5K37C FLORA MALESIANA LIBRARY APR 26 1972 NEW YORK BOTAN/CAL GARDEN/ SERIES I - SpERMATOPHYTA Flowering Plants Vol. 7, part 1 INDEX TO REVISED FAMILIES Aceraceae . 4: 3, 592 Elatinaceae 4: 203 Pentaphragmataceae . 4: 517 Actinidiaceae s.str . . 4: 37 Epacridaceae . 6: 422 Pentaphylacaceae 5: 121 Aizoaceae 4: 267 Ericaceae . 6: 469 Philydraceae . 4: 5 Alismataceae . 5: 317 Erythroxylaceae . 5: 543 Phytoiaccaceae 4: 229 Amaranthaceae 4: 69, 593 Ficoidaceae . 4: 267 Pittosporaceae 5: 345 Ancistrocladaceae 4: 8 Flacourtiaceae 5: 1 Plumbaginaceae 4: 107 Aponogetonaceae 4: 11, 7: 213 Fiagellariaceae 4: 245 Podostemaceae 4: 65 Basellaceae . . 5: 300 Geraniaceae . 6: 445 Polemoniaceae 4: 195 Batidaceae 5: 414 Gnetaceae . 4: 336 Pontederiaceae . 4: 255 Betulaceae 5: 207 Gonystylaceae 4: 349 Portulacaceae . 7: 121 Bixaceae s. str. 4: 239 Goodeniaceae 5: 335 Primuiacea . 6: 173 Burmanniaceae 4: 13, 592 Haemodoraceae 5: 111 Proteaceae 5: 147 Burseraceae 5: 209 Haioragaceae 7: 239 Punicaceae 4: 226 Butomaceae . 5: 118 Hamamelidaceae . 5: 363 Restionaceae . 5: 416 Byblidaceae . . 7: 135 Hippocrateaceae . 6: 389 Rhizophoraceae 5: 429 Callitrichaceae 4: 251 Hydrocaryaceae . 4: 43 Salicaceae . 5: 107 Campanulaceae 6: 107 Hydrocharitaceae . 5: 381 Salvadoraceae. 4: 225 Cannabinaceae 4: 223 Hydrophyllaceae . 4: 207 Sarcospermaceae 4: 32 Capparidaceae 6: 61 Icacinaceae 7: 1 Saururaceae 4: 47 Capri foliaceae 4: 175, 598 Juglandaceae . 6: 143 Scyphostegiaceae 5: 297 Cardiopteridaceae 7: 93 Juncaceae . 4: 210 Simaroubaceae . . 6: 193 Celastraceae . . 6: 227, 389 Juncaginaceae. 4: 57 Sonneratiaceae 4: 280, 513 Centrolepidaceae 5: 421 Lemnaceae 7: 219 Sparganiaceae . 4: 233 Ceratophyllaceae 4: 41 Loganiaceae . 6: 293 Sphenocleaceae . 4: 27 Chenopodiaceae 4: 99, 594 Lophopyxidaceae 7: 89 Stackhousiaceae 4: 35 Clethraceae . . 7: 139 Malpighiaceae 5: 125 Staphyleaceae 6: 49 Cochlospermaceae 4: 61 Martyniaceae . 4: 216 Stylidiaceae 4: 529 Combretaceae 4: 533 Moiluginaceae 4: 267 Styracaceae 4: 49 Connaraceae . 5: 495 Moringaceae . 4: 45 Thymelaeaceae 4: 349, 6: 1 Convolvulaceae 4: 338, 599 Myoporaceae . 4: 265 Trigoniaceae . 4: 59 Corynocarpaceae 4: 262 Myricaceae . 4: 277 Turneraceae . . 4: 235 Crassulaceae . 4: 197 Najadaceae 6: 157 Typhaceae 4: 243 Datiscaceae . . 4: 382 Nyctaginaceae 6: 450 Umbelliferae . 4: 113,595 Dichapetalaceae 5: 305 Nyssaceae . 4: 29 Valerianaceae . 4: 253 Dilleniaceae 4: 141 Ochnaceae 7: 97 Violaceae . 7: 179 Dioscoreaceae 4: 293 Oxalidaceae . 7: 151 Xyridaceae 4: 366, 598 Dipsacaceae . 4: 290 Papaveraceae . . 5: 114 Zygophyllaceae 4: 64 Droseraceae . . 4: 377 Pedaliaceae . 4: 216 ICACINACEAE (H. Sleumer, Leyden) Trees, or whether or not cUmbing shrubs, or lianas. Leaves spirally arranged, rarely opposite, simple, entire or lobed (in Mai. never crenate or serrate), penni- or palmatinerved, exstipulate. Inflorescences mostly axillary, sometimes terminal, rarely extra-axillary, or from old wood, in spikes or spike-like racemes, or often in cymes, both spikes and cymes not rarely collected to panicles or heads, very rarely reduced to few-flowered fascicles or to a solitary flower. Flowers bi- or unisexual, in the latter case at least functionally so, i.e. the plants dioecious, actinomorphic, (4-)5(-6)-, by reduction rarely in part 3-merous, cyclic (with sepals or calyx lobes and petals) or rarely spiral (with petals only in Pyrenacantha, or without petals in the !2 flowers of Platea and some spp. of lodes and Gomphandra). Pedicels, if any, articulated with the calyx. Sepals 4-6, free or mostly connate below to various degree to a 4-6-lobed calyx, the lobes imbricate or valvate, generally persistent. Petals 4-6, free or connate below to various degree, sometimes to a tube, the lobes valvate, very rarely subimbricate, tip inflexed, mostly caducous, sometimes persistent. Stamens as many as sepals or petals, episepalous, inserted basally or sometimes in the upper part of the tube; filaments subulate, fleshy, often flattened, or filiform, not rarely with clavate subglandular elongate hairs distally; anthers 2-celled, cells often diverging below, basifixed, latrorse or introrse, in Polyporandra dismissing the pollen from numerous operculate pores. Disk whether or not present, either annular or cup-like, free or adnate to the ovary, or a unilateral fleshy scale. Ovary free, 1 -celled (in Pseudobotrys, Gonocaryum and Citronella 2-celled with an empty tube-like unilateral cell) (in Mai.); ovules 2 (rarely 1 abortive), apical, pendent, anatropous, apotropous, unitegmic; style 1 or none; stigma punctiform, subcapitate or peltate, entire or slightly 2-5-lobed or -crenate, often depressed to one side. Drupe ellipsoid to globose, often laterally compressed and almond-like; exocarp generally thin-fleshy; endocarp thin-crustaceous to thick-woody, sometimes spongious or fibrous, often veined or ribbed lengthwise or reticulate-lacunose outside, smooth or with tubercles or blunt aculei inside, the seed pitted then. Seed 1, exarillate, generally with abundant endosperm, which rarely is ruminate; embryo straight; cotyledons whether or not foliaceous. Distribution. About 56 genera with c. 300 spp., all woody, predominantly in the tropics, rapidly decreasing in number towards the subtropics; 5 genera with part of their species in the temperate zones of Africa, Asia, Australia and S. America. In Malesia a total of c. \(.){) spp. in 21 genera, of which 3 are strictly endemic, viz Cantleya (W. Malesia), Harileyci and Pseudobotrys (both in New Guinea); 8 other genera find their main area of distribution and generally their greatest number of species in Malesia, but occur also in parts of S. and SE. Asia, viz Gonocaryum, Plaiea (both also in New Britain), Codiocarpus, Stcmonurus (also in the Solomon Is.), Miquelia, Noihapodyles, Phytocrene, and Sarcostif^nw, 3 genera, viz Apodyles, lodes and Pyrenacantha, are found in Africa (also Madagascar), SE. Asia and Malesia. The genus Citronella is amphipacilic (Malesia, E. Australia, Melanesia, New Caledonia, New Hebrides, Fiji, Samoa, Tonga, Costa Rica, Venezuela, Brazil, Paraguay, N. Argentina, Bolivia, and Central Chile); 6 genera, viz Gomphandra, Medusanlhera, Merrilliodcndron, Polyporandra, Rhylicaryum, and Stcmonurus, extend from Malesia into Micronesia and Melanesia or even W. Polynesia; of these only Gomphandra and Rhyticaryum are also found in NE. Australia. Whitmorea, so far known, is limited to the Solomon Is., but might occur also in SE. New Guinea. Icacinaceac of Malesia show a strong afTinity with those of tropical South Asia (SW. India, Ceylon) and .Southeast Asia (Sikkim to Assam, Bengal, Burma, Thailand, Indo-China, W., Central and S. China incl. Hainan and Formosa, S. Japan). Most of the members of the family in Australia point to an Asiatic-Malcsian origin; lrvinf(hailcya, limited to Queensland, is taxonomically very close to Codio- (1) Flora Malesiana [ser. 1, vol. 7^ carDus and Medusanthera. Pennantia in New Zealand and on Norfolk I. is more distinct from the Asiatic- Mrsianb^co genera. New Caledonia has, besides CitroneUa, 2 endemic genera nz Amsomallon Sn ed to Apodvtes) and Gastrokpis (related to the group of genera around Medusanthera) Eco ogy Most Malesian sp/ occur in the substage of the primary or secondary ram-forest a few are canopy trees (S/^nc>.»n/., Platea, Cantleya, Citronella p.p.). They are mainly found m the lowlands, hut mav ascend to the montane and even the mossy forest {Platea p.p., Rhyticaryump.p., Gomphandra Tv , UP to 2400 m (highTrai itudes mentioned on collector's labels being rather doubtful); several genera re^chthdr highest altitude on Mt Kinabalu or in New Guinea. The distribution is scattered in general for the t?ee specie , even more so for the climbers, and can cover enormous areas as for instance Platea iVfoliaTS^SSrLg^ng from Sikkim, Himalaya, to New Guinea. Several species iStemonurus, Gom- Zndra- ^/a^ea even with stilt roots) are clearly limited to peat swamp forest. Some Icaanaceae are found also in coastal forest, and such species may possibly derive their wide range of distribution from the 'Tl^lSetUs a': r^e'ched only by a few species of Platea, Cantleya, Stemonurus, Citronella, Con. nhandra Apodvtes, or raTe\y by such of Nothapodytes and Medusanthera. Among the rain-forest species none is reported to be dominant in any vegetation type, though oc- casionallv single species may be locally frequent. ,, ^ , ^ ■ u i, ,- Few species occur in the drier, mainly seasonally dry parts of Malesia, all of them being shrubs or '' Di'spersal. Little is known of the dispersal of fruits, part of which are eaten by wild animals (e.g. of Gonocaryum, Cantleya, Gomphandra, Medusanthera). Buoyancy of fruits is certain for Merrdhoden- dron less so for Gonocaryum and Stemonurus, and may have contributed to their dispersal Gals No galls specific for a distinct genus or species are hitherto known from Ua\^s^an Icaanaceae (c/ DOCTERS VAN Leeuwen, Zoocecidia, 1926, 332, f. 597-599: Gomphandra, Phytocrene, Platea; Ned. Kruidk Axdn. 5\,\9A\,\15,f. A%: Stemonurus). • . . i x/i i •■• Anatomy. Wood. Structure, properties and identification: Den Berger, Determinatietabe Ma lesie Veenman Wageningen (1949) (hand lens), Balan Menon, Res. Pamphl. For. Dept Malaya 18 (1955) iT6T;«SrCHATTAWAY, Trop. Woods 102 (1955) 55-74, ibid. 104 (1956) 100-124 (crystals) dIch Mai For Re^^^^^^^ (1941) 233 {Cantleya, Stemonurus), Fei-Tan & Chu, Malayan Forester 32 0969)' 2^7 293"(fibres; StemomLs), Gosh, Rao c.s. Indian Woods 2 (1963) New Delhi {Apodytes Gomphandra, Nothapodytes as ^Mappia^), Janssonius, Key to Javanese woods, ^eyden (1952) and Moll & Janssonius Mikr. 2 (1911) 234-254 (Apodytes, Gomphandra, Platea, Stemonurus) Va^ der Walt c.s. T. Nat. Wetensk. Suid-Afr. (1970) 173-199 {Pyrenacantha, anomalous growth)^ For |f"er/J ^"[^5^^ also comprehensively covering the older literature see Solereder^ Syst. Anat^ D.cot Stuttgart (1899) 227-237 and ibid. (1908) 81-83, and Metcalfe & Chalk, Anat. Dicot. Oxford (1950) 367-377. The vegetative anatomy of the Icacinaceae is extremely diverse and no character from leaf twig or wood anatomy is consistent or even typical for the family. Because of the enormous range m structure, rnatomkal Characters have been extensively used in discussing the relationships and subdivisions within the family. Engler's wood-anatomical characterizations of the tubes Icaaneae, lodeae, Sar- wumu uic laiiuy. i- ^ ,c;t, Rf>r Prpn<;<; Akad Wiss Berlin 1 8, 1893, 1-23) were not based on costigmateae and Phytocreneae (Sitz. Ber. I'reuss. Akaa. wiss. oerim lo, 107.?, i , enough material and therefore inaccurate. Bailey & Howard (J. Arn. Arb. 22, 1941 125-132, 171-187 432442, 556-568) distinguished three groups based on a combination of the nodal anatomy and the type of vessel perforation. Group I with trilacunar nodes and exclusively scalariform vessel perforation includes Apodytes, Citronella, Pittosporopsis and Platea (all Icacineae),Gron^ II with t"lacunar nodes and a mixfure of simple and scalariform vessel perforations xncXxxdes Cantleya, ^^"^f'^""'' ^^f^^"' thera, Gomphandra (as ^Stemonurus) and Stemonurus (as 'Urandra) (all ^cacneae) Gjoup III with unilacunar nodes and simple vessel perforations includes lodes, Mappianthus, Merrdliodendron Miquelia, Natsiatum, Nothapodytes, Phytocrene, Polyporandra, Pyrenacantha, Phyticaryian and Sar^^'^lSnia (from all four tribes sensu Engler). Other xylem characters such as vessel distribution vessel member length, type of fibres, parenchyma arrangement and ray structure are more or less related to these groups of increasing xylem specialization. Codiocarpus, Hartleya, Pseudobotrys and Whitmorea were not in- cluded in the studies of Bailey & Howard. Whitmorea has a trilacunar node and mixed vessel perlo- rations and therefore belongs to group II. This observation supports Sleumer s view that ^^^^^'norea is related to Stemonurus (Blumea 17, 1969, 264). All data on wood anatomy provided by Bailey & Howard suggest a more or less continuous range of xylem characters within Icacmaeceae rather than the presence of distinct boundaries. .... ■ , , u • ♦ ^o tViot Hairs. Heintzelmann & Howard (Amer. J. Bot. 35, 1948, 42-52) distmguish ten hair types that occur in floral parts or on leaves and young twigs of Icacinaceae. Noteworthy are the two-armed or Malpighiaceous hairs in Nothapodytes and some genera from outside Malesia. This hair type intergrades with the so-called Icacinaceous hair with one much reduced and one well developed arm occurring in a majority of the genera. Rhyticaryum has clustered hairs; Platea stellate-peltate hairs (called scales in the taxonomic part) (see also Uphof, Hummel & Staesche, Handb. Pflanzenanat. IV, 5 1962). Stomata are very imperfectly known in Icacinaceae. Both anomocytic and paracytic stomata have been recorded in literature, but actinocytic {Merrilliodendron) and anisocytic {Medusanthera and Ste- monurus) types also occur. 1971] ICACINACEAE (Sleumer) 3 The vascularization of the petiole and midrib would also merit further investigation. Species of Apodytes, Citronella, Medusanthera, Phytocrene, and Stemonuriis have central strands with or without latero-dorsal bundles, but the range in the whole family must be much greater since Pennantia shows a very complex pattern icf. Blumea 18, 1970, 217). Crystals occur as druses, solitary rhomboids and crystal sand in various combinations (c/. Heintzel- MANN & Howard, I.e.). The fine translucent dots of the leaves of Merrilliodendron are caused by large intercellular spaces in the spongy tissue of the mesophyll. Domatia have been recorded for several Icacinaceous genera but are absent from all Malesian species. — P. Baas. Phytochemistry . The few chemical data available about the chemistry of this family were summariz- ed in my "Chemotaxonomie der Pflanzen' 4 (1966) 275-277, 494. In the meantime, oleanolic acid was isolated from the bark of Apodytes dimidiata E. Meyer (= A. beddomei Mast.). The scanty chemical information available about Icacinaceae at this time, prevents a chemotaxonomie discussion. — R. Hegnauer. Uses. In Malesia but a few tree species grow to big dimension with a clear bole {Cantleya, Stemonurus, Platea). Of these only Cantleya corniciilata (Becc.) Howard has a marketing value and is exported from Sarawak and Brunei. Icacinaceae have a hard or mostly rather soft, often whitish or cream, sometimes aromatic wood, and are apparently only locally used by the natives, as can be deduced from the many vernacular names known. The leaves of Rhyticaryum species are eaten as a vegetable. Medicinal use is recorded for several species but needs confirmation. The seeds of Cantleya, Phytocrene, Stemonuriis and Sarcostigma are edible, but of a poor quality. The stems of lianas (Miquelia, Phytocrene) hold fresh edible water. Taxonomy. Four of the genera found in Asia and Malesia were revised by R.A.Howard (Cantleya: J. Am. Arb. 21, 1940, 479; Codiocarpus: Brittonia 5, 1943, 60; Medusanthera: Lloydia 6, 1943, 133; Nothapodytes: J. Arn. Arb. 23, 1942, 66); these revisions were based on rather scarce materials as far as Malesia is concerned. A precursory paper with revisions of most of the genera concerned and based on practically all Asiatic and Malesian specimens available today was published by myself in Blumea 17 (1 969) 1 8 1 -264, supplementing my previous studies on the family (Notizbl. Berl.-Dahl . 15,1 940, 228-257 ; ibid., 1942, 359-365; in E. & P. Natl. Pfl. Earn. ed. 2, 20 b, 1942, 322-396). key to the genera 1. Trees or (not climbing) shrubs. 2. Sepals essentially free at least in their upper Yi and imbricate. 3. Flowers bisexual. 4. Petals free, valvate, or subimbricate in their upper part 1. Citronella 4. Petals connate below into a tube, their upper part free and valvate in bud. . 2. Pseudobotrys 3. Flowers unisexual. (Petals connate below.) 5. Filaments free, fixed to the very base of the petals. Leaves with a layer of rounded to star-shaped appressed scales underneath at least in the young state 3. Platea 5. Filaments adnate for almost their entire length to the lower tubular part of the petals. Scales absent 4. Gonocaryura 2. Sepals connate into a cup-like calyx, its upper free part (or lobes), if any, short and not imbricate. 6. Flowers unisexual (or functionally so). 7. Drupe ovoid-ellipsoid or oblongoid, without a fleshy lateral appendage. Flowers in rather short cymes 5. Gomphandra 7. Drupe laterally compressed (almond-like). 8. Drupe without a fleshy lateral appendage. Flowers in spikes (very rarely in panicles composed of spikes, or almost fascicled) 6. Rhyticaryum 8. Drupe with a thick fleshy, laterally borne, practically entirely adnate appendage. Flowers in cymes. 9. Disk unilateral, thick-squamular 7. Hartleya 9. Disk absent. 1 0. Filaments glabrous. Fleshy appendage of drupe covering two pronounced ribs of the endocarp. 8. Codiocarpus 10. Filaments with apical, longish, club-shaped hairs at least in the fertile stamens. Prominent ribs under the appendage of the drupe less pronounced or absent. . . 9. Medusanthera 6. Flowers bisexual. II. Ovary with a lateral swelling which in the fruit developcs into a thick succulent appendage. Disk absent 10. Apodytes II. Ovary and fruit without such an adnate appendage. Disk whether or not present. 12. Connective surpassing the anther cells as a marked glabrous apiculus. Outer part of the endocarp finally spongious-corky and deeply irregularly lacunosc. Disk absent. II. Merrilliodendron 4 Flora Malesiana [ser. I, vol. 7^ 12. Connective, if any, hardly or not surpassing the anther cells. Outer part of the endocarp fibrous, slightly ribbed or grooved lengthwise, or smooth outside. 13. Peduncle of inflorescence with numerous small knob-like bracts which form alveoles. Stigma peltate. Disk absent 12. Cantleya 13. Peduncle quite smooth. Stigma small, subcapitate or point-like. Disk ± cup-shaped. 14. Inflorescence usually terminal. Anthers glabrous 13. Nothapodytes 14. Inflorescence axillary. Anthers with an apical tuft of penicillate hairs. 15. Flowers sessile. Petals up to 6 mm, free to almost the base. Stigma point-like at the top of the ± attenuate (sometimes shortly style-like) part of the ovary 14. Stemonurus 15. Flowers 1-2 mm pedicelled. Petals (12-) 13-1 5 mm, free in the distal part only. Stigma small on one side of the inverted, i.e. cup-like distal part of the ovary. ... 15. Whitmorea 1. Climbing shrubs, or lianas, sometimes with tendrils. Flowers unisexual or functionally so. 16. Leaves opposite. 17. Anthers broadly club-shaped to subglobular, many-celled, with numerous pollen-bearing alveoles. 16. Polyporandra 17. Anthers, as usual, with 2 cells 17. lodes 16. Leaves spirally arranged. 18. Flowers in elongate spikes or spike-like racemes, these solitary or sometimes composed to panicles. 19. Leaves markedly prominently tessellate on both faces. Sepals persistent. Albumen absent; cotyledons thick-fleshy 18. Sarcostigma 19. Leaves with rather lax and but slightly raised reticulation. Sepals absent. Albumen thick, ru- minate; cotyledons foliaceous 19. Pyrenacantha 18. Flowers in peduncled heads or umbels, these solitary or composed to racemes or panicles. 20. Style absent, i.e. stigma sessile, thick-peltate 20. Miquelia 20. Style (very) shortly thick-columnar, with 2-4 stigmatic lobes 21. Phytocrene 1. CITRONELLA D. Don, Edinb. New Phil. J. 13 (1832) 243; Sleum. Blumea 17 (1969) 186. — Villaresia R. & P. Fl. Per. Chil. 3 (1803) 9, t. 231, non R. & P. 1793. —Pleuropeta- lon Bl. Mus. Bot. Lugd.-Bat. 1 (1850) 248, non Hook. f. 1845. — Chariessa MiQ. Fl. Ind. Bat. 1 (1856) 794. — Fig. 1. Trees or shrubs, bisexual (Mai.). Leaves spiral, entire and slightly re volute at the edge (Mai.), petiolate. Inflorescences terminal and/or (supra-)axillary, panic- ulate or thyrsoid, cymes of secondary branches scorpioid, whether or not forked, in Mai. greyish to rusty hirsute. Flowers 5-merous, sessile, subtended by a minute bract. Calyx imbricately lobed V3 to 1/2, persistent. Petals free, valvate, or subim- bricate in the upper part, apex inflexed, midrib inside much prominent, sometimes wing-like. Stamens 5, free; filaments fleshy, subulate; anthers elliptic, or ovate, or subcordate, introrse, basifixed. Disk 0. Ovary subgibbous, 1 -celled, rarely 2-celled by the presence of a pseudoloculus. Style 1 (rarely 2), slender, stigma small, capitate, subbilobed. Drupe moderately fleshy; endocarp woody, rather thin. Seed 1, longitudinally plicate around the vertical pseudoloculus, hippocrepiform; embryo in the copious endosperm small. Distr. About 21 spp. of which c. 12 occur in tropical Central and South America, and 6 spp. in Australia (New South Wales, Queensland), Solomon Is., New Caledonia, Loyalty Is., New Hebrides, Fiji, Samoa and Tonga Is.; in Malesia 3 spp. Fig. 2. Ecol. Understorey of lowland and lower montane rain-forest. Taxon. The genus was revised by R. A. Howard (Contr. Gray Herb. 142, 1942, 60-89, t. 4-6). The inflorescence and floral characters used there to separate the Malesian spp. have been found of little value after a reinvestigation on the base of much more material than seen by Howard. This is the reason why in the key fruit characters have been preferred for the discrimination of species. Howard keeps the New World spp. apart from the Old World ones on the sectional level; the characters given for the dis- crimination of these sections are rather vague or do not hold. ICACiNACEAE (Sleumer) Fig. 1. Ciironella suaveulens (Bl.) Howard, a. Inflorescence, •' '/j, h. flower bud, x5, c. petal from the inside. /IVi, d. ovary, x7'/2, e. stamen, /7'/2,/. habit and infructescence, x '/j, g. drupe, x 1 ia-e ACHMAD 1 181,/-^ A( HMAD 220). KEY TO THE SPECIES 1, C suavcolcns 1. Drupe essentially cllipsoid-oblongoid, (l.6-)l.8 2.2( 2.4) by 0.9-1. 2(-l. 3) cm. I. Drupe ovoid, or ellipsoid-ovoid, or broadly ellipsoid, 2-.3 by 1.8-2 cm. "> Leaves broadly ovate, base h obtuse to rounded. Solomon Is., Samoa, Tonga, New Hebrides. C. samoensis (A. Gray) Howarij 2. Leaves oblong- to obovate-elliptic, base '• cuneatc. 3. Leaves r- obtuse or shortly and rather abruptly acuminate, tip biuntish. • • 2. C. liHifolia 3. Leaves rather gradually and ; acutely acuminate at apex 3. C". phillppinensis Flora Malesiana [ser. I, vol. 7^ 1. Citronella suaveolens (Bl.) Howard, J. Am. Arb. 21 (1940) 475; Contr. Gray Herb. 142 (1942) 82; Back. & Bakh./. FI. Java 2 (1965) 59; Sleum. Blumea 17 (1969) 187. — Pleiiropetalon suaveolens Bl. Mus. Bot. Lugd.-Bat. 1 (1850) 248. — Cluiriessa suaveolens (Bl.) Miq. Fi. Ind. Bat. 1 (1856) 794; Becc. Malesia i (1877) 118; Sleum. Notizbl. Berl.-Dahl. 15 (1940) 229; Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 2. — Villaresia suaveolens (Bl.) Valet. Crit. Overz. Olacin. (1886) 199, t. 5, f. 32 a-g; Back. Schoolfl. Java (1911) 228. — C. brassii Howard, Contr. Gray Herb. 142 (1942) 81. — Fig. 1. Tree, (8-) 15-36 m, trunk up to Vi m 0, rarely with thin buttresses; bark grey-brown, irregularly fissured. Branchlets smooth or striate, older parts lengthwise set with elongate lenticels. Leaves elliptic to oblong- or ovate-elliptic, variable in shape and size, apex generally short-acuminate (tip subacute and often plicate then), or obtuse- rounded, base broadly attenuate to rounded, often subequal, coriaceous, shining above, (6-)10-24(30) by (3-)5-13(-17) cm, nerves (4-)5-6 curved- ascendent and anastomosing pairs, lower l(-2) rather close to the base of the lamina, slightly impressed above, much raised beneath, retic- ulation fine to rather coarse, a little prominent above, more distinctly so beneath; petiole 6-15 by 2-3 mm. Panicles terminal, greyish-rusty hirsutu- lous, mostly solitary and spike-like initially, some- times 2 or 3 forming a ± pyramidal inflorescence, lateral branches of the single panicles of ± the same length (0.5-2 cm), mostly 2-(rarely more-) forked, each branch recurved and bearing several flowers arranged in a cyme. Flowers bisexual, fragrant. Calyx c. 1.8 mm, lobes ciliate. Petals elliptic-oblong, white to cream, or some- times suffused with red, (4-)5(-6) mm at full anthesis, midrib inside raised considerably, some- times almost wing-like in the lower '/2-% part. Filaments thick-subulate, c. 3 mm; anthers ovate- elliptic or subcordate, 1 mm. Ovary ovoid, glabrous, style 1-2 mm, stigma small, a little oblique. Drupe ellipsoid to oblongoid (subcylin- drical), sometimes a little oblique, (1.6-)1.8-2.2 (-2.4) by 0.9-1. 2(-l. 3) cm, smooth or irregularly lengthwise, very slightly many-ribbed in the lower part, purple when ripe, blackish when dry. Distr. Malesia: NW. Sumatra (Simalur I.), W. & Central Java, N. & E. Borneo, Celebes, S. Moluccas (Kai Is.), New Guinea. Ecol. Primary rain-forest on well drained soil, from the lowland up to 1600 m, apparently scattered. Uses. Wood light brown, hard and heavy, not used. Known by numerous vernacular names. Vern. Simalur: awa iseum-iseum, iseum batu, i. iseum fatuh, i. pajo, i. sito bulung, sitenheur delok, tutun surimanu, M; Java: jerukie, S; Celebes: keker, Menado (Tt), balula lembo, Tobela, lenguru tanru, pano, Bugin., sokko, Makassar; New Guinea: garus, Dumpu, mara, Faita, pomak, Armat, sakohukwa, Manikiong, sibeer, Hattam, soromma rinde, Depapre, tambeu, Iko (Hol- landia), ulumon, Amele, yewel, Bilia. Fig. 2. Distribution of Citronella. 2. Citronella latifolia (Merr.) Howard, J. Arn. Arb. 21 (1940) 472; Contr. Gray Herb. 142 (1942) 80, t. 6, f. 9-12; Sleum. Blumea 17 (1969) 188. — Villaresia latifolia Merr. Philip. J. Sc. 14 (1919) 415; En. Philip. 2 (1923) 488. — Chariessa latifolia (Merr.) Sleum. Notizbl. Berl.-Dahl. 15 (1940) 229. Large tree. Branchlets dark brown, with sparse lenticels. Leaves obovate or oblong-elliptic, apex obtuse or rounded, the bluntish tip sometimes abruptly short-acuminate and folded, base cu- neate, a little decurrent, coriaceous, shining above, (12-)14-18 by (6-)8-ll cm, nerves 5-6 anasto- mosing pairs, reticulation rather dense, minutely raised on both faces; petiole l-1.5(-2.5) cm by 2(-3) mm. Infructescence terminal, consisting of 1 or 2, sometimes 3 almost pyramidal panicles, up to 15 cm, rusty hirsute; each panicle with 1971 ICACiNACEAE (Slcumer) numerous racemosely arranged branches which generally are 2(-3)-forked and bear several flowers crowded at the curved end of the branches, the latter up to 5 cm in the lowest, and c. 1 cm (and usually not forked) in the uppermost part of the panicle. Drupe ovoid to broadly ellipsoid- ovoid, c. 1 by 1.8 cm in submature state, brown when dry, very slightly and irregularly many- ribbed lengthwise in the lower -3; endocarp hard, c. 1 mm. Distr. Malesia: Philippines (Samar, in the Catubig R. area), one collection in fruit. Ecol. Damp forest near river at low altitude. Vern. Malaampipi, S. L. Bis. 3. Citronella philippinensis (Merr.) Howard, J. Arn. Arb. 21 (1940) 474; Contr. Gray Herb. 142 (1942) 85, t. 6, f. 1-8; Dahl, J. Arn. Arb. 33 (1952) 260, f. 20 «& 20 A (pollen); Sleum. Blumea 17 (1969) 188. — Villaresia philippinensis Merr. Philip. J. Sc. 14 (1919) 414; En. Philip. 2 (1923) 488. — Chariessa philippinensis (Merr.) Sleum. Notizbl. Berl.-Dahl. 15 (1940) 229; Hatus. Mem. Fac. Agr. Kagosh. Un. 5, 3 (1 966) 38. Tree. 4-5 m. Branchlets smooth. Leaves elliptic to elliptic-oblong, apex rather gradually acu- minate for 1-1.5 cm and subacute, base cuneate and a little decurrent on the petiole, ± inequal, coriaceous, shining on both faces, 6-12 by (2-)4— 6.5 cm, nerves 4-5 rather steeply curved-ascending pairs, reticulation dense, distinctly raised mainly on the undersurface; petiole 2(-3) cm by 2 mm. Panicles terminal, solitary or in twos, 5-8(-10) cm, lateral branches numerous and of equal length (c. 1 cm), rather crowded, each bearing at apex 5-8 crowded cymosely arranged flowers. Calyx 1.5 mm, lobes ciliate. Petals oblong, white, gla- brous, 5 (-6) mm, the midrib extending as a keel in the lower half of the petal. Filaments c. 5 mm; anthers broadly ovate-cordate, 1 mm. Ovary ovoid, glabrous, 5 nun, style slender, 3^ mm, stigma capitate, minute. Drupe ovoid, rather oblique, blackish, 2.5-3 by 2 cm; endocarp shallow - ly and irregularly many-ribbed lengthwise, 1-1.5 mm 0. Distr. Malesia: Philippines (N. Luzon). Ecol. On forested slopes, c. 1000 m, apparently rare. Excluded Villaresia scandens Hassk. Nat. Tijd. N. I. 10 (1856) 152 = Dichapetalum timoriense (DC.) BoERL. (Dichapetalaceae). 2. PSEUDOBOTRYS MOESER in Fedde, Rep. 10 (1912) 310; Sleum. in E. & P. Nat. Pfl. Fam. ed. 2, 20 b (1942) 360, f. 101, A & B; Blumea 17 (1969) 248. — Fig. 4. Trees. Leaves spirally arranged, coriaceous, entire, penninerved. Flowers bi- sexual, born in short fascicles or panicles on the trunk. Pedicel articulated with the calyx, subtended by 2 small bracteoles. Sepals 5, connate at base, imbricate. Petals 5, linear, valvate, tips a little inflexed inside, connate into a tube in the lower Vi-y^, caducous. Stamens 5, almost equalling the petals in length; filaments filiform, dilated towards the base and fixed there to the petals; anthers linear, subsagittate, introrse, almost basifixed. Ovary ovoid-subglobose; style filiform; stigma small, depressed-subcapitate. Drupe: exocarp thin, fleshy, early dissoluted; endocarp woody, verrucose outside, forming a pseudoseptum along the funiculus inside. Seed 1; embryo small. Distr. Malesia (New Guinea), 2 spp. Fig. 3. KEY TO THE SPECIES 1. Ovary densely brownish-pubescent. Petals 2.8-3.2 cm. Anthers 5-6 mm 1. P. dorae I. Ovary glabrous. Petals c. 2.2 cm. Anthers 3-4 mm 2. P. caullHora 1. Pseudobotrys dorae Mof.ser in Fcddc, Rep. 10 (1912) 310; SuLM. Notizbl. Berl.-Dahl. 15 (1940) 235; m E. & P. Nat. Pfl. Fam. cd. 2, 20 b (1942) 360. r. 101. A & B; Blumea 17 (1969) 249. Trcclct or shrub, 1.5-6 m, glabrous; bark grey, squamular. Branchlets terete, striate, slender. Leaves lanceolate to oblong or elliptic-oblong, apex gradually more shortly or long acuminate, tip blunt, base broadly cuncatc to rounded. coriaceous, of a dull olivaceous-green colour and a minutely tuberclcd undersurface when dry, greenish yellowish in fresh specimens, entire. l6-32(-44) by 5-l4(-l8) cm, midrib slightly raised above, strongly so beneath, nerves (6-)8-IO(-l2) pairs, lower ones curved, upper ones generally more straight, faintly inarching before the edge, hardly raiscdorobscurc above, slightly prominent bciieuth, veins lax, generally rather inconspicuous; petiole Flora Malesiana [ser. I, vol. V Fig. Yseudobotryscauliflora (Pulle) Sleum. a. Habit, x 1/2,6. inflorescence, x I, c. young infructescence XI, rf. flower. XI 1/2.^. ovary, x 1 1/^./. endocarp of mature drupe, xl (a <& c Docters van Leeuwen 9127, b,d&e ditto 9307, f ditto 1 1243). 1971 ICACINACEAE (Slcumer) rugose, l-1.5(-2) cm by 2-3 mm. Flowers in short fascicles or corymbs from swollen parts of the trunk (occasionally also from the axils of leaves?). Pedicels thickish, short. Sepals subovate, blunt, dull purple, c. 3 mm. Petals linear, blunt, thin, connate irregularly in the lower part, i.e. early splitting from top after the bud stage, spreading or recurved at full anthesis, white all over, or pur- plish at tube, 2.8-3.2 cm by 2-2.5 mm. Filaments 2-2.4 cm; anthers 5-6 mm. Ovary 3-4 mm, densely brownish-pubescent; style 2.5-2.7 cm. Drupe only known in submature state, obliquely obovoid, apiculate, laterally a little compressed, c. 2 by 1.7 by 1.3 cm, subtended by the reflexed sepals, brownish-hairy, finely verrucose outside; endocarp hard; pseudoseptum protruding for c. 1 mm. Seed 1. Distr. Malesia: New Guinea (Morobe, Central and Northern Distr.). Fig. 3. Fig. 3. Distribution of Pseudobotrys. P. dorae MOESER (crosses), P. caulijlora (Pulle) Sleum. (dots). Ecol. Primary (also swampy) forest, 30-1525 m, apparently very local. Fl. May-Sept., fr. Nov. 2. Pseudobotrys cauliflora (Pulle) Sleum. Notizbl. Berl.-Dahi. 15 (1940) 235; Dahl, J. Arn. Arb. 36 (1955) 160, f . 1 & 1 A (pollen); Sleum. Blumea 17 (1969) 249. — Chariessa cauliflora Pulle, Nova Guinea 8 (1912) 657; Schellenb. Bot. Jahrb. 58 (1923) 159; Birnie, Nova Guinea 14 (1926) 275. — Fig. 4. Tree, (4-)6-18 m; bark dark grey-green; wood white, with prominent rays. Leaves very similar to those of P. dorae, though sometimes more ovate, up to 36 by 19 cm. Flowers massed on dome-like swellings of l-3(-10)cm o on lower trunk, numerous, set in small fascicles. Pedicels 1-3 mm, minutely bracteolate at base. Sepals ovate, ciliate, purple, 2(-3) mm. Petals linear, apex acute, in bud stage coherent in a tube which splits down- wards at full anthesis to the lower third or less, white, or white inside and purplish outside, with 3 longitudinal darker lines, c. 2.2 cm by 2 mm. Filaments c. 1.6 cm; anthers 3-4 mm. Ovary gla- brous, c. 3 mm; style slender, 16-18 mm. Drupe ovoid, attenuate-apiculate, 2.5-3 by 2 cm, on thickish pedicel 8-12 mm; exocarp fleshy, dark blue at full maturity, wrinkled in dry fruit; en- docarp woody (2 mm), outside with knob-like emergences in numerous longitudinal rows, knobs smaller towards apex of the fruit; inside with a large woody pseudoseptum around which the seed is formed. Distr. Malesia: New Guinea. Fig. 3. Ecol. Rain-forest substage, river flood-plains, or hillsides, 3-175 m (once said to be collected at 1645 m), scattered in general, though sometimes locally common. Fl.fr. Jan.-Dec. 3. PLATEA Bl. Bijdr. (1826) 646; Sleum. Blumea 17 (1969) 242. — Platystigma R. Br. in Wall. Cat. (1832) n. 7523, nom. nud.; ex Benth. in Benth. & Hook. f. Gen. PI. 3 (1880) 283, nota; H(X)K./. Ic. PI. 18 (1887) t. 1707. — Fig. 6. Trees, generally tall and with straight, angular or ± deeply grooved bole, rarely treelels or shrubs, dioecious. Leaves spiral, almost distichous, entire, penninerved, at least initially covered with fascicled or star-shaped, early going hairs, and/or subpersistent star-shaped, flat, finally silvery scales with a distinct darker centre, as are branchlcts and inflorescences; petioles not rarely transversely wrinkled as in Gonocaryum. Flowers axillary, the o ones in interrupted, glomcrulose, mostly to panicles arranged spikes, sessile, subglobular in bud, each subtended by a small persistent bract, the ; ones in few-flowered short cymes, each on a distinct pedicel and subtended by a subpersistent bract immediately below the calyx lobes. Calyx lobes (4-)5, slightly imbricate, small, persistent. Petals (4-)5, small, valvalc, glabrous, inflexcd at apex, shortly connate at base, absent in the r flowers. — '^ Flowers: Stamens (4-)5, inserted with short filaments at the tube of the petals; anther cells ovoid-2-cellcd, basifixed, laterally (almost cxtrorsely) dehiscent. Ru- diment of an ovary generally absent. — V Flowers: Ovary thick-cylindric to ob- 10 Flora Malesiana [ser. I, vol. 7^ conical, blunt, crowned by a large sessile discoid stigma. Drupe elongate-ovoid, crowned by the large stigma, on short pedicel; exocarp thin-fleshy, red-orange to finally purplish blackish; endocarp woody, thin, with 1 or 2 slight longitudinal grooves, reticulately wrinkled; embryo small, in the apex of the albumen. Distr. About 5 spp., one endemic in Hainan, another extending from Sikkim into SE. Asia and Malesia; in Malesia 4 spp., of which 2 endemic in Borneo and the other 2 from the Malay Peninsula and Sumatra to New Guinea and New Britain (not known yet from the Solomon Is.). Fig. 5. Ecol. Mainly rain-forest, from the lowland up to c. 2200 m (on Mt Kinabalu up to 2895 m?), often on wet soil, scattered. Fl.fr. Jan.-Dec. Uses. The wood is whitish cream, available in big dimensions due to the large size of the trees, but soft and suitable only for inside house constructions. Note. The distinction of species in this genus is mainly based on the type of indument, clearly rec- ognizable only with a lens. This can only well be examined on young leaves, where the full set of in- dument characters is still present, as hairs and/or scales go rather early, and old leaves may become practically glabrous. The scales give the undersurface of the leaves a metallic silvery lustre similar to that found in leaves of certain Myristicaceae, Euphorbiaceae, Sapotaceae, and even Lauraceae, with which they are sometimes confused in sterile sheets. Young leaves may show fine, round, translucent dots against strong light (Endert, Med. Proefst. Boschw. Ned. Ind. 20, 1928, 193). KEY TO THE SPECIES 1. Indumentum on the undersurface oi young leaves giving a woolly appearance to the naked eye, and under a lens representing two distinct strata, i.e. an upper layer of partly flattish, though mostly oblique- ly erect star-shaped or fascicled hairs (with a very small or generally invisible dark centre), and a lower layer of scales. 