Washington P.H. Raven, St. Louis M.A. Rifai, Bogor S.H. Sohmer, Fort Worth P. Srivastava, Lae 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: P.O. Box 9514 2300 RA Leiden The Netherlands Publication Department Rijksherbarium / Hortus Botanicus Fax (3.1):@1) 273511