2. Leaves shortly cordate at base, almost peltate, lateral nerves, veins and veinlets ± distinctly impressed above, the lamina ± bullate, hairs persistent 1. P. bullata 2. Leaves never properly cordate at base, nerves sometimes, veins and veinlets never impressed above, hairs generally early disappearing except along midrib and nerves 2. P. latifolia 1 . Indumentum on the undersurface oi young leaves giving a scaly appearance to the naked eye, seemingly representing a single layer of scales, but under the lens appearing to consist of two layers of flat star- shaped scales, i.e. a lower layer of ± persistent, small and densely arranged scales, and an upper layer of similar, though larger and more scattered, rather early caducous ones. 3. Leaves (elliptic or ovate-elliptic, rarely ovate) coriaceous, rigid 3. P. sclerophylla 3. Leaves (of various shape) coriaceous to subcoriaceous, ± flexible 4. P. excelsa 1. Platea bullata Sleum. Blumea 17 (1969) 243. Tree 13-14 m, c. 15 cm 0; bark smooth, grey- brown. Branchlets rufous-brownish tomentose and slightly many-grooved lengthwise as are petioles and rachises in dry specimens. Leaves lax, sub- obovate-elliptic or -oblong, apex subacutely acu- minate for c. 1.5 cm, base rounded in general, the very base short-cordate and the lamina almost peltate, firmly chartaceous, olivaceous-brownish Fig. 5. Distribution of Platea. and dull above, more rufescent beneath in dry specimens, ± deeply bullate by nerves, veins and partly also veinlets above, these well pro- minent beneath, glabrous above except the short- tomentose midrib and maybe nerves, brownish- rufous-tomentose and soft to the touch by obliquely erect stellate or fascicled longish hairs all over the undersurface especially on midrib, nerves and veins, moreover covered all over beneath with a coherent greyish silvery layer of small star-shaped, flat, appressed scales each with a minute dark centre (similar to stellate flattish hairs), 14-32 by (6-)7-13cm, nerves 7-10(-12) pairs moderately curved-ascending and looping, reticulation fine and dense, minutely raised above, rather obscure beneath; petiole 2-2.5 cm by 2-3 mm. few- 32 Flora Malesiana [ser. I, vol. 7^ to many-flowered cymes, together on a rather stoutish peduncle (1-2 cm by 1mm), fulvous- puberulous. Calyx widely cup-shaped, subentire, mostly laxly hairy, 1-1.5 mm. Petals 5, remaining connate to a subcampanulate tube for their lower % part, greenish-whitish, glabrous (maybe with a few hairs at apex), (3-)4-5 mm. Stamens 5, exserted for 1-2 mm; filaments set with few to numerous longish hairs vcntrally below the anther cells, much less or not so dorsally at connective. Rudiment of ovary glabrous. — ? Inflorescences with 3 or 4 one-flowered branches (3-6 mm) on top of a common peduncle (c. 1 cm). Calyx and petals as in cJ flowers. Staminodes 5, laxly or not hairy. Ovary glabrous. Drupe subelliptic- oblongoid, slightly asymmetrical, ± obtuse at both ends, base sometimes slightly (or uni- laterally) swollen, pale yellowish green, sometimes with reddish or orange tinge, 2-2.5(rarely -3.2) by (0.9-)l-1.2(-1.5) cm, ribs of endocarp marked and down to or almost to the very base. Distr. Melanesia (New Britain, New Ireland, Solomon Is. and Santa Cruz Group); in Malesia: New Guinea. Ecol. Primary lowland and montane, also swampy, sometimes secondary or devastated forest, usually in understorey, locally not rare, from sea-level up to 1675 m. Uses. Wood creamy to whitish, rays prominent, no use known, apparently due to its small di- mensions. Vern. Aiyalo, Kwara'ae. Note. Difficult to separate from G. australiana F. v. M. in leaf and fruit characters, though cer- tainly diff"erent in flower structure. 22. Gomphandra parviflora (Bl.) Valef. Grit. Overz. Olacin. (1886) 218, t. 4, f. 10 & 11; Sleum. Blumea 17 (1969) 200. — Stemonuriis parviflorus Bl. Mus. Bot. Lugd.-Bat. 1 (1850) 250; Walp. Ann. 2 (1851) 182; Miers, Ann. Mag. Nat. Hist. II, 10 (1852) 36, repr. Gontr. Bot. 1 (1852) 87; O. Ktze, Rev. Gen. PI. 1 (1891) 112. — Lasian- thera parviflora (Bl.) Miq. F1. Ind. Bat. 1 (1856) 791; Suppl. (1860) 137. Treelet. Branchlets with minutely pubescent tips. Leaves oblong to elliptic-oblong, rarely lan- ceolate or oblanceolate-oblong, apex shortly rather abruptly acuminate for 1-2 cm, base broadly cuneate and slightly inequilateral, firmly membranous to subchartaceous, glabrous, green- ish-greyish to olivaceous and dull in dry speci- mens, (10-) 15-22 (-30) by 4-8 cm, nerves (8-) 10-12(-14) rather straight and spreading pairs, ± parallel to each other and looping before the edge, the lowest pair in part of the leaves (in the same specimen!) ascending in a more acute angle from the midrib, reticulation none; petiole often yellowish as is the midrib beneath in dry speci- mens, 1-1.3 cm by 1.5 mm. — x cupular, 0.5 mm, dense- ly strigose. Petals 5, connate in the lower 2/,, greenish, laxly strigose at apex outside, 1.5-2 mm. Rudiment of ovary subcylindric, hairy. — ? In- florescence only known in fruiting stage: rachis subdensely strigose, 3-4(-5) cm, slender (hardly I mm 0), drupes laxly arranged. Remnants of calyx densely, of petals laxly strigose. Drupe ovoid-ellipsoid, much compressed laterally, lateral crests rather sharp, laxly setulose, 1.5 by 1-1.3 by 0.8-1 cm in dry, said to measure 4 by 2 cm in fresh state; exocarp fleshy, yellow with orange hue, or reddish. Distr. Malesia: W. New Guinea (Vogclkop Peninsula). f col. Primary forest, also along creek in pe- .lodically flooded forest dominated by Inocarpus, from sca-lcvcl up to 45 m. 11. Khylicaryum fasciculatum Beer. Malesia I (l«77) 121. t. 4, f. 24 & 25 m. V); Valet. Crit. Over/ Olacin. (1H86) 257; ScHELLENB. Bot. Jahrb. 58 (1923) 168; Sleum. Blumea 17 (1969) 252. Shrub, 1-2 m. Branchlets slender, youngest parts densely yellowish-substrigose. Leaves ob- long to subovate-oblong, apex abruptly acuminate for 1, rarely up to 2 cm, tip rather blunt, base broadly cuneate, membranous to chartaceous, laxly to very laxly strigose on midrib and nerves, rarely also on veins beneath, glabrous otherwise, green or yellowish when dry, shining on both faces, 12-24 by 5-9 cm, nerves 8(-10) curved- ascending and looping pairs, prominent mainly on the undersurface, reticulation fine and rather dense, moderately raised on both sides; petiole strigose, 6-12 by 1.5 mm. — c? Inflorescence un- known. — 2 Flowers 3-5(-7) very close to each other along a stout c. 6 mm long rachis, seemingly fascicled, densely strigose in all parts. Calyx cup- shaped, 1 mm incl. the short teeth. Petals fleshy, connate almost halfway, 1.8 mm. Ovary sub- conical, densely strigose, 1 .5 mm. Drupe ± oblique- ly subovoid-ellipsoid, much flattened laterally and distinctly crested, laxly strigose, red, (1.3-) 1.7-2 by (1-)1.5 by 0.7-0.9 cm; endocarp ir- regularly coarsely reticulate-alveolate. Distr. Malesia: W. New Guinea (Vogelkop Peninsula). Ecol. Lowland forest, apparently rare. 12. Rhyticaryum longifolium K. ScH. & Laut. F1. Schutzgeb. (1900) 415; Nachtr. (1905) 306; Sleum. Blumea 17 (1969) 252. — /?. oncocarpum K. ScH. & Laut. F1. Schutzgeb. (1900) 416; Schellenb. Bot. Jahrb. 58 (1923) 171. — /?. ? oxvcarpum {non K. Sch. & Laut.) Valet. Bull. D6p. Agr. Ind. Neerl. 10 (1907) 30. — Pocillaria pubescens Ridl. Trans. Linn. Soc. H, Bot. 9 (1916) 28. — R. bullatum Schellenb. Bot. Jahrb. 58 (1923) 169. — R. ohlongum Schellenb. I.e. 169. — R. elongatum Schellenb. I.e. 170. — R. ovale Schellenb. I.e. 171. — R. racemosum {nan Becc.) Schellenb. I.e. 168; Birnie, Nova Guinea 14, 2 (1926) 276. — Antidesma megalo- carpum Sp. Moore, J. Bot. 61 (1923) Suppl. 46. — R. pubescens (RiuL.) Sleum. Notizbl. Berl.- Dahl. 15 (1940) 250. -- Fig. 12. Sprawling shrub or small tree, 1-5 (rarely up to 15) m; bark greyish-brownish. Branchlets with laxly to subdensely strigose tips. Leaves oblan- ceolate, or oblong, or obovate-, sometimes elliptic- oblong, apex rather suddenly acuminate for 1-2 cm by 5 mm at base, tip acute or mostly bluntish, base cuneate to almost rounded, firmly charta- ceous to subcoriaccous (more stilV at higher al- titudes), usually olivaceous-grecn and somewhat shining on both faces in the dry state, glabrous except maybe some scattered apprcssed hairs on midrib and nerves of the undersurface, (6.5-) 8 17(-25) by (2.5-)3 7(-IO) cm, nerves 6 «( -10) pairs, curved-ascending and generally markedly looping, a little raised above, more distinctly so beneath, not rarely raised within a shallow de- pression above (the lamina not properly bullatc, as the veins are never sunken above too), re- ticulation rather dense, considerably or usually 40 Flora Malesiana [ser. I, vol. 7^ Fig. 12. Rhyticaryuin longifolium K. Sch. & Laut. a. Habit, c? specimen, X V2. i>. S flower bud, x20, c. petal from inside, x 20, d. ^ more developed flower, x 20, e. stamen, x 30,/. habit of ? specimen with fruits, X 1/2, g- fruit, x 1 1/2 (a-e BW \0666,f-g Van Royen & Sleumer 6136). 1971] ICACiNACEAE (Slcumer) 41 more slightly raised on both faces; petiole (4— )6-17 by c. 1.5 mm. — o Spikes solitary or sometimes few from the same axil, rather dense-flowered, (2-)4-10(-18) cm, in juvenile state all over pale rusty tomentose, in later stages densely short- strigose; rachis slender to more stoutish. Calyx cupular, 5-dented, low. Petals 4 or 5, connate halfway, yellow or greenish, sparsely to more densely strigose in the upper half outside, c. 2 rrmi. Stamens 4 or 5; filaments short. Rudiment of ovary with erect hairs. — 2 Spikes generally shorter than the o ones, and the rachis more stoutish. Calyx and petals as in o flowers. Ovary ovoid-conical, densely strigose. Drupe ovoid-el- lipsoid, much compressed laterally and crested. 2-2.3(-2.5) by 1.5(-1.7) by 0.8-1 cm, red to orange; endocarp hard, coarsely lacunose-reti- culate. Distr. NE. Australia (Queensland: Cape York Peninsula), Melanesia (Solomons); in Malesia: New Guinea. Ecol. Substage of primary-, sometimes also secondary forest, from lowland up to 1800 (-2500?) m, scattered. Uses. The leaves are cooked and eaten by the natives in the Solomon Is. Vern. Aicaruar, Wandammen, badzehkiziwo, Orokaiva, beniros, Wagau, hahamen, Kole, kame game, Kaugel, kamokum, Enga, namsi, Orne. In the Solomon Is.: aiavea, Kwara'ae. 7. HARTLEYA Sleum. Blumea 17 (1969) 218. — Fig. 13. Dioecious tree. Leaves spirally arranged, entire, coriaceous, penninerved, petioled. Inflorescences axillary, 3-5 peduncled cymes arranged to a panicle. — D Flowers unknown. — 2 Flowers subsessile, crowded at the apex of the ultimate branches of the inflorescence; bracts minute, caducous. Calyx cup-shaped, 4-6- lobed halfway, lobes valvate. Petals 4, valvate, smooth inside. Staminodes 4, of the habit of apparently normal stamina, but without pollen; filaments flattened, glabrous; anther cells elliptic, medifixed, the connective between not produced beyond them. Ovary with a lateral gibbosity; stigma sessile, subcapitate, papillose, clearly set off from the ovary, subbilobed or depressed to one side. Disk unilateral, free, opposite the gibbosity of the ovary, a semiorbicular low truncate or sub- crenate fleshy scale. Drupe curved, ovate-acuminate when seen from one side, exocarp thin, endocarp with numerous prominent nerves on the concave, and a thick median rib on the convex side, the latter covered with a fleshy appendage, the very apex excepted, and with a low semiorbicular fleshy free scale opposite the appendage, subtended by the remnants of the calyx. Seed not yet known. Distr. Monotypic; in Malesia: New Guinea. Ecol. Montane forest. Note. Closely allied to Gastrolepis Tiegh., a monotypic genus from New Caledonia, from which it differs by the subcapitate stigma, clearly set off from the ovary (in Gastrolepis rather small and on top of the gradually attenuate ovary), the large midrib on the concave side of the endocarp (which is not present in Gastrolepis), the squamular disk which is adnate only to the very base of the ovary, and otherwise free (adnate at base and both sides such as to form a pocket in Gastrolepis), and the unisexual flowers and g/a- brous filaments (bisexual and with numerous hairs in Gastrolepis). 1. Hartleya inopinata Sleum. Blumea 17 (1969) 218. Fig. 13. Tree, 20-35 m, spreading, 60-120 cm 0; bark grey-brown, fissured lengthwise. Branchlets rather short, irregular, early corticate, 3 8 mm 0. Leaves elliptic or oblong-clliptic, apex short-acuminate, or obtuse, base cuncatc to the petiole, subcqual, coriaceous, brittle and brownish-blackish in dry specimens, dark shiny green above, ycllowish- grccn or pale glaucous beneath in fresh state, gla- brous. 5-8 by 2.5 4.5 cm, edge a little rcvolutc, midrib slightly impressed above, bold beneath, nerves 5-6 pairs rather obscure on both faces, no reticulation; petiole 1 cm by 2 mm. Inflorescences panicled, branches (c. 1 cm) rather straight- horizontal, each ending with a cluster of cymosely arranged flowers, 3-4 cm, the short peduncle in- cluded, subdcnsely yellowish-pubescent especially apically. I lowers subsessile, 4 only known. ( alyx c. I mm. Petals oblong, glabrous, yellowish green. c. 3 mm. Stamens seemingly well-developed, but anther cells without pollen, I mm; filaments gla- brous, r. 3 mm. Dvary ovoid, glabrous, with a lateral gibbosity, and a fleshy subtruncatc semi- orbicular free scale. Drupe curved, ovate-acu- minate seen from one side, c. I cm by 4-5 mm; 42 Flora Malesiana [ser. I, vol. 7^ IT -69 Fig. 13. Hartleya inopinata Sleum. a. Habit, X I/2, b. flower bud, X 7, c. ditto, with 2 petals removed, X 7, d. developed ovary, showing the vertical fleshy appendix and the basal scaly disk, > 5, e. sterile stamen from adaxial side, e'. ditto, laterally seen, x7, /. submature fruit, adaxial view with large fleshy appendix,/', ditto, dorsal view,/", ditto, laterally seen, x3 {a-e Hartley 12501 (type), / Hartley 11836). exocarp thin, becoming blackish; endocarp with numerous longitudinal prominent nerves on the convex, and a strong mediane rib on the concave side, the latter almost completely covered with a fleshy flattened appendix, subtended at base by the lateral scale and the remnants of the calyx. Distr. Malesia: New Guinea (Morobe Distr., twice found; possibly present in the Vogelkop Peninsula (sterile specimens only known)). Ecol. Montane (also mossy) rain-forest, 1830-2400 m. 8. CODIOCARPUS Howard, Brittonia 5 (1943) 53, f. 2, 3 e-f; Sleum. Blumea 17 (1969) 188. — Fig. 14. 1971 ICACiNACEAE (Slcumer) 43 Dioecious tree. Leaves spirally arranged, entire, penninerved. Inflorescence axillary, generally solitary, the one-bracteate peduncle with several dichotomously branched distal branches, each branchlet with a several-flowered compact cyme of sessile small flowers, which are articulated at the base of the calyx. Calyx cup-shaped, shortly 5(-6)-dented. Petals 5(-6), valvate, apex minutely inflexed, shorter in the 2 flower. — 3 Flowers: Stamens 5(-6), exserted; filaments filiform, glabrous; anthers versatile, the cells oblong, slightly diverging below, dehiscing introrsally. Rudiment of ovary present. — $ Flowers: Staminodes 5(-6); filaments shorter than ovary; anthers much reduced in size, cells empty. Ovary subcylindric (upper part curved and contracted), with a lateral oblong swelling; stigma disk- like, obscurely 3-lobed. Drupe oblong, attenuate upwards, flattened, slightly curved; sarcocarp thin; endocarp thin, hard, with 3-5 (or more) ridges running the length of the convex surface, and a longitudinal mediane furrow, i.e. the funi- culus with a pair of woody ribs on either side of the funiculus on the concave surface, both funiculus and ribs surmounted by an oblong fleshy appendix extending over the whole length of the fruit, or even surpassing it a little. Seed 1, with copious endosperm and a minute embryo. Distr. Two very closely related spp., one in the Andaman and Nicobar Is., the other in Malesia: Philippines (Mindoro, Palawan). Fig. 15. Ecol. Lowland primary forest. 1. Codiocarpus merrittii (Merr.) Howard, Brittonia 5 (1943) 56, f. 2, 3 e-f (wood anat.); Dahl, J. Arn. Arb. 33 (1952) 267, f. 47 & 47 A (pollen); Sleu.m. Blumea 17 (1969) 189. — Ste- monurus merrittii Merr. Philip. J. Sc. 3 (1908) Dot. 240. — Tylecarpiis merrittii (Merr.) Sleum. Notizbl. Berl.-Dahl. 15 (1940) 237. — Medu- santhera merrittii (Merr.) Sleum. ibid. (1941) 364. — Fig. 14. Tree, 5-15(-30)ni, up to 25 cm 0, sometimes fluted. Branches horizontally spreading. Branch- lets and petioles (1-2 cm) shortly appressedly yellowish-pubescent, glabrate. Leaves oblong- elliptic to elliptic, apex shortly rather abruptly acuminate, tip obtuse, base broadly cuneate to rounded, sliglitly inequilateral and decurrent on the petiole, thin-coriaceous to almost charta- ceous, glabrous above, finely appressedly yel- lowish-hairy beneath, ultimately glabrate, lower surface often minutely tuberculatc in dry, ap- parently ycllowish-grecn in fresh specimens, with numerous fine pellucid points against strong light, 15-20(-28) by 5-10 cm, midrib grooved above, much raised beneath, nerves 8-10 rather straight to curved-ascending pairs, obscurely inarching before the edge, generally prominent beneath; petiole (1.5-)2-2.5 cm by 2-2.5 mm. Inflorescence with a rather thick peduncle 1 2.5 cm, 2-3- chotomous, the branches short, thickish, each with several cymosely and compactly arranged sessile flowers, all over covered with yellowish- greyish and : apprcssed hairs. Calyx c. 1.5 mm, lobes very short. Petals oblong, a little fleshy, white, hairy outside and at the thickened inflexed apex inside, otherwise glabrous, 4 5 mm in the J, r. 3 mm in the 1 flowers. J Flowers: /'ilaments ' 5 mm; anthers c. I mm. Rudiment of ovary elongate-conical, c. 2 mm. — 9 Flowers: Stamino- des a little shorter than the ovary; filaments weak; anther cells much reduced in size, empty. Ovary subcylindrical, glabrous. Drupe oblong, though narrowed gradually upwards, (1.8-)2-2.5 by 0.6-0.8 cm, greenish-whitish, with 3-5 prominent longitudinal and several oblique short ribs on the dorsal convex, and a creamy-white fleshy oblong appendage on the much deepened ventral surface. Distr. Malesia: Philippines (Mindoro, Pa- lawan). Fig. 15. Fig. 15. Distribution of the species of Medusan- ihera ( ) and Codiocarpus ( ). 1. A/. gracilis (KiNG) Sleum., 2. M. laxijhra (Miers) Howard, 3. M. samoensis (Reinecke) Howard and A/, ovata Howard, 4. M. viliensis Seem., 5. C. andamanicus (KuRZ) Howard, 6. C. merrittii (Merr.) Howard. Hcol. Dense lowland forest, under partial shade up to 200 m, on clayey soil or fertile humus. II. Jr. Jan. -Dec. Uses. Wood dense and hard, yellowish-white, used for house posts. Vern. Palawan: apitunif, kalas-kalas, panahd, tihalao, Tagb. 44 Flora Malesiana [ser. I, vol. 7^ Fig. 14. Codiocarpus merrittii (Merr.) Howard, a. Habit, x 1/2. b. flower bud, x7'/2, c. petal inside, x7'/2, d. anther, dorsal side, x 10, d'. anther, ventral side, x 10, e. stamen, xlYj,/. ovary, 71/2, g. drupe, dorsal side, xl, g'. drupe, lateral view, xl, g". drupe, ventral side with fleshy appendage, xl, /?. endocarp (exocarp removed) showing the two free ribs on the ventral side, x II/2, /. drupe in cross-section, showing the two free ribs of the ventral side covered by the appendage, x 1 1/2 («-/ Eimer 12622, ^-; Elmer 12842). 1971 ICACINACEAE (Sleumer) 45 9. MEDUSANTHERA Seem. J. Bot. 2 (1864) 74: Howard. J. Am. Arb. 21 (1940) 469; Lloydia 6 (1943) 133; Sleum. Blumea 17 (1969) 226. — Txlecarpus Engl, in E. & P. Nat. Pfl. Fam. 3, 5 (1893) 247; K. Sch. & Laut. F1. Schutzgeb. (1900) 413 {'Tilecarpus ); Sleum. Notizbl. Berl.-Dahl. 15 (1940) 246; in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 362. f. 103. A & B. — Fig. 16. Fig. 16. Medusanthera laxiflora (MiERS) Howard, a. Habit, X '/j, h. ^ flower bud, y.lVii c. stamen, dorsal view, x 10, c'. ditto, ventral view, X 10, d. drupe in dry state, dorsal view, showing the prominent ribs of the endocarp, /.lYz, d'. ditto, ventral view with large appendage, x2'/4, d". ditto, cross-section, showing a low rib on either side of the median depression of the ventral side covered by the appendage, X 2>/2 {a-c BSIP 61 Walker & White, d-d" BW 3686 Kalkman). Dioecious trees. Leaves spirally arranged, subdistichous, entire, penninerved. Panicles 1 -2(-3) per axil, 2-3-chotomized, the branches cymose, peduncled; pedicels short, articulated with the calyx. Calyx cupular, small, obscurely 5- toothed. Petals 5, valvate, lanceolate-oblong, apex inflexed. — c?/r;4^-» r.— --ti K/- \ * ^^^ 4 i-^ri-A Si3^ ^ ^^it^' t;;^"! 'i 1 !C7 i Fig. 18. Distribution of Apodytes. A. dimidiata E. Meyer ex Arn.: approximate area in Africa and SE. Asia, • localities in Malesia. X A. brachystylis F.v.M. Java (extreme western part). Lesser Sunda Is. (E. Sumba, Flores, W. Timor), N. Borneo (Mt Kinabalu, Mt Trusmadi), Philippines (Mindanao: Bukidnon), Moluccas (Batjan: Mt Sibela). Fig. 18. Ecol. Primary rain-forest, on steep slopes or in ravines, edge of stream, on rich maybe volcanic soil, rare, 800-2000 m. Fl. Febr.-Sept.,//-. April- Nov. Uses. The leaves are of a bitter and adstringent taste. Wood light brown or yellowish, used in Indo-China for cabinet work, described by Moll & Janssonius, Mikr. 2 (1908) 244, f. 107 {A. javanica). Vern. Ki badak, S, kakomba, Sumba, simcirbi- lalang, Batak. Note. The species is conceived here in a broad sense. The special form occurring in Malesia, described above, is almost matched by forms or varieties found in Africa and Madagascar. Pending a revision of the genus, it seems advisable not to give a proper name on the varietal level to the Malesian specimens. 11. MERRILLIODENDRON Kanffi. Bot. Mag. Tokyo 48 (1934) 920, f. 7; I.e. 956 (Japon.); Sleum. Blumca 17 (1969) 228. — Peekeliodendron Sleum. Notizbl. Bcrl.-Dahl. 13 (1937) 509. — Fig. 19. Shrub or generally tree. Leaves spiral, entire, penninerved. Flowers bisexual, arranged in lax cymes to elongate panicles, lhe.se 1-3 in foliate or defoliate axils, or from older branches resp. from trunk. Pedicels articulated with calyx. Sepals (3 )4 5, connate to a cup in the lower, free (and not imbricate) in the upper half. Petals (4-)5, free, valvate, tip indexed, villous inside, rellexcd in anthesis, early caducous. Stamens 5; filament dilated, short, glabrous; anthers elliptic, cells in- trorscly and longitudinally dehiscent, exceeded by the connective in form o{ subulate glabrous apiculus. Ovary ovoid, tapering to a thick-liliform style; stigma punctiform. Drupe large; cxocarp thin-coriaceous, smooth, early decaying; cndocarp first coriaceous and hard, becoming corky and irregularly -]-: longi- 50 Flora Malesiana [ser. I, vol. 7^ tudinally lacunose in its outer, and woody (not hollowed) in its inner part in later stages. Seed 1, large; endosperm copious, fibrous, starchy; embryo foliaceous. Distr. Monotypic, scattered over Micronesia, Melanesia, and E. Malesia. Fig. 20. 1. Merrilliodendron megacarpum (Hemsl.) Sleum. Notizbl. Berl.-Dahl. 15 (1940) 243; in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 366, f. 104 (fr.); Balgooy in Steen. & Balgooy, Pac. PI. Areas 2 (1966) 176 (distr. map); Sleum. Blumea 17 (1969) 228. — Stemonurus ? megacarpiis Hemsl. in Hook. Ic. PI. 24 (June 1895) t. 2398; Kew Bull. (June/ July 1895) 133; K. Sch. & Laut. F1. Schutzgeb. (1900) 414; SCHELLENB. Bot. Jahrb. 58 (1923) 164. J.T.'68 Fig. 19. Merrilliodendron megacarpum (Hemsl.) Sleum. a. Inflorescence, b & b'. infructescence, the drupes still covered with the thin-leathery exocarp, c. old weatherbeaten fruit, showing the spongious endocarp, d. ditto, transverse section, all x Yj (.a BSIP 2017, b BSIP 4827, b' NGF 27224, c-rfNGF 10925). — Mangifera xylocarpa Laut. Bot. Jahrb. 56 (1921) 354. — M. rotense Kaneh. Bot. Mag. Tokyo 48 (1934) 920, f. 7; I.e. 956 (japon.); J. Dep. Agr. Kyushu Imp. Un. 4 (1935) 359; Kaneh. & Hatus. Bot. Mag. Tokyo 54 (1940) 435, f. 82 & 83 (phot., fl. & fr.); Hosokawa, J. Jap. Bot. 16 (1940) 540; Merr. & Perry, J. Arn. Arb. 27 (1946) 324; Dahl, ibid. 33 (1952) 265 (pollen). — Peekeliodendron missionariorum Sleum. Notizbl. Berl.-Dahl. 13 (1937) 510. — Fig. 19. Shrub, or generally tree, (3-)6-15(-25) m, bole straight, sometimes fluted, up to 55 cm o; bark green-grey to light brown, smooth, shed in ir- regular soft flakes; blaze yellow with orange streaks. Leaves oblong to oblong-ovate, apex shortly attenuate, mostly blunt, rarely acutish, base broadly cuneate to rounded, thin-coria- ceous, glabrous, showing numerous minute pel- lucid points against strong light, (15-)20-30 by 7-11 (-16) cm, in suckers or young shoots up to 36 by 17 cm, entire, midrib strongly obtusely prominent beneath, nerves (8-) 10-1 2 curved- ascendent pairs faintly looping before the edge, veins obliquely transverse, reticulation rather in- distinct; petiole 1.5-2 cm. Panicles composed of short lateral cymes, laxly many-flowered, 1-3 in foliate or already defoliate axils, or from the wood of old branches and/or trunk, 7-20 cm, lower branches up to 9 cm, upper ones 2-4 cm; peduncle 2-5 cm, with a basal swelling, glabrous or sparsely appressedly and coarsely hairy, slender in anthesis, stout (2 mm 0) in fruiting stage as is the rachis. Pedicels slender and 4-6 mm in anthesis, stoutish and c. 1 cm in fruit. Sepals suborbicular, connate in the lower half, lobes 1-1.5 mm. Petals ovate-oblong, a little fleshy, dull cream or yellow to purplish, 3-4 by 1.5 mm, glabrous out- side, lb densely yellowish-villous inside. Stamens 2-3 mm; filaments c. 0.7 mm; connective hairy, exceeding the oblong anther cells (1.5 mm) by c. 0.8 mm. Ovary 2.5 mm. Drupe pendulous, ob- longoid to ellipsoid, rarely ovoid-oblongoid, slight- ly laterally or subquadrangularly compressed, (4-)6-10 by (2-)3-6 cm, exocarp coriaceous, whitish or yellow to purplish when fresh and al- ready fully ripe, becoming purplish to blackish in older stages or when dry, smooth, thin, early dis- soluted; endocarp remaining woody and massive in the inner part (1-2 mm), becoming corky or spongious and irregularly longitudinally grooved and/or lacunose-excavated in the outer part (3-5 mm) when exposed to sea-water. Seed starchy, 4-6 by 2-2.5 cm. Distr. Micronesia (Marianas: Rota & Guam; Carolines: Kusaie), in Melanesia scattered on New Ireland, New Britain, the Solomons, and the Santa Cruz Is. (Vanikoro I.); in Malesia: Philip- pines (once in Palawan), SE. Celebes (Muna I.), 1971 ICACiNACEAE (Slcumer) 51 Fig. 20. Distribution MerrilUodendron megacarpum (Hemsl.) Sleum. and scattered in Northern New Guinea. Fig. 20. Ecol. Generally coastal and lowland partly swampy rain-forest, at 3-30 m (on Japen I. said to occur at 700 m), on the inland edge of mangrove, on sandy or coral beaches, also on volcanic soil, in moist places, often near streams or in freshwater swamp forest, locally not rare, though apparently scattered in general. Fl. fr. Jan.-Dec. Disp. The corky or spongious part of the endocarp gives the fruit its power of buoy- ancy, and dispersal is obviously effected or facilitated by sea-currents and freshwater streams. Uses. Wood yellowish, straight-grained, ap- parently not used. For wood anatomy cf. Sleum. Notizbl. Berl.-Dahl. 13 (1937) 511. Fruit said to be edible. Vern. Celebes: pake-saukatibu, Mima; New Guinea: manggaresi, Japen (Ambai), semna wen, Sekoli (Kemtuk); New Britain: vabilii; New Ire- land: idalu, Pala, varunatun, Uquana; Solomon Is.: abii, aibo, aiebo, aiembuembe, sasa sti, Kwa- 12. CANTLEYA RiDL. Fl. Mai. Pen. 1 (1922) 436; Howard, J. Arn. Arb. 21 (1940) 479, t. 2, f. 1-7; Sleum. Blumea 17 (1969) 186. — Fig. 21-23. Tree. Leaves spirally arranged, entire, inconspicuously penninerved. Cymes axillary and solitary; peduncle completely covered with obtuse small thick spirallly arranged bracts forming alveoli, from the upper 2 or 3 of which rise short branches, these articulated with the peduncle and branched dichotomously, each branch bearing at the end a cluster of 3-5(-8) sessile bisexual flowers. Calyx thick-cu- pular, shortly subimbricately 5-parted. Petals (4-)5, valvate, oblong, with a longitudinal ridge and the apex inflexed inside. Stamens (4-)5; filaments fleshy, broadened upwards, with a dense clavate-penicillate pubescence distally; anther cells almost horseshoe-shaped, diverging at the base, introrsely and longitudinally dehiscent, affixed at the apex. Ovary elongate-conical; stigma a fleshy reversed cup; ovules 2, pendent from near the apex of the cavity. Fruit fusiform; exocarp thin, pulpy; endocarp rather thin though firm, fibrous-corky, extending as a thick longitudinal pad (similar to a second cavity) inside, finally with a lengthwise split from the base up to the middle. Seed 1. Distr. Monotypic; in Malesia. Fig. 24. 1. Cantleya corniculata (Becc.) Howard, J. Arn. Arb. 21 (1940) 479. t. 2, f. 1-7; Dahl, ibid. 33 (1952) 270. f. 61 & 61 A (pollen); Browne, For. Tr. Sarawak & Brunei (1955) 199; Smythies, Common Sarawak Trees (1965) 71; Sleum. Blumea 17 (1969) 186. — Platea corniculala Becc. Malesia I (1877) 117; Valet. Crit. Ovcrz. Olacin. (1 886) 254. — Urandra sp.. Foxw. Philip. J. Sc. 4 (1909) Bot. 492. 542, f. 47. — Urandra cor- niculala (Becc.) Foxw. Philip. J. Sc. 6 (1911) Bot. 179; Merr. En. Born. (1921) 356; Mead, Emp. For. J. 4 (1925) 95; Den Berger, Hand. 4c N. I. Wet. Congr. Batavia 1926 (1927) 401 (wood anat.); Foxw. J. Mai. Br. R. As. Soc. 5. 2 (1927) 339; Mai. For. Rec. 3 (1927) 173; ibid. 8 (1930) 13; BuRK. Diet. (1935) 2205; Hhyne. Nutt. PI. 1 (1950) 986. — C. johurica Ridl. Fl. Mai. Pen. 1 (1922) 436. — Stenionurus cornicii- latus (Becc.) Ridl. ibid. 5 (1925) 297; Burgess, Timbers of Sabah (1966) 320. — Mg. 21-23. Tree, 15-40m. trunk straight, slender, terete, 30-60(-l50) cm o, expanded at base, buttresses generally not distinct, rarely up to 150 by 50 cm; crown high, dense, with numerous small branches; bark grey or greenish to brownish, sometimes with brightly coloured patches, rather smooth, 52 Flora Malesiana [ser. 1, vol. 7^ Fig. 21. Cantleya corniculata (Becc.) Howard, a. Habit, x V2. b. peduncle with bracts, X I'/^, c. petal inside, x 10, d. stamen, front side, x 10, d'. ditto, back side, x 10, e. calyx and ovary, x 7|^ {a-e Endert 56 E. 2 P. 664). finely, shallowly and rather irregularly fissured lengthwise. Branchlets striate-angular when dry. Leaves elliptic to subobovate-, rarely almost oblong-elliptic, shortly rather abruptly sub- acutely acuminate, base cuneate to rounded, slightly inequilateral, coriaceous, glabrous, mi- nutely papillose beneath, drying dark brown, edge slightly revolute, 5-9(-13) by 3-4(-5) cm, midrib narrowly sunken above, strongly raised beneath, nerves 8-10 pairs, straight and parallel to each other, rather indistinct; petiole 8-13(-20)mm. Inflorescence 1-3.5 cm; flowers crowded at the end of short dichotomous branches, these 2 or 3 on top of a l-2(-3) cm long peduncle which is covered all over with short obtuse thickish bracts (each forming an alveole), sessile, small, greenish to white or yellowish, pale puberulent except the petals. Calyx c. 1 mm. Petals oblong, green, 4—5 by 2 mm, fragrant to almost fetid and of a bitter taste. Stamens c. 4 mm, finally a little exserted, caducous as are the petals; filaments 3^ mm, glabrous below, upper part penicillate by sub- clavate white hairs; anther cells c. 1 mm. Ovary glabrous, 2 mm. Fruit fusiform, more attenuate distally, first green, later brown to blackish, c. 3.5 by (0.6-)0.8-l cm; exocarp thin, pulpy, of an acid and adstringent taste; endocarp fibrous- corky, c. 1 mm, finally splitting from below on one side up to the middle, extending as a thick pad for the whole length inside. Seed 1, oblong, c. 2.5 cm. Distr. Malesia: Sumatra, Malay Peninsula (Johore, Singapore), Riouw & Lingga Arch., Banka, Borneo (not yet known N of the line Labuan-Tarakan). Fig. 24. Ecol. Drier parts of primary freshwater swamp forest, or drier hill forest, from sea-level to c. 300 m, apparently local and scattered, on marshy or sandy soil. Fl.fr. Jan.-Dec. Uses. Timber highly valued, much sought and marketed, heavy and hard, brownish-yellow, close grained, fragrant and therefore used as a sub- stitute for sandalwood, very durable in the ground. Used for house and ship building, heavy con- structions, likely to shrink and warp and suitable for furniture or turnery only if fully seasoned. The fruit can be eaten, but is said to be of a rather poor quality. Fig. 22-23. 1971 ICACINACEAE (Slcumer) 53 J^ll^p n ^ f^' ■■■■ ,._^»;, 7 1 i »MS«i-.. , on Banka known as mendarii; Borneo: kaddo, mirore, Dajak, kakal, Sampit, penipok, Iban, sainala, Brunei (Kedayan),5iV««rt/f/!"/irfa/7a, Berau. Fig. 23. Cantleya cornicuUita (Becc.) Howard. Stem-base. S. Sumatra (Palembang) (Photogr. Endert). Fig. 22. Cantleya corniculala (Becc.) Howard. Tree-shape, with buttresses. S. Sumatra (Palem- bang) (Photogr. Endert). Vcrn. Generally known as hi'daru, darn, dr'daru, I'ndaru, ffaru, pi'daru, (enipilai, M; local names in Sumatra: f(aru bi'lina, f(. huaju, larai pahanx, Indragiri, tulang, Kedongong, lusaiv; Fig. Distribution of Cantleya corniculala (Becc.) Howard. 13. NOTHAPODYTKS Bi.. Mus. Hot. Lugd.-Bai. 1 (1850) 248; Howard, J. Arn. Arb. 23 (1942) 66; Si.F-UM. Blumca 17 (1969) 232. — Mappia Jacq. sect. Trichocratcr MiERS, Ann. Mag. Nat. Hist. II, 9 (1852) 395, rcpr. Conlr. Bot. 1 (1852) 64. — Mappia Jacq. 54 Flora Malesiana [ser. I, vol. 7^ sect. Nothapodytes (Bl.) O. Ktze in Post & Ktze, Lex. Gen. Phan. (1904) 350. — Neoleretia Baehni, C. R. Soc. Phys. Hist. Nat. Geneve 53, 1 (1936) 33; Can- dollea 7 (1936) 177. — Fig. 25. /f^« Fig. 25. Nothapodytes montana Bl. a. Habit, x 1/2, b. inflorescence, x 1, c. flower bud, x 5, d. petal, x 5, e. ovary, v.lViJ- stamen, adaxial side, Y-lViJ'- ditto, dorsal side, X71/2 (^-/Kostermans & Kuswata 53). Trees or treelets. Branches usually angular, with large leaf-scars. Leaves spirally arranged or partly (the upper ones) subopposite, entire, appearing very finely pellucid-punctate against strong light, penninerved. Flowers bisexual, in terminal, rarely axillary cymes collected to corymbs, articulate with the short ebracteate pedicels. Calyx cupular, 5-lobed or sinuate-dented to about 1/3, persistent. Petals 5, valvate in bud, linear-oblong, a little cohering at base, apex slightly inflexed, short-strigose without, villose or rarely almost glabrous within. Stamens 5, free, zb as long as the petals; filaments fleshy, somewhat flattened below, more slender distally, glabrous; anther cells subovate- to elhptic-oblong, diverging at base, in- trorse, with a basal abaxial reflexed pulviniform appendage ± cohering to the filament, medifixed; connective as long as the cells. Disk cup-shaped, shallowly 5- or 10-crenulate, fleshy to foliaceous, glabrous without, generally with some hairs within and on the edge. Ovary symmetrical, hirsute, ovoid to subconical, tapering to a short stoutish or more slender, shghtly elongate, hairy or glabrous (apparently shghtly dimorphic) style; stigma subcapitate, truncate or mostly 1971 ICACINACEAE (Slcumer) 55 oblique, sometimes slightly bilobed or concave. Drupe ±. ellipsoid, exocarp thin, juicy; endocarp thin-crustaceous, smooth or rugose. Seed 1, albuminous; embryo large, almost as long as the albumen. Distr. Yonr spp., 3 of which in S. India, Ceylon, Assam, Burma, Thailand, Cambodge, Tonkin, SW. to SE. China incl. also Hainan, on Lan Yu. I. {=-- Bote! Tobago or Kotyosyo) SE. of Formosa, Ryu Kyu Is. (Yaeyama group); in Malesia 2 spp., of which N.foetida has a wide distribution in SE. Asia, and A^. moniana is very close to A', pittosporoides (Oliv.) Sleum. in SW. and Central China. Fig. 26. Ecol. Primary and secondary rain-forest, also mixed deciduous forest, from sea-level to c. 2000 m. KEY TO THE SPECIES 1. Leaves ± unequal at base; petiole (2-)3-6 cm. Petals densely to more sparsely whitish-villose within. 1. N. foetida 1. Leaves — equal at base; petiole 1-2 (rarely -2.5) cm. Petals glabrous or with few hairs in the lower half within 2. N. montana 1. Nothapodytes foetida (Wight) Sleum. Notizbl. Berl.-Dahl. 15 (1940) 247; Howard, J. Arn. Arb. 23 (1942) 70; Dahl, ibid. 33 (1952) 273 (pollen); Sleum. Blumea 17 (1969) 232 (with extra-Ma- lesian synonyms and literature); Fl. Thail. 2 (1970) 82. — Stemonurus foeiidus Wight, Ic. 3 (1845) t. 955. — Neoleretia foetida (Wight) Baehm, Candollea 7 (1936) 177, t. 4, f. 2, D-F. Tree or treelet, (5-)IO-15m, 10-20 cm 0. Branchlets conspicuously angled, early covered with straw cork (wrinkled in dry specimens), large leaf-scars, youngest parts laxly to densely covered with yellowish appressed — substrigose or more softer hairs as are the petioles, the undersurface of the leaves, the inflorescences and the fruit, the tomentum however, variable in density or fading early. Leaves variably elliptic-ovate to -oblong, apex short-acuminate, base broadly cuneate to rounded, unequal, membranous (and subglabrous in the Philippines), or chartaceous (and tomen- tulose beneath in Sumatra), 9-22 by 4-12 cm, nerves (5-)6-IO(-l2) pairs rather steeply curved- ascendent, raised beneath, reticulation ± ob- scure; petiole slender to stoutish, (2-)3-6 cm. Corymbosely arranged cymes long peduncled, many-flowered, 5-10 cm across, peduncles and axes — flattened or angled, all over crisped- to sub- strigose-hairy. Pedicels I (-2) mm. Calyx c. 1.5 mm mcl. the short subacute lobes. Petals oblong- linear, subacuminate, white, densely to more sparsely hairy within, 4-5(-6) by c. 1.5 mm. Filaments (2-)3-4 mm, apparently accrescent during anthcsis; anther-sacs 1-1.3 mm. Ovary ovoid to pear-shaped, hirsute, gradually narrowed to the stouter or more slender, short or up to 3 mm elongate style, which sometimes is com- pletely glabrous. Drupe oblong-ovoid or ellipsoid, 1-2 by 0.8-1 cm, black, generally short-pubes- cent; endocarp thin-woody, shallowly rugose to lacunosc-rcticulatc; endosperm with a fetid odor. Distr. S. India, Ceylon, Assam, Burma, Cam- bodge. Thailand, Lan Yu I. (SE. of Formosa), Ryu Kyu Is. (Yaeyama group); in Malesia: W. Sumatra (once on Mt Sago near Pajakumbuh), Philippmcs (N. Luzon: llocos Norte & Isabela \*Tos .). Fig. 26. -v-vx^:- Fig. 26. Distribution of Nothapodytes. N. foetida (unbroken line and dots), N. montana (squares), N. ohtusifolia (...), A', pittosporoides (- . - . -). Ecol. Rain-forest, on Mt Sago at c. 1000 m, in the Philippines between 100 and 300 m (out- side Malesia up to 2000 m, occurring also in mixed deciduous forest). Fl. Febr. -March, fr. June (Malesia). 2. Nothapodytes montana Bl. Mus. Bot. Lugd.- Bat. I (1850) 248; Sleum. Notizbl. Berl.-Dahl. 15 (1940) 247; Howard, J. Arn. Arb. 23 (1942) 67; Amsh. in Back. Bekn. Fl. Java (cm. ed.) 6 (1948) fam. 135. p. 6; Back. & Bakh. /. Fl. Java 2 (1965) 61; Sleum. Blumea 17 (1969) 233; PI. Thail. 2 (1970) 83. — Mappia montana (Bl.) MiERS. Ann. Mag. Nat. Hist. II, 9 (1852) 398, repr. Contr. Bot. I (1852) 67; Miq. Fl. Ind. Bat. I (1856) 790; Suppl. I (I860) 136; Becc. Malesia I (1877) 118; Valet. Crit. Ovcrz. Olacin. (1886) 183, I. 5. f. 28 (fl.); K. & V. Bijdr. 5 (19(X)) 143; Ba(K. Schoolfl. Java (1911) 226; Koord. Exk. Fl. Java 2 (1912) 532. Fljj. 25. 56 Flora Malesiana [ser. I, vol. 7^ Tree or treelet, 5-8(-20) m, 10-15(-20) cm 0; white, glabrous or with few hairs in the lower half bark smooth, light brown. Branchlets slender, within, 4-5 by 1.5-2(-3)mm. Filaments linear obtusangular, early covered with pale brown cork below, filiform above, 3-4 mm; anther cells el- and wrinkled in dry specimens. Leaver lanceolate liptic, 1.5 mm. Ovary ovoid, gradually tapering to obovate-oblong, apex obliquely acuminate (1-2 to a shorter or more slender style (c. 2 mm) cm), subacute, base cuneate, ± equilateral, her- which at least in its lower half is appressedly hairy baceous to chartaceous, shining above, practically as is the ovary; stigma oblique. Drupe ellipsoid, glabrous, often laxly tubercled beneath, 8-16 greenish-brown when fresh, blackish when dry, by 3-6.5 cm, midrib raised beneath, nerves in rather smooth, c. 1.5 by 1cm. 6-9 pairs, curved or more strict, widely patent, Distr. Thailand (Prachinburi); in Malesia: rather indistinctly looping before the edge, re- Sumatra (Eastcoast: Karo Highlands), extreme ticulation of veinlets rather dense, finely raised on W. and SW. Java, Lesser Sunda Is. (W. Sum- both faces; petiole slender, 1-2 (rarely -2.5) cm. bawa, W. Flores). Fig. 26. Corymbs rather few-flowered, 2-4 cm across, pe- Ecol. Primary and secondary forest, almost duncle slender, 2-4 cm, densely appressedly yel- from sea-level to c. 1650 m, also on coral lime- lowish hairy in all outer parts. Calyx 1.5-2 mm stone. F/. Jan.-Aug.,//-. April, incl. lobes. Petals oblong-linear, subacuminate Vern. Kihadji,S. distally, finally recurved, a little fleshy, greenish- 14. STEMONURUS Bl. Bijdr. (1826) 64S, p.p. lectotyp.; Hassk. Cat. Hort. Bog. (1844) 214 CStemo- miray, Bl. Mus. Bot. Lugd.-Bat. 1 (1850) 249, /?./?.; em. Becc. Malesia 1 (1877) 108, 109, 111; Sleum. Blumea 17 (1969) 255. — Urandra Thw. in Hook. J. Bot. Kew Misc. 7 (1855) 211. — Lasianthera (non P. Beauv.) Miq. F1. Ind. Bat. 1 (1856) 790, p.p.; King, J. As. Soc. Beng. 64, ii (1895) 116. — Fig. 28-31. Trees, sometimes with buttresses or with pneumatophores, glabrous except a fine subglandular puberulence on the inflorescences, the petals excluded. Branch- lets and buds often vernicose. Leaves spiral, entire, coriaceous to various de- grees, dark green to yellowish green above, paler beneath, sometimes tubercled on the undersurface, midrib narrowly sunken above, bold beneath, nerves pinnate, looping; petiole robust. Umbels axillary and generally solitary, peduncled, each branch of the umbel bearing the flowers either distally and crowded, or uni- or biseriately (scorpioid) along its upper, mostly zh recurved part; bracteoles at the base of the umbel conspicuous, sometimes persistent. Flowers bisexual, sessile, white to yellowish, fragrant. Calyx cup-shaped, base i truncate, limb truncate or 5-lobed to various degree, persistent. Petals (4-)5, valvate in bud, connate at base only, oblong, apex inflexed, midrib or keel often raised on both faces, ca- ducous. Stamens (4-)5; filaments fleshy, flat, widened upwards, shortly bearded be- low the anther on the ventral side, and long-penicillate distally on the backside, the penicilli inflexed in bud, at full anthesis erect and much exserted; cells ovate- elliptic, basifixed, a little diverging downwards. Disk rim-like or patellar, or mostly a short, whether or not shallowly lobed cup, surrounding the very base of the ovary, the latter ovoid-conical much attenuate towards the very short slender style and the punctiform stigma, unilocular, with 2 pendant ovules. Drupe ovoid, ellipsoid or oblongoid-fusiform; exocarp thin, the lower 1/3 part dark red to purple, the upper part white to cream or greenish; endocarp firm, coriaceous to woody, its outer part fibrous. Seed 1; embryo small, in the top of the albumen. Distr. About 12 spp., 2 of which in Cochinchina and Annam, 1 in Ceylon; in Malesia 9 spp., one of which extends to the Solomon and Palau Is. Fig. 27. Ecol. In dry hilly land or in peat swamp forest (part of the species apparently restricted to the latter), mostly scattered, though widespread in lowland, rarely up to 1750 m. 1971] ICACINACEAE (Sleumer) 57 Disp. The ripe fruits with their fibrous outer and hard inner endocarp are apparently able to float. This may explain the wide area of distribution of 5. ammiii (Palau Is., northcoast of New Guinea, New Britain, Solomon Is.) which occurs both in lowland hillside and in coastal swamp forest, even in the mangrove. Uses. Wood aromatic, not durable and no commercial use known in Malesia (Endert, Med. Proefst. Boschw. Ned. Ind. 20, 1928, 198, 199, 201). Note. The genus comprises two groups of species, one with uni- or biseriately arranged flowers around 5. secundiflorus Bl. (\V. Malesia incl. Celebes), the other with flowers crowded at the end of the branches of the umbel around S. umbellata Becc. (W. to E. Malesia. Solomon and Palau Is.). Within these two groups the species must be distinguished mainly by calyx and fruit characters, as their leaf and other flower characters overlap to rather great extent. KEY TO THE SPECIES 1 . Flowers uni- or biseriate along the whole or at least the upper (in S. grandifoUiis thickened) part of the branches of the umbel, generally 5-12 per branch. 2. Bracts at the base of the umbel persistent for a while during anthesis, (3-)4-5 by c. 1 mm. 3. Leaves (18-)26-36 by 7-ll(-13) cm, very coriaceous, generally a little shining above only in the dry state, nerves 15-20 pairs generally very slightly or hardly raised on both faces, veins — obscure. Drupeellipsoid-ovoid, 2-2.4by 1.2-1.5cm 1. S.grandifolius 3. Leaves ll-23(-30) by 6-9(-ll)cm, coriaceous, generally shining on both faces in the dry state, nerves 1 5-20 pairs slightly though markedly raised on both faces as are the rather lax veins. Drupe ovoid-ellipsoid, 4.5-5. 5(-6.5) by 2-2.5(-3) cm 2. S. scorpioides 2. Bracts at the base of the umbel early caducous, 1-2 by c. 0.5 mm. 4. Inflorescences robust, peduncle c. 3 mm o. Calyx shallowly lobed or almost entire. Petals 5-6(-7) mm. Drupe ovoid to oblong-ovoid, 2.5-3(^) by (1.6-)2cm 3. S. celebicus 4. Inflorescences relatively slender, peduncle 1-2 mm 0. Calyx lobed ± halfway. Petals c. 4 mm. Drupe oblong-ellipsoid, 4.5-5.5 by 1-1.5 cm 4. S. secundiflorus 1. Flowers clustered at the end of the branches of the umbel or even at the end of the peduncle when the branches of the umbel are very short. (Bracts at the base of the umbel generally small, fugacious or rarely subpersistent). 5. Drupe subovoid-oblongoid to almost fusiform, at least 2 times as long as (or mostly longer than) across. (Leaves generally smooth on the undersurface, maybe except some tubercles found close to and along the midrib). 6. Calyx rather deeply obtusely lobed. Petals (2.5-)3(-4) mm. Disk cup-shaped. Drupe 4.5-5 by (l-)l.5-2.2 cm 5. S. malaccensis 6. Calyx shallowly lobed. Petals (4-)5 mm. Disk low, almost patellar. Drupe (4.3-)4.5-5 by 1.5-1.7 cm. 6. S. ammui 5. Drupe ovoid to ellipsoid-ovoid, up to c. 2 times as long as across. 7. Leaves practically smooth on the undersurface; petiole (2-)2.5-3 cm. Calyx rather distinctly lobed. 7. S. gitingensis 7. Leaves generally rather ± densely tubcrcled all over the undersurface or at least in a broad stripe along the midrib; petiole 1-2 (very rarely up to 3) cm. 8. Calyx ± distinctly obtusely lobed. Petals 3(-4, very rarely -4.5) mm. . . 8. S. monficolus 8. Calyx truncate or very shallowly (apiculately) lobed. Petals (4-)5(-6) mm. . 9. S. umbellatus 1. .Stemonurus grandifolius BEf;c. Malesia 1 (1877) (18-)26-36 by 7-ll(-13) cm, the extreme margin 1 14. t. 5, f. 1; Valet. Grit. Overz. Olacin. (1886) slightly and narrowly revolutc, nerves 15-20 pairs, 235; Merr. En. Born. (1921) 356; Sleum. Blumea generally very slightly raised on both faces, veins 17 (1969) 257. — Urandra grandifolia (Becc.) 4- obscure; petiole rugulose, 1. 5-2(-3) by 0.3-0.5 O. Ktze. Rev. Gen. PI. 1 (1891) 113. — 5. doli- cm. Peduncle of the umbel 3-5 cm by 2-3 mm, chophyllus Merr. Un. Calif. Publ. Bot. 15 (1929) with 4-8(-12) shorter or longer (r. 10 mm) robust 171. — Urandra dolichophylla (Merr.) Howard. and a little flattened branches, which bear 3-4(-5) J. Arn. Arb. 21 (1940) 468; Dahl. ibid. 33 (1952) flowers crowded at the thickened end of each 270 (pollen). branch, or. as normally more (up to 8) flowers Tree. (7-)25-30(-40) m, bole (6-)10-15m, arc present, these close together and uni- or 10-30(-40)cm 0; crown dense; bark greyish to biseriate on the recurved and thickened upper brownish, smooth or superficially fissured, yellow- part of the respeclive branch. Bracteoles at base of brown mottled. Branchlcts robust (5-10 cm o). the umbel broad-subulate, (3 )4-5 by I mm, ± often finely wartcd and vcrnicosc. Leaves Ian- persistent. Calyx cup-shaped. 2 mm, lobed to Vy. ceolate or elliptic-oblong, apex very shortly acu- Petals c. 5 by 2 mm, light yellow or white. Drupe minatc, or merely • abruptly apiculatc. base ellipsoid-ovoid, apex gradually attenuate, obtuse, broadly cuneate to rounded, very coriaceous, stiff. 2-2.4 by 1.2 -1.5 cm. a little shining above in the dry state, dull beneath, Distr. Malesia: Borneo. 58 Flora Malesiana [ser. I, vol. 7^ Fig. 27. Distribution of Stemonurus. The figure above the hyphen indicates the number of en- demic species in the island or area, the figure below the hyphen the number of non-endemic species. Ecol. Primary lowland (Dipterocarp) forest, on hills or low ridges, not rare on sandstone or sandy soil, or coastal kerangas, never in peat swamp, from sea-level-100(-680, a form with narrower leaves in Tristania forest on Mt Kina- balu at 1220) m. Fl. fr. Jan.-Dec. Uses. The pale brown wood is apparently not used. Vern. Baru tiias, Bulungan, kambong, Kina- batangan, kapal-kapal, Kedayan, katak, Dusun, inersaut, Kutei. 2. Stemonurus scorpioides Becc. Malesia 1 (1877) 113, t. 6; Valet. Crit. Overz. Olacin. (1886) 234; HocHR. Bull. Inst. Bot. Btzg 22 (1905) 6 {var.y, Merr. En. Born. (1921) 356 Cscorpiurus'); Ridl. Fi. Mai. Pen. 5 (1925) 296; Burk. Diet. (1935) 2074; Heyne, Nutt. PI. 1 (1950) 986; Anderson, Gard. Bull. Sing. 20 (1963) 166; Sleum. Blumea 17 (1969) 257. — Urandra scorpioides (Becc.) O. Ktze, Rev. Gen. PI. 1 (1891) 113 Cscorpiodes'): Dahl, J. Arn. Arb. 33 (1952) 270, f. 58 & 58 A (pollen). — Lasianthera secundiflora [non (Bl.) MiQ.] King, J. As. Soc. Beng. 64, ii (1895) 177, p.p. — S. secundiflorus {non Bl.) Ridl. Fl. Mai. Pen. 1 (1922) 430, p.p. — Fig. 28. Tree 10-30(-35) m, clear bole up to 24 m and 15-30(-60) cm 0; bark thin, whitish to greyish brownish, smooth or pustular. Blunt asparagus- like pneumatophores sometimes present. Branch- lets robust, often vernicose as are the buds. Leaves elliptic-oblong, apex shortly and ir abruptly acuminate, or apiculate, tip bluntish, base broadly cuneate, coriaceous, stiff", dark green above, paler beneath, glossy on both faces also in the dry state, the very margin often slightly re- volute, ll-23(-30) by 6-9(-ll) cm, nerves 15-20 subregular pairs, slightly though distinctly raised on both faces as are the rather lax veins, veinlets finely and prominently reticulate above only; petiole 1.5-2 cm by c. 3 mm. Scorpioid cymes 4-6 in an umbel on a robust 3-5(-6) cm long axillary peduncle, branches of the umbel 3-5 cm, each with 6-10 sessile flowers biseriate on each side; bracts at base of umbel several, subulate, Fig. 28. Stemonurus scorpioides Becc. Seedling, a. with the first leaves, b. preceding stage with the cotyledons still enclosed by the longitudinal fibres of the endocarp, both x 1/3 (KEP 99938). rather persistent, (3-)4-5 by 1 mm. Calyx cup- shaped, 1.8 mm, obtusely lobed in the upper spreading W->,. Petals (4-)5 by 1.5 mm, white to cream or greenish. Drupe ovoid-ellipsoid, obliquely attenuate towards the apex, 4.5-5.5(-6.5) by 2-2.5(-3) cm, whitish above, dark green- purplish in the lower third, with numerous slight vertical grooves, able to float when dry. Distr. Malesia: Sumatra (incl. also Simalur I.), Lingga and Riouw Arch., Banka, Malay Peninsula, Borneo. Ecol. In Shorea albida peat swamp forest in Borneo, generally in maybe seasonally swampy, also in secondary vegetation, on peaty or sandy soil, locally frequent, in lowland up to 45 m. Fl.fr. Jan.-Dec. Uses. Timber soft, not durable, cracking. 1971] ICACiNACEAE (Slcumer) 59 locally used for planks and household utensils. Pounded bark used as medicine. The fruit is said to be edible. Vern. S. scorpiuides has remarkably numerous native names in the Malay; part of them, how- ever, are used also for other species of the genus, and the name bidant is commonly in use for Cantleya corniculata (Becc.) Howard outside North Borneo. — Sumatra: kaju longgalui, k. pasir, sitobal, Eastcoast, sebungku, sibunik, P. Bengkalis, sibungkiik, Indragiri, setebal, Djambi, bahuhu-etim, Simalur, hanol, Banka, darn darn dauri lebar, P. Singkep; Malay Peninsula: biingkok, peripat paya, Pahang; batang torong, Manggu; Borneo: ampasir, W. Borneo, bedaru, hantii burok, katok; iiiedang katiik, m. telor (Benuni); N. Borneo & Brunei, Sarawak: entaburok, Iban, jenimii, Milanau, katop, Dusun, semburok; kaju malam, Kuching, posir-pasir, Pontianak. 3. Stemonurus celebicus Valet, in Koord. Minah. (1898) 394; Ic. Bog. 1 (1901) t. 88; Hochr. PI. Bog. Exs. (1904) 45; Bull. Inst. Bot. Btzg 22 (1905) 48; Sleum. Blumea 17 (1969) 258. — Urandra celebica (Valet.) Howard, J. Arn. Arb. 21 (1940)468. Tree, 20-30 m tall, up to 45 cm 0. Branchlets robust, smooth or finely rugulose, vernicose, 5-12 mm 0. Leaves elliptic to obovate-elliptic, apex broadly subacuminate-attenuate or obtusely apic- ulate, sometimes rounded, base broadly cu- neate, coriaceous, a little glossy above, dull be- neath when dry, (10-)12-16(-22) by 5-8(-10.5) cm, edge somewhat revolute in dry specimens, nerves 10-14(-16) erecto-patent pairs, slightly or hardly raised on both faces, veins + obscure; petiole 1.5-2 cm by c. 2.5 mm. Umbels solitary, robust, on peduncle 3-5 cm by 3 mm, branchlets of the umbel (4-)5-7, more slender, though thickened distally, (2-)3^ cm, their basal bracts ovate, obtuse, 2 mm, early caducous. Flowers sometimes only 3 or 4 and crowded at the apex, generally 5-6(-8), close together and biseriate in the upper half of each branchlet of the umbel, secund. Calyx broadly cup-shaped, 2 mm, shal- lowly lobed or almost entire. Petals 5-6(-7) by 2.5 mm, keeled, white-greenish. Drupe ovoid to oblong-ovoid, apex gradually attenuate, obtuse, 2.5-3(-4) by (l.6-)2cm, darker in the lower 2/3. Seed oblong, a little flattened. Distr. Malesia: NE. and Central Celebes. Ecol. Primary forest, rarely in swamps, 20-1000 m. Fl. April-July, fr. June-Nov. Vern. Bilulang, Luwu, palonlanggasl, Padu, simhelan, Minahasa (Tl.), simpuga lanru, Bugin., woliauMi, Tobcla. 4. .Stemonurus secundiMoru-s lii . Bijdr. (1826) 649; Walp. Rep. I (1X42) 378; Hassk. Cat. Hort. Bog. (1844) 214; Bi.. Mus. Bot. Lugd.-Bat. I (1850) fig. 45; .MiER-S. Ann. Mag. Nat. Hist. II, 10 (1852) 36. rcpr. Contr. Bot. I (1852) 86; Schni/i . Ic. 3 (1866 70) t. 223; Becc. Malesia 1 (1877) 112, t. 4. f. 16-17; Valet. Crit. Ovcrz. Olacin. (1886) 234; K. & V. Bijdr. 5 (1900) 145; \\<>< mh I'I, Hog, Exs. (1904) 46, incl. var. valetonii Hochr.; Bull. Inst. Bot. Btzg 22 (1905) 41, incl. var. valetonii Hochr.; Moll & Janss. Mikr. 2 (1908) 234, f. 105 (wood anat.); Back. Schoolfl. Java (1911) 226; Koord. Exk. Fl. Java 2 (1912) 531; Koord. - ScHUM. Syst. Verz. 1, Fam. 162 (1912) 3; Koord. Atlas 1 (1913) t. 121; Ridl. Fl. Mai. Pen. 1 (1922) 430, p.p.; Baker/. J. Bot. 62 (1924) Suppl. 21; RiDL. Fl. Mai. Pen. 5 (1925) 296; Burk. Diet. (1935) 2074; DocT. v. Leeuwen, Ned. Kruidk. Arch. 51 (1941) 175, f. 48 (gall); Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 5; Back. & Bakh. /. Fl. Java 2 (1965) 60; Sleum. Blumea 17 (1969) 258. — Croton laurifolius Nor. Verh. Bat. Gen. 5 (1790) ed. I Art. IV, 13, nom. niid. — Lasianthera secundiflora (Bl.) Miq. Fl. Ind. Bat. 1 (1856) 792, incl. var. sumatrana MiQ. I.e. (1858) 1097; Suppl. 1 (1860) 137, incl. var. sumatrana; ibid. (1861) 342, incl.f. sumatrana Miq. I.e., t. 2; King, J. As. Soc. Beng. 64, ii (1895) 177, p.p. — Urandra secundiflora (Bl.) O. Ktze, Rev. Gen. PI. 1 (1891) 113; Engl, in E. & P. Nat. Pfl. Fam. 3, 5 (1893) 248; Heyne, Nutt. PI. 1 (1950) 987. — 5. pauciflorus Ridl. Kew Bull. (1926) 61, nan Bl. 1826; Howard, J. Arn. Arb. 21 (1940) 481, in te.xt. — S. ridlevanus Sleum. Notizbl. Berl.-Dahl. 15 (1940) 243. KEY TO THE VARIETIES 1. Leaves (subcoriaceous to coriaceous) generally oblong to elliptic- or obovate-oblong, nerves (8-)IO-12(-14) pairs, i distant from each other, slightly or hardly raised, veins ± obscure. 1. var. secundiflorus 1. Leaves with 14-16(-18) pairs of nerves, these i close together and generally a little raised on both faces as are the lax veins. 2. Leaves elliptic, rigidly coriaceous. 2. var. hosei 2. Leaves lanceolate-oblong or -elliptic, co- riaceous 3. var. lanceolatus 1. var. secundiflorus. Tree, 10-24(-32) m, trunk cylindric, up to 40 cm 0, sometimes with pneumatophores in swampy places; bark rather smooth, whitish grey-green. Branches t horizontal. Tips of branch- lets and buds vernicose. Leaves rather variable in shape and size, oblong-obovate, sometimes ellip- tic-oblong or oblong, apex obtusely apiculate or short-acuminate, base acutely or more rarely ob- tusely cuncatc, subcoriaceous to ;;oriaccous, shining above, much paler beneath, (6-) 10- 1 7 (-20) by (3-)4-8.5cm. nerves (8-)IO-l2(-l4) slender and ! distantly subparallcl, rather in- distinctly inarching pairs, slightly or hardly raised on both faces, veins i obscure; petiole l.5-2(-3) cm. Cymes arranged in an solitary umbel, rarely in two umbels per axil, peduncle rather slender, c. 3 cm by 1-2 mm, branchlets of the umbel 3-5, slender, 1 recurved. 2-2.5 cm, bearing in the upper part 3-5(-7) flowers close together in a row on each side; bracts at base of the umbel ovate or oblong, 60 Flora Malesiana [ser. I, vol. 7^ minute, early caducous, leaving blunt scars. Calyx cupuiar, 2-2.5 mm, obtusely lobed almost half- way. Petals c. 4 by 1.5 mm, white or yellowish, fragrant. Drupe oblongoid-ellipsoid, attenuate towards the bluntish apex, 4.5-5.5 by 1-1.5 cm. Distr. Malesia: Sumatra (incl. also Simalur, Mentawei & Enggano Is.), Banka, Biiliton, Malay Peninsula, W. Java. Ecol. Rain-forest, 30-700(-1000) m, sometimes in peat swamp forest or seasonal swamp forest near sea-level, scattered. Fl. fr. Jan.-Dec. Uses. Timber light brown, of inferior quality, used in places under the house-roof. Medicinal use for the cure of swellings doubtful. Seeds have a bitter taste and are edible only after repeated cooking. Vern. Sumatra: lokan, Westcoast, sebungku, ^SiiicodiSi, ganggaram, P. Bengkalis, /carj/r, sebonku, si benjiet, Indragiri, schring, Bencoolen, bahiihu uding, suwahar uding, Simalur, mangguguk, Mentawei, kaju bemban, Enggano, gelam kataju, Minangkabau; saber bubu, Banka, krikis ayer, Biiliton; Malay Peninsula: berlat, Pahang, M, kalapa siong, kimeong, sintok hintjang, S. 2. var. hosei Sleum. Blumea 17 (1969) 259. — S. labuanensis Staff, Kew Bull. (1906) 71, p.p.; Merr. En. Born. (1921) 356, p.p. Leaves elliptic, very shortly acuminate, stiff- coriaceous, nerves close together and numerous (c. 18 pairs), slightly prominent on both faces as is the net of veins. Inflorescence as in var. lanceo- latus. Branchlets of the umbel 3 or 4. Fruit not known. Distr. Malesia: Borneo (Baram R. area in Sarawak), once found. 3. var. lanceolatus (Becc.) Sleum. Blumea 17 (1969) 259. — S. lanceolatus Becc. Malesia 1 (1877) 114, t. 5, f. 2-4; Valet. Crit. Overz. Olacin. (1886) 235; Merr. En. Born. (1921) 356. — Urandra lanceolata (Becc.) O. Ktze, Rev. Gen. PI. 1 (1891) 113; Howard, J. Arn. Arb. 21 (1940) 468, t. 1, f. 1-6. — Urandra - secundiflora\ Dahl, J. Arn. Arb. 33 (1952) 270, f. 57 & 57 A (pollen). — S. umbellatus (non Becc.) Anderson, Gard. Bull. Sing. 20 (1963) 166. Tree; bark greyish brown. Leaves lanceolate- oblong to narrowly elliptic, coriaceous, nerves 14-16 pairs, slightly raised on both faces as is the lax network of veins. Umbels with 2 or 3 branches. Drupe similar to that of var. secundiflorus, ob- longoid, 3.5-4.5 by 1-1.5 cm, apex attenuate and slightly curved. Distr. Malesia: Borneo. Ecol. Generally mixed swamp or peat swamp forest, or border of swamp forest with Agathis forest, low country. Fl. fr. Jan.-Dec. Vern. Entaburok, Iban, kepot bedjuku, Dajak, semanten, Sampit, ensurok, semburok, M. 5. Stemonurus malaccensis (Mast.) Sleum. No- tizbl. Berl.-Dahl. 15 (1940) 243; Blumea 17 (1969) 260; Fl. Thail. 2 (1970) 85. — Lasianthera apicalis (Thw.) Benth. in Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 584, as to the Labuan plant only. — Lasianthera malaccensis Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 584; Becc. Malesia 1 (1883) 257; King, J. As. Soc. Beng. 64, ii (1895) 118. — S. capitatus Becc. Malesia 1 (1877) 114, t. 5, f. 7-11; Valet. Crit. Overz. Olacin. (1886) 236; Merr. En. Born. (1921) 355; Ridl. Fl. Mai. Pen. 1 (1922) 431; Craib, Fl. Siam. En. 1 (1926) 272; BuRK. Diet. (1935) 2074. — Urandra ca- /7//ara(BECC.)0. Ktze, Rev. Gen. PI. 1 (1891) 113. — S. labuanensis Staff, Kew Bull. (1906) 71, p. p. lectotyp.; Merr. En. Born. (1921) 356, p. p. — Urandra nitida Howard, Lloydia 6 (1943) 147, t. 1, f. 3-6. — Fig. 29. Treelet or tree, (2.5-)6-15(-20) m, bole up to 10 m and 20 cm 0; bark smooth, brownish-grey, shallowly lengthwise fissured. Branchlets with pustules on older parts. Leaves variable in shape and size, oblong to subobovate-oblong, sometimes obovate, short-acuminate or obtusely apiculate, rarely rounded, base cuneate, subcoriaceous to coriaceous, smooth, i.e. practically not tubercled, a little shining above, dull beneath both in the fresh and dry state, (6-)8-12.5(-14) by (2-)2.5-4 (rarely -7) cm, nerves c. 15 pairs, with other shorter ones between, rather straight and sub- parallel, very slightly prominent on both faces, or not rarely almost obscure beneath, similar to those of 5. secundiflorus var. secundiflorus, re- ticulation of veins faintly visible above only, or obscure at all; petiole 1-1.5(-1.8) cm by (l-)1.5-2 (rarely -2.5) mm. Peduncle of the umbel-like in- florescence generally rather slender (1 mm, rarely up to 2 mm 0), 1-2 cm long, with 3-5 branchlets of the umbel (2-)3-10mm long, each bearing (3-)4-6 flowers crowded at the very end, the flowers seemingly on top of the peduncle when the branches of the umbel are very short; bracts at the base of the umbel ovate-oblong or broadly linear, obtuse, persisting during anthesis only, c. 2 by 2 mm. Calyx cup-shaped, c. 2 mm, rather deeply and obtusely lobed. Petals (2.5-)3(-4) mm, white- yellowish to dull pink, fragrant. Disk cup-shaped. Drupe subovoid-oblongoid, attenuate and some- times a little curved distally, 4.5-5 by (l-)1.5-2.2 cm, pink or pale brown below, merging into pale green in the upper part. Distr. Lower Burma, southern Peninsular Thailand; in Malesia: Malay Peninsula, Borneo. Ecol. Primary rain-forest, also bamboo forest, generally in lowland up to 150 m, more rarely on hillside or ridge tops up to 1370 m, never in peat swamp forest, apparently on fertile soil, but once reported from Slwrea materialis heath forest in Trengganu; scattered. Fl. fr. Jan.-Dec. Uses. Apparently not used, possibly due to the small size of the trees, contrary to most other species of the genus, which reach fair dimensions. Vern. Alau daan, Murut Bokan, gigi buntol, Selangor, baruas, pasir pasir, Kinabatangan, peropot, Dusun, petom, Pahang, tarung pelandok^ Perak, M. 6. Stemonurus ammui (Kaneh.) Sleum. Blumea 17 (1969) 260. — Urandra elliptica Schellenb. Bot. 1971] ICACiNACEAE (Sleumcr) 61 JT;69 Fig. 29. Stemonunis malaccensis (Mast.) Sleum. a. Habit, x 14, b. flower, X 5, c. petal inside, X 5, d. stamen, :' 5, e. calyx and ovary, y.lVi-f- fruit, X ^/4 (a N. Borneo For. Dep. 7685, b-e SAN 30985, / KEP 76124). Jahrb. 59 (1924) 17, non Merr. 1910. — Urandra ammui Kaneh. Bot. Mag. Tokyo 45 (1931) 293; Fl. Micron. (1933) 199, f. 86; J. Dep. Agr. Kyushu Imp. Un. 4 (1935) 359. — S. e///pr/c/«(ScHELLENB). Sleu.m. Notizbl. Berl.-Dahl. 15 (1940) 242. Tree, 10-24(-33) m, sometimes with buttresses or slenderly cylindrical pneumatophores, bole straight, 10-25(-30) cm o; bark smooth or with fine longitudinal fissures or pustules, whitish-grey to pale or dark brown. Leaves oblong- to obovate- elliptic, apex shortly (1-2 cm) ± abruptly and obtusely acuminate, sometimes rounded, very rarely slightly emarginate, coriaceous, shining above, rather dull beneath in dry specimens, margin slightly rcvolute, 9-l6(-19) by 4-8(-9) cm, smooth on both faces or practically so (a few tubercles maybe present along the midrib under- neath), nerves 12-14(-18) slightly curved pairs, generally a little raised or at least visible above, much less or hardly so beneath, reticulation of veins faint and above only; petiole 1.5-2 cm by c. 2 mm. Peduncles of umbel-like inflorescence 3-5 cm, stoutish, bearing 5-7 (rarely -9) branches, these 1-1.5 cm long, each with a subulate subpcrsistcnt basal bract c. I by 0.5 mm. Calyx cup-shaped, 2 mm, shallowly subacutely lobed. Petals (4-)5 by 2 mm, slightly keeled outside, ycllowish-whitc, with a strong lemon scent. Disk low, almost patellar, slightly or hardly lobcd. Drupe oblongoid-subfusiform, much attenuate at both ends. (4.3-)4.5-5 by 1.5 1.7cm. Distr. Micronesia (Patau Is.), Melanesia (New Britain, Solomons); in Malesia: eastern part of New Guinea. Ecol. Primary forest, generally on well-drained hilly slopes up to 200 m, occasionally in fresh- water swamp forest or mangrove along the coast. Fl. fr. Jan.-Dec. Vern. Aikunu, ailalo, aimarako, ainunnra, mala sata, Solomon Is. (Kwara'ae), animiii, Palau Is. 7. Stemonurus gitingensis (Elm.) Sleum. Blumea 17 (1969) 261. — Urandra gitingensis Elm. Leafl. Philip. Bot. 4 (1912) 1476. — 5. seciindijiuriis (non Bl.) Merr. En. Philip. 2 (1923) 489. Tree, 13 m or higher, up to 60 cm o; bark thick, yellowish or greyish, brittle, rough with lenticels. Leaves elliptic or obovatcly so, apex rounded, base broadly attenuate and obtuse, or sometimes rounded, dark green and shining above, dull and much paler beneath, coriaceous and rather stiff, smooth, i.e. practically not tubercled beneath, edge a little revolutc, 7-13 by 4-7 cm, nerves (I2-)15-I7 (rarely more) pairs, rather straight, subparallel and close together, slightly raised above, hardly so beneath, reticulation obscure; petiole stout, (2-)2.5-3 cm. Inflorescence not known. Infructcscence an umbel of 3-4 branches (c. 1 cm) on a robust peduncle (2 4 cm), bracts at base of the umbel according to the scars small and apparently early caducous. Flowers, as seen from the left scars, 2-4 clustered at the end of each branch of the umbel, certainly not in a row as in .v. sectindillora. Calyx an immature fruit 62 Flora Malesiana [ser. I, vol. 7^ cup-shaped, 2 mm, rather distinctly lobed. Drupe ovoid-ellipsoid, short-apiculate, 3.2-3.7 by 2-2.3 cm. Seed said to be surrounded with a soft white sourish tasting layer. Distr. Malesia: Philippines (Sibuyan: Mt Gi- tinggiting; after Merrill also on Panay). Ecol. Primary forest at 750-1750 m. Fr. March-April. Vern. Tugbdk, P. Bis. 8. Stemonurus monticolus (Schellenb.) Sleum. Notizbl. Berl.-Dahl. 15 (1940) 243 {' monticola' ); Blumea 17 (1969) 261. — ? Urandra scorpioides [non (Becc.) O. Ktze] Pulle, Nova Guinea 8 (1912) 659; Schellenb. Bot. Jahrb. 58 (1923) 165. — Urandra monticola Schellenb. Bot. Jahrb. 58 (1923) 164. — Urandra umbellata {non Becc.) Pulle, Nova Guinea 8 (1912) 658; Schellenb. Bot. Jahrb. 58 (1923) 165. — Urandra brassii Howard, Lloydia 6 (1943) 146, t. 1, f. 1 & 2; Dahl, J. Arn. Arb. 33 (1952) 270 (pollen). Tree, 10-30(-40) m, bole up to 18 m and 10-30 (-60) cm 0; bark grey to pale brown, smooth, maybe with a few longitudinal fissures, a little or not peeling; buttresses sometimes present, up to 1.5 by 1 by 0.1 m. Leaves elliptic to obovate-, sometimes lanceolate- or oblong-elliptic, apex shortly obtusely acuminate, or mostly blunt, some- times rounded and maybe a little emarginate, base ih broadly attenuate, coriaceous, dark green to yellowish green above, paler beneath, generally rather dull on both faces, smooth above, i dense- ly tubercled all over the undersurface, or at least so in a broad stripe along the midrib, very rarely smooth beneath, the edge a little recurved in dry specimens, (5-)6-10(-18) by (2.5-)3-6(-9) cm, nerves (12-)14-16 rather straight pairs, looping before the edge, very slightly or hardly raised or visible above and/or beneath, generally obscure beneath; petiole (1-) 1.5-2 cm by 1-2 mm. Umbel- like inflorescences peduncled (2-4, rarely -4 cm by 1-1.5 mm), branches 4-6 rather slender, each bearing 3-4(-5) flowers crowded distally; bracts at the base of the umbel ovate-subacuminate, 1-2 by 1 mm, subpersistent during anthesis. Calyx cup-shaped, 2 mm, ± distinctly obtusely lobed. Petals 3 (-4, rarely -4.5) by 1.5 mm. Disk a flattish slightly 5-lobed cup. Drupe subellipsoid-ovoid, 3-3.5 by 1.6 cm, apex subabruptly attenuate or apiculate. Distr. Malesia: New Guinea. Ecol. Primary (and occasionally also seconda- ry) rain-forest or forest substage, on slopes, from sea-level to 850(-1000) m, on clayey or sandy soil, sometimes on limestone with clay cover, locally common. Fl. fr. Jan.-Dec. Uses. Wood yellowish or light orange, appar- ently not used. Scrapings of bark heated in hot ashes are used for wounds in the Vogelkop Peninsula. Vern. Beram, bramsos, iinpam, keferan maper, manseran, sandawer, sapuduper, Biak, duwili, Kapauku, harihoh, korappi, korowapi, ukipi, wapapi, Japen, karmuk, karmu-u, lakobiliem, lobak, Mooi, kuwat, Amberbaken, nikoboa, Kebar, permeh, pirmir, Asmat, sakokohom, sakokom, Manikiong, lafus, Berik, tifus, Mander, weijan, Meosnum (Wanapi). 9. Stemonurus umbellatus Becc. Malesia 1 (1877) 115, t. 5, f. 5 & 6, incl. var. ovulifolius Becc. I.e. t. 4, f. 14 & 15; Valet. Crit. Overz. Olacin. (1886) 235; RiDL. Fl. Mai. Pen. 1 (1922) 431, p.p.; BuRK. Diet. (1935) 2074, p.p.; Sleum. Blumea 17 (1969) 262. — Urandra umbellata (Becc.) O. Ktze, Rev. Gen. PI. 1 (1891) 113; Dahl, J. Arn. Arb. 33 (1952) 270, f. 59 & 59 A (pollen). — Lasianthera umbellata (Becc.) King, J. As. Soc. Beng. 64, ii (1895) 117, e.xcl. specim. cit. (= S. malaccensis); Merr. En. Born. (1921) 355, incl. var. ovalifolia (Becc.) Merr. — S. evenius Staff, Kew Bull. (1906) 71; Merr. En. Born. (1921) 355. — Urandra hallieri Merr. Philip. J. Sc. 5 (1910) Bot. 194; Dahl, J. Arn. Arb. 33 (1952) 270 (pollen). — S. hallieri (Merr.) Merr. En. Philip. 2 (1923) 489. — Urandra evenia (Staff) Howard, J. Arn. Arb. 21 (1940) 468. — S. in- tercedens Heine in Fedde, Rep. 54 (1951) 237; Pfl. d. Samml. Clemens Kinabalu (1953) 57, 114. — Fig. 30-31. Tree, (8-)12-24(-37) m, (15-)20-35cm 0; bark smooth or finely cracked or hooped, some- times patchy, whitish-greyish to greenish or greyish-brownish. Branchlets dark by resinous Fig. 30. Stemonurus umbellatus Becc. Fruiting twig, Mt Blumut, Johore (FRI 8841) (Whitmore, 1968). 1971 ICACINACEAE (Sleumer) 63 excretion. Leaves oblong to obovate, sometimes elliptic, apex generally rounded or obtusely apic- ulate, sometimes slightly retuse, rarely shortly bluntly acuminate, base cuneate, coriaceous, some- times very stiffly so, shining above, rather dull beneath, the whole undersurface or at least the parts closer to the midrib set with fine epidermal tubercles, margin a little revolute. (6-)7-12(-16) by (3-)'l-7(-9) cm, nerves numerous (20-25) straight subparallel pairs, generally slightly raised above only, rather obscure though visible be- neath, reticulation, if any, remarkable above only; petiole 1.2-2 (very rarely -3) cm by 2(-3) mm. Cymes arranged in an umbel, its peduncle robust, l-2(-3) cm by 2 mm, the (4-)5-6(-8) branches (3-)5-8 mm long and c. 1.5 mm o, each branch bearing (l-)3^(-5) distally crowded flowers; bract at base of umbel ovate-subulate, c. 2 mm, sub- persistent. Calyx broadly obconical, 3 mm, base rugulose, margin truncate or very shallowly (apiculately) lobed, the very edge almost pellucid and ciliolate (lens!). Petals broad-oblong, slightly keeled, (4-)5(-6) by 2(-2.5) mm, white or light brown, scented. Disk cup-shaped and shallowly lobed. Drupe ovoid, very apex rather abruptly attenuate, lower half glossy green, upper one dull grey, 3-3.5(-4) by 1.5-2.2 cm, the fibrous part of the endocarp 1 mm thick. Distr. Malesia: Malay Peninsula, Borneo. Ecol. Primary forest, generally on hillside or high level alluvium, occasionally in swamp between hills, also in peat swamp forest, in coastal and high inland kerangas, on padang and on cliffs, generally in lowland up to 900 m, in the Cameron Highlands up to 1465 m, on Mt Murud in Sarawak at 1700 m, and on Mt Pagon Periok in Brunei at 1800 m; on Mt Kinabalu up to 1220 m, on sandy, maybe waterlogged soil (Agathis forest), sandy peat soil or sandy loam, rarely on sandstone. Fl. fr. Jan. -Dec. Uses. Wood pale orange-brown or honey- coloured, apparently not used, the value of the timber not known. Vern. Sampin kris, sampit kris, M; Borneo: djadjar, Dajak (Tajan), inlabiirok, st'mbiirok, Iban, kakidi, Dusun; Philippines: malafadii, Mbo. Fig. 31. Sleinuniinis unibcllatiis Becc. Stem, Mt Blumut, Johore (FRI 8841) (Whitmore, 1968). 15. WHITMOREA Sleum. Blumea 17 (1969) 263. — Fig. 32. Tree. Leaves spirally arranged, entire, coriaceous, penninerved, petioled. Flowers bisexual, generally pedicelled, uni- or biseriate towards the ± recurved end of secondary peduncles, the latter 3 or 4 as a cross-like umbel on top of a common axillary and solitary peduncle. Calyx cup-shaped, entire, persistent and ± flattened below fruit. Petals (4-)5, oblong, valvate, free in the uppermost part, remaining connate below at full anthesis, apex inflexcd, with a conspicuous midrib inside, caducous. Stamens (4-)5, inserted at the very base of the petals; lilamcnls linear, fleshy, with longish subclavate white hairs below the anther cells inside and at the connective outside; anther cells linear, thecae diverging below, basifixed. Disk annular, flat, rather obscure. Ovwr;- ovoid-conical, apex truncate, i.e. inverted there and forming a kind of cup, with the small hardly bilobed unilateral stigma on its inner edge. Drupe oblong-ellipsoid, smooth and reddish in the lower )4» 64 Flora Malesiana [ser. I, vol. 7^ Fig. 32. Whitmorea grandiflora Sleum. a. Habit, X VS, b. inflorescence, x 1/2, c. open flower, xV/i, d. petal inside, x2, e. stamen, x4,/. ovary and calyx, x5,/'. upper part of ovary obliquely seen from above to show the stigma turned inside the apical cavity of the ovary, x 10, g. submature fruit, x % (a-/ Brass 341 8A, g BSIP 5212 Whitmore, type). 1971 ICACiNACEAE (Slcumer) 65 papillose and greyish-yellowish in the upper V4 part, exocarp thin, with an outer fibrous, and an inner crustaceous layer. Seed 1, ellipsoid-subquadrangular, with a lengthwise depression around the middle; embryo small, in the apex of the copious albumen. Distr. Monotypic, hitherto only known from the Solomon Islands. 1. Whitmorea grandiflora Sleum. Blumea 17 (1969) 264. — Urandra itmbellalc [non (Becc.) O. Ktze] Walker, For. Fl. Br. Solomon Is. (1948) 127. — Fig. 32. Spreading canopy tree, 6-25(-30) m, bole straight, up to 12 m, up to 50 cm o, with slender and weak pneumatophores in wet places; bark grey to brown, smooth or cracked. Branchlets robust, often covered with resinous exudation as are the inflorescences. Leaves elliptic or oblong- elliptic, apex obtuse or slightly emarginate, base broadly cuneate to rounded, coriaceous, firm, shining green above, dull and greyish or yellowish green beneath, glabrous, 14-22 by 4-8(-12) cm, nerves in 20-25 pairs, straight and parallel to each other, slightly raised on both faces, reticu- lation dense, visibly raised above only; petiole 2-3(-4) cm by 2(-3) mm, minutely transversely wrinkled. Inflorescences axillary, solitary, with 3 or 4 secondary peduncles '1-3 cm) crosswise and umbellately spreading from top of main peduncle (2.5-4 cm by 1.5-2 mm), flowers several, close together, scorpioid, uni- or biseriate, on the upper i recurved part of the secondary peduncles, thick-pedicelled for 0.5-2(-3) mm; bracts at base of secondary peduncles knob-like, rather obscure. Calyx obconical-cupular, maybe suddenly nar- rowed at base for 1-2 mm to a kind of foot for 1-2 mm, rugose, persistent and more spreading or almost patellar under the ripe fruit, 4-5 by 6-7 mm. Petals chartaceous, glabrous, veined length- wise, becoming free only in uppermost part, remaining connate for the rest, oblong, white with cream centre or greenish-yellowish, with sweet smell, (12-)13-15 by 3-4 mm. Filaments linear, fleshy, 9-10 by 1 mm, with longish white hairs below the anthers inside and at connective out- side; anthers linear, thecae spreading below, 4 by 1 mm. Ovary ovoid-conical, truncate, hollow at apex, its edge folded in dry specimens, 3.5-4 by 2-2.5 mm. Drupe oblong-ellipsoid, apiculate, reddish and smooth in the lower Y^, papillate and greyish-yellowish in the upper |4 part, 3.5-4 by 2-2.5 cm. Distr. Melanesia: Solomon Is. (Bougainville to San Cristobal). Ecol. Primary and secondary rain-forest, both on well drained slopes or ridge top and in swampy valley bottoms, though locally not rare, from lowland up to 950 m. Uses. Wood rather hard, moderately heavy, cream to brownish, used for planting sticks, suitable for not too heavy constructions. Vern. Aialo, aikitma, aikuni, aikimu, arifanatia, Kwara'ae, diileke, Kolombangara, porutolo. Faro I. 16. POLYPORANDRA Becc. Malesia 1 (1877) 125; Sleum. Blumea 17 (1969) 248. — Fig. 33. Dioecious climber, whether or not with extra-axillary or intrapetiolar tendrils. Leaves opposite, entire, pcnnincrvcd. Panicles axillary, composed of short cymes. Flowers small, 5-6(-7)-mcrous. Pedicels articulated with calyx. — 3 Flowers: Calyx cup-shaped, shortly dented. Petals valvate, inflexcd appcndiculatc distally inside in bud. Stamens sessile; anthers thick, clavalc-subglobosc, with numerous small pollen-bearing alveoles, each of which is covered with a caducous membrane. Rudiment of ovary conical. — ^i? Flowers: Calyx deeply dented. Petals very short, connate in the lower '/^ Stamens 0. Ovary subglobose-cylindric, stigma large, disk- shaped, the centre depressed, crcnulate or very shortly many-lobcd in later stages. Fruit compressed-ellipsoid; exocarp thin; endocarp firm, irregularly foveolate- reticulatc outside when the exocarp is gone, wrinkled inside. Seed 1; cotyledons foliaccou.s. Distr. Monotypic, Solomon Is. and East Malesia. Fig. 34. 66 Flora Malesiana [ser. I, vol. 7^ J.T.'68 Fig. 33. Polyporandra scandens Becc. a. Habit, X y2, b. S inflorescence, X V2> f- c? flower, X 5, rf. ? flower xlO, e. anther, x20,/. drupe, xl {a, c, e Van Royen & Sletjmer 6696, b ditto 6830, d Carr 12610 f ditto 16435). I. Polyporandra scandens Becc. Malesia 1 (1877) 125, t. 7; Pulle, Nova Guinea 8 (1912) 660; Schellenb. Bot. Jahrb. 58 (1923) 176; Birnie, Nova Guinea 14, 2 (1926) 277; White, J. Arn. Arb. 10 (1929) 235; Sleum. in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 379, f. 109; Dahl, J. Arn. Arb. 33 (1952) 275, f. 70 (pollen); Steen. Nova Guinea, Bot. n. 23 (1965) 495; Sleum. Blumea 17 (1969) 248. — ? lodes ovalis (non Bl.) Warb. Bot. Jahrb. 13 (1891) 299. — P. hanseman- nii Engl. Bot. Jahrb. 16, Beibl. 39 (1893) 13 Chausemanni'); in E. & P. Nat. Pfl. Fam. 3, 5 (1895) 253; K. Sch. & Laut. F1. Schutzgeb. (1900) 418. — lodes sogerensis Baker/. J. Bot. 61 (1923) Suppl. 10. — Fig. 33. Scandent shrub or branched liana, climbing to i 1971] ICACINACEAE (Sleumcr) 67 35 m, stem up to 8 cm o. Branchlets slender, divaricate, pubescent initially; nodes thickened; internodes 4— 10 cm. Leaves opposite, ovate to elliptic, apex shortly — abruptly acuminate to cuspidate, base broadly attenuate to rounded, slightly inequilateral, chartaceous to subcoria- ceous, glabrous above, covered with scattered short appressed hairs all over beneath, 7-15(-17) by 3.5-9 cm, margin slightly revolute, entire, mid- rib a little impressed above, strongly prominent beneath, nerves (4-)6-8 curved-ascendent pairs, prominent beneath only as is the rather dense network of veins; petiole 8-15(-20) mm; tendrils whether or not present at the upper axils, elon- gate, bifurcate. Panicles 1-3 from the same axil, few- to many-flowered, (2-)3-9 cm, ascending or recurved, all over covered with short appressed hairs. Peduncle 1-3 cm, bearing several few- flowered cymes. Pedicels slender in the S, thicker in the 2 flowers. — 3 Flowers: Calyx tube short, lobes ovate-acuminate, c. 1 mm. Petals ovate- caudate, — expanded, white to cream or yellow, 4-5 (-6) mm, acumen abruptly narrowed, ± in- flexed, glabrous, 2 mm, tip blunt. Anthers c. 1 mm 0. Rudiment of ovary hardly 1 mm. — • $ Flowers: Calyx as in o flowers. Petals only 1-1.5 mm. Stamens 0. Ovary densely hairy, c. 1.3 mm. Fruit compressed-ovoid-ellipsoid, c. 2 by 1.5 by 1.5 cm, appressed-pubescent; exocarp rather soft and smooth, 1 mm, apparently going early; en- docarp hard, 0.5 mm, coarsely reticulate-foveolate. Fig. 34. Distribution of Polyporandra scandens Becc. Distr. Solomon Is.; in Malesia: New Guinea, Moluccas (Halmahera, Batjan). Fig. 34. Ecol. Climber over shrubs or high liana in primary and secondary rain-forest, from sea-level to 700 (possibly to 1000) m. Fl. fr. Jan.-Dec. Uses. On Bougainville young leaves are cooked and eaten or added to taros to give these a flavor. Vern. Dodofu, Halmahera. 17. lODES Bl. Bijdr. (1825) 29 Clodes'); Sleum. Blumea 17 (1969) 219. — Mappianthus Hand.-Mazz. Anz. Ak. Wiss. Wien M.-N. Kl. 58 (1921) 150; Sleum. Blumea 17 (1969)225. — Fig. 36. Dioecious climbing shrubs or lianas. Branchlets hairy, generally bearing shortly bifid tendrils at or closely to the nodes (but not axillary!). Leaves decussate, entire, apiculate by a somewhat projecting midrib, penninerved, with ± appressed and strigose and/or softer, erect or obliquely spreading coarse hairs on midrib, nerves, veins and veinlets, and sometimes besides with a tomentum of very short fine substellate hairs which covers the intervenium beneath. Inflorescences in pedunclcd cymes which may be combined to panicles, axillary, supra-axillary or a little lateral from the nodes, sometimes terminal, solitary, the ; ones often elongate and many-flowered, the ^^ ones generally shorter and rather few-flowered; lower peduncles often sterile and becoming tendrils. Flowers small, white to cream or yellowish, fragrant. Pedicels arliculalc with the calyx and often dilated distally; bracts and bractcolcs minute. Calyx cup-shaped, 4- or 5-lobed halfway or up to •*4of its total length (Malesia), persistent in V- Petals^ or 5, or sometimes wanting in ?, valvalc in bud, generally connate at base only, rarely and only in ," connate to Vi to % of their length into a lube, only the rcflcxcd lobes free, a little keeled iaside, tips sometimes narrowed and inflexed already in bud, persistent in S^ — J Flowers: Stamens (3 )4 or 5; filaments very short and fixed cither to the base of the (then shortly connate) petals or to the end of the corolla tube between the lobes, 68 Flora Malesiana [ser. 1, vol. 7^ or elongate and fixed to the base of the corolla tube; anther cells almost basifixed, introrse. Ovary rudimentary. — I Flowers: Staminocles sometimes present. Ovary sessile; stigma sessile, discoid, cut on one side, or faintly 2-5-lobed. Drupe ovoid- ellipsoid or oblong-obovoid (Malesia), or almond-like, generally laterally com- pressed and longitudinally grooved or crested; exocarp thin, fleshy, yellow to red; endocarp crustaceous to thin-woody, irregularly ribbed or reticulately lacunose, or warty outside, smooth inside. Seed 1, albuminous; cotyledons large, foliaceous. Distr. About 16 spp. in tropical Africa (and Madagascar), 3 spp. in SE. Asia (Assam and Burma to SW.-SE. China, Hainan, Indo-China, and Thailand), throughout Malesia (7 spp.), but in New Guinea only questionably known from a single old collection; 2 Malesian spp. also known from SE. Asia. Fig. 35. Anat. Robinson (Ann. Inst. Bot. Btzg 8, 1890, 111, t. 18 & 19, f. 1-5) has investigated the anatomy of the stem of/, cirrhosa Turcz. Vascular strands are formed here also in the pith, though the bast and wood elements are inversed. Notes. For the determination of the various species the indumentum plays an important role. The use of a lens is necessary, and only leaves of — the same state of maturity should be investigated or compared. Mappianthus differs from lodes only by rather vague characters such as filaments rather long in pro- portion to the length of the petals and a drupe with a finely grooved and tubercled exocarp. It is regarded here as congeneric, a point of view already expressed by Baillon, who, however, created for /. hookeriana a proper sect. Lasiodes (Adansonia 10, 1872, 268). KEY TO THE SPECIES 1. Petals in the o flowers practically free, i.e. united only at the base. 2. Indument on the undersurface of i mature leaves consisting almost exclusively of rather sparse to more dense manifestly appressed stiff hairs. 3. Flowers 5-merous. Pedicels of the cJ flowers swollen and ± plate-like dilated distally when the flower has gone. ? Flowers without staminodes. Veinlets of leaves rather faintly reticulate. 1. I. ovalis 3. Flowers 4-merous. Pedicels of c? flowers not swollen distally. ? Flowers whether or not with 4 staminodes. Veinlets of leaves strongly reticulate 2. I. yatesii 2. Indument on the undersurface of ± mature leaves consisting exclusively either of numerous fine and soft iz erect hairs, or of both such hairs and more robust, stiff and generally ± appressed (rarely obliquely spreading) hairs of various density. 4. Flowers 4-merous. Hairs on undersurface of leaves rather sparse. Petals present in 2 flowers. Drupe 2.5-3 by 1.5 by 1 cm 3. I. reticulata 4. Flowers 5-merous. Hairs on undersurface of leaves (rather) dense. Petals absent in $ flowers. Drupe up to 2 by 1.2(-1.5) by 1 cm 4. I. cirrhosa 1. Petals in the ^ flowers united into a tube for about their lower 2/3 (the tube at least 3 mm long). 5. Drupe oblong-ellipsoid, ± obtuse at both ends, 1.2-1.5 by 0.8-1.2 by 0.8-1 cm. 5. 1, phillppinensis 5. Drupe larger. 6. Drupe oblong-ovoid, apex broadly obtuse, base narrowed, 2.8-3.3 by 1.3-1.5 by 0.8-1.2 cm; crustaceous endocarp laxly coarsely lacunose 6. I. velutina 6. Drupe almond-like, 2.3-3.5(-4.5) by 1.6-2 by 0.7-1 cm; crustaceous endocarp with several longi- tudinal shallow furrows and numerous flattish warts 7. I. hookeriana 1. lodes ovalis Bl. Bijdr. (1825) 30; Miq. F1. Planch, in Mast, in Hook./. FI. Br. Ind. 1 (1875) Ind. Bat. 1 (1856) 795, p.p.; Baill. Adansonia 3 597; King, J. As. Soc. Beng. 64, ii (1895) 128; (1863) 364 in text, p.p.; in DC. Prod. 17 (1873) Ridl. F1. Mai. Pen. 1 (1922) 435. 22, incl. var. gemdna Baill. I.e. 23, p.p.; Becc. Slender shrub, scandent with tendrils, up to Malesia 1 (1877) 124, p.p.; Scheff. Ann. Jard. 5 m. Leaves oblong-lanceolate to -elliptic or Bot. Btzg 1 (1876) 14; Hall./. Med. Rijksherb. -obovate, apex acuminate to rounded, though 1 (1910) 13; Back. Schoolfl. Java (1911) 230 & minutely cuspidate by the protruding midrib, base Addenda; Koord. Exk. Fl. Java 2 (1912) 532; rounded to subcordate, slightly inequilateral, thin KooRD.-ScHUM. Syst. Verz. 1, Fam. 162 (1912) chartaceous, glabrous above the midrib excepted, 5; Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 which is covered on both faces with a pale rusty (1948) fam. 135, p. 7; Back. & Bakh. / Fl. Java short villous tomentum equally found on the 2 (1965) 61; Sleum. Blumea 17 (1969) 220. — branchlets and petioles, undersurface ± densely /. horsfieldii Baill. Adansonia 10 (1872) 267; covered with stiff appressed hairs on nerves, veins in DC. Prod. 17 (1873) 24. — Polyporandra jung- and veinlets, the proper intervenium glabrous, huhnii Koord. Kon. Ak. A'dam Versl. Verg. (5-)6-15 by (2.5-)3-6.5 cm, midrib and nerves Wis- & Natuurk. 17 (1909) 780. — /. oblonga generally slightly impressed above, prominent 1971] ICACINACEAE (Slcumer) 69 Fig. 35. Distribution of lodes (incl. Mappiantlnis). The tigure above the hyphen indicates the number of endemic species in the island or area, the figure below the hyphen the number of non-endemic species. beneath, nerves 6-8 pairs, curved-ascending and inarching, reticulation of veins rather coarse, that of the veinlets more dense though rather faintly raised beneath: petiole 3-15 mm. Inflorescences all over covered with both rusty short strigose and softer hairs, the S ones repeatedly dichotomous, many-flowered, up to 10 cm, the 2 ones con- sisting of 8-16-flowered cymes and only 1.5-4 cm (incl. peduncle). Pedicels slender, swollen distaliy, the very end finally plate-like dilated when the flower has gone, 1-3 mm. Cfl/y.v 5-lobed to Vi-Ji-) c. 1.5 mm. Petals 5, oblong, white or light yellow, tip extended and inflcxed, c. 2.5 mm in the o, lanceolate and acute (not inflcxed) and c. 2 mm in the 2 flowers, laxly appressed-strigose outside as is the calyx. — o Flowers: Stamens 5; filaments slender, c. 0.6 mm; anthers subovate-oblong, c. 1 mm, cells with crenate-lobed margin. Rudiment of ovary hairy. — V Flowers: Staminocles 0. Ovary thick-cylindric, all over densely appressedly hairy, 2 mm; stigma discoid, wider than the ovary, deeply incised on one side, margin crenulatc, gla- brous. Drupe inequally ovoid-compresscd, said to be scarlet and to attain 1.5-2.5 cm in length (seen in immature state only up to 1 cm); cndo- carp with slightly prominent, not properly keeled ribs. Distr. Malesia: Sumatra (Eastcoast), Malay Peninsula (Pcnang to Singapore), W. Java, Central Java (Pckalongan: Margasari, once). Ecol. Along border of woods, in shrubberies, along roads or riversides, often in secondary veg- etation, once in teak-forest (Margasari), some- times on limestone, 50-300(-800) m. Fl. Jr. J m.-Dec. Notes. What has been understood as /. ovalis in numerous floras of SE. Asia and Malesia, belong.s partly to other species of the genus, mainly to /. cirrlwsa Turcz., /. pliilippincnsis Mf.HK. and /. vitixinea Hance. /. ovalis Bl. was mentioned by K. Sch. & Laut. Fl. Schutzgeb. (1900) 417 and Schellenb. Bot. Jahrb. 58 (1923) 176 for New Guinea (Morobe Distr., once found by Warburg), but the specimen is lost and its identity remains uncer- tain. There is no recent record of lodes from New Guinea. 2. lodes yatesii Merr. Pap. Mich. Ac. Sc. 19 (1934) 165; Sleum. Blumea 17 (1969) 221. var. yatesii. Scandent shrub. Branchlets weak, sparsely pu- bescent; tendrils up to 15 cm. Leaves ovate- or oblong-elliptic, apex abruptly acuminate for 1-1.5 cm, tip acute, base rounded to subcordate, char- taceous, ± shining on both sides, glabrous above, the midrib excepted, subdensely covered beneath with appressed strigose hairs on veins and veinlets, and besides with softer hairs on midrib and nerves, the proper intervenium hairless, (5-)7-12 by 3-6 cm, midrib a little sunken above, much pro- minent beneath, nerves 5-6 curved-ascending and inarching pairs, raised beneath, veins coarsely re- ticulated and but little prominent beneath, re- ticulation of veinlets dense, finely raised on both surfaces; petiole slender, subdensely hairy, 0.5-1.2 cm. Cymes 3-8 cm incl. peduncle, few- to many- flowered, very slender, all over covered with short substrigose and brownish softer hairs. Pedicels slightly dilated distaliy in the v, hardly so in the J flowers, very slender, 2-5 mm. Calyx c. 0.8 mm, 4-lobed to 1 3. lobes spreading. Petals 4, oblong- acuminate, tip not extended and hardly inflexed, appressedly strigose outside as is the calyx, gla- brous inside, c. 1.2 mm. — o Flowers: Stamens 4; filaments 0.6 mm; anther cells ovate, margin not crenulate, c. 0.4 mm. Rudiment of ovary sub- globular, hairy, minute. — ? Flowers: Sepals and petals as in cj flowers, persistent. Staminodes 4. Ovary ovoid, densely hairy; stigma flat, subgla- brous. Drupe elongate-oblong-ovoid, laterally compressed, brownish-substrigose-iomentose, 2.5- 3 by 1.5 by 1.3 cm, slightly keeled; endocarp shallowly and coarsely reticulate-impressed on both faces. Distr. Malesia: Sumatra (Eastcoast; Riouw: Indragiri Uplands). Ecol. Forest (also secondary) along river or roadside. Fl. fr. March-Nov. var. glabresccns (Ridl.) Slf.um. Blumea 17 (1969) 221. - /. reticulata King var. glabrescens Ridl. Kcw Bull. (1931) 35. Appressed hairs on the undcrsurfacc of leaves sparse. + Flowers without staminodes. Distr. Malesia: Borneo (Sarawak: Kuching), once found. 3. lodes reticulata King. J. As. Soc. Beng. 64. ii (1895) 126; Ridl. Fl. Mai. Pen. 1 (1922) 434; SiruM. BUimcu 17 (1969) 221. Woody climber, up to 9 m. Branchlets slender, striate, with a broad line of dilutcly fcrrugineous tomcnlum on one side, changing sides at the nodes; 70 Flora Malesiana [ser. 1, vol. 7^ tendrils up to 15 cm. Leaves ovate-elliptic, some- times subobovate, apex abruptly acuminate for 0.5-1.5 cm, acute, base generally slightly cordate, a little inequilateral, thin-coriaceous, glabrous and shining above, the hairy midrib excepted, all over the undcrsurface covered with rather sparse obliquely or patently spreading hairs, soft to the touch, 5-12 by 3-6.5 cm, midrib and nerves slightly impressed above, boldly prominent be- neath, nerves 4—5 curved high-ascending and looping pairs, reticulation of veins and veinlets finely raised above, strongly so beneath; petiole densely hairy, 1-1.5 mm. Cymes lax, the S many-flowered and usually exceeding the leaves in length, the i ones few-flowered and generally shorter, ± patently soft-rusty-pubescent in all outer parts. Pedicels very slender in the S, thicker and swollen distally only in the ?. — cj Flowers: Calyx cup-shaped, 1 mm, 4-lobed to '/2-%- Petals 4, ovate-acuminate, a little fleshy, densely hairy outside, glabrous inside, 0.8 mm. Stamens 4, subsessile; anther cells subcordate, 0.2 mm. Ru- diment of ovary minute, glabrous. — 5 Flowers: Calyx as in ;5 flowers. Petals 4, subovate-oblong, united at the very base, tip acute, hairy outside as is the calyx, reflexed, 2-2.5 mm. Staminodes 0. Ovary thick-cylindrical or barrel-shaped, all over densely appressedly hairy, 1.5 mm; stigma discoid, deeply cut on one side, margin hardly crenulated. Drupe ellipsoid, compressed, base narrowed, fer- rugineous-velvety, 2.5-3 by 1.5 by 1 cm, on stoutish pedicel, 5-8 mm; endocarp bony, coarsely furrowed outside. Distr. Malesia: Malay Peninsula fPerak, Pa- hang, Negri Sembilan). Ecol. Open jungle, at low altitudes up to 250 m. Fl. May-Nov., fr. May-June. 4. lodes cirrhosa Tlrcz. Bull. Soc. Nat. Moscou 27, ii (1854) 281; Sleum. Notizbl. Berl.-Dahl. 15 (1940) 250; Heine, Pfl. d. Samml. Clemens Ki- nabalu (1953) 58; Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 7; Heine in Fedde, Rep. 54 (1951) 238; Back. & Bakh./. Fl. Java 2 (1965) 62 Ccirrosa); Sleum. Blumea 17 (1969) 221; Fl. Thail. 2 (1970) 86. — /. ovalis inon Bl.) Hassk. Cat. Hort. Bog. (1844) 172; R. Br. in Benn. & Brown, PI. Jav. Rar. (1852) 243, t. 48; MiQ. Fl. Ind. Bat. 1 (1856) 795, p.p.; Baill. Adansonia 3 (1863) 364, in text, p.p.; in DC. Prod. 17 (1873) 22, ind. var. genuina Baill. I.e. 23, p.p. et var. miquelii Baill. I.e. 23; Mast. in Hook. /. Fl. Br. Ind. 1 (1875) 596; Becc. Malesia 1 (1877) 124, p.p.; King, J. As. Soc. Beng. 64, ii (1895) 128; Hochr. PI. Bog. Exs. (1904) 18; Bull. Inst. Bot. Btzg 19 (1904) 40; ibid. 22 (1905) 106 (as var. genuina); Brandis, Ind. Trees (1906) 154; Gagnep. Fl. Gen. I.-C. 1 (1911) 844, incl. var. cochinchinensis Pierre ex Gagnep. I.e. 845; Merr. En. Born. (1921) 356; RiDL. Fl. Mai. Pen. 1 (1922) 435; Craib, Fl. Siam. En. 1 (1926) 276; Gagnep. Fl. Gen. I.-C. Suppl. (1948) 757; Dahl, J. Arn. Arb. 33 (1952) 274 (pollen). — Natsiatum oppositifolium Planch, in Hook. Lond. J. Bot. 5 (1846) 247, nom. nud. — /. tomentella MiQ. Fl. Ind. Bat. 1 (1856) 796; Kurz, J. As. Soc. Beng. 44, ii (1875) 137; For. Fl. Burma 1 (1877) 243; Becc. Malesia 1 (1877) 124, p.p.; KooRD. Minah. (1898) 393; Back. Schoolfl. Java (1911) Addenda ad p. 229; Koord. Exk. Fl. Java 2 (1912) 532; Koord. -Schum. Syst. Verz. 1, Fam. 162 (1912) 5; Dahl, J. Arn. Arb. 33 (1952) 274 (pollen). — /. brandisii Kurz, J. As. Soc. Beng. 41, ii (1872) 298; ibid. 44, ii (1875) 156; Mast, in Hook. / Fl. Br. Ind. 1 (1875) 596; Kurz, For. Fl. Burma 1 (1877) 243. — /. hors- fieldii Baill. Adansonia 10 (1872) 267; in DC. Prod. 17 (1873) 24. — /. floribunda Merr. Pap. Mich. Ac. Sc. 19 (1934) 166, t. 29; Dahl, J. Arn. Arb. 33 (1952) 274, f. 65 (pollen). — Fig. 36a-d. Woody climber, 5-10(-18) m tall, stem up to 8 cm 0. Branchlets slender, rusty-tomentellous. Leaves ovate to ovate-elliptic, rarely oblong- or obovate-elliptic, apex shortly acuminate and acute by the slightly protruding midrib, base rounded or subcordate, firmly chartaceous, upper surface glabrous except midrib and nerves which bear rather dense soft yellowish erect or obliquely spreading hairs as are found all over the undcr- surface on midrib, nerves, veins and veinlets (the proper intervenium hairless), forming a ve- lutinous tomentum, 5-15(-18) by 3.5-8(-11.5) cm, nerves 4—6 curved and rather steeply ascending pairs, prominent beneath as are midrib and the transverse veins, reticulation of veinlets dense, a little raised beneath only; petiole l-2(-3.5) cm, tomentellous. Inflorescence all over covered with a short soft rusty tomentum, the c-c'. Woody climber, up to 6 ni. Branchlcts slender, striate, soft villous by Icrrugincous fine rather short hairs as are the petioles; tendrils slender, up 72 Flora Malesiana [ser. I, vol. 71 to 15 cm. Leaves ovate-elliptic to -oblong, apex short acuminate, the midrib hardly projected beyond the lamina, base rounded or mostly slightly cordate, thin chartaceous, upper surface finally glabrous except the midrib, undersurface all over soft tomentose, more densely so on midrib and nerves, the hairs being of two markedly dif- ferent types, i.e. stoutish substrigose — appressed, and much finer, soft and i erect ones, the latter sometimes rather scarce, the space between the veinlets hairless, (3-)5-9 by 2.5-5(-7) cm, midrib and 4-6 pairs of ascendent and — looping nerves plain above, — strongly raised beneath, less so the veins, reticulation of veinlets not very distinct; petiole 0.6-1 (-1.5) cm. — o Inflorescences: Pan- icle; composed of several cymes, rather lax and relatively few-flowered, 2-4f-5) cm incl. the long peduncle, covered with — crisped and appressed rusty substrigose hairs. Pedicels slender, 1-2 mm, the end glabrous and a little dilated when the flower is gone. Calyx cup-shaped, 1.5 mm, acutely 5-lobed halfway. Petals subglobose in bud, rapidly accrescent to 6-7(-8) mm at full anthesis, united into a narrow tube for the lower 2/3, free at the 5 lobes, the latter acuminate, narrow, tip inflexed in bud stage, ultimately reflexed or spreading, hairy outside and at the mouth inside, white or cream. Stamens 5, inserted at the end of the tube, alternating with the lobes; filaments flattened, very short; anther-sacs narrowly sub- ovate-oblong, not crenulate, 0.8-1 mm. Rudiment of ovary cylindric, glabrous, c. 1.5 mm. — 1 Inflorescences 3-5 Crarely -12)-flowered, 2-3 cm incl. the peduncle, hairy as are the(^ ones. Pedicel slender, 3 (-4) mm, not swollen distally. Calyx cup-shaped, 1.8 mm, 5-lobed halfway or a little more. Petals (4-)5, narrow-oblong, connate at the base, white or cream, thin, finally reflexed, glabrous inside, 2.5-3 by 0.6-0.8 mm. Staminodes 0. Ovary cylindric, all over clad with stiff" forward directed hairs, 2 mm; stigma thick, bilobed, glabrous. Drupe obliquely oblongoid-ellipsoid, laterally compressed and rather sharply crested, — obtuse at both ends, red to orange, 1.2-1.5 by 0.8-1.2 by 0.8-1 cm, densely rusty hairy; endocarp coarsely reticulate-lacunose; pedicels stoutish, 2-5 mm, a little swollen distally. Sepals and petals persistent on the fruit for a long time. Distr. Malesia: N. Borneo, Philippines (all islands). Central E. Celebes (once found), Mo- luccas (Talaud, Morotai, Halmahera, Ternate, Ambon). Ecol. Edge of forests or thickets, also seconda- ry growths, in the lowland up to 200 m, but ascending in the foothills of Mt Kinabalu up to 1525 m. often near rivers. Vern. Kogopusut, Dusun (Sambunan). 6. lodes velutina King, J. As. Soc. Beng. 64, ii (1895) 127; Ridl. FI. Mai. Pen. 1 (1922) 434; BuRK. Diet. (1935) 1245; Sleum. Blumea 17 (1969) 224. var. velutina. Climber, similar in leaves and flowers to /. philippinensis. Branchlets shortly rufous-pubescent especially so on one side; tendrils glabrescent. Leaves ovate, apex short acuminate and subacute by a glandular tip, i.e. the protruding midrib, base rounded or generally subcordate, firmly chartaceous, upper surface initially — densely short pubescent, soft to the touch, glabrescent except the midrib, undersurface velvety by nu- merous — appressed substrigose and even more numerous softer — erect hairs, the tomentum rusty-olivaceous in dry, said to be whitish in fresh specimens, the intervenium, i.e. the proper space between the veinlets glabrous and thus green, (3-)4-10 by 2-5.5(-8) cm, nerves 4-5 rather steeply curved-ascendent pairs, a little raised be- neath only, veins and reticulation of veinlets ~ hidden by the tomentum underneath; petiole slender, tomentose, 0.5-1. 5(-2) cm. — !^ In- florescences: Cymes few- to rather many-flowered, all over rusty-pubescent, 2-4 cm (incl. peduncle). Pedicels slender, 2-3 mm, finally a little swollen distally and glabrous at the very tip. Calyx cup- shaped, c. 1.3 mm, 4- or 5-lobed for V3 to almost '/2- Petals 4 or 5, bud club-shaped, at full anthesis 5-7 mm and connate to a tube in the lower 2/3, lobes finally reflexed, their tip inflexed, sub- densely — appressedly hairy outside as is the calyx, long-hairy at the throat inside. Stamens 4 or 5, inserted at the throat; filaments laxly hairy, c. 1 mm; anther-sacs ovate-oblong, 0.8-1 mm. Rudiment of ovary conical-cylindric, glabrous, 0.5 mm. — ^ Inflorescences: Cymes 2-5-flowered, 2-3 cm incl. the rather robust peduncle. Pedicels rather slender and 3-5 mm in anthesis, a little accrescent and more robust in fruit. Calyx cup- shaped, c. 1 mm, 4- or 5-Iobed halfway, appressed- ly hairy as are the petals outside. Petals 4 or 5, subobovate-oblong, free to almost their base, 4-5 mm. Staminodes 0. Ovary cylindric, densely hairy; stigma slightly 4- or 5-lobed. Ripe drupe oblong- obovoid, apex broadly obtuse, base attenuate, much compressed laterally and manifestly crested, with a less obvious crest on each of the flat sides, tomentose initially, finally subglabrous, 2.8-3.3 by 1.3-1.5 by 0.8-1.2 cm, the thin exocarp blackish when dry; endocarp laxly coarsely lacunose; stigma slightly 5-lobed, 2 mmo. Seed soViiary, compressed. Distr. Malesia: Malay Peninsula (Perak, Ne- gri Sembilan, Malacca, Singapore). Ecol. Woods and hedges, at low elevation. Vern. Akar china bukit, a. salupat, M. var. subvillosa Sleum. Blumea 17 (1969) 224. — /. tomentella (non MiQ.) Schellenb. Bot. Jahrb. 58 (1923) 176. Undersurface of leaves covered with a felt of small substellate hairs (and thus greyish) on the space between the veinlets. In flower and fruit characters not distinguishable from var. velutina. Distr. Malesia: Sumatra (Eastcoast, Ben- coolen), Borneo (Amai Ambit and area S. of Kuching). Questionable for New Guinea by an old specimen said to have been collected by Zip- PELIUS. Ecol. Primary forest both on sand- and lime- 1971 ICACINACEAE (Sleumer) 73 stone in Borneo, 100-1000 m. Fl. fr. Aug. 7. lodes hookeriana Baill. Adansonia 10 (1872) 268; in DC. Prod. 17 (1873) 23; Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 596; KuRZ, J. As. Soc. Beng. 44, ii (1875) 157; For. Fl. Burma 1 (1877) 244; Brandis, Ind. Trees (1906) 154; Kanjilal c.s. Fl. Assam 1 (1936) 254. — /. thomsoniana Baill. Adansonia 10 (1872) 270; in DC. Prod. 17 (1873) 25; Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 596; Brandis, Ind. Trees (1906) 154. — L. reticulata King \ar. glabrescens Ridl. Kew Bull. (1931) 35. — Mappianthiis borneensis Merr. Webbia 7 (1950) 317. — Mappianthus hookerianus (Baill.) Sleum. Blumea 17 (1969) 225. Fairly large woody climber, sometimes with tendrils, branchlets, undersurface of leaves along nerves and midrib, petioles and inflorescences sub- densely clothed with substrigose, appressed hairs. Leaves elliptic- to ovate-oblong, apex abruptly shortly (maybe bluntly) acuminate, or almost cus- pidate, base — rounded, rarely subcordate, thin- coriaceous, shining on both faces, margin slightly revolute, entire (seemingly crenulate or denticulate in dry specimens with crisped edge), (8-) 10-1 5 (-20) by (3.5-)5-8 cm, midrib narrowly impressed above, strongly raised beneath, nerves 7-8 (-9) pairs, moderately curved-ascending and inarching before the edge, faintly raised above, distinctly so beneath as are the — transverse veins, reticulation of veinlets dense and finely raised on both faces; petiole 1-1.5 cm. Inflorescences slightly extra- axillary dichasia, the o ones rather many-, the ? ones few-flowered, often reduced in the lower axils to tendrils. Pedicels c. 1 mm, slender in the o, stoutish in the 2 flowers. Calyx cup-shaped, c. 1 mm. edge minutely 5-dented to almost entire. — o Flowers: Petals oblong, fleshy, c. 3 mm, con- nate to c. \i, densely and shortly yellowish- substrigose outside, glabrous inside. Filaments fili- form, c. 2 mm; anther cells linear-oblong, c. 1.8 mm. Rudiment oi ovary cylindric, all over shortly strigose-tomcntose. — s^ Flowers unknown. Drupe almond-like, 2.3-3.5(^.5) by 1.6-2 by 0.7-1 cm, on thick pedicel c. 2 mm; stigma rather small, faintly 5-lobed; exocarp fleshy, orange red, ap- pressedly short-strigose; endocarp crustaceous, outside with several longitudinal shallow furrows and numerous flattish warts, inside smooth. Distr. SE. Asia (Assam, E. Bengal); in Malesia: Sumatra (once found, Westcoast) and Borneo (Sarawak: Kuching area; N. Borneo: foothills of Mt Kinabalu, c. 900 m, twice found). Ecol. Forests. Fr. Sept.-Nov. Note. Closely related to and possibly con- specific with Mappianthus iodoides Hand.-Mazz. (N. of Tonkin and adjacent China, eastwards to Hunan and Fukien). Excluded lodes ferruginea K. ScH. & Laut. Fl. Schutzgeb. (1900) 418; SCHELLENB. Bot. Jahrb. 58 (1923) 176, nota sub I. tomentella; Sleum. Blumea 17 (1969) 225 = Alangium villosum (Bl.) Wang. (Alangiaceae). 18. SARCOSTIGMA W. & A. Edinb. New Phil. J. 14 (1833) 299; Sleum. Blumea 17 (1969) 253. — Fig. 38. Dioecious climbers. Leaves spirally arranged, entire, penninerved, strongly reticulate; petiole curved or twisted, lengthwise striate and transversely wrinkled, leaving a raised elliptic white scar. Flowers sessile or sometimes pedicellate (? only), clustered (or fascicled) or solitary along the rachis of long, pendulous interrupted or spike-like racemes, these simple or composed to panicles, (4-)5(-6)- merous. Calyx small, cupular, shortly denied, persistent. Petals shortly connate at base, valvate in bud, afterwards recurved, oblong, persistent in ? flowers. — Flowers: Stamens 5, exserted; filaments linear; anthers small, elliptic, medifixed, cells slightly divergent downwards. Rudiment of ovary hirsute. — 9 Flowers: Staminodes filiform, whether or not present. Ovary ovoid-oblongoid, hirsute; stigma almost sessile, thick, umbonate. Drupe: exocarp coriaceous, thin; endocarp woody, thin, smooth or slightly verrucose and covered with a white membrane inside, sometimes a little reticulate-foveolate outside in fully mature fruits. Seed 1, testa thin; albumen none; cotyledons fleshy. Distr. Two spp., India (Western Ghats from N. Kanara to Travancorc), Lower Burma, Andamans. Cochinchina, Annam; in Malesia: Sumatra, Malay Peninsula. Java, Borneo, Philippines. Fig. 37. KEY TO the species 1. Flowers in interrupted spikes; rachis stout. Petals 3-4 mm \. S. kicinii I. Flowers in slender panicles. Petals c. 2.5 mm. 2. S. pnniiiiliitu 74 Flora Malesiana [ser. I, vol. 7^ 75' 80' 85" 90" 95' 100' 105 110 115 120 125 130 Fig. 37. Distribution of Sarcostigma. 1 and dots S. kleinii, 2 and crosses S. paniciilata. 1. Sarcostigma kleinii W. & A. Edinb. New Phil. J. 14 (1833) 299; Miers, Ann. Mag. Nat. Hist. II, 10 (1852) 116, repr. Contr. Bot. 1 (1852) 103, t. 18; Sleum. Blumea 17 (1969) 254 (with further synonyms). — S. horsfieldii R. Br. in Benn. & Brown, PI. Jav. Rar. (1852) 241, t. 47; Miers, Ann. Mag. Nat. Hist. II, 10 (1852) 118, repr. Contr. Bot. 1 (1852) 105; MiQ. Fl. Ind. Bat. 1 (1856) 795; Schnizl. Ic. 3 (1857/65) t. 172, f. 32 & 33; Hassk. Retzia ed. nov. (1858) 360; Baill. in DC. Prod. 17 (1873) 16; Becc. Malesia 1 (1877) 125; Hochr. Bull. Inst. Bot. Btzg 19 (1904) 39; PI. Bog. Exs. (1904) 16; Koord. Jungh. Gedenkb. (1910) 175; Back. Schoolfl. Java (1911) 229; Koord. Exk. Fl. Java 2 (1912) 533; Koord. -ScHUM. Syst. Verz. 1, Fam. 162 (1912) 5; Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 7; Dahl, J. Arn. Arb. 33 (1952) 276 (pollen); Back. & Bakh. /. Fl. Java 2 (1965) 62. Much branched liana, up to 26 m, stem up to 8 cm 0, sometimes leafless during anthesis; wood hard. Branchlets slender. Leaves oblong to ob- long-obovate or -elliptic, rarely lanceolate, apex shortly rather abruptly and generally bluntly acuminate, or obtuse, base broadly cuneate to rounded, subcoriaceous to coriaceous, glabrous on both sides, or rarely with some hairs beneath, yellowish-green when dry, (7-)9-25 by 4-7 (-11.5) cm, midrib slightly depressed above, strongly raised beneath, nerves (4-)6-8 pairs, curved- ascendent and inarching before the edge, pro- minent beneath, reticulation dense and very prominent (sometimes almost foveolate) on both faces, or more distinct beneath; petiole l-1.5(-2) cm, often recurved, striate as is the midrib beneath and transversely wrinkled in dry specimens. Spikes usually l-3(-5), fascicled, slightly supra-axillary from partly already defoliate axils, or from old wood, pendent, densely shortly brown-hairy to -hirsutulous, .^ 1 5-25, $ up to 40 cm, the flowers in numerous clusters or fascicles with flowerless parts of the rachis between, generally sessile, rare- ly (only in the 4) pedicelled (0.5-1.5, in fruit up to 5 mm), fetid. Calyx 1-1.5 mm, obtusely lobed halfway, hairy below. Petals linear-oblong, sub- acute, densely brown-hairy outside, glabrous and with a slight median ridge inside, greenish- yellow, 3-4 mm. — S Flowers: Filaments strap- shaped, glabrous, c. 5 mm; anthers elliptic, c. 0.8 mm. Rudiment of ovary elongate-conical, hirsute. — S Flowers: Staminodes whether or not present. Ovary ellipsoid-obovoid, hirsute; stigma a reversed subconical or flattish cup. Drupe unequally sub- ovoid-ellipsoid or -oblongoid, laterally compressed and almond-like, 2.5-3.5 by 1.7-2 by 1-1.5 cm, very base suddenly contracted and hidden by the persistent calyx and petals, apex crowned by the thickish small (1 mm) stigma; exocarp very thin, with numerous vertical thick lines standing out in the dry fruit, bright orange red or yellow, brownish-hirsute, glabrescent at length; endocarp thin, 0.5 mm, hard, a little reticulate-lacunose out- side, smooth or faintly verrucose and shiny white in the fresh state inside. Seed 1, with a thin edible sweetish pulp round the nut inside. Distr. India (Western Ghats S. to Travancore), Lower Burma, Andamans, Indo-China (Annam); in Malesia: Malay Peninsula, W. &. E. Java, Borneo (Sarawak: Semengoh F. R.). Fig. 37. Ecol. Forests (also teak-forests), scattered, in Java largely under seasonal climatic conditions, up to 550 m. Fl. fr. Febr.-Oct. Uses. In India the oil of the seeds is highly esteemed in the treatment of rheumatism; this use is not known from Malesia. Vern. Kamaras, tangkilan, J, bua/i pelay tedong, M. 1971] ICACINACEAE (Sleumer) 75 2. Sarcostigma paniculata Pierre, Bull. Soc. Linn. Paris 2 (1897) 1319; Gagnep. F1. Gen. I.-C. 1 (1911) 841. f. 102, 1-3, incl. var. angustifolia (Pierre) Gagnep. I.e. 842; Ridl. F1. Mai. Pen. 5 (1925) 298; Sleu.m. Blumea 17 (1969) 254. — S. angustifolia Pierre, Bull. Soc. Lin. Paris 2 (1897) 1320. — S. philippinensis Merr. Philip. J. Sc. 10 (1915) Bot. 276; En. Philip. 2 (1923) 492; Dahl, J. .Am. Arb. 33 (1952) 276, f. 71 (pollen). — S. surigaoensis Elm. Leafl. Philip. Bot. 10 (1939) 3749; Dahl, J. Am. Arb. 33 (1952) 277 (pollen). — Fig. 38. Climbing to 30 m, stem up to 10 cm o, crooked, branched toward top and forming hanging masses; wood very soft, coarsely porous, i yellow in the centre, watery, with a large whitish pith; bark grey to brown, smooth, rather thin. Branches divaricate, laxly rebranched, the free ends pendent. Leaves scattered along the slender branchlets, oblong or ovate- to elliptic-oblong, rarely lanceolate, apex broadly attenuate, bluntly subacuminate or obtuse, base broadly cuneate to rounded, thin-coriaceous, paler (greyish) green beneath, glabrous, shining on both faces, (8-) 12-20 by (2.5-)4-9 cm, midrib slightly raised above, prominent and longitudi- nally striate beneath, nerves 4-6 pairs, steeply curved-ascending, rather obscurely looping, raised beneath, reticulation dense, finely tessellate on both sides; petiole 1.5-2(-4) cm. ■ — S Panicles axillary or terminal, descending, slender and flexible, much-branched and many-flowered. tomentose initially, early glabrate, 30-60 cm, the lower branches up to 16 cm; flowers 3-5 per cluster, these spicately arranged at intervals of 5-15 mm, with brownish appressed hairs on rachis and more densely around the clusters. Calyx w^idely cup-shaped, membranous, indistinctly den- ticulate, 0.6 mm. Petals narrowly oblong, apex attenuate and slightly inflexed inside, thin, whitish or yellow, practically glabrous, with 3 longi- tudinal veins, c. 2.5 mm. Filaments slender, gla- brous, 1-1.5 mm; anthers elliptic-ovate, 0.7 mm. Rudiment of ovary columnar, hairy. — ? In- florescences (PNH 14563) panicled, glabrate. Flowers sessile, spaced along a rather slender rachis. Calyx cup-shaped, hardly dented, 0.8 mm. Petals oblong, subglabrous, 3 mm. Staminodes 1 mm, no anthers. Ovary oblongoid, densely hairy, 2 mm; stigma a large pad depressed in centre. Infructescence paniculate, stoutish, hairy, c. 10 cm, lower branches up to 4 cm; pedicels c. 2 mm long and across. Drupe (subovoid-)ellipsoid, may- be somewhat compressed, laxly hairy, glabrescent, 3-3.5 by 1.5-2 cm; exocarp blackish-brownish and wrinkled on dry fruit; endocarp thin, woody, smooth inside. Seed 1, white-coated. Distr. Cochinchina; in Malesia: Sumatra (Riouw), Malay Peninsula (Perak), Borneo, Philippines (Leyte, Samar, Mindanao). Fig. 37. Ecol. Primary lowland forest, from sea-level up to 500 m (twice found at 1065 m in Mindanao), on sandy loam, scattered. Fl. Jr. Jan. -Dec. Fig. 38. Sarcostigma paniculata Pierre, a. Habit, X '/2, h. r^ inflorescence, X '/2, r. cJ flower, XlO, d. rj flower bud, x 10, <-. fructification with almost mature fruits in dry stale, X '/2 (o-^SAN 24459, e SAN 17107). 76 Flora Malesiana [ser. I, vol. 7^ 19. PYRENACANTHA Wight in Hook. Bot. Misc. 2, 4 (1830) 107, Suppl. t. 9 & 10, nom. cons.; Sleum. Blumea 17 (1969) 249. — Freeria Merr. Philip. J. Sc. 7 (1912) Bot. 292. Dioecious scandent shrubs or lianas. Leaves spirally arranged, glandular-repand- dentate and penninerved. Flowers small, sessile, bracteate, in lax supra-axillary spikes. Sepals or calyx 0. Petals connate at base, valvate in bud, persistent in ? flowers. — 6 Flowers: Stamens (3-)4(-5); filaments very short; anthers minute. Rudiment of ovary present. — $ Flowers: Staminodes very short or absent. Ovary sessile; stigma sessile, thick, truncate. Drupe laterally compressed; exocarp fleshy; endocarp crustaceous, punctate outside, verrucose-papillose inside, the papillae projecting into the interstices of the deeply ruminate and pitted albumen. Seed 1 . Distr. About 20 spp., most of them in tropical and subtropical Africa & Madagascar, 1 sp. {P. voliibilis Wight) in S. India and Ceylon, Indo-China and Hainan; in Malesia: 1 sp. in the Philippines. 1. Pyrenacantha repanda (Merr.) Merr. En. strigose-pubescent, (2-)4-8 cm; flowers scattered Philip. 2 (1923) 492; Sleum. Blumea 17 (1969) in the upper half or less, each flower subtended by 249; Shaw, Kew Bull. 23 (1969) 113. — P. sp., an oblong acuminate small bracteole. Petals 4, Ceron, Cat. PI. Herb. Manila (1892) 46. — ovate-oblong, pubescent outside, 1.5-2 mm. — Freeria repanda Merr. Philip. J. Sc. 7 (1912) Bot. S Flowers: Filaments very short; anther cells ovate- 292. — Tragia luzoniensis Merr. Philip. J. Sc. 16 elliptic, hardly 0.5 mm. Rudiment of ovrt/-y hirsute, (1920) 564. subglobose. • — $ Flowers: Staminodes minute. Woody vine, up to 8 m. Branchlets slender, Ovary ellipsoid, densely pubescent, 1 mm; stigma minutely verrucose, the growing parts short- round, truncate, minutely papillate. Drupe elliptic- pubescent. Leaves oblong to oblong-lanceolate, oblong, laterally compressed to rather sharp apex slenderly acuminate, bluntish, base narrowed, edges, glabrous, yellowish or red, 1.8-3 by 1-1.5 very base cordate, subcoriaceous, glabrous and by c. 0.8 cm; exocarp pulpy or fleshy, rather thin, smooth above (though very finely papillate under wrinkled in dry fruits. Seed I; endosperm ruminate, the lens), set with scattered appressed strigose hairs yellow, oily, externally deeply pitted by the pro- (whose rounded bases are partly pellucid against tuberances on the inside of the thin endocarp. strong light) and scabrid to the touch beneath, Distr. Malesia: Philippines (Luzon: Ilocos 6-14 by 1.5-4(-5) cm, margin ± repand with a Norte, Bulacan, Tayabas, Sorsogon; Samar, Min- short glandular tooth at the excurrent end of each danao). lateral nerve, midrib and nerves prominent on Ecol. Primary forest at low altitudes, scattered, both faces, the latter in 5-6 curved-ascendent and Fl. March-May, fr. Sept. anastomosing pairs, reticulation rather lax and Note. Apparently close to P. volubilis Hook. slightly raised on both sides; petiole usually and to certain species from Madagascar in leaf- strongly curved, glabrescent, 5-15(-30) mm. characters, distinguishable from the first by its Spikes supra-axillary, slender, minutely sub- larger fruits. 20. MIQUELIA Meisn. pi. Vase. Gen. 1 (Sept. 1838) 152; ibid. 2 (Sept. 1838) 109; Sleum. Taxon 17 (1968) 449 et ibid. 18 (1969) 479, nom. cons, prop.; Blumea 17 (1969) 229, non Bl. June 1838 (= Stauranthera Benth. 1835). — Jenkinsia Griff. Calc. J. Nat. Hist. 4 (1843) 231; ibid. (1844) t. 12, non Hook. Gen. Fil. (1842) text opp. t. 75 b. — Fig. 39. Dioecious climbing or twining shrubs, the wood with large porous vessels. Leaves spirally arranged, entire or coarsely lobed (Mai.), generally minutely papillose-tubercled on both faces, nerves palmate to pinnate. Flowers sessile, arranged in heads or umbels; these on ± elongate, distally i thickened peduncles, from defohate or foliate axils; inflorescences solitary or several in lax racemes, panicles or fascicles. Calyx small, + deeply 4- or 5-lobed. — d* Flowers: Petals 4 1971 ICACiNACEAE (Slcumer) 77 Fig. 39. Miquelia celebica Bl. a. Habit, X 1/2, b. ^ flower bud, X 5, c. petal, X7V2, d- & d'. stamen, x7VS, e. submature infructescence, x Vz,/- mature fruit, with remnants of the petals at the very base, X 1. — .\f. caudata King. g. Fruit, x 1 {a-d Elmer 9894, e Kruyff 23,/Koorders 19594, g Clemens 27608). or 5. united into a short or mostly elongate-filiform tube adnate to the flower axis, upper part or lobes free, valvate in bud, apex a little inflexed, patent or ~ reflexed in anthesis. Stamens 4 or 5; filaments short-linear; anthers elongate- linear-oblong to (sub)sagittate, exceeding the filaments in length, dorsifixed a little above the base, introrse. Rudiment of ovary minute. — + Flowers: Calyx and lobes of the petals either distant from each other by the elongate flower axis, or close together, i.e. practically no flower axis present. Staminocies flattish, very short. Ovary thick-cylindric, hairy; stigma sessile, disk-shaped, its centre a little depressed. Drupe oblongoid to subovoid-ellipsoid, laterally compressed, whether or not narrowed into a stalk-like base, either sessile or on a pedicel- like flower-axis; mesocarp thin; endocarp crustaceous, rugose or shallowly la- cunose, i manifestly verrucose inside, the warts causing small pits in the flattened ovoid seed which is surrounded by a juicy pulp; testa thin; albumen fleshy, ruguiose; cotyledons elliptic, thick-foliaceous. Distr. About 8 spp. in SE. Asia (India: Western Ghats; Assam, Burma?, Thailand, Laos); in Malesia: A spp. (Sumatra, Malay Peninsula, Borneo, Philippines, Celebes). I ig. 40. Ecol. Mainly lowland forest, rarely up to 1980 m (Mt Kinabalu). KEY TO THE SPECIES 1. Calyx and lobes of the petals distinctly spaced both in the V and J flowers by the elongate flower-axis (— the basal connate tube-like part of the petals). Drupe 'pseudopedicclled" by this axis (and besides much narrowed towards the base into a kind of stalk). 2. Leaves oblong to subovate-oblong, (10 )i2 17 by 4-6.5 cm; inner basal pair of nerves ascending to the lower third of the lamina. Umbels c. lO-flowercd I- M. rcticiilalu 2. Leaves ovate, 7-11 by 5-9 cm; inner basal pair of nerves ascending to V2 or J/i the length ol the lamina. Umbels c. S-flowcrcd 1. M. phllippinensis 78 Flora Malesiana [ser. I, vol. 7^ 1. Calyx and lobes of the petals close to each other in the ? flowers and drupe, spaced in the ^ flowers. Drupe not 'pseudopedicelled' (but much narrowed towards the base into a kind of stalk). 3. Drupe 2.3-3.3 by 1 .3-1 .8 by 0.7-1 cm, narrowed at apex into a shorter or longer rather slender beak; endocarp coarsely lacunose 3. M. caudata 3. Drupe (0.8-)l-1.2(-1.7) by 0.7-0.8(-1.2) by 0.6 cm, shortly and rather gradually narrowed at apex; endocarp more finely lacunose 4. jyt. celebica 1. Miquelia reticulata Merr. Philip. J. Sc. 9 0914) Bot. 312; En. Philip. 2 (1923) 493; Sleum. Blumea 17 (1969) 230. Scandent. Branchlets slender, sparingly hirsute with short hairs, as are the petioles. Leaves oblong to subovate-oblong, apex gradually attenuate or subacuminate, base rounded and a little oblique, very base shortly cordate, membranous, glabrous and smooth above, set with scattered hairs all over the undersurface and somewhat rough to the touch, entire, (10-)12-17 by 4-6.5 cm, midrib prominent beneath, nerves 2 basal pairs, the outer pair short, the inner one ascending to the lower third of the lamina, other higher nerves from the midrib 3-4(-5) pairs curved-anastomosing, slightly depressed above, raised beneath, reticulation coarse and faint above, a little prominent beneath; petiole (l-)1.5-2.5(-5.5) cm by 1-2 mm. —^ In- florescences unknown. — $ Flowers umbellately arranged on top of a slender peduncle, the latter either solitary and axillary, or mostly 2-4 pe- duncles from a short thickened rachis on defoliate stem, 3-10 cm, with about 10 flowers per umbel. Calyx 4-lobed to almost the base, lobes hairy, c. 1 mm. Petals 4, greenish-yellow, fleshy, lobes elongate-oblong, subacuminate, laxly puberulous outside, glabrous inside, c. 3 by 1.5 mm, tubular basal part of the petals 5-7 mm at full anthesis. Stamens 4, flattened, very short. Ovary hairy, crowned by a thick pad-like stigma. Drupe narrowly ovoid in general, somewhat compressed, c. 2 by 1.3 by 0.8 cm, wrinkled when dry, suddenly attenuate at the very base into a kind of stipe (5-8 mm), on the base of which the lobes of the petals persist for some time; flower-axis below these lobes 6-8 mm, slightly more slender than the basal stipe of the drupe; exocarp thin, densely hairy; endocarp thin, coarsely foveolate outside, finely tubercled inside. Distr. Malesia: Philippines (Camiguin and Bohol Is.), twice found. Fig. 40. Ecol. Forest at low altitudes, up to 600 m. 2. Miquelia philippinensis Merr. Philip. J. Sc. 14 (1919) Bot. 416; En. Philip. 2 (1923) 493; Sleum. Blumea 17 (1969) 230. Scandent shrub. Branchlets slender, laxly hir- sute with shorter and longer hairs, as are the petioles. Leaves ovate, apex broadly subacuminate- attenuate, base rounded and very shallowly or hardly cordate, membranous, glabrous and oliva- ceous in dry state above, rather laxly and finely short-hairy on the undersurface, entire, somewhat repand or shortly (c. 5 mm) 1-2-lobed on each side, 7-11 by 5-9 cm, midrib and nerves faint above, a little raised beneath, 2 pairs of basal nerves, the outer pair rather short, the inner one Fig. 40. Distribution of Miquelia. 1. M. kleinii, 2. M. reticulata, 3. M. philippinensis, 4. M. cau- data, 5 and 2 dots M. celebica. ascending to Vz-Vi the length of the lamina, upper lateral ones c. 3 pairs, curved-anastomosing, re- ticulation coarse and faint, less obvious than in M. reticulata; petiole 2-3 cm by 0.5-1 mm. — (5 Inflorescence unknown. — 2 Inflorescence axillary, solitary peduncles c. 3 cm, each bearing c. 5 umbellately arranged flowers. Calyx lobes ovate-acuminate, hairy, 1-1.5 mm. Petals united below into a c. 5 mm long tube, free lobes oblong, subglabrous, 2.5 mm. Submature drupe oblongoid- ellipsoid, slightly hairy, 1-1.3 cm long, narrowed downwards to a slender stalk as long as the fruit proper. Distr. Malesia: Philippines (Luzon: Ilocos Norte Prov.), once found. Fig. 40. Ecol. Forest at low altitude. 3. Miquelia caudata King, J. As. Soc. Beng. 64, ii (1895) 124; Ridl. F1. Mai. Pen. 1 (1922) 433; Burk. Diet. (1935) 1478; Sleum. Blumea 17 (1969) 230. — Fig. 39g. A slender climber up to 6 m. Branchlets pale, striate, early glabrescent and corticate, 2-5 mm o. Leaves oblong-lanceolate to elliptic-oblong, apex shortly, sometimes rather abruptly acuminate, base broadly cuneate to rounded, thin-charta- ceous to membranaceous, glabrous above, laxly puberulous all over the surface and soft to the touch beneath, glabrescent, entire, (10-)12-20 by 1971 ICACINACEAE (Sleumcr) 79 (4-)5-8 cm, midrib and nerves moderately raised beneath, nerves 1 or 2 basal and 5-6 upper pairs, equally curved-ascending and rather obscurely anastomosing, reticulation generally lax and but slightly prominent: petiole pubescent, 1.2-3(— 4) cm. — Umbels of o inflorescences solitary or most- ly several, in fascicles or along a short rachis from hair>' tubercles of foliate or defoliate axils of the branchlets; peduncle filiform, 1.5-3 cm, the whole inflorescence set with short almost scabrid hairs. Calyx cup-shaped, 1 mm, 4(-5)-lobed halfway. Petals 4 (or 5) united to a filiform elongate basal tube-like (4-10 mm), free in the upper oblong and rr spreading part (2 mm), white. Stamens 4 or 5; filaments short; anther cells elongate-linear, subsagittate, 1.5 mm. Rudiment o^ ovary hirsute. — i Flowers 8-15 together in capitules on top of axillary, solitary stoutish peduncles, these gla- brescent and accrescent in fruiting stage up to 11 cm. Calyx as in o flowers. Petals without a marked- ly elongate basal part, i.e. close to the calyx, some- what leathery and glabrous, 2.5 mm. Ovary to- mentose; stigma discoid, wider than the ovary. Drupe broadly ovoid, laterally much compressed, rounded at base in general, though abruptly nar- rowed to a slender stipe for 5-7 mm, at the base of which the remnants of the petals and calyx are found, tapering upwards to a tail or beak for (5-)6-IOmm, crowned by a small stigma, 2.3-3 (-3.3) by 1.3-1.8 by 0.7-1 cm; exocarp thin, rusty-pubescent, ripening to a tomato or orange red; endocarp bony, rather deeply (1-2 mm) and coarsely reticulate-lacunose outside, tubercled all over inside. Distr. Malesia: Malay Peninsula, Borneo. Fig. 40. Ecol. Primary forest, also mossy forest, from the lowland up to 1980 m on Mt Kinabalu. Uses. The stem holds a reservoir of potable water. Vaughan-Steffens stated that the red fruiti are added to Antiaris latex in making 'sakai (pangan)' dart-poison (Ridl. Agr. Bull. Mai. Pen. 8, 1898, 209), but no confirmation of this has been available. Vcrn. Pisang-pisang bulu, M, sHowung, Pangan. 4. .Miquelia celebica Bl. Rumphia 4 (1849) 37; Mus. Bot. Lugd.-Bat. 1 (1849) 42, f. 8; Walp. Ann. 2 (1851) 22; Miq. FI. Ind. Bat. I (1856) 798; ScHMZL. Ic. 3 (1857/65) t. 172. f. 22-31; Baill. in DC. Prod. 17 (1873) 14; Becc. Malesia I (1877) 124; KfK>RD. Minah. (1898) 393; Slelm. Blumca 17 (1969) 231. — A/, cumingii Baill. Adansonia 10 (1872) 278; in DC. Prod. 17 (1873) 14; Vidal, Phan. Cuming Philip. (1885) 103; Rev. PI. Vase. Filip. (1886) 86; Merr. En. Philip. 2 (1923) 492; Dahl, J. Arn. Arb. 33 (1952) 275, f. 74 (pollen). — M. rostrata Merr. PI. Elm. Born. (1929) 172. — Fig. 39a-f. High climbing shrub, much ramified. Branchlets slender, green-olivaceous when dry, striate, sub- glabrous. Leaves elliptic-, sometimes ovate-, rarely lanceolate-oblong, apex shortly acuminate, tip submucronate by the protruding midrib, base a little oblique, broadly cuneate to truncate-rounded, sometimes shallowly cordate, thin-chartaceous, entire, or rarely obsoletely repand to coarsely few- lobed, initially with scattered rather scabrid short hairs (also found on the inflorescence) underneath, early glabrescent, 9-15(-20) by 4-7.5(-10) cm, midrib and nerves slightly raised beneath only, lateral nerves 1-2 basal and 4-6 higher pairs, all curved-ascendent and rather obscurely in- arching before the edge, no proper reticulation; petiole slender, l.5-5(-7) cm. — o Umbels sub- globose, many-flowered, slenderly long-peduncled, axillary or slightly supra-axillary, rarely cauline, either solitary or 2-3 very laxly racemosely ar- ranged on a rachis of 1-3 cm by c. 2 mm. Calyx shortly 4(-5)-lobed, very small. Petals 4(-5), united for 5-7 mm to a filiform tube below, free for the uppermost 2 mm, greenish yellow. Stamens 4(-5); filaments 1 mm; anther cells linear-oblong, base slightly divergent, 1.5 mm. Rudiment oi ovary 4-gibbous, glabrous. — 2 Flowers arranged to many- and dense-flowered subglobose heads, these generally solitary from the axil on a slender pe- duncle 5-10 cm, rarely 2 or 3 laxly racemosely arranged along a short rachis, each head then very slenderly elongately peduncled. Calyx very small. Petals 4, not elongate into a tube below as in the cJ, oblong, thickish, 2 mm. Staminodes 4. Ovary subcylindric, densely hairy, crowned by the thick stigmatic pad. Drupe obliquely ellipsoid-ovoid, laterally compressed, (0.8-)l-1.2(-1.7) by 0.7- 0.8(-1.2) by 0.6 cm, apex shortly and rather grad- ually attenuate, base suddenly narrowed to a kind of stipe 5-7 by 1 mm (bearing the persistent petals on its base); exocarp tiiin, laxly set with short strigose hairs, yellow to reddish; endocarp crustaceous, outside with numerous vertical low ridges and transverse connections, shallowly lacunose-reticulate in the dry fruit, inside with numerous low warts which leave their mark in the thus pitted seed. Distr. Malesia: W. Sumatra (Padang, once found), Borneo, Philippines (Luzon to Min- danao), Celebes (Minahasa, Buton I.). Fig. 40. Ecol. Primary forest, sometimes along streams, generally in the lowland, rarely up to 700 m, in Buton I. on coralline limestone, apparently scattered or rare. Vern. Lalai in lasik, sumhilan makanteh, Minahasa (Tt.). 21. PHYTOCRENE Wai.i.. pi. As. Rar. 3 (1831) II, I. 16; Sleum. Taxon 17 (1968) 448 et ihicl. 18 (1969) 479, twm. cons. prop.\ Blumca 17 (1969) 234. — Gynocephalum Bl. Bijdr. (1825) 483; Endl. Gen. (1837) 281 rGynocephalium'): Rriciin. Norn. (1841) 80 Flora Malesiana [ser. I, vol. 7^ 67 {"Gyrocephaliumy, Hassk. Cat. Hort. Bog. (1844) 78 {'GynaecocephaliunC); Bl. Rumphia 4 (1849) 36, tiota {"Gynocephaliufn); Benth. in Benth. & Hook./. Gen. PI. 1 (1862) 354 {'Gynocephala'). — Fig. 42. High climbing dioecious shrubs or lianas, sometimes with a robust stem, younger parts of the latter usually hairy and prickly, youngest parts slender and striate, sometimes rolled-in, and simulating tendrils, which are wanting. Leaves remotely spirally arranged, entire or palmately 3-7-lobed, subpalmately 3-7- nerved; petiole from a thickened and depressedly cup-like part of the stem, ± torted, never swollen basally or distally. — o Flowers in globose clusters, these either solitary at the end of an axillary peduncle, or several spaced and racemosely arranged then, or generally very numerous in long repeatedly branching spike-like racemes or panicles both axillary and/or stem-borne. Calyx segments 3-5, rz free. Petals 3^, dn united below, valvate, alternating with the calyx lobes. Stamens 3^ (rarely -6), finally exserted; filaments filiform, ± connate at base into a column which bears the hirsute rudiment of the ovary; anther cells ovate-elliptic, small. — $ Flowers in large solitary either cauhne or mostly axillary peduncled globose heads. Calyx segments 3-5, free to almost the base. Petals 3-4(-5) as in the 6 flowers. Stamuiodes absent. Ovary subovoid-columnar, tapering to an equally thick and short columnar style with several 2-lobed stigmas which form a kind of funnel. Drupes numerous, collected in generally large, globose heads, each drupe tipped by the style, ± reversely setulose by brittle irritant hairs; exocarp thin, larger than the endocarp and i hollow in the upper extended part at full maturity; endocarp rather thick, hgneous, pitted to deeply lacunose outside, smooth or warty inside. Seed 1, large, with a thin almost juicy coat which exudes a kind of clear gum; endosperm fleshy, strongly furrowed or lobulated; cotyledons very large, flat, foliaceous, double-folded or tortuous. Distr. About 11 spp., of which 4 both in SE. Asia and Malesia, and 7 confined to Malesia from Sumatra to New Guinea. Fig. 41. Ecol. Scattered in primary and secondary lowland forest, occasionally up to 1200 m, also in open places, as rarely found on limestone rocks. Uses. The wood has very large porous vessels which contain water, used by the natives, hence the generic name; for its anatomy see the comprehensive study made by Timmermans, Ann. Jard. Bot. Btzg 41 (1931) 65-104, t. 23-34, and the review given by Sleumer in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 332-336, f. 90-93. Note. Leaves of Phytocrene are similar in shape and nervation to those of various genera in the Menispermaceae; besides the petiole emerges from a shallow cup-like thickening of the stem which also occurs often in Menispermaceae. In the latter, however, generally the petiole is swollen in the uppermost and maybe also in its basal part, which is never the case in Phytocrene. KEY TO THE SPECIES 1. S Flower heads solitary, each on top of an axillary or slightly supra-axillary peduncle (as is always the case with the 2 ones) on younger foliate parts of the branchlets 1. P. anomala 1. (J Flower heads several together, composed to an inflorescence, the latter either from foliate or de- foliate axils, or from tubercles on the stem. 2. c? Flower heads rather few, composed to a true raceme, i.e. the peduncled heads spaced along a rachis 2. P. racemosa 2. S Flower heads numerous, composed to a spike-like panicle. 3. S Flower heads in fascicles or glomerules, these markedly spaced at least in the lower half of the rachis. 4. Leaves broadly oblong, base slightly cordate, flat in dry specimens 3. P. intemipta 4. Leaves ovate, base rounded, ± bullate by impressed nerves and veins in dry specimens. 4. P. malacothrix 3. c? Flower heads in short-peduncled cymes, secondary racemes or panicles, these generally close together (rarely spaced) along the rachis. 1971] ICACINACEAE (Slcumer) 81 5. Bracts in the £ inflorescences persistent, markedly plumed by i spreading hairs. 6. Basal part of bracts hardly or not widened 5. P. trichura 6. Basal part of bracts widened up to 5 mm 6. P. borneensis 5. Bracts in the S inflorescences wanting (already gone in full anthesis), or if persistent, filiform to narrow-subulate, and not markedly plumed. 7. ^ Inflorescences rather short, 2.5-5 by (0.5-)l-1.5 cm. Leaves normally oblong or oblong- lanceolate, early glabrescent 7. P. oblonga 7. 3 Inflorescences elongate to caudate, i.e. markedly longer and at least 2 cm 0. Leaves oblong to ovate-cordate, or palmate, generally tardily or hardly glabrescent. 8. Leaves normally palmate, i.e. deeply and narrowly (3-)5(-7)-lobed in flowering specimens. 8. P. palmata 8. Leaves normally entire or ± broadly 3-lobed in flowering specimens (sometimes palmate in sterile juvenile specimens). 9. o Inflorescences slender, c. 2 cm o. Bracts — persistent, numerous, much exserted beyond the partial inflorescences. Drupe 4.5-8 by 1.5-2.3 cm incl. the hollow beak of the exocarp. 9. P. bracteata 9. (J Inflorescences thickish, (2.5-)3 cm or more 0. Bracts, if persistent, rather few and less ex- serted. 10. Mature drupe 3^(-4.5) by 1.5-2 cm incl. the hollow short beak of the exocarp; hard endo- carp (2-)2.5-3(-3.5) cm long 10. P. hirsuta 10. Mature drupe (5.5-)6-10(-13) cm long incl. the hollow elongate beak of the exocarp. 11. P. macrophylla s.l. 1. Pbytocrene anomala Merr. J. Str. Br. R. As. Soc. n. 86 (1922) 325; Sleum. Blumea 17 (1969) 235. Woody vine. Branchlets (slender, up to 5 nrmi 0, their distal part often curved and similar to a tendril), petioles, peduncles and leaves (mainly on the nerves of the undersurface) hirsute by short greyish and longer fl-2rrun) rufous and more spreading hairs, ultimately glabrescent. Leaves ob- long to broadly oblong, or lanceolate, apex short- acuminate, subacute, base slightly cordate, char- taceous, olivaceous, glabrous or almost so except the hirsute midrib above, the tomentum on the undersurface rather soft to the touch, denticulate by protruding veins in young, entire in mature leaves, l(>-22f-30) by (2.5-)3-10 cm, nerves 6-8(-10) pairs, two of them ± basal, short and spreading, the other ones curved-ascending, raised beneath only as is the coarse net of veins and veinlets; petiole l-2(-3)cm by 2-3 (-4) mm. — ^ Inflorescences in globular peduncled heads, solitary, from a little above the axils of subsequent leaves of the distal part of a branchlet; peduncle slender, c. I cm; heads covered with a pale fcr- rugincous tomentum of short bristly hairs, 5-8 mm 0. Calyx segments 4 or 5, linear-spathulate, c. 2.5 mm. Petals 4, clongate-obovate, connate halfway to V*, hirsute dorsally, c. 3 mm. Stamens 4; filaments 1.5-2 mm; anther cells subovatc- elliptic, I mm. — i Inflorescences as the ^ ones on young branchlets in axillary solitary heads, on peduncle c. 2 cm by 2 mm. Calyx segments and petals as in J. Ovary ovoid, short-attcnuatc to a conical very shortly lobcd stigma, hirsute. Drupes collected to heads of c. 4 cm 0, each drupe ovoid, hirsute by simple bristly hairs all of the same type, known up to 2.5 by 1.5 cm, apparently larger at full maturity, accrescent petals seen up to 6 mm. Form, size and type of pits of the cndocarp not yet known. Distr. Malesia: Borneo (Upper Baram R. and Sandakan area, Bcrau). Fig. 41. Distribution of Phytocrene. The figure above the hyphen indicates the number of en- demic species in the island or area, the figure below the hyphen the number of non-endemic species. Ecol. Primary and secondary lowland forest, apparently rare. 2. Phytocrene racemosa Sleum. Blumea 17 (1969) 236. — Fig. 42. Tall woody climber, up to 40 m, stem c. 1 .5 cm o covered with thin longitudinally striate and finely fissured brown bark which bears a few transverse linear Icnticcis; branchlets and petioles with a short fawn tomentum. Leaves oblong-obovate, apex abruptly (c. 1.5 cm) acuminate, acutish. base broadly cuncate and shallowly cordate, charta- ccous to subcoriaccous, glabrous except the nerves which are hairy in their basal part above, • den- sely set with short subapprcssed or more spreading pale rusty hairs especially on nerves and veins and soft to the touch beneath, edge slightly sinuate- denticulate, or entire, 18-32 by 7-20 cm, nerves 82 Flora Malesian [ser. I, vol. 71 (a, e MuRTHY & AsHTON S 22325,' jt;69 by 2-4 mm. — ^ Racemes from tuberclp<: «f fh^ heads « m /f "''• 'rS"'°''' P'luncled (1 cm) sPa.h.,a,e, Cg^C^,' fl'/XrV'.S; 1971 ICACiNACEAE (Sleumer) 83 oblanceolate, free to almost the base, long-haired dorsally, 2.5 mm. Stamens 4; anther cells 0.4 mm. — i Racemes from c. 1 cm above defoliate axils, with about 5 laxly arranged heads, fulvous-hirsute at peduncles and the c. 4 cm long rachis. Flower heads globose, c. 5 mm o, on thick peduncles 1.5-2 cm. Calyx segments and petals as in c? flowers. Ovary cylindric, hirsute, c. 1 mm. De- veloped ovary or immature drupe (c. 1 cm long) densely covered with short and shortly hirsute emergences in the upper, and longer and longish- hirsute ones in the lower half. Mature drupe ar- ranged to a globose head of c. 8 cm o, each drupe showing the same shorter and longer emergences of the exocarp as already found in the very young state, subelliptic-oblongoid, attenuate at both ends, 4-5 by 2-3 cm; endocarp oblongoid-ellipsoid, ob- tuse at both ends, much compressed laterally, smooth, with numerous almost linear rather deep pits, which appear on the inner side as flat warts, c. 3 by 1.5 by 1.3 cm. Distr. Malesia: Borneo (Sarawak), 3 col- lections. Ecol. Mixed Dipterocarp forest, from low elevation up to 250 m, apparently very scattered. Vern. Bua/i pisang munsang, Kapit. 3. Phytocrene interrupta Sleum. Notizbl. Berl.- Dahl. 15 (1940) 252; Blumea 17 (1969) 236. Climber. Branchlets slender (5-7 rrma o), deeply striate and brownish-hirsute, older parts laxly set with hardly pungent small emergences. Leaves broadly oblong, apex for 3-10 mm subabruptly acuminate, base slightly cordate, firmly charta- ceous, glabrous above except nerves, covered with soft hairs all over the undersurface, and with stiffer ones on midrib and nerves, soft to the touch, entire, 7-17 by 4-8(-9) cm, midrib and nerves flat or a little impressed above, markedly prominent beneath, nerves 2 basal pairs (of which the outer pair is short, the inner one ascending to the lower Yi of the length of the lamina), other pinnate nerves 6-8 pairs, rather straight and sub- parallel to each other and the inner basal pair, reticulation of veins and vcinlcts fine and hardly raised above, more coarse and much raised beneath; petiole 2-3.5 cm by 2-3 mm, hirsutulous. — S Inflorescences spike-like, several in a fascicle from a tubercle of the stem (c. 1.5 cm 0), pendent, the heads of flowers sessile or very shortly pe- duncled and (2-)3-6 of them fascicled, these fasci- cles spaced in the lower, less or not so in the upper half of the rachis (10-30 cm by 2 mm), the whole inflorescence covered with shorter soft and longer stifl'-brownish hairs; peduncle of flower heads slender, up to 4 mm, the basal bract subulate, 2-3 mm, apparently early going. Calyx segments 4, ovate, fleshy, concave, hirsute, 0.8 mm. Petals 3 cr 4, connate to the lower Vi to V2> ovatc- oblong, subacuminutc, green, J: densely substrigosc dorsally, r. 1.5 mm. Stamens 3 or 4; anther cells ovatc-clliplic, 0.4 mm. - V Inflorescences and drupe unknown yet. Distr. Malesia: New Guinea (Central Distr. in ascent from Port Moresby to Mt Victoria), once found. Ecol. Forest at c. 450 m. 4. Phytocrene malacothrix Sleum. Notizbl. Berl.- Dahl. 15 (1941) 361; Blumea 17 (1969) 237. "Scandent. Branchlets soft-olivaceous-tomentose in the younger parts, glabrescent below. Leaves ovate, apex subabruptly acuminate for I cm, tip obtuse, base rounded, thin-chartaceous, glabrous above, softly brownish-tomentose all over beneath especially on midrib and nerves, 16-18 by 8-10 cm, midrib raised on both faces, nerves curved-as- cendent, ± distinctly impressed as are the veins above (the lamina buliate in dry specimens), much raised beneath, reticulation rather dense, slightly prominent beneath only; petiole tomentose, 3-4 cm. (J Spikes only known in a poorly developed state, brownish-tomentose, solitary from the axils of the upper leaves, the flower heads composed to glomerules, the latter spaced on a rachis c. 5 cm long." Distr. Malesia: New Guinea (Morobe Distr.: Boana), at 1030 m, once found. Note. Imperfectly known species, said to be related to P. interrupta Sleum. in the original diagnosis, from which the description given above was made; type material lost. 5. Phytocrene trichura Ridl. F1. Mai. Pen. 1 (1922) 433; Sleum. Blumea 17 (1969) 237. Tall woody climber. Stem of branchlets shallow- ly striate, patently rufous-setose distally as are the petioles, corky and with numerous roundish len- ticels (no prickles) downwards, 4-6 mm 0. Leaves ovate in circumference, whether or not 3-lobed to Vi-Vi, lateral lobes acutish, base shallowly cor- date, thin-coriaceous, subdensely short-tomentu- lose on both faces initially, glabrescent and a little rough above, soft to the touch beneath, 12-19 by 8-1 1 cm, palmately 5-nerved, midrib and inner pair of basal nerves more prominent than the other basal and the few upper lateral ones beneath, reticulation coarse, strongly raised be- neath; petiole 3-6 cm by 2-3 mm. — cj Panicles from bare lower part of stem (1.5 cm 0), sessile, pendulous, grcy-grccn, squirreltail-like, 12-23 by 3.5-4 cm, formed by numerous pcduncled dicha- sia, the ultimate pedicels (3-6 mm) bearing globular heads consisting of numerous flowers, all densely shortly greyish-pubescent; bract of each globule partly connate with pedicel, slender, almost fili- form, persistent, spreading, its basal part hardly or not widened, short-pubescent, upper part plumose by spreading rufous 2-3 mm long hairs. Calyx segments 4 or 5, irregularly obcuncatc, I mm, free, apex fringed with long hairs. Petals 4, ovate-oblong, connate to the lower Y), sparsely short-hairy or papillose outside, 1.5 mm. Stamens 4, slightly cxscrtcd; anther cells elliptic, 0.5 mm. — $ Iriflorescences and drupe not yet known. Distr. Malesia: Malay Peninsula (Perak, Pa- hang, Selangor). Ecol. Dense forest, ascending the top of lofty trees, apparently very rare. 84 Flora Malesiana [ser. I, vol, 7^ 6. Phytocrene boraeensis Becc. Malesia 1 (1877) 128; Merr. En. Born. (1921 ) 356; Sleum. Blumea 17 (1969) 237. — P. porphyrea Staff, Kew Bull. (1906) 72. Big woody climber. Stem robust (seen up to 2.5 cm 0), longitudinally ribbed (2-3 mm) and with distant transverse linear lenticellar cracks. Branchlets ~ patently rufous-setose and striate at distal part (3-6 mm 0). Leaves ovate in cir- cumference, simple, or subtrilobed, or deeply obtusely 3-lobed, base cordate, edge very shortly denticulate by protruding veins, chartaceous to subcoriaceous, young ones soft-tomentulose all over both faces, matureones finally glabrous above, ± densely covered with appressed to suberect rather stiff hairs especially on nerves, and soft to the touch beneath, 15-25(-30) by (8-) 10- 18 (-22) cm, nerves slightly sunken above, strongly raised beneath, basal palmate nerves 7-9 pairs, branched mainly outward, upper ones from the midrib Ti-A pairs of entire, and 4-6 ones in the central lobe of lobed leaves, all these nerves curved-ascending, connected by numerous straight and ± transverse veins and a coarse reticulation markedly prominent underneath only; petiole subappressedly rufous to ferrugineous-setulose, (3-)4-8(-12) by 2-4 mm. — c? Inflorescences pendent from old wood, tail-like as in P. trichura, sessile, compact, consisting of numerous repeatedly branched and shortly pedicelled dichasia, all over covered with greyish to brownish softer and coarse, in part hooked hairs, (12-) 15-25 (-50) by 3-4(-5) cm, most of the ultimate pedicels bearing a narrowly to broadly subulate-caudate, (1.2-)2-3.5 cm long and at base (l-)2-5 mm broad persistent bract, the latter covered with the same short to- mentum and besides with longer, spreading, often dark red or brownish hairs which give the bracts a ± plumose appearance; flowers in globular pe- dicelled heads (3-4 mm 0). Calyx segments 4 or 5, irregularly obcuneate to spathulate, distally densely hairy outside, glabrous inside, 1 mm. Petals 4, oblong-obovate, acutish, fleshy, slightly to densely hairy outside only especially below, 1.5 mm. Stamens 4; anther cells oblong-elliptic, 0.5 mm. — ? Inflorescence and drupe unknown. Distr. Malesia: Borneo. Ecol. Lowland forest, up to 1065 m. Uses. Water from the stem is used for drinking. Vern. Bills akar, S. Kinabatangan. 7. Phytocrene oblonga Wall. PI. As. Rar. 3 (1831) 12; Cat. (1832) n. 4948; Walp. Rep. 1 (1842) 98; Baill. in DC. Prod. 17 (1873) 13; Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 592; King, J. As. Soc. Beng. 64, ii (1895) 122; Gagnep. Fl. Gen. L-C. 1 (1911) 839, f. 102, 4-8; Ridl. Fl. Mai. Pen. 1 (1922) 432; Craib, Fl. Siam. En. 1 (1926) 275; Gagnep. Fl. Gen. L-C. Suppl. (1948) 757; Sleum. Blumea 17 (1969) 237; Fl. Thail. 2 (1970) 90. — Gynocephaluin oblongiim (Wall.) Trecul, Ann. Sc. Nat. Ill, 8 (1847) 149. — Miquelia cancellata KuRZ, J. As. Soc. Beng. 44, ii (1875) 201. Climber to top of tall trees, with rather slender stem (up to 1.5 cm 0); bark brownish, rather rough, striate, not prickly. Branchlets densely set with fine retrorse prickly lenticels, tip short- pubescent. Leaves oblong or oblong-lanceolate, i abruptly and shortly acuminate, acumen rather slender and slightly curved, subacute, base cu- neate to rounded, coriaceous, glabrous and shining above, rather densely stiffly short-hairy and a little rough to the touch beneath, early glabrescent, entire or undulate-subdentate, (10-) 12-22 by (4-)5-9 cm, nerves 5-7 pairs, one of which basal, ascending and ± manifestly in- arching before the edge, prominent as is the rather dense reticulation beneath; petiole rugose, 1-2.5 cm by 2-3 mrn. — cJ Panicles axillary, or mostly clustered on woody tubercles on the stem or larger branches, consisting of numerous pedicelled few- flowered, more laxly or densely arranged um- bellules, all over patently rusty to rufous-tomen- tose, 2.5-5 cm by (0. 5-) 1-1. 5 cm; pedicels slender to more stoutish, 2-5 mm, each subtended by a subulate bract as long as or shorter than the pedicel, which is generally hidden in the cluster of flowers. Calyx segments 4 or 5, oblong-ovate, free, 1 mm. Petals 4, almost free, oblong-ovate to obovate, glabrous and yellow-green inside, rufous- sericeous outside, c. 1.8 mm. Stamens 4; anther cells 0.3-0.5 mm. — ? Flowers (often diseased) in peduncled globose heads (c. 8 mm 0), borne on stem and branches, peduncle stout, set with subulate bracts, c. 1 cm, all parts of the inflores- cence brownish-rufous-tomentose as are the c? ones. Ca/>'x segments linear, 2 mm. Pe/a/^ subspathulate, base slightly connate, 3 mm. Ovary cylindric, apex somewhat contracted into a very short thick style, tomentose; stigmas (2-)3-4, flattish, glabrous, spreading. Drupes in globular orange coloured clusters 6-9(-ll)cm 0, each drupe ovoid to ovoid-oblongoid, obtuse, the very apex only ± apiculate by the style, covered especially in the upper part with stout straight reddish- brown subulate prickles (5-8 mm) which after falling leave numerous circular pits (c. 1 mm 0) in the endocarp, (3-)3.5(-4) by 2-2.5 cm incl. the prickles; exocarp coriaceous; endocarp crusta- ceous, slightly tubercled inside, c. 2.5 by 1.5 cm. Distr. Lower Burma, Thailand (Pattani), Indo-China (Annam, Cochinchina); in Malesia: Malay Peninsula (Penang to Malacca and Johore). Ecol. Lowland forests, up to 150 m. Uses. The leaves are used by Malay natives for fever headache. The caustic seed is reported to be edible. Vern. Akar chumprai, M. 8. Phytocrene palmata Wall. PI. As. Rar. 3 (1831) 12; Cat. (1832) n. 4949; Walp. Rep. 1 (1842) 98; MiQ. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 248; Baill. in DC. Prod. 17 (1873) 11; Mast, in Hook./. Fl. Br. Ind. 1 (1875) 592; Becc. Malesia 1 (1877) 127; King, J. As. Soc. Beng. 64, ii (1895) 123; Ridl. Fl. Mai. Pen. 1 (1922) 433; Burk. Diet. (1935) 1722; Sleum. Blumea 17 (1969) 237; Fl. Thail. 2 (1970) 91. — Gynocephalum palmatum (Wall.) Trecul, Ann. Sc. Nat. Ill, 8 (1847) 149. 1971] ICACiNACEAE (Slcumer) 85 — p. stvlocarpa Griff. Not. (1854) 320; Ic. (1854) t. 489, I. Liana; stem generally not branched, younger parts with prickles (1 mm), striate, tips rufous- hispid. Leaves limited to the upper part of the branches, orbicular to reniform in outline, deeply palmately (3-)5-(rarely -7)-lobed, lobes oblong to oblong-obovate, acute or acuminate, base of lamina deeply cordate, entire or shallowly wavy, upper surface glabrous and shining, lower sur- face densely covered with longish coarse sub- appressed rufous to yellowish hairs especially on midrib and nerves, soft to the touch, 1 5-30 cm long, breadth about the same, main nerves 5 (-7), palmate, lateral nerves per lobe 3-4 pairs, curved- ascending and looping, with more numerous coarse transverse veins in a — right angle from the main nerve, reticulation coarse, — deeply impressed above, much raised beneath as are all nerves, the lamina not rarely almost bullate in dry specimens; petiole stout, often twisted, rufous- to rusty-tomentose, 7-12.5 cm. — Panicles of o flowers subcylindric, axillary or from lower part of stem, (6-)15^W) by c. 3.5 cm, all over covered with a tomentum of short soft rufous to ferrugineous hairs, the ultimate branches (c. 1 cm) bearing small, 12-15-flowered, pedicelled (2-4 mm) umbellules or heads (c. 2.5 mm 0), part of them subtended by a zh caducous subulate bract (2-4 mm). Calyx segments 3 or 4, obovate, hairy, 0.8 mm. Petals (3-)4, oblong-lanceolate, pale green, glabrous inside, hairy outside, c. 1.5 mm. Stamens 4; anther cells linear-oblong, 1 mm. — S Flowers in peduncled (c. 6 mm) axillary ovoid heads, c. 1 cm 0. Ovary ovoid, hirsute. Drupes numerous, collected in subglobular heads 7-10 cm 0, elongate-ellipsoid, or slightly obovoid, much tapering to the base, apiculate by the conical style, all over — densely covered with yellowish shining stiff hairs, (3.5-)4-5.5 by 1.6-2(-2.5) cm; exocarp thin; endocarp bony, rather smooth outside where the hairs are gone, laxly pitted outside, low-papillate inside respec- tively. Distr. Thailand (Pattani); in Malesia: Malay Peninsula, Sumatra (Palembang; Riouw Arch.: P. Bintan). Ecol. Hilly lowland forest, up to 500m. Vern. Akar pisang pisang bulu, Malay Penin- sula, kipajang akar, Palembang, M. 9. Phytocrene bracteata Wall. PI. As. Rar. 3 M83U 12; Cat. n832) n. 4947; Walp. Rep. I (1842) 98; Baill. in DC. Prod. 17 (1873) 12; Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 592; Becc. Malesia I (1877) 127; Merr. En. Born. (1921) 357, p.p.; Ridl. Fl. Mai. Pen. i (1922) 432; Baker/. J. Bot. 62 (1924) Suppl. 21; Merr. PI. Elm. Born. (1929) 172; Dahl. J. Arn. Arb. 33 (1952) 275 (pollen); Slelm. Blumca 17 (1969) 238; Fl. Thail. 2 (1970) 91. — Gynocephaliim bractealum (Wall.) Tr^cul, Ann. Sc. Nat. Ill, « (1847) 149. — P. macrucarpa Grifk. Not. (1854) 322; Ic. (1854) t. 487 (fl. o & t. 488 (fr.). High climbing liana; stem up to 2.5 cm o, with spiny tubercles 1 mm. Branchlets — deeply striate, setose and minutely prickly in younger parts. Leaves broadly ovate, often 3-lobed and obscurely dentate then, apex acute, base cordate, coriaceous, glabrous and a little asperulous above, the nerves excepted, densely covered all over the undersurface with short brownish hairs and soft to the touch, and the nerves moreover set with bristle-like stiffer hairs beneath, 10-20(-25) by 7-15(-20) cm, 7 pairs of basal palmate nerves and 2-3 pairs of upper nerves, curved-ascending, slightly sunken above, prominent beneath, re- ticulation low though distinct on the undersurface; petiole hairy and bristly, ~ torted, 3-7(-9) cm by 2-3 mm. — Panicles of cj flowers mainly axillary, rarely from older branches, solitary or 2-3, spike- like, all over shortly greyish to brownish tomentose, 10-20 by c. 2 cm, rachis and peduncles of main branches slender, ultimate branches consisting of minute clustered, (3-)6-10-flowered pedicellate (2-5 mm) umbellules, each pedicel subtended by a filiform appressed-hispid, not properly plumed — persistent bract (1 cm), which is partly connate to the pedicel and exserted from the mass of flower heads. Calyx segments 3-5, free, obcuneate, villous outside, hardly 0.6 mm. Petals generally 3 (rarely 4), ovate-oblong or lanceolate, slightly hairy outside only, practically free, finally re- flexed, 1-1.2 mm. Stamens 3(-4); anther cells oblong-elliptic, 0.6 mm. — 2 Flowers in oblong to spherical peduncled heads, either solitary or 2-3 per axil. Calyx segments 4, spathulate. Petals 4, short-strigose outside as are the calyx segments. Ovary strigose; style very short, 3-4-partite. Drupes in pendulous globose clusters up to 20 cm 0, each drupe ovoid-oblongoid, tapering to both ends, densely covered with appressed long soft yellowish bristles, 4.5-8 by 1.5-2.3 cm at full maturity; exocarp fleshy especially in the upper part of the drupe which bears a short beak of the former style; endocarp ligneous, ovoid-oblongoid, up to 4 by 2 by 1 .5 cm, rather smooth and minutely pitted outside, faintly tubercled inside. Seed testa sparsely and shallowly pitted. Distr. Thailand (Surat, Nakawn Sritamarat), in Malesia: Sumatra (Eastcoast, Palembang), Malay Peninsula, Lingga Arch., possibly in Borneo. Ecol. Lowland hillside woods and hedges. Vern. Akar pisang pisang, M (as P. palmata). Note. All investigated Bornean specimens ap- parently belonging to P. bracteata bear immature mflorcscences, or are sterile, and could possibly belong to P. borneensis. 10. Phytocrene hlrsuta Bl. Rumphia 4 (1849) 37; Mus. Bot. Lugd.-Bat. I (1849) 42; Walp. Ann. 2 (1851) 22; MiQ. Fl. Ind. Bat. I (1856) 797; Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 247; Baill. in DC. Prod. 17 (1873) 12; Becc Malesia I (1877) 127; K(K)Ri). Minah. (1898) 629; Hall. /. Med. Rijkshcrb. I (1910) 14; Sleum. Blumca 17 (1969) 238. nan Teysm. & Binn. Hort. Bog. (1866) 207. — P. crinipes Baii L. ex Biirfal;. Mon. Bign. (1864) 146, in text, nom. mul. — /'. minuhassac 86 Flora Malesiana [ser. I, vol. 7^ KooRD. Minah. (1898) 394, nom. nud. — P. dasy- carpa (non MiQ.) Dahl, J. Arn. Arb. 33 (1952) 275 (pollen). Climber; stem (1 cm or more 0) smooth. Branchlets slender, set with longish rufous sub- setose and — spreading hairs, older parts also laxly with very short pale retrorse prickles. Leaves ovate, or ovate- or elliptic-oblong, apex acuminate, the midrib characteristically protruding up to 8 mm beyond the leaf margin as a hirsute breakable thread in young leaves, base rounded to slightly cordate, firmly chartaceous, finally glabrous above except the nerves, remaining hirsute or tomentose by shorter and longer rufous to ferrugineous hairs and soft to the touch beneath, entire or shallowly sinuate-dented, or obscurely repand especially in the lower half, 12-17 by 6-9 cm, midrib and nerves very slightly immersed or flat above, prominent beneath, nerves 2 pairs palmate- ly from the base of the lamina (outer pair short, inner one rather straight ascending to tz half the length of the blade), other 3 (-4) pairs of nerves from the upper 2/3 of the midrib generally more curved, reticulation dense and finely raised above, coarser and more prominent beneath; petiole thickened at base, rufous-hirsute, 3.5-4 cm by c. 2 mm. — S Inflorescences in spike-like panicles, solitary in the axils of lower leaves, up to 25 by 3 cm, composed of numerous shortly peduncled cymes, the ultimate peduncle (2-3 mm) bearing numerous flowers in a head (c. 4 mm 0) and provided at base with a subulate ± persistent bract (1-1.5 mm), the whole inflorescence cov- ered with shorter greyish and longer rufous, bristle-like and — spreading hairs. Calyx seg- ments 3 or 4, connate below, obcuneate, hirsute, 1.3 mm. Petals 3 or 4, almost free, oblong, dirty yellow, with a few hairs inside, hirsute outside, c. 2 mm. Stamens 3 or 4; anther cells 0.5 mm. — ^ Flowers in heads c. 8 mm0, on stout peduncle 2-3 cm, generally from old wood. Calyx segments and petals as in o flowers. Ovary glabrous below, hirsute above; stigmas 4 or 5, short, acuminate. Drupes collected in subglobular masses, each drupe ovoid-oblongoid to -ellipsoid, laterally much compressed, apex — broadly attenuate, densely set with retrorse setose bristle-like stiff" hairs, 3-4(-4.5) by 1.5-2 by 1.5 cm, incl. the short hollow beak of the exocarp; endocarp hard, with minute scattered roundish pits, (2-)2.5-3.5 by 1.5-2 cm. Distr. Malesia: Celebes (incl. Buton I.), Mo- luccas (Buru). Ecol. Primary and secondary lowland forest, up to 800 m, also in rather dry places, even on limestone rocks. Vern. Kunet ri sasap, Minahasa (T\.), pitji pitji lantjeh, Makassar, sapet kulo, Minahassa (Tt.). 11. Phytocrene macrophylla (Bl.) Bl. Rumphia 4 (1849) 36; Mus. Bot. Lugd.-Bat. 1 (1849) 41, f. 7; Walp. Ann. 2 (1851) 22; Zoll. Syst. Verz. 2 (1854) 112; MiQ. Fl. Jnd. Bat. 1 (1856) 796; Suppl. 1 (1860) 137; Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 247; Baill. in DC. Prod. 17 (1873) 10; Hist. PI. 5 (1874) 282 (fl.); Becc. Malesia 1 (1877) 127; C. B. Rob. Bot. Z. 47 (1889) 645 (wood anat.); Engl, in E. &. P. Nat. Pfl. Fam. 3, 5 (1893) 255, f. 140; Racib. Flora 87 (1900) 13 (foliar biology); HocHR. PI. Bog. Exs. (1904) 17; Bull. Inst. Bot. Btzg 19 (1904) 39; ibid. 22 (1905) 114; Back. Schoolfl. Java (1911) 229 {'macrocarpd'); KooRD. Exk. Fl. Java 2 (1912) 533; Koord.- ScHU.M. Syst. Verz. 1, Fam. 162 (1912) 6; Warb. Pflanzenwelt 2 (1916) 348, f. 231 A, 1-9, f. 232 (phot.); DocT. v. Leeuwen, Zoocecidia (1926) 332 (galls); UiTTiE.N, Rec. Trav. Bot. Neerl. 25 (1929) 464 (veg.); Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 8; Back. & Bakh. /. Fl. Java 2 (1965) 62; Sleum. Blumea 17 (1969) 239. — Gynocephalum macropliyllum Bl. Bijdr. (1825) 483; Trecul, Ann. Sc. Nat. Ill, 8 (1847) 148. —P. gigantea Wall. Phil. Mag. N.S. 3 (1828) 223, nom. nud.; PI. As. Rar. 3 (1831) 11, t. 216; Cat. (1832) n. 4946; Walp. Rep. 1 (1842) 98; Bl. Rumphia 4 (1849) 36; Ann. Mus. Bot. Lugd.-Bat. 1 (1849) 41; Zoll. Syst. Verz. 2 (1854) 112; Griff. Not. (1854) 329; Ic. (1854) t. 490, f. II (fl. e & anat.); Miq. Fl. Ind. Bat. 1 (1856) 796; ScHNizL. Ic. 3 (1857/65) t. 172, f. 1-21; Baill. in DC. Prod. 17 (1873) 9; Mast, in Hook./. Fl. Br. Ind. 1 (1875) 591; Kurz, J. As. Soc. Beng. 44, ii (1875) 156; For. Fl. Burma 1 (1877) 241; Becc. Malesia 1 (1877) 127; Brandis, Ind. Trees (1906) 152. — Kadsura blancoi Azaola in Blanco, Fl. Filip. ed. 2 (1845) 594; ed. 3, 3 (1879) 118. — Gynocephalum giganteum (Wall.) Trecul, Ann. Sc. Nat. Ill, 8 (1847) 149. — P. calicarpa Griff. Not. (1854) 327; Ic. (1854) t. 490, f. I (fr.). — Gynocephalum luzoniense Llanos, Rev. Progr. Cienc. 15 (1865) 55. — P. dasycarpa Miq. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 248, t. 7, as to fruits only. — P. luzoniensis (Llanos) Baill. Adansonia 10 (1872) 281; in DC. Prod. 17 (1873) 10; Hist. PI. 5 (1874) 283 (fr.); F.-Vill. & Naves in Blanco, Fl. Filip. ed. 3, 4 (1880) 86; F.-Vill. Nov. App. (1880) 46; Ceron, Cat. PI. Herb. Manila (1892)46. — P. blancoi (Azaola) Merr. Philip. J. Sc. 2 (1907) Bot. 432; Spec. Blanc. (1918) 237; En. Philip. 2 (1923) 492; Dahl, J. Arn. Arb. 33 (1952) 275, f. 75 (pollen). — P. obovoidea Merr. Philip. J. Sc. 10 (1915) Bot. 322; En. Philip. 2 (1923) 492. — P. forbesii Baker/. J. Bot. 62 (1924) Suppl. 21. See for further synonyms under the varieties. key to the varieties 1. Drupe (7-)7.5-10(-13) by 3-3.5 cm; endocarp (4— )5-6 by 2-2.5 cm. . . 1. var. macrophylla 1. Drupe (5.5-)6-6.5(-7) by 1.5-2 cm. 2. Endocarp 4-4.5 by c. 1.5 cm. 2. var. caudigera 2. Endocarp c. 2.5 by 1 cm. . 3. var. dasycarpa 1. var. macrophylla. High climbing, often much branched shrub or liana, up to 25 m; stem up to 10 cm 0, not spiny. Branchlets striate, younger parts with appressed or somewhat spreading shorter and longer rufous or ferrugineous hairs, and besides ~ densely set 1971] ICACiNACEAE (Slcumer) 87 with pale conical prickles (0.5 mm), older parts glabrescent and smooth, the cork with transverse ienticellar cracks. Leaves of young shoots often orbicular with rather deeply cordate base, and broadly and — obtusely 3-5-lobed, such of older parts generally ovate-oblong from a subcordate base and subacuminate, tip obtuse, firmly sub- coriaceous, entire, glabrous above, the nerves excepted, the whole undersurface with a tomentum of short, weak rather pale rusty (partly almost stellate) hairs, and less numerous longer strigose rufous hairs especially on nerves and veins, soft to the touch, — glabrescent at full age, 14— 30(-33) by 9-12(-20) cm, palmately 5-7-nerved, outer pair(s) of nerves short, inner one(s) ascending to V2 and even ?4 of the length of the lamina, other 3-4 pairs from upper part of the midrib, all nerves almost flat above, well raised beneath, reticula- tion dense and very fine above, moderately raised beneath; petiole 5-15 cm by 2-3 mm. — o Inflorescences spike-like panicles, from old branches or stem, generally several together, cylindric, dense or rarely more loose, all over short- ly greyish-brow nish-tomenfose, up to 25 cm long and (2.5-)3(-5) cm o, composed of lateral racemes or panicles up to 4 cm long which bear the flowers in heads 4-6 mm o. Calyx segments 3-5, — nar- rowly obcuneate, apex truncate, hirsute outside, 0.7-1.2 mm. Petals 3 or 4, light green to yellowish, connate — halfway, lobes recurved, densely ap- pressedly hairy outside, 1.5-2 mm. Stamens 3 or 4 (in the same inflorescence!); anther cells ovate- elliptic, c. 0.4 mm, often devoid of pollen. — X Inflorescences collected in large, solitary or fascicled heads, each head on a thick peduncle (3-1 cm) and c. 1.5 cmo, allovershort-tomentose. Calyx segments as in o flower. Petals 2.5-3.5 mm. Ovary columnar, tomentose with forward directed hairs, c. 4 mm, stigma 2-4-lobed. Fruit heads 20-25 cm 0. Drupe obovoid-oblongoid, (7-)7.5-10(-l 3) cm long incl. the hollow beak and the basal narrowed part of the exocarp, 3-3.5 cm wide, slightly variously compressed or round in 0, rusty-tomentose by normally forward directed hairs (sometimes reversely so in specimens from Sumatra and the Philippines), these hairs -jz ap- pressed, stifi", shining, easily going; endocarp oblongoid, bony, laterally flattened, the crests less marked on the flattened side, with numerous roundish pits (1 mm), (4-)5-6 by 2-2.5 by (l.5-)2cm. Distr. Burma; in Malesia: Sumatra, Java, Philippmcs (Luzon, Polillo, Samar, Biliran, Leyte, Bohol, .Mmdanao, Palawan). Ecol. Lowland thickets and forests from sca- Icvcl to c. 1200 m, also in secondary and in swamp forest, in the Philippines also in Diptero- carp forest, scattered. Galls. DocTERS VAN Leeuwen mentions a leaf-gall caused by a gall-midge. Uses. Fruits said to be used on Palawan as 'talang', i.e. fish poison. In Java used against stomach pain and tumors. Vern. Akar pisang, Sumatra Westcoast, M, labii kumbung silai, olor delos, Simalur, areuj pitjiing tjeleng, hondje bu-itt areuj, undjewut, S; Philippines: olo-olo. Bag. 2. var. caudigera (Sleu.m.) Blumea 17 (1969) 240. — P. caudigera Sleum. Notizbl. Berl.-Dahl. 15 (1940) 253; Heine, Pfl. d. Samml. Clemens Kinabalu (1953) 57. Differs from var. macrophylla by generally smaller leaves 7-15 by 5-8 cm, and mainly by the oblongoid, slightly 4-6-angled minor drupes, (5.5-)6-6.5(-7) by c. 2 cm; endocarp 4-4.5 by c. 1.5 cm, the pits much smaller than in var. macrophylla. The hairs on the drupe are strigose and retrorse. Distr. Malesia: Borneo, only known from the Mt Kinabalu area, in forest around Dallas at 915-1370 m. 3. var. dasycarpa (MiQ.) Sleum. Blumea 17 (1969) 240. — P. dasycarpa MiQ. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 248, t. 7, the fruit excluded (which is var. macrocarpa); Baill. in DC. Prod. 17 (1873) 11; Becc. Malesia 1 (1877) 127; Koord. Minah. (1898) 394; Hochr. PI. Bog. Exs. (1904) n. 92; Bull. Inst. Bot. Btzg 22 (1905) 1 14; Ic. Bog. 2 (1906) t. 108; Koord. -Schum. Syst. Verz. 3 (1914) 74; Koord. Minah. Suppl. 2 (1922) t. 49; Suppl. 3 (1922) 25. — P. hirsuta (non Bl. 1849) Teysm. & BiNN. Hort. Bog. (1866) 207. — P. ovalifolia Koord. Minah. (1898) 394, nam. nud., 629, descr.; Koord. -Schum. Syst. Verz. 3 (1914) 74; Koord. Minah. Suppl. 2 (1922) t. 50; Suppl. 3 (1922) 26. — P. tinosporifolia KoORD. Minah. (1898) 394, 629; Koord. -Schum. Syst. Verz. 3 (1914) 74; Koord. Minah. Suppl. 2 (1922) t. 51; Suppl. 3 (1922) 26. Diff"ers from var. macrophylla by its smaller drupes, (5.5-)6-6.5 by 1.5-2 cm; endocarp c. 2.5 by 1 cm. Distr. Malesia: Celebes. Ecol. Lowland forest, up to 350 m. Vern. Apo, Tontembuan, samadan, Tonsea, sapet kulo, Tt., s. putih, Tl., tali tali utan, Bt. Excluded P/iylocrene loheri Merr. Philip. J. Sc. 7 (1912) Bot. 293; En. Philip. 2 (1923) 492, mixtum, tiom. illeg. The leaves belong to a Menispermacea (probably Hypserpa), and the loose fruits to Phytocrene macrophvlla var. macrophylla, cf. Sleum. Blumea 17 (1969) 241. Excluded Leucocorema Ridl. Trans. Linn. Soc. II. Bot. 9 (1916) 29 = Trichadenia Tiiw. (Flacourtiaceae), cf. SiEts. Bull. Jard. Bot. Brux. 27 (1957) 114. Peniavira RiDi.. Trans. Linn. Sot. II, Bot. 9 (1916) 27 ■--- Dichapetalum Thou. (Dichapetalaceae), cf. LiENHouts. Fl. Mai. I, 5 (1957) .105. LOPHOPYXIDACEAE (H. Sleumer, Leyden) 1. LOPHOPYXIS Hook. f. Ic. PI. 18 (1887) t. 1714; Sleum. Blumea 16 (1969) 322. — Combretopsis K. ScH. in K. Sch. & Hollr. Fl. Kais. Wilh. Land (1889) 69. — Treubia Pierre ex BoERL. Handl. 1, 2 (1890) 445. — Fig. 1. Scandent or liana-like shrubs or small trees, with interxylary bast elements, watch-spring tendrils (modified leaves or subtending leaves of inflorescences), and umbrella-like branching. Leaves spiral, simple, serrulate to crenulate, stip- ulate. Flowers small, regular, monoecious, in glomerules on the branches of loose axillary panicles. Sepals 5, valvate, shortly united basally, persistent. Petals 5(-6). much smaller than the sepals, free. Stamens 5(-6), opposite to the sepals, with filiform filaments and subglobose introrse, almost basifixed anthers, alter- nating with 5(-6) oppositipetalous cordate glands, these in the o i adnate to the subtending petal, and in the $ dz concrescent into a 5(-6)-lobed disk (the glands or lobes opposite the ovary cells); pollen grains ± ellipsoidal, tricolporate, exine reticulate. Ovary superior, conical, shallowly 5-ribbed, 5(-4)-celled, with 5(-4) sessile subulate stigmas; ovules 2 per cell, pendulous, apical, axile, anatropous, epitropous, bitegmic, each surmounted at the micropyle by a small obturator- like appendage coming from the funicle. Fruit obovoid or ellipsoid, indehiscent, fusiform, 1-locular, 1 -seeded, with 5 broad stramineous wings. See^ oblong, with endosperm; embryo erect, with oblong cotyledons and a short erect radicle. Distribution. Monotypic, in Malesia, Melanesia, and Micronesia. Fig. 2. Anatomy. Stem. Anomalous secondary growth: Engler, Sitz. Ber. Preuss. Ak. Wiss. Berl. 18 (1893) 265-266; Handa, Bot. Mag. Tokyo 54 (1940)41-47; Pfeiffer, Rev. Sudam. Bot. 10 (1951) 3-6; Sleumer in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 393, f. 1 18. The young stem has five ribs with a continuous xylem cylinder enclosing a pentagonal pith with a central portion of thick-walled parenchyma cells and a marginal area of thin-walled cells, erroneously referred to by Engler (t.c.) as intraxylary phloem. This thin-walled tissue may become caducous, at least in herbarium specimens. Between the ribs the secondary xylcm has numerous vessels; in the ribs the vessels are narrower and scarcer. Vessel perforations are simple. Through anomalous activity of the cambium, 5 phloem strands become enclosed within xylem in the young shoot. Later interxylary phloem is formed as continuous bands alternating with secondary xylem. The phloem is stratified into soft and fi- brous portions. Axial xylem parenchyma is scarce and paratracheal (only seen in young twigs). The ground tissue of the xylem is composed of fibres with numerous minutely bordered pits. Rays vary from 1-6-seriate in the young stem. The outer phloem is surrounded by a cylinder of fibres and stone cells. Cork arises in the layer below the epidermis. The anatomy of the leaf has hitherto never been described. The petiole shows a strongly incurved arc of separate vascular bundles and two additional latcro-dorsal bundles as seen in transverse section through the distal end. The vascular system forms a closed flattened cylinder in the midrib. The stomata, confined to the abaxial surface, arc paracytic. Crystals are present as solitary rhomboids and clusters. The hairs are unicellular. To evaluate the taxonomic significance of the vegetative anatomy of Lopliopy.xis with regard to the affinities of the genus more research is still needed. The only straightforward conclusion to be drawn at present is that Lophopyxis is anatomically entirely dilTercnt from Gouania (Rliaiiinaceae), to which Airy Shaw (in Willis Diet. ed. 7, 1966, 668) related it. Gouania differs e.g. in having anomocytic stomata, sty- loids, unlignificd perivascular fibres and exclusively narrow rays. Dr. C. R. Metcalfe (Kew) kindly put slides and anatomical information of Lophopyxis and Gouania at my disposal. -P. Baas. Taxonomy. This genus was tentatively ascribed to the I. uphorhiacvae by Hooki r/., but removed from this family by Pax (1890). FIngier (1893) accommodated it as a distinct subfamily Lophopy.xidoidcae within Icacinaceae, from which I rejected it in 1942. Hutchinson (f-am. Flow. PI. 1959) placed it in the Celaslraceae. its gross morphology, wood anatomy, embryology and pollen morphology is well known and it is (89) 90 Flora Malesiana [ser. I, vol. 7^ Fig. 1. Lophopyxis maingayi Hook. / a. Habit, x 1/2, a', showing domatia in the axils of lateral nerves, ><2'/2,*- ? flower, x 7, c. ovary, cross-section, x 7, J. ovary, longit. section, x 7, e. cj flower buds, X21/2, f. (S flower, X 5, g. stamen, front view, x 15, g'. stamen, back view, x 15, h. infructescence, x^/j, /. fruit, x«/5 ia-d, h-i Kanehira (1938), partly altered, e-g Sleumer (1942)). 1971 LoPHOPYXiDACEAE (SleuiTier) 91 now apparent that it should be placed within the Geraniales-Sapindales-Celastrales. It seems, however, that it does not fit in any of the established families of these orders. Its relation to Rhamnaceae, suggested by Shaw (1966) rests on a superficial habit similarity with Gouania, as shown by Bakhuizen & Van Steenis, F1. Mai. Bull. 21 (1966) 1426 (see also sub Anatomy). The best solution is to regard it as the type of a family of its own, as has been casually proposed by vanTieghem (1897) and Pierre (1897), and formally by Pfeiffer (1951). I have this more fully explained in my precursory paper in Blumea 16 (1969) 320. 1. Lophopvxis maingayi Hook./. Ic. PI. 18 (1887) t. 1714; Fl. Br. Ind. 5 (1888) 673; Pax in E. & P. Nat. Pfl. Fam. Ill, 5 (1890) 117; Engl. ibid. (1893) 238, 257; Hall. /. Med. Rijksherb. 1 (1910) 9; RiDL. Fl. Mai. Pen. 1 (1922) 435; Slcx)T. Bull. Jard. Bot. Btzg III, 7 (1925) 364; HoLTH. & H. J. Lam, Blumea 5 (1942) 205, f. 7; Sleum. in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 393; Fl. Males. I, 5 (1954) 63; Blumea 16 (1969) 322, \\ith extensive bibliogr. — Combretopsis pentapiera K. Sch. in K. Sch. & Hollr. Fl. Kais. Wilh. Land (1889) 69. — Treubia combretocarpa Pierre ex Boerl. Handl. 1, 2 (1890) 445. — L. pierrei Boerl. I.e. 673, noni. ill.; Heyne, Nutt. PI. 1 (1950) 987. — L. sehumannii Boerl. Handl. 1, 2 (1890) 674, nom. ill. — L. eombretoearpa (Boerl.) Engl, in E. & P. Nat. Pfl. Fam. Ill, 5 (1893) 257. — L. pentaptera (K. Sch.) Engl. Sitz. Ber. Kon. Preuss. Ak. Wiss. (1893) 265, t. 2, f. 6 & 7; Sleum. in E. & P. Nat. Pfl. Fam. ed. 2, 20b ( 1 942 ) 393, f. 1 1 7 (habit) & 1 1 8 (wood anat., embryol.); Dahl, J. Arn. Arb. 36 (1955) 160, 161 f. 2 & 2A (pollen); Peekel, Fl. Bismarck Arch. MS p. 1083, fig. — Homalium gilgianum Laut. in K. Sch. & Laut. Nachtr. (1905) 320. — Sinapistrum RuMPH. Herb. Amb. 5 (1747) 73, t. 39, f. 1. — Fig. 1. Vigorous woody vine or climbing shrub, 3-8(-30> m; stem up to 7 cm 0, bark whitish. Branches virgate, youngest parts longitudinally grooved, with elliptic lenticels in the grooves, pu- berulent. Leaves ovate to oblong, apex -± acu- minate, acutish, base cuneate to obtuse, rarely rounded or subcordate, subequal, chartaceous to subcoriaceous, caducous-pubcrulous especially at the nerves beneath, the hairs persisting in the nerve axils as domatia, scrrulate-crenulate or crenate, sometimes subcntire, 8-18(-24) by 4-8(-10) cm, nerves l(-2) basal or slightly supra- basal, and 3-4 upper pairs, all arched and steeply ascending, veins and vcinlets densely reticulate, slender but prominent on both faces; petiole r. 10 by I mm. Stipules small, knob-like. Axillary branchlels metamorphosed into strong woody tendrils coiled only at the end, often bearing a bud. Panicles loose, composed of a few axillary or terminal spike-like racemes, pendent, puberulous, 10-25 cm. I lowers solitary, or mostly crowded into glomerulcs. these spaced along the slender to filiform rachis. Pedicels very slender, up to 2 mm. Bract at the base of the inflorescences often meta- morphosed into a weak flat completely coiled tendril, .'sepals ovate, greenish white or yellowish, hairy on both sides, c. 1.5 mm. Petals ovate, thin, c. I mm. Disk yellowish. — 3 Flowers: filaments hairy, 2 mm; anthers subglobose, 0.5 mm; ru- diment of ovary subglobose, shallowly 5-ribbed, hairy. — 5 Flowers: ovary ovoid, whitish-yellowish puberulent, 2 mm. Capsule obovoid-ellipsoid in outline, 5-winged, green, later dark brown, caducous-pubescent, 2.2-3(-3.5) by 1.3-1.8 cm; wings chartaceous, 5-8 mm wide, with irregularly crenulate margin; pedicel short, subtended by the non-accrescent calyx. Seed 1, subcylindrical, acuminate, lengthwise grooved, 12-15 by 5-6 mm. Fig. 2. Distribution of Lophopyxis maingayi Hook. /. Distr. Micronesia (Palau Is.), Melanesia (New Ireland, Duke of York I., New Britain, Solomon Is.), in Malesia: Malay Peninsula (Pcnang to Malacca), North Borneo, W. Central and N. Celebes, Moluccas (Talaud Is., Ceram, Sula Is., Ambon), New Guinea. Fig. 2. Ecol. Straggling climber in canopy or edge of primary lowland forest, both in well drained and in swampy riverine forest, in littoral forest and even sometimes in the mangrove, also in disturbed gully forest or forest regrowths, seaside scrub; scattered, though locally not too rare, from sea- level up to c. 300 m, often on alluvial soil. Uses. The rather hard stem splits easily into pieces, and apparently for this reason is used in New Britain for tying thatch. Rumi'iiius under his "Sinapistrum' says, that crumpled leaves give a strong smell of mustard, and are used to cure ulcers. Vern. Akar hHimhing hulan, Brunei (Kina- batangan), simpuru, Celebes, tahurulu Talaud. /(/// sasuwi, Anihoi) (sa. Ri nhmiius). CARDIOPTERIDACEAE (H. Sleumer, Leyden) 1. CARDIOPTERIS [Wall, ex] Royle. 111. Box. Himal. Mts (1834) 136, in text; em. Bl. Rumphia 3 (1847 or 1849) 205; ibid. 4 (1849) t. 177; Hassk. Flora 30 (1847) 110. in adnot.; Nat. Tijd. N. I. 10 (1855) 64; Engl, in E. & P. Nat. Pfl. Fam. 3, 5 (1893) 239, 257 CCardiopteryxy, Bakh. /. & Steen. F1. Mai. Bull. 15 (1960) 725; Taxon 11 (1962) 28; Back. & Bakh./. F1. Java 2 (1965) 62. — Sioja Buch.-Ham. ex Lindl. Nat. Syst. ed. 2 (1836) 82, nom. nud. — Peripterygium Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 142; Cat. Hort. Bog. (1844) 235; Sleum. in E. & P. Nat. Pfl. Fam. ed. 2. 20b (1942) 400; Amsh. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 9. — Fig. 1. Sinistrorsely twining herbs with white milky juice. Leaxes spirally arranged, simple or lobed to varying degrees, cordate, palmatinerved, long-petioled, glabrous as is the stem, exstipulate. Flowers bisexual, or polygamous (andromonoecious), small, subsessile, in unilateral repeatedly forked cincinni, composed of loose axillary panicles, ebracteate. Calyx deeply (4-)5-partite, lobes imbricate in bud, whether or not slightly accrescent in fruit, persistent. Petals (4-)5, caducous, lower half forming a widely funnel-shaped corolla, lobes imbricate in bud. Stamens (4-)5 inserted on the upper part of the corolla tube, alternate with its lobes; filaments very short, glabrous; anthers medifixed, introrse; pollen grains oblate, triangular in polar view, tricolporate. Disk absent. Ovary oblong-ovoid, subquadrangular (rudimentary in o), 1-celled; styles 2, one with a deep longitudinal groove, and two inequal ovate rather obtuse distal divisions, accrescent in fruit and becoming linear and succulent then, the other short-curved, capitate at apex, deciduous; ovules 2 (often one of them aborted), pendent from apex of cavity, anatropous, with dorsal raphe. Fruit indehiscent, compressed, with 2 longitudinal broad and transversely striate stramineous wings, obovate-elliptic to orbicular in outline, apex emarginate, crowned by the columnar accrescent soft and green stigma, base very shortly or hardly (Mai.), sometimes elongately contracted into a kind of stipe. Seed 1, linear, sulcate; testa thin; embryo minute, conical, in top of granular fleshy albumen. Distr. Two spp., one in SE. Asia and W. Malesia, and one in E. Maiesia. Ecol. Climber on edge of (mostly secondary) forest and thickets, in open places as limestone rocks. Uses. The leaves are eaten as a vegetable. Taxon. Cardiopteris was conceived as type of a monogeneric family by Blume (1847 or 1849) and R. Brown (1852); it was considered to constitute a subfamily of the Icacinaceae by Engler (1893) and Post & Kuntze (1904;, and again as a distinct family within the Ceiastrales by King (1893), Williams (1915, as Peripterygiaceae), Gagnepain (1910, 191 1 ), Sleumer (1942, as Pcriplerygiaceae), Hutchinson (1959) and Takhtajan (1966). The pollen of the genus shows no distinctive features against Icacinaceae and resembles very much that of the Afro-Malagasian genus Cassinopsis Sonder. Note. The family name Cardiopteridaceac is derived from the original spelling Cardiopteris used by Wallich and later by Bi.ume and others. Engler changed the name to Cardioplcryx, which would be niorc correct as an allusion to the winged fruit, a substitute name, however, which cannot be used ac- cording to the Code; for this reason, the family name 'Cardiopterygaceae', proposed recently, has not been used. (93) 94 Flora Malesiana [ser. I, vol. 7^ Fig. 1. Cardiopteris moluccana Bl. a. Habit, withe? inflorescence, x i/^, b. cj flower, corolla and stamens, X 15, c. ? flower, corolla and ovary, x\5, d. flower, lateral view with calyx and corolla, x7, e. flower calyx, X 10,/. infructescence, x V2 (<2-/Blume). 1971 Cardiopteridaceae (Sleumer) 95 KEY TO THE SPECIES 1. Leaves membranous, generally — deeply and ± acutely 3-5(-9)-lobed, very rarely (sub)entire. In- florescence l-2(-3)-forked, cincinni rather few-flowered 1. C. quinqueloba 1. Leaves firmly chartaceous, entire. Inflorescences with more numerous forks, cincinni rather many- flowered 2. C. moluccana 1. Cardiopteris quinqueloba (Hassk.) Hassk. Nat. Tijd. N. 1. 10 (1855) 64. — Peripierygium quin- quelobum Hassk:. Tijd. Nat. Gesch. Phys. 10 (1843) 142; Cat. Hort. Bog. (1844) 235; Sleum. in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 400, f. 120 F-G; A.MSH. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 9. — C.javanica BL.Rumphia 3 (1847 or 1849) 206, nom. illeg.; ibid. 4 (1849) t. 177, f. lA; Back. & Bakh./. Fl. Java 2 (1965) 63. — C. lobara R. Br. [in Wall. Cat. (1847) //. 8033] in Benn. & Br. PI. Jav. Rar. (1852) 246, t. 49, nom. illeg.: MiQ. Fl. Ind. Bat. 1 (1856) 799; Mast, in Hook. /. Fl. Br. Ind. 1 (1875) 597; KuRZ, J. As. Soc. Beng. 44, ii (1875) 157; Becc. Malesia 1 (1877) 131; King, J. As. Soc. Beng. 64, ii (1895) 131; Gagnep. Fl. Gen. I.-C. 1 (1912) 849; Back. Schoolfl. Java (1911) 230; KooRD.-ScHLM. Syst. Verz. 1, Fam. 162 (1912) 6; Merr. En. Born. (1921) 357; Ridl. Fl. Mai. Pen. 2 (1923) 464; Craib, FI. Siam. En. 1 (1926) 276; Blrk. Diet. (1935) 456; Kanjilal c.s. Fl. Assam 1, 2 (1936) 254; Henders. J. Mai. Br. R. As. Soc. 17 (1939) 59; Gagnep. FI. Gen. I.-C. Suppl. (1948) 759; Heyne, Nutt. Pi. 1 (1950) 987; Wu & Wang, Act. Phytotax. Sin. 6 (1957) 284; Larsen, Dansk Bot. Ark. 23 (1963) 71. — C. rumphii Baill. Adansonia 10 (1872) 280, nom. illeg., incl. var. lobata Baill. et var. subhamata Baill. I.e. 281; DC. Prod. 17 (1873) 26; Scheff. Ann. Jard. Bot. Btzg 1 (1876) 14. — C. platycarpa Gagnep. Not. Syst. 1 (1910) 198; Fl. Gen. i.-C. 1 (1911) 847, f. 104, p.p., f. 105. — Peripierygium platycarpum (Gagnep.) Sleum. Notizbl. Berl.- Dahl. 15 (1940) 257; in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 400. — C. moluccana (non Bl.) Wu & Wang, Act. Phytotax. Sin. 6 (1957) 284. Much branched with twining terete stems, which often preserve their freshness after the fall of the leaves and are loaded with fruits then, 2-5(-9) m. Leaves widely spaced, polymorphous, broadly ovate in outline, with a cordate base, generally rb deeply or irregularly 3-5(-9)-lobed, apex generally acutish, lateral lobes acute or obtuse, very rarely entire or almost so, herbaceous, caducous, pale green, (4-)6-12(-17) by (3-)4-7(-l6) cm, main nerves 7-9 radiating from apex of the petiole, slightly raised beneath, veins rather obscure; petiole (3-)5-l2 cm. Panicles solitary on 3-5(-10) cm long peduncles, l-2(-3)-forkcd, cincinni rather few-flowcrcd, glandular-pubcrulous or almost gla- brous, 5-10 cm, accrescent in fruiting stage. Flowers distant from each other along rachis, on short pedicels (1 mm, accrescent to 5 mm in fruit). Calyx lobes 2 mm. Petals white, 2.5(-3.5) mm. Fruit obovatc-clliptic in outline, the wings in- cluded, apex cmarginatc, base cuncatc and sudden- ly contracted to a kind of foot up to 2 mm, on the very base of which the persistent calyx is found, wings glossy yellow or light stramineous colour, tipped by the accrescent style which remains green and fleshy for a long time. Distr. SE. Asia (W. Bengal and Assam to Burma, Thailand, Indo-China and Yunnan); in Malesia: N. & NE. Sumatra, Malay Peninsula (Kedah, Perlis, Perak), SE. Borneo (twice found), Java (incl. also Bawean & Kangean Is. and Ma- dura), Lesser Sunda Is. (Lombok, Sumbawa, Sumba, Alor), Celebes (incl. also Saleijer Is.), Moluccas (Tenimber Is.). Ecol. Primary and secondary (also seasonal, deciduous) forest edge, scrub jungle, bamboo forest, hedges around villages, open waste places, even on open limestone rock, generally at low elevations, rarely up to 1000 m, scattered. Uses. Locally the leaves are used as a vegetable. Vern. Angi anginan, iri iri, kanjar kawang, parianom, rindengan, serintil, sobowengi, wiwi wiwo, J, oke oke. Mad., gambas kawaong, riru ruruan, saburung, S, kokrah, Alor. - y .- .;. .- .. r>*. ^l hly /Pl '%^ 1 tT^It^'^TlT^Nj =^ ... . .. .r^"^^) \ ^ Fig. 2. Distribution of Cardiopteris. 1. C quinque- loba (Hassk.) Hassk., 2. C. moluccana Bl. 2. Cardiopteris moluccana Bl. Rumphia 3 (1847 or 1849) 207; /A/W. 4 (1849) t. 177, f. 1 B, f. 2 A & B, incl. var.: MiQ. Fl. Ind. Bat. 1 (1856) 799; Becc. Malesia 1 (1877) 131; K. ScH. Notizbl. Berl. 2 (1898) 130; K. Sch. & Laut. Fl. Schutzgeb. (1900) 418; Merr. Philip. J. Sc. 2 (1907) Bot. 423; SCHELLENB. Bot. Jahrb. 58 (1923) 176; Merr. En. Philip. 2 (1923)493; Heyne, Nutt. PI. I (1950)987. — Dioscorea sativa L. Amoen. Ac. 4 (1759) 133, pr. pi. Amhoin. — C. rumphii Baill. var. hlumeana Baill. Adansonia 10 (1872) 281, incl. var. inte- grifolia Baill. I.e. 280; DC. Prod. 17 (1873) 26. — C. lobata R. Br. var. moluccana (Bl.) Mast, in Hook./. Fl. Br. Ind. I (1875) 597. — C. lobata {non R. Br.) Becc. Nuov. Giorn. Bot. Ital. 9 (1877) 100, t. 8; F.-ViLL. Nov. App. (1880) 46; 96 Flora Malesiana [ser. I, vol. 7^] F. M. Bailey, Queensl. Agric. J. 24 (1910) 20. — C. celebica R. Br. ex Koord. Minah. (1898) 392, nom. mid. — Aspidocarya kelidop/iylla K. Sch. & Laut. Fi. Schutzgeb. (1900) 313, sec. Diels, Menisp. (1910) 320. — Peripterygium moluccanum (Bl.) Sleum. Notizbi. Berl.-Dahl. 15 (1940) 257; in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 400, f. 120 A-E; Dahl, J. Arn. Arb. 36 (1955) 161 (pollen). — Oliis sanguinis Rumph. Herb. Amb. 5 (1747) 482, t. 180. — Fig. 1. Leaves ovate-cordate, entire, acutely acuminate, base ± deeply cordate, firmly chartaceous, edge often slightly recurved in dry specimens, up to 24 by 22 cm. Panicles generally 3- or more- forked, cincinni rather rich-flowered, flowers usually close together. Otherwise as C. quinqueloba. Distr. Malesia: Celebes, Philippines (Luzon, Panay?, Negros, Samar, Bohol, Catanduanes, Mindanao), Moluccas (Buru, Ambon (type), Ternate, Ceram), New Guinea, and New Britain (Gazelle Peninsula). Ecol. Climber in tall rain-forest or forest edge, also in secondary vegetation and in native gardens, generally in the lowland, ascending to 1460 m in New Guinea, scattered. Uses. A decoction of the stem is used against hepatitis in Ternate. The leaves are eaten as a vegetable. Vern. Malta malta, iamatta, Makassar, telin teli, Menado, uge jabba, Ternate, iita lala, u. turi, Ambon; Philippines: bangogan, Bik., gurisan, sagumati. Bag., Ida, Buk., tagidauai, P. Bis.; New Guinea: foyomangeni, Wapi (Miwaute), kehunghe, Mekeo (Maipa), swalow, Buang. Note. According to Miquel also in Bali, which is apparently erroneous. OCHNACEAE (A. Kanis, Leyden) Woody plants, very small undershrubs to tall trees. Leaves distichous or spirally arranged, stipulate, simple, glabrous; midrib prominent on either side. Inflorescen- ces I- to many-flowered, cymose, racemose, or thyrsoid, bracteate; pedicels articulate. Flowers actinomorphic, bisexual (rarely functionally polygamous). Sepals 5, free or a little connate at base, quincuncial, persistent. P^r^/^ 5-10, free, contort, caducous. Staminodes 0-c/:. Stamens 5-10-C/5; anthers basifix, ± latrorse and dehiscing lengthwise, or with 1-2 apical pores. Carpels 2-5-10(-15), superior, free with 1 ovule, or fused with 2-^ ovules per carpel; styles fused, basigynous or epigynous; stigmas free or ± fused. Fruit{s) a drupe(s), berry, or capsule. Seeds \--r., small or large, sometimes winged, with or without albumen. Taxonomy. There is little doubt that the family of the Ochnaceae represents a natural one among the more primitive in the Gutiiferales (= Chisiales or Theales s. /.). Nonetheless, there are striking differences between the genera, even at first sight. It is not difficult to arrange them in a few distinct, supra-generic taxa. A supposed natural system, as far as relevant to the Malesian genera, is as follows: Subfamily Ochnoideae Tribe Ochneae Subtribe Ochninae 1. Ochna 2. Brackenridgea Subtribe Ouratinae 3. Gomphia Subfamily Sauvagesioideae Tribe Euthemideae 4. Euthemis Tribe Sal\ agesieae Subtribe Sauvagesiinae 5. Neckia 6. Indovethia 7. Schuurmansiella 8. Schuurmansia Distribution. About 30 genera and c. 250 spp. through the tropical, rarely subtropical countries (S. Africa, N. India), chiefly in S. America and Africa. In Malesia 8 genera and 13 spp.; absent from Java and the Lesser Sunda Islands. In subfam. Ochnoideae, the monogeneric tribe Elvasieae and the genus Ouratea are restricted to S. America. Of the 3 genera of the Ochneae mentioned above, only Brackenridgea is chiefly Malesian. Ochna and Gomphia are principally African, both reaching the western part of Malesia with one species. In subfam. Sauvagesioideae, the monogeneric tribe Lophireae occurs only in Africa, whereas the sub- tribe Luxemburgiinae is restricted to S. America. The 5 relevant genera mentioned above are almost purely Malesian. Of the subtribe Sauvagesiinae, Sinia Diels and Indosinia Vidal arc found in continental SE. Asia, whereas the majority of the genera is restricted to S. America, only Sauvagesia L. occurring both in S. America and Africa. There is an interesting parallelism between some of the Sauvagesioideae, which arc more or less restric- ted to sandstone areas in W. Malesia, and related genera found in similar areas in northern S. America (Roraima flora). Ecology. The Malesian genera are restricted to everwet areas, except Ochna which is adapted to a seasonally dry climate and is deciduous. They are more or less adapted to poorer, sandy or peaty soils in relatively undisturbed areas, except Schuurmansia which is a genus of pioneer species. Pollination. There is no literature on this subject. Pollination probably takes place by insects, because of the brightly coloured and (sometimes?) scented flowers. The colours of the petals arc yellow in Ochna and Brackenridgea sect. NotochncUa, mostly white, creamy, or tinged purple in other genera, sometimes dark purplish red in Schuurmansia. Only Ochna has flowers c. 3 cm across. Flowers of other genera usually do not exceed I cm, but they are often combined to conspicuous inflorescences. The undershrubs of Neckia, however, have solitary flowers with relatively small, early caducous petals. Dispersal. The bluish or black, 1-seeded fruits of the Malesian Ochneae are probably mainly dis- persed by birds. In Ochna and Brackenridgea they are contrasting with a purple calyx and torus. The f.uihemideac have red or white berries on a dark red calyx, which are probably also dispersed by birds ' cc RiDi>.y, Disp. 1930). The Malesian Sauvagesieae have many-seeded capsules. Their seeds do not liow adaptations to a special mode of dispersal, except those of Schuurmansia with two wings like propeller blades. The latter characteristic points to wind dispersal, which tits the pioneer-like nature of the genus concerned. The sepals in Neckia are also turning dark purple in fruit; the meaning of this phenomenon is not understood. (97) 98 Flora Malesiana [ser. I, vol. 7^ Morphology. From the description of the family and the key to the genera it will be clear that the morphological differences between subfamilies and tribes are considerable. A discussion on these dif- ferences I gave in my thesis (Blumea 16, 1968, 8-15). A short note should be made here on the inflores- cence types, as these may not always be easily understood. The inflorescences in subfam. Ochnoideae are all considered to be of a thyrsoid nature, viz racemes with cymose branches. Those of Gomphia serrata are lateral and terminal, bearing terminal flowers which makes sympodial growth of the vegetative branches necessary. Those of Ochna integerriina have also terminal flowers, but they are terminal on short side branches and monopodial growth of the main branches remains possible. The inflorescences oi Brackenridgea spp. are terminal, sometimes also lateral. A terminal flower is lacking and monopodia! growth of the rachis to a vegetative shoot is still possible, although not equally frequent in all species. The cymose branches are very much shortened here, espe- cially in sect. Brackenridgea where flowers are almost sessile on the rachis. In some species there are several branches per inflorescence, each bearing 3 (-5) flowers. In other species there are only a few branches, each bearing 7 or more flowers in pseudo-umbels. In sect. Notochnella the cymose branches are not shortened so much, whereas the bracts on the rachis sometimes have a more leaf-like appearance. The inflorescences in subfam. Sauvagesioideae are more of a paniculate nature. They are profusely branch- ing in Schuurmansia, but in the other genera most branches are very much shortened, flowers and fruits of diff"erent age standing closely together. In Neckia the rachis is bearing several bracts, but only one (terminal?) flower. Anatomy. The Ochnaceae are characterised by the presence of cortical bundles without resin canals (GiLG, Ber. Deut. Hot. Ges. 11, 1893, 20-25). The subfam. Ochnoideae shows typical 'cristarque' cells in branches and leaves (van Tieghem, Bull. Mus. Hist. Nat. Paris 8, 1902, 266-273). For other data, see Metcalfe & Chalk, Anat. Dicot. 1 (1950) 104, 108, 333-338, 340, f. 76, 77, and Decker, Phytomor- phology 16 (1966) 39-45. Palynology. The pollen of the Ochnaceae only show interesting diff"erences at generic or higher level, but they do not diff'er fundamentally. See Erdtman, Pollen Morph. PI. Tax. (1952) 290, and Muller, Rev. Palaeobot. Palynol. 9 (1969) 149-173. Phytochemistry. No data available. Some species are used locally for medicinal purposes, because of bitter components of unknown nature. Uses. No economically important applications of Malesian spp. have been recorded. For properties of the wood of some species, see under Ochna integerrima and Gomphia serrata. Note. The present revision is based on my precursory treatment of Indo-Pacific Ochnaceae in Blumea 16 (1968) 1-83. KEY TO THE GENERA 1. Stamens 10-cv). Carpels (3-)5-10(-15), free. Fruits 1-5, 1-seeded drupes on a swollen torus. Leaves distichous. 2. Anthers opening with 2 apical pores. Stipules intrapetiolarly united. Inflorescences with (l-)3-oo, ± remote flowers, usually thyrsoid, sometimes simple cymes. 3. Stamens 12-cvi; ovaries (3-)5-10(-15); embryo straight. Leaves without an intra-marginal nerve. 1. Ochna 3. Stamens 10; ovaries 5; embryo curved. Leaves with a distinct intra-marginal nerve. 3. Gomphia 2. Anthers opening with 2 longitudinal slits. Stipules free. Inflorescences of umbelloid appearance with cv) flowers in conferted, cymose clusters of 3 or more 2. Brackenridgea 1. Stamens 5. Carpels 2-5, fused. Fruit a more-seeded berry or capsule; torus not distinctly enlarged in fruit. Leaves alternate, not distichous. 5. Ovary 5-carpelled, 5-celled. Fruit a berry. Anthers opening by 1 apical pore. Leaf margin always stiffly, though sometimes very finely, denticulate 4. Euthemis 5. Ovary 3-carpelled, 1 -celled. Fruit a capsule. Anthers opening by 2 longitudinal slits. Leaf margin entire or (bi)serrulate. 6. Seeds not winged. Fruit opening with 3 valves. Inflorescences simple, or compound and (nearly) all branches shortened. Leaves evenly spaced. Leaf margin (bi)serrulate. Shrubs or undershrubs. 7. Inflorescences axillary; the rachis bearing a varying number of bracts, but only 1 flower. Under- shrubs, up to 1 m, but often much smaller 5. Neckia 7. Inflorescences terminal (or pseudo-axillary by sympodial growth), many-flowered. Shrublets or shrubs, up to 7 m high. 8. Staminodes 10, in 1 whorl. Fruit subglobose. Leaf blades oblanceolate, up to 35 cm long. 6. Indovethia 8. Staminodes oo, in more than I whorl, those of the inner whorl larger. Fruit fusiform. Leaf blades linear oblong, up to 17 cm long 7. Schuurmansiella 6. Seeds winged. Fruit opening with 3 longitudinal slits under the persistent style. Inflorescences much-branched panicles. Leaves distinctly tufted. Leaf margin entire, glandular dotted. Treelets, usually not exceeding 15 m 8. Schuurmansia 1971] OcHNACEAE (Kanis) 99 1. OCHNA LiNNE, Gen. PI. ed. 5 (1754) 229; Sp. PI. 1 (1753) 513; Kanis, Blumea 16 (1968) 22, 83. — Diporidium Wendl./. in Bartl. & Wendl. Beitr. Bot. 2 (1825) 24; O.K. Rev. Gen. PI. 1 (1891) 104, emend, illeg.. incl. typ. Ochna. — Discladium Tiegh. Bull. Mus. Hist. Nat. Paris 8 (1902) 214, nom. illeg. — Polythecium Tiegh. Ann. Sc. Nat. Bot. VIII, 16 (1902) 196, 366. — Pleopetalum Tiegh. Bull. Mus. Hist. Nat. Paris 9 (1903) 163. — Polvthecanthum Tiegh. Ann. Sc. Nat. Bot. IX, 5 (1907) 160, 175. — Fig. 1. Shrubs or treelets, sometimes undershrubs. Stipules small, intrapetiolarly united, caducous. Leaves shortly petioled, chartaceous or subcoriaceous; nerves curved upward, especially near the margin, not joining; veinlets — at right angles to the nerves near the midrib and joining in irregular secondary nerves, ^z transverse near the margin. Inflorescences lateral or terminal thyrses with a terminal flower; peduncle — persistent, bearing many, small, distichously conferted, caducous bracts at base, leaving a distinct annulus of scars; pedicels filiform, articulate. Flowers with — hemispherical torus, distinctly tumid and red in fruit. Sepals 5, greenish, accrescent and turning red in fruit. Petals 5-10, in 1-2 whorls, yellow. Stamens v: in 2 or more whorls; filaments subterete; anthers opening with 2 apical pores. Ovaries 5-10(-15), obovoid; ovule atropous; stigmas as many as ova- ries, on short branches or zb united. Fruits l-3(-5), greenish, turning black when ripe. Distr. The majority of the species is found in Africa, south of the Sahara, and in Madagascar. In India and Ceylon 4 spp. occur, one of these ranging from Assam to Indo-China, Hainan, the Nicobar Is., and Malesia: in the North of the Malay Peninsula. Ecol. Adapted to a seasonal climate, on poorer soils, below 1500 m. Dispersal by birds because of conspicuous black fruits on red torus and calyx (Ridley, Disp. 1930, 419). 1. Ochna integerrima fLouR.) Merr. Trans. Am. fruit up to 5 cm, the basal 2-8 mm persistent. Phil. Soc. n.s. 24, 2 (1935) 265, emend.; Kanis, Torus '/2-I mm high, 1 '/2-2 1/2 mm 0, in fruit up Blumea 16 (1968) 36; Fl. Thail. 2 (1970) 25. — to 6 mm high, 10 mm 0. Sepals 5, ovate to ovate- Elaeocarpus inregerrimus Lour. Fl. Cochinch. oblong, 10-16 by 4-9 nun. Petals 5-6(-10), obo- (1790) 338. — O. wallichii Planch, in Hook. vate, 15-25 by 8-15 mm, tapering at base or Lond. J. Bot. 5 (1846) 650; King, J. As. Soc. subunguiculate. Stamens (25-)30-60(-75); fila- Beng. 62, ii (1893) 231; Ridl. J. Str. Br. R. As. ments 2'/2-7 mm, unequal, the outermost longest; Soc. n. 59 (1911) 83; Craib, Fl. Siam. En. 1 anthers 4-6 by 0.4-0.8 mm. Ovaries 6-10(-15), (1931) 244; Burk. Diet. 2 (1935) 1569. — O. 0.7-1.1 by 0.5-0.7 mm; style 10-15 by r. 1/2 mm, crocea Griff. Not. PI. As. 4 (1854) 463; Kurz, in fruit up to 20 mm; stigmas sometimes on up to J. As. Soc. Beng. 40, ii (1871) 49, err. in syn. 1 mm long branches. Fruits mostly 2-3, up to 11 Gomphia sumatrana Jack; King, ihid. 62, ii (1893) by 8 mm. 233. — O. grandis RiDL. J. Str. Br. R. As. Soc. Distr. NE. India, E. Pakistan, Burma, Anda- n. 59 (1911) 83; Fl. Mai. Pen. 1 (1922) 365. — man and Nicobar Is., Thailand, Laos. Cambodia, Ouratea crocea (Griff.) Blrk. Diet. 2 (1935) Vietnam, Hainan, in Malesia: Malay Peninsula \614; Kew Bull. (]935) '^]», p.p., quoad typus. — (Peninsular Thailand, Perils, Kcdah, Langkawi FiR. 1. Is). Deciduous undershrub, shrub, or treclet up to Ecol. From sea-level up to 1200m in hilly 12 m and 45 cm 0. Stipules 5-8 by 2-3 mm. country, in moist or dry, deciduous forests, often Leaves mostly obovatc-oblong or (obovatc-) of a mixed Diptcrocarp type, on loamy, sandy, or lanceolate, rarely obovatc or lincar-lanccolatc, rocky soils. Tall specimens are found near river 6-20(-25) by 2-7 cm, mostly acuminate, some- banks, small shrubs near seashores. Flowering times acute or obtuse at apex, mostly acute, shortly beforcor during development of new leaves: sometimes obtuse at base, margin fmcly dcnticu- in the northern part of the area mainly in Febr. late; petiole 2-5 mm. Inflorescences compound, and March, in the southern part generally a numy-flowcrcd; rachis Vi-^Vi^-^) cm; branches little earlier, but less restricted, especially in the 1-3-flowcrcd, monochasial; pedicels 2-4 cm, in Malay Peninsula. 100 Flora Malesiana [ser. I, vol. 7^ Fig. 1. Ochna integerrima (Lour.) Merr. a. Fruiting twig, xVj, b. inflorescence, x^/s, c. stamen, x4, d. gynoecium, x4, e. fruit, x2 (a & e Ridley 15746, b-d Garrett 1346). 1971] OcHNACEAE (Kanis) 101 Uses. The bark tastes bitter and yields a diges- Tiegh. Ann. Sc. Nat. Bot. VIII, 16 (1902) 356 = tive tonic (Cochinchina). The wood is recorded O. mauritiana Lamk, fide Kanis, Blumea 16 as used for huts in the Andamans. U968) 80. Properties of wood. Light brown, hard, close- A species based on one collection by Riedle, grained and brittle. erroneously recorded for Timor. The specimen must have been collected during Capt. Baudin's Excluded expedition on I'lle de France (Mauritius) on the Ochna decaisnei TiEGH. Bull. Mus. Hist. Nat. way to Timor. It must have been mislocalized later Paris 8 (1902) 47-49. — Diporiditini decaisnei on. 2. BRACKENRIDGEA A. Gray, New Gen. PI. (1853) 5, preprint of Proc. Am. Ac. Arts Sc. 3 (1857) 51; Kanis, Blumea 16 (1968) 41. — Campylopora Tiegh. Bull. Mus. Hist. Nat. Paris 8 (1902) 547. — Notochnella Tiegh. I.e. 549. — Fig. 2. Trees or treelets. Stipules small, free, often more or less laciniate, caducous. Leaves shortly petioled, chartaceous, glossy above, nerves strongly curved to the apex, often some of the lower ones partly parallel to the margin, the higher ones joining successively, veinlets branching, ± transverse. Inflorescence thyrsoid, but of umbelloid appearance, made up of simple or compound, distichously arranged, shortened cymes, the rachis often growing on vegetatively after flowering; bracts small, broadly linguiform to triangular, i laciniate, ± caducous, often many at base of inflorescence, leaving a distinct annulus of scars; pedicels filiform, 4: ac- crescent and turning red in fruit, articulate at base. Flowers with i hemispherical torus, distinctly tumid and red in fruit. Sepals 5, accrescent, fleshy and red in fruit. Petals 5(-10), white or yellow. Stamens 10 (or tn); filaments subterete; anthers dehiscing from the apex downwards by longitudinal slits. Oxaries 5{-\0), ohoyo\6.\ ovule camptotropous, epitropous, ± annularly curved around 2 connecting intrusions of the endocarp; stigma small. Fruits l-2(-5), greenish, turning black or almost so when ripe. Distr. 2 or 3 spp. in tropical eastern Africa and Madagascar, probably forming a distinct section; in Malesia 4 spp. in 2 sections arc found; a related species occurs in NE. Queensland and Fiji. Ecol. Confined to the everwct tropical areas, up to c. 1000 m. Dispersal mainly by birds because of conspicuous, black fruits on red torus and calyx (Guppv, Obs. Nat. Pacif. 2, 1906, 569; Ridley, Disp. 1930, 265). The fruits arc also capable of floating because of two air-filled spaces between exocarp and endocarp; this was recorded from the Kapuas R. (Beccari, Wand. Borneo 1904, 187) and from the New Guinea seadrift (Hemsley, Bot. Chall. Exp. 3, 1885, 289, t. 54). No specimen has ever been collected in beach forest. key to the species I. Petals 5. Stamens 10. Ovaries 5. (Sect. Brackenridgea). 2. Inflorescences never axillary, but sometimes terminal on short side-branches. 3. Inflorescences made up of many-flowered cymes, the pedicels in 2 or more tiers. Leaves mostly 71/2-20 cm long I. B. hookcri 3. Inflorescences made up of 3(-5)-flowered cymes, the pedicels in 1 tier. Leaves mostly 4-12'/: cm long 2. B. palustrls 2. Axillary inflorescences always present when in fertile state 3. B. forbcsii I. Petals 5-7-(IO). Stamens 10-45. Ovaries 5-10. (Seel. Notochnella) 4. B. fascicularls 1. Section Brackenridgea Brackenridgea A. Gray. - Campylopora Tiecjh. Cymes much confcrted; peduncle and branches much shortened. Flowers of 1 102 Flora Malesiana [ser. I, vol. 7^ cyme flowering simultaneously. Corolla regularly 5-merous, white. Stamens 10, in 1 whorl. Ovaries 5. Distr. Chiefly Malesian with 2 spp. in the Malay Peninsula and Borneo, one reaching Sumatra, Palawan, and Celebes, the other reaching the Andamans and Ko Chang; another sp. in New Guinea. Also 1 sp. in NE. Queensland and Fiji. 1. Brackenridgea hookeri (Planch.) A. Gray, New Gen. PI. (1853) 6, preprint of Proc. Am. Ac. Arts Sc. 3 (1857) 51; Furtado, Gard. Bull. Sing. 19 (1962) 182; Kanis, Blumea 16 (1968) 45; Fl. Thail. 2 (1970) 27. — Gomphia hookeri Planch, in Hook. Lond. J. Bot. 6 (1847) 3; MiQ. Fl. Ind. Bat. 1, 2 (1859) 675, err. 'G. gla- berrima Planch.'; Benn. in Hook./. Fl. Br. Ind. 1 (1875) 525; King, J. As. Soc. Beng. 62, ii (1893) 233, excl. var. corymbosa; Ridl. Fl. Mai. Pen. 1 (1922) 366. — Ochna hookeri (Planch.) O.K. Rev. Gen. PL 1 (1891) 106. — B. perakensis TiEGH. Ann. Sc. Nat. Bot. VIII, 16 (1902) 396. — Gomphia corymbosa (King) Ridl. J. Str. Br. R. As. Soc. n. 54 (1910) 33, p.p. excl. typus; Fl. Mai. Pen. 1 (1922) 367. — Oiiratea hookeri (Planch.) Burk. Kew Bull. (1935) 318; Diet. 2 (1935) 1615. — B. denticiikita Furtado, Gard. Bull. Sing. 19 (1962) 183. — Fig. 2b». Tree, up to 33 m, 1 m 0. Leaves oblong to lanceolate, 7'/2-20 by 2 1/2-6 cm, obtuse to acute, sometimes acuminate at apex, acute, often — tapering at base, margin entire, ~ acicular denticulate in young treelets; petiole 5-12 mm. Inflorescences terminal, made up of many-flowered shortened, sessile cymes, flowering successively or simultaneously, the rachis often not growing on vegetatively after flowering; pedicels 10-15 mm, up to 20 mm in fruit, those of one cyme fixed close together in c. 3, + distinct tiers. Torus c. 1/2 mm high, 1 mm 0, in fruit up to 3 mm high, 5 mm 0. Sepals ovate to obovate, 4-5 by 1 '/2-2 mm. Petals ovate to obovate, 3'^-5 by IVi-i-A mm. Filaments 1-1 '/2 mm; anthers 1 '/2-2 by c. Vj mm. Ovaries c. 0.7 by 0.5 mm; style n/2-3 mm, in fruit up to 5 mm. Fruits up to 6'/2 by 51/2 rnm. Distr. India: Andamans, and Thailand: Ko Chang (NE. Gulf of Thailand); in Malesia: Malay Peninsula and Borneo. Ecol. From sea-level up to 750m, in kerangas forests on sandy soils, on dry hillocks in swampy forests, in primary lowland Dipterocarp forests and in hill forests. Vern. Mai. Pen.: bunga kelat merah, bunga maskam, kayu luru, Malacca; Borneo: empodat, Sarawak, semukau, P. Madjang. 2. Brackenridgea palustris Bartell. Malpighia 1 5 (1901) 165, t. 10; Furtado, Gard. Bull. Sing. 19 (1962) 183; Kanis, Blumea 16 (1968) 46; Fl. Thail. 2 (1970) 27. — B. hookeri (Planch.) A. Gray var. leucocarpa Scheff. Nat. Tijd. N. I. 32 (1873) 411. — Gomphia hookeri Planch, var. corymbosa King, J. As. Soc. Beng. 62, ii (1893) 233. — B. serrulata Bartell. Malpighia 15 (1901) 163, t. 9; Furtado, Gard. Bull. Sing. 19 (1962) 184. — B. kingii Tiegh. Ann. Sc. Nat. Bot. VIII, 16 (1902) 395. — B. corymbosa (King) Tiegh. I.e. 395. — B. rubescens Tiegh. I.e. 396. — Gom- phia corvmbosa (King) Ridl. J. Str. Br. R. As. Soc. n. 54 (1910) 33, p.p.; Fl. Mai. Pen. 1 (1922) 367. — Ochna forworthyi Elmer, Leafl. Philip. Bot. 5 (1913) 1823; Merr. En. Philip. 3 (1923) 68. — B. foxworthyi (Elmer) Furtado, Gard. Bull. Sing. 19 (1962) 184. — Fig. 2. Tree, up to 30 m, 1.2 m 0. Leaves (ovate-) oblong to (ovate-)lanceolate, A-MYi by 1 V2-S cm (up to 20 cm long on young treelets), mostly acute to acuminate, sometimes obtuse at apex, rounded to acute, often ~ tapering at base, margin entire, i acicular denticulate in young treelets; petiole 3-10 mm. Inflorescences terminal, made up of 3(-5)-flowered, shortened cymes, sessile, or with up to Vi cm long peduncle, flowering simul- taneously, sometimes 2 inflorescences of different stages in close succession, the 5-10(-25) mm long rachis growing on vegetatively, sometimes branch- ing in the lower parts; pedicels 8-15 mm, up to 20 mm in fruit, those of one cyme fixed close together in one tier. Torus c. I/2 mm high, 1 mm 0, in fruit up to 4 mm high, 6 mm 0. Sepals ovate to oblong, 3 1/2-6 by 1 V2-3 mm. Petals ovate to obovate-lanceolate, 3 Yi-l by 1 '/2-3 mm. Filaments (i/2-)l-2(-2i/2)mm; anthers 1 '/2-3 by c. 1/2 mm. Ovaries c. 0.7 by 0.5 mm; style (l-)2 1/2-4 mm, ri accrescent. Fruits up to 8 by 6 mm. Distr. Malesia: Sumatra, Malay Peninsula, Borneo, Philippines (Palawan), and Celebes. key to the subspecies 1. Anthers 1 '/2-2 mm long. Style 21/4-4 mm long during anthesis 1. ssp. palustris 1. Anthers 2-3 mm long. Style IV2-2V2 mm long during anthesis. 2. Filaments c. 2 mm long during anthesis. Ovaries c. 0.7 by 0.5 mm. . 2. ssp. foxworthyi 2. Filaments c. V2 mm long during anthesis. Ovaries c. 0.5 by 0.3 mm. . 3. ssp. kjellbergii 1. ssp. palustris. — All synonyms except Ochna foxworthyi Elmer. Cymes 3-flowered, sessile, up to 5-flowered and with up to 1/2 cm long peduncle in young treelets. Sepals 31/2-51/2 by 11/2-3 mm. Petals 31/2-6 by 11/^-3 mm. Stamens with 1-2 mm long filaments; anthers 1 1/2-2 mm long, up to 3 mm in young treelets. Ovaries c. 0.7 by 0.5 mm; style 21/^-4 mm during anthesis. Fruits up to 8 by 6 mm. Distr. Malesia: Sumatra, Mentawai Is., Banka, Billiton, Malay Peninsula, Borneo. Ecol. Usually found in the lowlands, but oc- 1971] OCHNACEAE (KaiUS) 103 Fig. 2. Brackenridgea palustris Bartell. a. Flowering twig, X 2/3, b. part of twig with scars of flowers and bracts arranged in simple cymes, x4 ib'. ditto of B. hookeri (Planch.) A. Gray, scars of many flowers arranged in shortened, cymose, partial inflorescences, - 4), c. petal from bud, d. flowerbud, sepals and petals removed, e. flower, petals and anthers shed, sepals removed, all y 6,/. fruiting flower, x 2 (a-b & e Rahmat si Toroes 4187, c-d ditto 4162,/Iboet 214; b' Paie 13589). casionally up to 1000 m, reported from peat- swamp forests and from kerangas forests on sandy, sometimes rather rocky soils, with humic podsols. Uses. The wood is reported twice as being used in house-building (Malacca, Sarawak). Vern. Sumatra: niajang majang, mampat, rampat dahan, st'niang, M, kaju barat laiit, k. galugus badak, k. ludt', k. saholat, k. to pa tupa, Kota Pinang Distr.; niadu Itiai, niensulung kaju, Banka; nn'nsolongang, Biliiton; Mai. Pen.: lidah mura, Pahang, pindurah, cheharahan, Malacca; Borneo: mala iindang, limur bt'sih. P. Madjang. 2. 5.1/7. foxworthyi (Elmer) Kams, Biumca 16 (1968) 48. — Ochna foxworthyi Elmer. Cymes .^-flowered, sessile. Sepals 5-6 1/2 by 2-3 mm. Petals 5-7 by 1 V2-- mm. Stamens with P/i-2Vi mm long filaments; anthers 2-3 mm long. Ovaries c. 0. 7 by 0.5 mm; style 1 Vi-'^Vi mm long during anthcsis. Fruits up to 5 by 4 mm. Distr. Malesia: Philippines (Palawan). Ecol. At sea-level, along river in forest, and from 150 m on rocky hillside near river-bank. 3. ssp. kjellbergii Kams. Biumca 16 (1968) 48. Cymes 3-flowcrcd, sessile. Sepals 4-5 Vi by 1 Y2-2 mm. Petals 4— 4V2 by 1 V4-2Y4 mm. Stamens with V^-V^ mm long filaments; anthers 2V2-3 mm long. Ovaries c. 0.5 by 0.3 mm; style 1-2 mm long during anthesis. Fruits up to 5 by 4 mm. Distr. Malesia: Celebes. Ecol. At sea-level in swamp and from 400 m at the edge of a lake. 3. Brackenridgea forbesii Tiegh. in Morot, J. Bot. 16 (1902) 46, nom. nud.; Ann. Sc. Nat. Bot. VIII. 16 (1902) 395, descr.; Pulle. Nova Guinea 8 (1912) 667; Rendle, J. Bot. (1923) Suppl. 7; Kanis, Blumea 16 (1968) 49. Tree, up to 30 m, 55 cm 0. Leaves oblong to lanceolate, 5-15 by 1 '/2-5 cm, acute to acuminate at apex, acute or a little tapering at base, entire; petiole 3-5 mm. Inflorescences terminal and axillary, made up of a varying number of mostly 3- to 5- flowered, shortened, sessile cymes, flowering simultaneously, the 2-5(-IO) mm long rachis mostly growing on vegetativcly when terminal, sometimes when axillary; pedicels r. V2 cm. up to I cm in fruit, those of one cyme fixed close to each other in one tier. Torus c. ' j mm high, V2-V* mm 0, in fruit up to 2'/2 mm high, 4 mm 0. Sepals ovate to elliptic, 3-4'/2 by I'/^-P/^ mm. 104 Flora Malesiana [ser. I, vol. 7^ Petals obovate-lanceolate, 3-4'/2 by 1-1 Va mm, acute at apex. Stamens with c. 1 '/a mm long filaments; anthers c. 1 '/a by '/s mm. Ovaries c. 0.6 by 0.5 mm; style 1 '/2-2 mm, in fruit up to 3 mm. Fruits up to 6 by 5 mm. Distr. Malesia: New Guinea. Ecol. Primary rain-forest on flat country to steep slopes, up to 750 m, on clay, sand, or peat, in sites which may be inundated during the wettest season. Vern. W. New Guinea: jobias, Je, obaisang, Mooi, serukdeho, Manikiong. 2. Section Notochnella (TiEGH.) Kanis, Blumea 16 (1968) 43. — Notochnella Tiegh. Cymes ± remote; peduncle ± distinct; branches ± shortened, sometimes unequal in length, the longer overtopping the central flower. Flowers of 1 cyme flowering successively. Corolla irregular, yellow. Stamens c/^, in more than 1 whorl. Ovaries 5-10. Distr. Malesia: Philippines (excl. Palawan), monotypic. 4. Brackenridgea fascicularis (Blanco) F.-Vill. Nov. App. (1880) 40; Kanis, Blumea 16 (1968) 43. — Ochna fascicularis Blanco, F1. Filip. ed. 2 (1845) 245; ibid. ed. 3, 2 (1878) 92; Vidal, Sinopsis (1883) 19, t. 27A; Rev. PI. Vase. Filip. (1886) 79; Bartell. Malpighia 15 (1901) 162; Merr. Gov. Lab. Publ. Philip. 27 (1905) 29; Sp. Blanc. (1918) 263; En. Philip. 3 (1923) 68. — Diporidium fasciculare (Blanco) O.K. Rev. Gen. PI. 1 (1891) 105; Tiegh. in Morot, J. Bot. 16 (1902) 203. — Notochnella fascicularis (Blanco) Tiegh. Bull. Mus. Hist. Nat. Paris 8 (1902) 549; Ann. Sc. Nat. Bot. VIII, 16 (1902) 403. — Ouratea mindanaensis Merr. Philip. J. Sc. 17 (1920) 287; En. Philip. 3 (1923) 68. Tree up to 25 m, 30 cm 0. Leaves oblong to lanceolate, 5-15 by 2-5 cm, sometimes obtuse, mostly acute to zt acuminate at apex, obtuse to acute, sometimes a little attenuate at base, margin entire or r: finely denticulate; petiole 4-8 mm. Inflorescences terminal, made up of a varying number of many-flowered, ~ shortened cymes, sometimes separate cymes in the axils of normal leaves; pedicels 1-2 cm, up to 3 cm in fruit, the basal 1-5 mm persistent. Torus c. 1 mm high, 1 V2 mm 0, in fruit up to 4 mm high, 7 mm 0. Sepals elliptic to obovate, 6-8 by 3-4 mm. Petals obovate to obovate-lanceolate, 6-8 by 2V2-4'/2 mm. Filaments 1 1/2-2 V2 mm; anthers 21/2-3 by c. 1/2 mm. Ovaries 0.7-0.8 by 0.5-0.7 mm; style c. 4 mm, in fruit up to 7 mm. Fruits up to 7 by 6 mm. Distr. Malesia: Philippines. Ecol. In primary forests at low and medium altitudes, once reported from logged Dipterocarp forest and once along a stream. key to the subspecies 1. Stamens 20-45 . Cymes with 2-5 mm long peduncles 1. ssp. fascicularis 1. Stamens 10-15. Cymes with 5-10(-20) mm long peduncles 2. ssp. mindanaensis 1. ssp. fascicularis. — Ochna fascicularis Blanco. Inflorescences with 1-21/2 cm long rachis; cymes with 2-5 mm long peduncles, up to 15-flowered. Sepals 5. Petals 5-7 (-10). Stamens 20^5. Ovaries 7-10. Distr. Malesia: Philippines (Luzon, N.Visayas). Vern. Philip.: aniatan, dirigkalin, bitas, mala- kiting kiting, masalisi, Luzon, Tag., bansilai, Visayas, Bis. 2, ssp. mindanaensis (Merr.) Kanis, Blumea 16 (1968) 44. — Ouratea mindanaensis Merr. Inflorescences with 2-6 cm long rachis, often indistinct by development of bracts to normal leaves; cymes with 5-10(-20) mm long peduncles, up to 7-flowered. Sepals (3-)5. Petals (3-)5. Stamens (8-)10-15. Ovaries 6-7. Distr. Malesia: Philippines (Mindanao). Doubtful Brackenridgea elegantissima (Wall.) Kanis, Blumea 16 (1968) 50. — Euthemis elegantissima Wall, in Roxb. Fl. Ind. 2 (1824) 305; in Hook. Bot. Misc. 2 (1830) 77, note; Planch, in Hook. Lond. J. Bot. 5 (1846) 647; ibid. 6 (1847) 2, err. in syn. Gomphia sumatrana Jack; Benn. in Hook. /. Fl. Br. Ind. 1 (1875) 526; King, J. As. Soc. Beng. 52, ii (1893) 235; Bartell. Malpighia 15 (1901) 160; RiDL. Fl. Mai. Pen. 1 (1922) 368. — Euthemis ? pulcherrima Wall, ex Benn. in Hook. /. Fl. Br. Ind. 1 (1875) 526, err. in syn. Gomphia suma- trana Jack; King, J. As. Soc. Beng. 52, ii (1893) 233; Bartell. Malpighia 15 (1901) 160. Based on one sterile, juvenile specimen from Singapore I. It can not be decided whether it belongs to B. hookeri (Planch.) A. Gray or to B. palustris Bartell., although it should be cer- tainly one of these species. Wallich's epithet is the oldest available for any of the species in Brackenridgea. 1971] OcHNACEAE (Kanis) 105 3. GOMPHIA SCHREB. Gen. PI. ed. 8 (1789) 29\, p.p.; Kanis, Taxon 16 (1967) 420, 422; Blumea 16 (1968) 51. — Ochna Linne, Sp. PI. 1 (1753) 513; Gen. PI. ed. 5 (1754) 229, p.p. e.xcl. typ. — Ouratea [non Aubl. Hist. PI. Gui. Fr. 1 (1775) 397] Baill. Hist. PI. 4 (1873) 367, emend., p.p. e.xcl. typ. — Meesia Gaertn. Fruct. 1 (1788) 344, nom. rej., non Hedw. Sp. Muse. (1801) 173, nom. cons. — Campylospernmm TiEGH. in Morot, J. Hot. 16 (1902) 40, 194, 197, nom. superfl. — Cercinia Tiegh. I.e. 198. — Campylocercwn Tiegh. Bull. Mus. Hist. Nat. Paris 8 (1902) 546. — Fig. 3. Shrubs or treelets. Stipules small, intrapetiolarly united, caducous. Leaves shortly petioled, chartaceous, nerves close, parallel, ± straight, curving upward near the margin, forming an inconspicuous marginal nerve and a conspicuous, somewhat wavy nerve parallel to the margin at some distance, veinlets reticulate, joining in irregular secondary nerves parallel to the primary ones. Inflorescences lateral and/or terminal thyrses with ± reduced branches; peduncle ± persistent with sometimes a few small bracts at base, not leaving a distinct annulus of scars; pedicels filiform, articulate at base. Flowers with a short, columnar, 5- ribbed gynophore, enlarging and sometimes turning subglobular in fruit. Sepals 5, tinged pinkish, accrescent. Petals 5, yellow, creamy or white. Stamens 10 in 1 whorl; filaments terete, very short; anthers opening with 2 apical pores. Ovaries 5, obovoid; ovule camptotropous, epitropous; stigma punctiform. Fruits l-2(-5), yellowish green, turning dark purple or blue-black when ripe. Distr. The majority of the species is found in Africa, S. of the Sahara, and in Madagascar. One sp. in SW. Peninsular India, Ceylon, E. Thailand, Indo-China, Hainan, and W. Malesia. Ecol. Confined to tropical areas with an everwet climate or with a moderately dry monsoon, up to 1 500 m. Dispersal possibly by birds, but the fruits are not as conspicuous as in Ochna and Brackenridgea, as calyx and torus are not coloured. I. Gomphia serrata (Gaertn.) Kanis, Taxon 16 (1923) 68; Ridl. Kcw Bull. (1930) 76. — Ochna (1967) 422; Blumea 16 (1968) 53; Fl. Thail. 2 angustifolia (Vahl) O.K. Rev. Gen. PI. I (1891) (1970) 28. — Meesia serrata Gaertn. Fruct. 1 106. — Ochna sumatruna (Jack) O.K. I.e. 106. (1788) 344, t. 70, f. 6. — G. angustifolia Vahl, — Ouratea sumatrana (Jack) Gilg in E. & P. Symb. Bot. Upsal. 2 (1791) 49; Scheff. Nat. Nat. Pfl. Fam. 3, 6 (1895) 142; Bartell. Mal- Tijd. N. I. 32 (1873)411; Benn. in Hook./. Fl. pighia 15 (1901 ) 160; Hall./. Beih. Bot. Centralbl. Br. Ind. 1 (1875) 525; F.-Vill. Nov. App. (1880) 34, 2 (1916) 35; Merr. J. Str. Br. R. As. Soc. n. 86 39; ViDAL, Sinopsis (1883) 19; Rev. PI. Vase. (1921) 387; Ridl. Kew Bull. (1930) 76. — Owra/fa Filip. (1886) 79; Phan. Cuming. (1895) 101; feornee-nj/j Bartell. Malpighia 15 (1901) 1 56, t. 6; Backer, Schoolfl. (1911) 194. — G. sumatrana Merr. J. Str. Br. R. As. Soc. n. 86 (1921) 387; Jack, Mai. Misc. 1, 5 (1821) 29; in Hook. Bot. Ridl. Kcw Bull. (1930) 76. — Ouratea nerii/olia Misc. 2 (1830) 77; Planch, in Hook. Ic. PI. II, 4 Bartell. Malpighia 15 (1901) 158, t. 7, sphalm. (1845) t. 712; in Hook. Lond. J. Bot. 6 (1847) 2; • neerifolia \ Merr. J. Str. Br. R. As. Soc. /;. 86 Mio. Fl. Ind. Bat. 1,2 (1859) 675; Sumatra (I860) (1921 ) 387; Airy Shaw, Kcw Bull. (1940) 249. 209, 534; Scheff. Nat. Tijd. N. I. 32 (1873) 41 1; — Ouratea hcccariana Barieli. Malpighia 15 Benn. in Hook./ Fl. Br. Ind. 1 (1875) 525, err. (1901) 159, t. 9; Merr. J. Str. Br. R. As. Soc. /;. 'sumatrensis'; King, J. As. Soc. Beng. 62, ii (1893) 86 (1921 ) 387; Airy Shaw, Kcw Bull. (1940) 249. 232; Ridl. Trans. Linn. Soc. II, Bot. 3 (1893) — C'ampylospermuni sunuiiranuiii (Jack) Tiegh. 285; Fl. Mai. Pen. 1 (1922) 365; Rendie, J. Bot. in Morot. J. Bot. 16 (1902) 197; Ann. Sc. Nat. Bot. (1924) Suppl. 16; Ridl. Kcw Bull. (1925) 79; Vlll, 16 (1902) 298; /Ak/. 18(1903)21. Campylo- II. Mai. Pen. 5 (1925) 296; Merr. J. Arn. Arb. spermum hurnccnse (Bartell.) Tiegh. in Morot, 13 (1952) 226. - Ouratea angustifolia (Vahi ) J. Bot. 16 (1902) 197; Ann. Sc. Nat. Bot. Vlll, 16 Baill. ex Laness. PI. Util. Col. Fr. (1886) 607; (1902) 301. — Campylospernwm hcccarianum KrK)RD. Exk. Fl. Java 2 (1912) 107; Hail./ Bcih. (Bartell.) Tieoh. Ann. Sc. Nat. Bot. Vlll, 16 Bot. Centralbl. 34. 2 (1916) 34; Merr. J. Sir. (1902) 301. — Campylocercum neriifolium Br. R. As. Soc. n. 86 (1921) 387; Ln. Philip. 3 (Bariell.) Tiegh. I.e. 304. - Campylocercum 106 Flora Malesiana [ser. I, vol. 7^ Fig. 3. Gomphia serrata (Gaertn.) Kanis. a. Flowering twig, x^i, b. part of leaf, vAVi, c. flower, x2, d. petal, /A, e. flower, sepals and petals removed,/, stamen, abaxial view, g. gynoecium with gynophore, all X 6, h. fruiting flower, x 2 ia-g Ampuria 41444, h SAN 25040). 1971] OcHNACEAE (Kauis) 107 zoUingeri Tiegh. I.e. 305. — Campyhspermiim wallichianum TiEGH. Bull. Mus. Hist. Nat. Paris 9 (1903) 76; Ann. Sc. Nat. Bot. VIM, 18 (1903) 17. — Campy lospenmun plicatum Tiegh. Bull. I.e. 78; Ann. I.e. 19. — Campylospermum strictum Tiegh. Bull. I.e. 79; Ann. I.e. 20. — Campylo- spermum kingHTiEGH. Bull. I.e. 79; Ann. I.e. 21. — Campylospermum perakense Tiegh. Bull. I.e. 80; Ann. I.e. 21. — Campylospermum abbreviatum Tiegh. Bull. I.e. 80; Ann. I.e. 21. — Campylo- spermum eumingii Tiegh. Bull. I.e. 80; Ann. I.e. 22. — G. mierophylla RiDL. Fl. Mai. Pen. 1 (1922) 365, f. 38. — G. oblongifolia Ridl. Kevv Bull. (1925) 281; Fl. Mai. Pen. 5 (1925) 296. — Ouratea areta Craib, Kew Bull. (1926) 341; Backer & Bakh./. Fl. Java 1 (1963) 327. — Ouratea megaearpa RiDL. Kew Bull. (1930) 76. — Ouratea miero- phylla (Ridl.) Craib, Fl. Siam. En. 1 (1931) 245. — Ouratea sumatrana (Jack) Gilg var. nervosa Craib, I.e. 245. — Ouratea eroeea (Griff.) BuRK. Kew Bull. (1935) 318, p.p. e.xel. typ. — Fig. 3. Shrub or tree, up to 25 m, 40 cm 0. Branchlets of young plants sometimes scrambling. Lfarei ovate- to obovate-lanceolate, 6-20 by 2-6 cm (up to 35 by 10 cm in young plants), chartaceous, mostly acute to acuminate, sometimes obtuse at apex, acute or a little tapering at base, margin finely denticulate; petiole 2 1/2-7 '/z mm. Infloreseenees many-flowered; rachis (2V2-)5-20(-35) cm, pri- mary branches of terminal inflorescences usually up to 10(-15) cm long, secondary branches or primary ones of lateral inflorescences usually reduced or up to 1 cm long, cymose, with (1-) 3-7(-:v), — conferted flowers; pedicels '/2-% cm, up to 1 cm in fruit. Torus 0.7-1 mm high, 0.7-1 mm o, in fruit up to 5 mm high, 5 mm o. Sepals mostly ovate to elliptic, sometimes obovatc, 4-7 by 2' 2-4 mm, mostly acute to obtuse, some- times rounded. Petals obliquely obovate to broad- spatulate, 4'/2-8 by 2V2-6 mm, obtuse, rounded or truncate, mostly yellow, sometimes creamy or white. Stamens subsessile or with up to '/2 mm long filaments; anthers 2'/2-5(-6) by 0.5-0.8 mm. Ovaries 0.7-1 by 0.4-0.6 mm; style 3-5 mm long, up to 7 mm in fruit; stigma minute. Fruits up to 8(-10) by 6(-8) mm. Distr. SW. Peninsular India, Ceylon, E. Thai- land, Indo-China, Hainan; in Malesia: Sumatra, Banka, Malay Peninsula, Karimundjawa Is., Borneo, Philippines, and Celebes. Ecol. From sea-level up to 1200 m, up to 1500 m on Mt Kinabalu, in primary and secondary vegetation: lowland and submontane, mixed Diptcrocarp forests, peat swamp forests, high kerangas forests, ridge forests, and open scrub veg- etation; on level land to steep slopes, also near river-banks and on cliffs near the sea. Soils arc rocky, sandy, loamy or clayish, but usually acid; limestone is reported occasionally, but it is very probable that the soils in the localities concerned have been leached out. fhc flower buds arc often galled by unidentified insects, especially in the Malay Peninsula, Su- matra, and the smaller islands cast of Sumatra. These buds are about ovoid, with small sepals and petals somewhat sunken in an enlarged torus, whereas gynoecium and androecium are strongly suppressed. Ripe fruits in normal flowers are often empty, which is probably also caused by insect damage, as perforations of the pericarp are sometimes clearly visible. It is recorded once that ants did attack a collector's press for these fruits (Clemens, Indo-China). Seeds are rare in herbariuni speci- mens of Malesian collections. Uses. The wood is reported as being used as poles or planks for the construction of houses in Ceylon, Vietnam, Sumatra, Malay Peninsula, and Sarawak. The lea\es are chewed by jungle tribes in Malaya. Roots and leaves are bitter and are decocted in S. India for a stomachic and anti- emetic tonic. Young branches are used against tooth-ache in Cambodia. Properties of wood: dull red or red-brown, hard and strong, apt to split in drying. Vern. Sumatra, North: kaju ndolak, Batak; West: kaiek djambak, st'balusi, Minangkabau; South: kaju mat, k. sepa/i, lakodjong, madjang madjang; Banka: mentungging, mesulung puti/i; Mai. Pen., Thailand: long pling; Malaya: bunga kclimbing bt'si, ehinta mula, lidah mura, mala kctam batu, membatu, nu'narah, mcndapor, meramong, murmagong, penarah, pengling, pokok Icbah, p. luas, lampang besi; several compound names were recorded occasionally with gironggang (= Crato.xylon), jambu and kt'lat (= Eugenia); Borneo, Sarawak: ki-ladang, kclutak, I ban, aam, Kenyah; compound names with ubah or ubar (= Eugenia) were recorded occasionally; Brunei: e/itUiaga lampang, Iban, pinis, Malay; Sabah: majang majang, quiiitalai, Kadazan, antimagas gimbaan, Murut, kolambang, Rungus Dusun, posoon, tulangkara, Kinabatangan, alas, tampala- nuk, Tidong, bakan bitanag, biobi, malatangor, ondogong, Malay; West Borneo: nn'lindingan, Da- yak; East Borneo: amir burgang, Dayak, batu batu, mulak, tingkedjing kcring, Malay; Philip., Pa- lawan: anduyong, bibingo, huisae; Luzon: salaetoe, Zambales, simahima, Bicol, sasa/iit, Tagalog; Visayas: buloeauan, earanan, Panay, minsaray, Mindoro; Celebes: It^bani, parasinga fjila, Bugi, mampa, Rangkas, morosisio, wulisi mapute, Tobcla. Notes. As this species grows under rather dif- ferent ecological conditions, the variation in its morphological characters is not surprising. Apart from these variations, some geographically more or less separated 'races' can be recognized. This is the reason why in certain Floras two or three difl"crent species have been recognized although the distinctive characters can hardly be used at a specific level. Over a large area it is impossible to make a key to these 'races', because the same dif- ferences occur in remote parts of the specific area. In Sumatra and the Malay Peninsula plants arc relatively uniform, the greatest variation occurring in Borneo, especially in Sarawak. In more ex- posed habitats, on clifls and on poor kerangas soils, specimens often have small leaves. On 108 Flora Malesiana [ser. I, vol. V limestone in the Langkawi Islands a race 'micru- phylla is found with small leaves and very reduced inflorescences. Similar, but less reduced forms are found elsewhere along the coasts of the Malay Peninsula. Excluded Gomphia magnoliaefolia Zipp. ex Span. Linnaea 15 (1841) 186, nam. nud. = Pycnarrheiia longifolia (Decne) Becc. (Menispennaceae), fide Diels, Pflanzenreich, Heft 46 (1910) 51. 4. EUTHEMIS Jack, Mai. Misc. 1, 5 (1821) 15; Kanis, Blumea 16 (1968) 62. — Fig. 4. Shrubs or shrublets, sparsely branched. Stipules free, caducous. Leaves coria- ceous, glabrous, denticulate, nerves numerous, parallel, from the midrib curving sidewards, straightly ascending to the marginal veins at an angle of c. 80'; petiole rt winged. Inflorescences terminal, many-flowered, compound racemes; bracts small, caducous. Flowers ^ or polygamous. Sepals 5, turning purplish red in fruit. Petals 5, white or pinkish. Staminodes 0(-5), filamentous. Stamens 5, free; anthers subsessile, rostrate. Ovary 5-celled; ovules 2 per cell, pendulous, axile; stigma minute. Fruit a berry with 5 pyrenes. Seeds 1 (2) per cell. Distr. SW. Cambodia, in Malesia: Sumatra, Malay Peninsula, Borneo. Ecol. Everwet tropical forest below 1250 m, in kerangas forests, on low ridges in peat-swamp forests, and in open ridge forests, on poor, mostly sandy soils. Dispersal probably by birds because of conspic- uous, white, rose-pink or red berries (Ridley, Disp. 1930, 410). KEY TO THE SPECIES 1. Inflorescence a panicle, branches well developed with scattered flowers. Leaves 8-40 cm long, margin distinctly denticulate. Mature fruit white 1. E. leucocarpa L Inflorescence a very slender, often cernuous raceme, nearly all branches reduced with conferted flowers. Leaves 4-15 cm long, margin faintly denticulate. Mature fruit red 2. E. minor 1. Euthemis leucocarpa Jack, Mai. Misc. 1, 5 (1821) 16; Wall, in Roxb. Fl. Ind. 2 (1824) 303; Jack in Hook. Bot. Misc. 2 (1830) 69; Planch, in Hook. Ic. PI. II, 4 (1845) t. 711; MiQ. Fl. Ind. Bat. 1, 2 (1859) 675; Sumatra (I860) 208, 533, incl. var. latifolia; Scheff. Nat. Tijd. N. I. 32 (1873) 411; Benn. in Hook. /. Fl. Br. Ind. I (1875) 526; King, J. As. Soc. Beng. 62, ii (1893) 234; RiDL. Trans. Linn. Soc. II, Bot. 3 (1893) 285; Bartell. Malpighia 15 (I90I) 167; Hall. /. Beih. Bot. Centralbl. 34, 2 (1917) 30; Merr. J. Str. Br. R. As. Soc. n. 86 (1921) 388; Ridl. Fl. Mai. Pen. 1 (1922) 368; Diels, Bot. Jahrb. 60 (1926) 311; BuRK. Diet. I (1935) 987; Atry Shaw, Kew Bull. (1940) 249; Merr. J. Am. Arb. 33 (1952) 224; Vidal, Adansonia 1 (1961) 60; Kanis, Blumea 16 (1968) 62; Fl. Thail. 2 (1970) 29. — E. robusta HoOK. /. Trans. Linn. Soc. 23 (1862) 163; Bartell. Malpighia 15 (1901) 168; Hall. /. Beih. Bot. Centralbl. 34, 2 (1917) 32; Ridl. Fl. Mai. Pen. 1 (1922) 368. Shrub up to 6 m. Branchlets stout, green. Stipules ovate, 4-6 by c. 2 mm, acute to acuminate, ciliate. Leaves oblong to linear oblong, 8-40 by 2-10 cm, acute at apex, tapering at base, margin distinctly and irregularly denticulate, nerves 1-2 mm apart; petiole 2-5 cm. Panicles erect, 8-20 cm; pedicels 4-10 mm, articulate at base; bracts 8-10 by 2-4 mm, lanceolate, acute. Flowers ^, erect, often in pairs. Sepals obliquely ovate to elliptic, unequal, 4-7 by 2-3 V^ mm, ciliate. Petals obliquely obovate to spatulate, 4—10 by 21/2-5 mm. Anthers 3-5 by c. 1 mm, yellow. Ovary ovoid to bottle-shaped, 2-4 by c. 1 mm, style 1 1/2-3 rnm. Fruit globular, up to 1 cm 0, fleshy, via red turning white. Seeds like sectors of a sphere, c. 4 by 2 mm. Distr. SW. Cambodia; in Malesia: Sumatra, Riouw & Lingga Is., Banka, Billiton, Malay Peninsula, Anambas Is., Borneo. Ecol. From sea-level up to 1000 m, on poor soils, preferably in moist, shady places (see also under the genus). Uses. Medical applications of the roots is reported from Malaya. In Brunei the fruits are used against eye-diseases. Vern. Sumatra: belusung putih, kaju padang, mata pelanduk, Banka; balong, Billiton; Malaya: pilawan berok; Borneo: tambu, Sarawak, ranggas hutan, Sabah, iur iiir, W. Borneo. 2. Euthemis minor Jack, Mai. Misc. 1, 5 (1821) 18; Wall, in Roxb. Fl. Ind. 2 (1824) 304; Jack in Hook. Bot. Misc. 2 (1830) 69; MiQ. Fl. Ind. Bat. 1, 2 (1859) 675; Sumatra (1860) 209, 534; Scheff. Nat. Tijd. N. I. 32 (1873) 412; Benn. in Hook. /. Fl. Br. Ind. 1 (1875) 526; King, J. As. Soc. Beng. 62, ii (1893) 235; Bartell. Malpighia 15 1971] OcHNACEAE (Kanis) 109 Fig. 4. Euthemis minor Jack. a. Fertile twig. ^ 2/,. h. detail of leaf margin. • 4, c. flowcrbud.y. open J flower, both x4, e. ditto, stamen, ad- and abaxial view,/, ditto, reduced pistil, both • 6. ;r- P'stil of + wcr, x4, h. fruit, /I, i. seed, -4 {a~c Mkijf.r 21807, '6, 7. fruit, x2, k. seed, x 10 (a after Ic. Bog. 1, t. 1, b-g BuwALDA 6256, A-y Teysmann HB 10893, k Brooke 9455). 7. SCHUURMANSIELLA Hall./ Rec. Trav. Bot. Neerl. 10 (1913) 344; Kanis, Blumea 16 (1968) 73. — Fig. 7. Shrubs with slender branchlets. Stipules acicular. Leaves short-petioled, nerves ascending at an angle of r. 80 . Injhrescences terminal, many-flowered, compound racemes; branches much shortened, with flowers of successive age; bracts small, those of one branch conferted. Fhnvers '. Sepals 5. Petals 5, white or pinkish, purplish at base, in anthesis soon surpassing the sepals in size. Staminoiles co, of 2 types, the inner ones mutually and with the stamens connate at base. Ovary 3-carpelled; stigma 1. Style of ripe fruits also splitting i nto 3 parts. .Sfc^/v not winged. Distr. Malesia: NW. Borneo. Monotypic. 114 Flora Malesiana [ser. 1, vol. 7' Fig. 7. Schuurmansiella angustifolia (Hook./.) Hall./, a. Fertile twig, X 2/3, b. flowerbud, x4, c. flower, petals shed, x4, d. part of androecium, inner view, x6, e. pistil, x8,/. fruit, g. open fruit, both x2, /i. seed, x6 (a-ZAbang Muas 5139, g Anderson 8368, h Bujang 12990). 1. Schuurmansiella angustifolia (Hook./.) Hall./ Rec. Trav. Bot. N^erl. 10 (1913) 345, t. 7; Merr. J. Str. Br. R. As. Soc. n. 86 (1921) 387; Kanis, Blumea 16 (1968) 73. — Schuurtnansia angustifolia Hook. / Trans. Linn. Soc. 23 (1862) 157. — Fig. 7. Shrub up to 7 m high. Stipules up to 12 by c. '/2 mm. Leaves linear-oblong, 8-17 by %-l'/4 cm, long acuminate at apex, ± tapering at base, chartaceous, margin serrulate, nerves V2-I rnni apart. Inflorescences c. 1 5 cm long; rachis c. 1 mm ; branches with up to 5 flowers; pedicels filiform, up to 12 mm in fruit; bracts acicular, 2-3 mm under the branches, smaller at base of pedicels. Flowers 1971 OcHNACEAE (Kanis) 115 erect. Sepals — linguiform, 1-1 '/i by '4-1 mm. Petals ovate, 3V2-4H by P/4-2',4 mm, distinctly reflexed. Outer siaminodes :^, small; inner ones 25-30, linear, 2-3 mm, purplish. Stamens with flattened filaments V2-I rnm long, purplish; anthers lanceolate, l'/2'-2l2 by 0.3-0.6 mm. Ovary ovoid, c. 1.2 by 0.7 mm; style c. 0.3 mm, purplish; stigma capitate. Fntit ellipsoid, sub- trigonous in cross-section, c. 8V2 by 3 mm. Seeds 0.5-1 by 0.2-0.5 mm, tomentose. Distr. Malesia: NW. Borneo (Sarawak, 1st and 4th Div.). Ecol. A lowland species, reported from up to 600 m, in moist places, especially in kerangas forests on poor soils, and on sandstone cliffs, even near the sea. 8. SCHUURMANSIA Blume, Mus. Bot. Lugd. Bat. 1 (1850) 177, t. 32; Kanis, Nova Guinea, Bot. 6 (1961) 63; Blumea 16 (1968) 74. — Fig. 8-10. Trees or treelets with stout, often hollow branches. Stipules entire. Leaves conferted at intervals (fig. 10), glandular along the margins, nerves almost straight- ly ascending at an angle of 60-70'. Inflorescences terminal, many-flowered, panicles often with many hypsophylls at base; peduncle and branches distinctly ribbed, often with con- or recaulescences. Flowers ^ or polygamous. Sepals 5. Petals 5, white, creamy or purplish. Siaminodes ;•:, in 1 or 2 whorls, the inner ones mutually and with the stamens connate at base. Ovary 3-carpelled; stigmas 1 or 3. Fruit opening with 3 longitudinal slits, the style usually not splitting. Seeds winged like a propeller with 2 blades. Distr. Solomons and Bismarcks; in Malesia: Borneo, Philippines, Celebes, Moluccas, and New Guinea. Ecol. Pioneer plants, from sea-level up to 3000 m, especially in natural or anthropogenous secondary vegetation. KEY TO THE SPECIES 1. Style as long as the ovary or longer; stigma capitate or punctiform 1. S. elegans 1. Style half as long as the ovary or shorter; stigma 3-lobed. 2. Filaments — twice the length of the anthers 2. S. vidalii 2. Filaments as long as the anthers or shorter 3. S. henningsii 1. Schuurmansia elegans Blume, Mus. Bot. Lugd. Bat. I (1850) 178, t. 32; MiQ. Fl. Ind. Bat. 1, 2 (1859) 118; Illustr. I (1871)66, t. 29; Capit. Bull. Soc. Bot. Fr. 57 (1910) 397; Hall./ Rec. Trav. Bot. N6erl. 10 (1913) 346; Merr. Philip. J. Sc. 11 (1916) Bot. 19; En. Philip. 3 (1923) 68; Heine in Fcddc, Rep. 54 (1951) 240; Pfl. Samml. Clemens Kinab. (1953) 63; Kanis, Nova Guinea, Bot. 6 (1961) 64, f. la; Blumea 16 (1968) 75. — S. par- viftora Ridl. Trans. Linn. Soc. II, Bot. 9 (1916) 18. — S. horneensis RiDL. Kcw Bull. (1930) 77. Trcelet or tree, up to l5(-30?) m, sometimes with low stiltroots. Stipules linguiform, I '/2-4 by l'/i-2'/2 mm, sometimes shortly ciliate. Leaves obovalc-oblong to -lanceolate, 1 0-30 by 2 '/2- 1 0cm, rounded or somewhat acuminate at apex, ± tape- ring at base, margins r involute, chartaccous, nerves 1-1 'Z: mm apart; surface of dried leaves very finely reticulate by protruding intercellular walls of epidermal cells, giving an impression of striation parallel to the nerves; petiole 1 Vi-f* cm. Inflorescences 10-25 cm; peduncle 2-5 mm 0; pedicels filiform, 3-6 mm; bracts broadly lin- guiform, up to 2 mm long, sometimes shortly ciliate. Flowers bisexual, erect. Sepals elliptic to obovate, 3-6 by 1 '/2-4 mm. Petals obovate, 4-8 by 2-5 mm. Outer staminodes 0-co, linear, 1-1 '/^ mm long; inner ones 15-25, linear to spatulate, 2'/^-5 mm long, with 1 distinct nerve. Filaments 1-2 mm; anthers 1 V^-l by c. Vi mm, connective distinctly protruding. Ovary subovoid, 1 '/2-3 by I '/4-2 mm, papillate, sometimes with a few glan- dular-capitate hairs; style I '/2-3 mm, growing in fruit, widening into the ovary; stigma small, capitate or punctiform. Fruit fusiform, up to 2'/2 by '/2 cm, acuminate. Seeds c. % by '/4 mm, with c. 3 mm long, slender wings. Distr. Malesia: Borneo, Philippines (Min- danao, only I collection of uncertain identity reported from Agusan), Celebes, Moluccas, and New Guinea. Ecol. From sea-level up to 2000 m, in primary and secondary forests, on level land to steep slopes, sometimes near river-banks or in swampy localities; on clay or more sandy or rocky soils. Probably not uncommon locally, but rarely flowering. Vcrn. Philip., Mindanao: tananf;, Munobo; 16 Flora Malesiana [ser. I, vol. 7^ Fig. 8. Schiiurmansia henningsii K. Sch. a. Fertile branch, X 1/2, b. ^ flower, X 5, c. ditto, part of androe- cium, inner view, d. ditto, reduced pistil, both x8, e. ? flower, x5,/. ditto, sepals and petals removed, X8, g. fruit, h. open fruit, both x 1 1/2, '• seed, x5 {a-d Hoogland & Pullen 5356, e-/ Brass 29335, g Hoogland 3507, h-i BW 4987). Celebes: labo labo, Toradja; Ambon: wat lopu; New Guinea, Vogelkop Pen.: hikselah, Tehid. 2. Schuurmansia vidalii (F.-Vill.) Merr. Philip. J. Sc. 11 (1916) Bot. 19; En. Philip. 3 (1923) 69; Kanis, Nova Guinea, Bot. 6 (1961) 69, f. Ic; Blumea 16 (1968) 76. — Calophyllum vidalii F.-ViLL. in Ceron, Cat. PI. Herb. Manila (1892) 229, plate. — S. parvifolia Merr. Philip. J. Sc. 1 1 (1916) Bot. 19. Treelet up to 7 m. Stipules 1-2 V2 by 1-2 mm, shortly ciliate. Leaves obovate-lanceolate, 5-11 by 1 1/2-^ cm, obtuse to obtusely acuminate at apex, tapering at base, margins somewhat involute, strongest nerves c. 1 mm apart, papyraceous to chartaceous; petiole V2-2 cm. Inflorescences 7-15 1971 OcHNACEAE (Kanis) 117 Fig. 9. Scliiiuriminsia lienningsii K. ScH. After anthcsis, Cycloop Mts (Photogr. Sleumer, July 1961). cm; peduncle l'/2-3'/2 mm o; pedicels 1-2 mm; bracts very small, semi-annular to triangular. Flowers 1, erect. Sepals obovate, 2-2V4 by I '2-2 mm. Petals obovate, c. 3 1/2 by 2'/2 mm, white or pink. Outer siaminodes 10-^, filiform, '/4-I nim long; inner staminodes c. 25, linear, I '/2-2 mm, r. Vi nim connate at base. Filaments c. \^/\ mm; anthers V^-\ by 0.4-0.5 mm. Ovary subglobular. - 3-lobed, 0.4-0.5 mm 0, glabrous; stigma 3- lobcd. subsessilc. Fruit fusiform, c. 10 by 3 mm, acuminate. Seeds c. 0.6 by 0.4 mm, with c. 2 mm long, basally connate wings. Distr. Malesiu: Philippines (Luzon: Aurora, Camarincs Sur & Sorsogon Prov., 3 collections). Ecol. In (mossy) forest, from 500 to well over l(X)0 m, possibly also at sca-lcvcl. 3. Schuurmansia hcnningsii K. ScH. Bot. Jahrb. 9 (1888) 210; K. Sch. & Hollr. FI. Kais. Wilh. Land (1889) 50; Warb. Bot. Jahrb. 13 (1891)283; F:. & P. Nat. Pn. Fam. 3. 6 (1895) f. 75; K. Sen. & Laut. FI. Schutzgcb. (1901) 448; Pulle, Nova Guinea 8 (1912) 667; Hall./. Rec. Trav. Bot. Nderl. 10 (1913) 346; E. & P. Nat. Pfl. Fam. cd. 2, 21 (1925) f. 41; Lane Poole. For. Res. (1925) 116; WiiitE & Francis. Proc. R. Soc. Qucensl. 38 (1927) 247; White, J. Arn. Arb. 10 (1929) 241; Kanis, Nova Guinea, Bot. 6 (1961) 65, f. lb; Blumca 16 (1968) 76. — S. bamleri K. ScH. & Laui. FI. Schutzgeb. (1901) 448; Nachtr. (1905) 318, incl. var. longifolia Laut.; Hall. /. Rec. Trav. Bot. Nierl. 10 (1913) 350. - S. gilidana Laui. in K. Sch. & Laut. FI. Schut/geb. Nachtr. 118 Flora Malesiana [ser. I, vol. 7^ (1905) 319; Hall. /. Rec. Trav. Bot. N^erl. 10 (1913) 348. — S. microcarpa Capit. Bull. Soc. Bot. Fr. 57 (1910) 398, t. II, 12, 17; Baker, J. Bot. 61 (1923) Suppl. 4. — S. theophrasta Hall./. Rec. Trav. Bot. N^erl. 10 (1913) 346; Holth. & Lam, Bliimea 5 (1942) 213. — S. pseiulopalnm Hall./. Rec. Trav. Bot. Neerl. 10 (1913) 347. — S. rauwolfioides Hall. / I.e. 349; K. Sch. Bot. Jahrb. 9(1888)211, siue nomen. — S. lungifoUa (Laut.) Gilg in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 80; A. C. Smith, J. Arn. Arb. 22 (1941) 524. — IS. lophiroides GiLG, I.e. 80. — ? 5. oreophila Gilg, I.e. 80. — ? S. seblecltteri Gilg, I.e. 80. — ? S. erassinervia Gilg, I.e. 80. — S. eoriaeea A. C. Smith, J. Arn. Arb. 22 (1941) 525. — S. montana A. C. Smith, /. c. 526. — S. grandiflora A. C. Smith, I.e. 527. — Fig. 8-10. Treelet or tree, up to 15(-20?) m, sometimes with stiltroots up to 1 m high. Stipules i/^-5 by 1/2-3 mm, often up to 3 mm long, ciliate. Leaves obovate-lanceolate, 6-85 by l'/^-15 cm, obtuse to acuminate at apex, tapering at base, margins some- what involute, nerves 2-7 mm apart, chartaceous or subcoriaceous; petiole up to 4 cm. Infloreseen- cej7-65cm; peduncle 1 '/2-8mm0; pedicels filiform, 2-5 mm; bracts usually very small, ± triangular, sometimes larger and transitional to leaves. Flowers ^ or functionally unisexual, erect. Sepals obovate to elliptic, 3-5 by 1 1/^-3 mm, greenish, sometimes purplish. Petals obovate-oblong, 4-7 V2 by 1 1/2-4 mm, white, creamy, pink or purplish red. Outer staminodes Q-00, filiform, e. 1 mm long; inner ones 5-30, filiform to linear, 1 1/2-2 mm long. Stamens in S flowers with %-2i/^ mm long filaments; anthers II/2-21/2 by V2-VA mm; in $ flowers with V4-V2 mm long filaments; anthers c. 1 by 1/3 mrn- Ovary subglobular to ovoid, ± 3-lobed, glabrous, in S flowers I/2-I by 1/3-% mm, in ? flowers c. 3 by 2 mm; style cylindric, up to 1 mm; stigma 3-lobed. Fruit fusiform, up to ^Vi by 34 cm, acuminate. Seeds c. 1 by '/3 mm with c. lYi mm long, slender wings. Distr. Solomons and Bismarcks; in Malesia: Moluccas (Talaud Is., Halmahera, Ceram), New Guinea. Ecol. From sea-level up to 3000 m, in primary and secondary forests, on landslides and in secondary grasslands, on level land to steep slopes, sometimes near river-banks or in swampy lo- calities, on clay or more sandy or rocky soils; scattered or locally common in open habi- tats. Vern. Moluccas, Talaud Is.: arisusii iirune; Tidore: maletope; Ambon: ut lapii, was lapii; W. New Guinea, Ajamaru: batsjevak, Maibrat; Biak I.: rambiian; NE. New Guinea, Wabag: opaga, orpaeh; Hagen: pappai, popai, porkai, piipai; Minj: biibar, bubus; Chimbu: akessa, u'ltlareh, hahessa, nienmen; Aiyura: arebi; Budemu: sipidund; Huon Pen., Sambui: pelip; Sattelberg: (m)beli; Andarora: haiwinge, Nauti, yatsiga. Manki; SE. New Guinea, Tari: obbo; Kutubu: karadewa; Mendi: op; Kairuku: engefukenge: Buna: kembiisa; Bausa: werawera. Notes. The striking variation in vegetative characters is for the greater part due to diff"erences in ecological conditions. The greatest influence is exercised by the altitude. Papuan material, ar- ranged according to increasing altitude, shows a very regular decrease in leaf size, especially be- tween 1000 and 3000 m. At the same time the leaves become more coriaceous and more distinct- ly petioled. Exposition and age of the plants play a less important role in the determination of the leaf size. Fig. 10. Schiiiirnuinsia henningsii K. Sch. Small- leaved, young plant, Edie Creek, Morobe Distr., e. 2000 m, Michael holding the twig (NGF 1 3949) (Photogr. Sleumer, 1961). The dimensions and colours of flower parts are also rather variable, but less easy to correlate with other data. These variations probably indicate genetical differences between local populations. More detailed local studies will be necessary to decide whether any infraspecific taxa can be dis- tinguished. A majority of the specimens has relatively large anthers and small pistils, whereas a minority has relatively small anthers and large pistils. It was observed once, that most specimens in a profusely flowering stand had proportionally large anthers and did not set fruit. It is not certain, how- ever, that the flowers are always functionally uni- sexual. Polygamy is likely to occur, as the reduc- tion of either stamens or ovary is never complete. Excluded Sauvagesia erecta Linne, Sp. PI. 1 (1753) 203; MiQ. Fl. Ind. Bat. 1, 2 (1857) 118; Koord. Exk. Fl. Java 2 (1912) 607; Backer & Bakh./. Fl. Java 1 (1963) 327; Kanis, Blumea 16 (1968) 80. 1971] OcHNACEAE (Kanis) 119 Erroneously recorded for Java by Miquel without reference to material or other source of information. His error might be caused by a collection of Sauvagesia erecta L. in G, which is labelled: 'Perrottet, Java, 1819". Although Perrottet visited the eastern part of Java in that particular year, it is clear that the speci- men concerned was mislabelled and in fact was collected in Cayenne on the same voyage. Tetramerista MiQ. Fl. Ind. Bat. Suppl. (1860) 534 belongs to the Theaceae, fide GiLG, Ber. Deut. Bot. Ges. 1 1 ( 1 893) 22, et aiict. div. PORTULACACEAE (R. Geesink, Leyden) Annual to perennial, erect or creeping, mostly branched herbs or shrubs, oc- casionally woody at the base, often with a tuberous or swollen main root, oc- casionally rooting at the nodes. Leaves spirally arranged to opposite, subsessile, occasionally with axillary hairs or scales (in Mai. only in Pornilaca), nervation pinnate or reticulate. Flowers bisexual, actinomorphous (occasionally cleistoga- mous), in axillary and/or terminal thyrsi, dichasia, in terminal capitules or solitary (terminal or axillary). Bracts leaf-like or membranous. Sepals 2 (4-8 in extra-Mai. Lew'isia and Grahamia), boat-shaped, deltoid to obovate at base shortly connate and confluent with petals and stamens. Petals (3-)4-6(-8), mostly obovate and unequal, shortly connate. Stamens (l-)3-c/3, in l-c/^ ± distinct whorls; filaments basally shortly connate; anthers 2- or 4-celled, dorsifixed, dehiscing lengthwise. Ovary superior or half-inferior, originally 2-20-celled, soon becoming 1 -celled; style with 2-20 mostly papillous arms. Ovules 4-c/: on a central, dendroid placenta, campylotropous. Capsule 3-7-valved or with a caducous operculum, occasionally surrounded by the persistent calyx. Seeds 1-c/^, smooth or ornamented, kidney- shaped to ~ globular, laterally compressed, mostly with a caruncle. Embryo curved, almost filling the ripe seed. Distribution. About 15 genera with possibly 200 spp. Cosmopolitan, with some tropical species occurring as adventives in temperate regions. In Malesia 4 genera with 1 1 spp. Ecology. The Malesian species are all more or less succulent herbs or semi-shrubs, the aquatic Montiafontana excepted. Only two genera are native, Portulaca and Montia. All Portulacas occur at low altitude, preferably in disturbed vegetation, waste places, or on the coast. Most are indifferent to climate, only P. macrorhiza and P. pilosa ssp. siindaensis are restricted to the seasonal climate of the Lesser Sunda Is. The sandy beach is tolerated by P. oleracea, P. liitea, and P. pilosa ssp. pilosa (race tuberosa). The occurrence of Montia fontana in the high mountain bogs of New Guinea is singularly interesting, as it is the only tropical montane occurrence in the Old World between its range in the temperate northern hemisphere and the southern counterpart in the SE. Australian Alps (at c. 36"^ SL), SE. Australia (Mt Lofty Ranges), Tasmania, and New Zealand, an almost bipolar type of distribution. Flower-biology. As far as known all Portulacaceae are self-pollinating. Whether cross-pollination occurs, deserves further study. Self-pollination results in genetically constant local populations, representing pure lines in nature. The production of seeds is profuse; seeds can usually stand a long time of drought. Man has doubtless been responsible for the dispersal of several Portulacaceae. Seed can be transported with cargo; some species are used as food or as ornamentals, and they easily escape as adventives and maintain in warm regions. For these reasons the native range of certain species cannot any longer be ascertained, notably of Portulaca oleracea, P. pilosa, P.quadrifida, Talinum paniculatum, and T. triangulare. The structure of the inflorescence in this family is mostly, possibly always, of a cymose nature. Through contraction and reduction several modifications arc represented; some derivations are schematically depicted in figures I & 2. Talinum has in principle a thyrse (fig. la, lb & 2) of which a full explanation is given in the notes under the genus. At first sight CaloncJrinia would seem to possess a true raceme (fig. Id) but this could also well be a reduced thyrse. Montia has also probably a reduced cymose inflorescence (fig. Ic). The inflorescences in Portulaca are in siihg. Portulacella (Australia) the most primitive and arc compound dichasia from which the structure in the capituliform inflorescences of the other subgenus can be derived, as I have explained in my precursor (Blumea 17, 1969, 277). Phytochemistry. Red betacyanins (replacing anthocyanins) and yellow betaxanthins form an outstanding chemical character of the family. Such nitrogen-containing pigments (chromoalkaloids) were demonstrated to be present in members of the genera Anacampseros, Calamlrinia, Claytonia, Montia, Portulaca, Spraf!uea, and Talinum. Saponins seem to occur frequently in the faniily; their chem- istry, however, is not yet known. The seeds store starch in the pcrispcrm and fatty oil in embryos; at the same time they contain appreciable amounts of protein. The chemical features known at present from this family confirm its intimate relationships with other members of the ccntrospcrmous alliance. A summary of phytochemical literature is to be found in Heonauer, Chcmotax. d. Pfl. 5 (1969) 383-387. - R. HFftNAUER. (121) 122 Flora Malesiana [ser. I, vol. 7^ Fig. 1. Scheme of the inflorescences of some Portidacaceae and their morphological relations, a. Thyrsus in Talinuin paniculatiim, b. corymboid thyrsus, resembling a pleiochasium, in Talinum triangulare, c. raceme-like inflorescence, cymose or racemose, in Montia fontana, d. raceme with terminal flower in Calandrinia grandiflora. Morphology. Von Poellnitz (in Fedde, Rep. 37, 1934, 240, and other papers) called the sepals "Involucralblatter", suggesting their homology with the involucral leaves surrounding the capituli, the latter being called by him "falsche Involucralblatter". Also Legrand (in Com. Bot. Mus. Hist. Nat. Montevideo 31,1, 1953, 1, and other papers) called the calyx lobes "pseudosepalos". Taking into account, that the calyx normally is considered as (metamorphosed) leaves and that in Portidacaceae the phyllotaxal position of the sepals is so distinctly set off" against the bracts, it seems to me that there is no morphological argument to accept the flower as being monochlamydeous. Anatomy. See also my precursory paper (Blumea 17, 1969, 276-279) on the axillary hairs; Metcalfe & Chalk, Anat. Dicot. 1 (1950) 153; Kowal, Monogr. Bot. 12 (1961); Chorinski, Oest. Bot. Z. 80 (1931) 308. Uses. See Heyne, Nutt. PI. (1927); Ochse & Bakh. Ind. Groent. (1931) 615; Burkill, Diet. Ec. Prod. Mai. Pen. (1935); Caius, J. Bomb. Nat. Hist. Soc. 41 (1939) 369. See also under the species. KEY TO the genera 1. Fruit dehiscing circumscissile with an operculum. Ovary half-inferior 4. Portulaca 1. Fruit valved or irregularly dehiscing. Ovary superior. 2. Leaves c. 4 mm long, opposite 3. Montia 2. Leaves more than 6 mm long, the middle cauline leaves spirally arranged. 3. Seed glabrous. Fruit globular, 3-5 mm 0. Calyx caducous or not 2. Talinum 3. Seed hairy. Fruit obpyriform, c. 10 by 8 mm. Calyx persistent, distinctly accrescent. 1. Calandrinia 1971] PoRTULACACEAE (Geesink) 123 1. CALANDRINIA H.B.K. Nov. Gen. Sp. 6 (1823) 77, ?iom. cons.; Poelln. in Fedde. Rep. 35 (1934) 161-173; D.C. Anon Suarez de Cullen, Bol. Soc. Arg. Bot. 5 (1953) 1-29. Mostly succulents, occasionally with short stems, occasionally unbranched. glabrous, pubescent or glandular. Leaves spirally arranged and/or in a basal rosette, occasionally with axillary hairs. Flowers arranged in mostly terminal, occasionally axillary thyrsi (corymboid or not), glomeruli or dichasia, or solitary. Bracts and bracteoles subulate or leaf-like, partly without an axillary axis. Calyx accrescent (caducous in the American sp. C. punae). Sepals glabrous, pubescent, spiny or glandular, occasionally with dentate, ciliate or glandular margin. Petals 5-7, persistent, twisted after anthesis. Stamens 1>-^, anthers linear to elliptic. Oxary superior; style 1 with 3-5 arms (seldom 3 simple styles). Fruit pyriform, globular, obpyriform or elleplic, 3-7-valved. Seeds at least occasionally without a caruncle. Distr. About \5Q spp., native in tropical and subtropical America and Australia. 1. Calandrinia grandiflora LiNDL. Bot. Reg. (1828) smaller. Sepals roundish, up to c. 1 'A ^rn o. t. 1194: G. Don, Gard. Diet. 3 (1834) 80, t. 18; acuminate in fruit. Petals 5-6, zn orbicular, up to L. H. Bailey, Man. Cult. PI. (1949) 336; Backxr c. 1 cm o, purplish. Stamens c. 75; filaments up to & Bakh./. FI. Java 1 (1963) 217. — Fig. Id. 1 cm; anthers c. 2 by 0.8 mm. Style c. 2'/: mm. Erect, up to c. 30 cm. Leaves spirally arranged, with several short, thick lobes. Fruit obpyriform, obovate, up to 13 by 5 cm, without axillary hairs; c. 10 by 8 mm, 3-valved. Seeds "v, c. 1 mm o; petiole semi-amplexicaulous; apex acute to acu- testa cells about hexangular, with hair-like ap- minate. Flowers in compound terminal racemes at pendages. least occasionally with a terminal flower; each Distr. Native of S. America, sometimes escaped raceme 1-6-fiowered. Bracts and bracteoles ovate from gardens, to elliptic, acute, up to c. 1 by '/i cr". apical ones 2. TALINUM JussiEU. Gen. (1789) 312, nom. cons, prop.; Poelln. in Fedde, Rep. 35 (1934) 1; Dandy, Taxon 18 (1969) 464. — Fig. la, b, 2. Herbs or semi-shrubs (in Malesia glabrous). Leaves spirally arranged (the lower- most sometimes opposite), linear to obovate. Flowers in terminal whether or not corymboid thyrsi, or cymosely arranged, seldom axillary or solitary. Sepals mostly caducous. Petals mostly 5. Stamens 5~y. Ovary superior; style mostly with 3 arms. Fruit globular; mostly 3-valved or irregularly caducous. Distr. About 50 spp., native in S. and Central America and S. Africa, the two treated species now pantropically naturalized. Morph. The inflorescences of the Talinums studied are explained in comparison with the vegetative ramifications, thus in agreement with C. Troll, who assumes that there is no essential ditVerencc between the vegctatise and the inflorcsccntial ramification. Their leaves arc spirally arranged, the lowermost sometimes excepted. Specimens of T. paniculutuni grown at Leydcn in winter under low light intensity and short day light produced all opposite leaves and opposite primary infloresccnlial branches. The flowers of T. paniculatum are arranged in a wide thyrsc with a terminal flower (fig. la). A change in phyllotaxis. in contrast with the rule of Troll mentioned, is found in the ultimate dichasial ramifications. Superficially. T. trianffulare is trichasial in the first ramification of the inflorescence (fig. lb. 2b). I have tried to bring this in agreement with the inflorescence of T. paniculatum by the following argu- mentation: the vegetative branches always have 2 basal cataphylls. each with a dormant axillary bud (fig. 2a C, B). The inflorescences lack these cataphylls and buds. The cataphylls easily fall, and mostly (he scars arc hardly to be found. The cataphylls are also present if the axillary axis is not developed (fig. 2a ). In the cultivated specimens these dormant buds developed after the axillary axis was pinched 124 Flora Malesiana [ser. I, vol. 7^ M B LiiiXi Fig. 2. Talinuin triangulare (Jacq.) Willd. Schematic demonstration of the homology of the vegetative ramifications and those of the inflorescence. In all three figures L means leaf bearing the lateral axis A. The main axis is indicated by M. Each lateral axis carries 2 cataphylls at the base indicated by C, each cataphyll having an axillary bud indicated by B, which remains normally dormant in the vegetative part (fig. a), but develops into an axis in the inflorescence (fig. b), but can by removal of the lateral axis also artificially be produced (as in fig. c). For further explanation see the text. out (fig. 2c, A), but this only succeeded near the inflorescence. Now the inflorescence may be explained as follows: The main axis (fig. 2a, b, c M)forms the central flower. The axillary axis (fig. 2a, c A; fig. 2b A) of the leaf (fig. 2a, c L; fig. 2b L, not visible, cf. arrow), is branched cymosely. From the dormant buds (fig. 2a, c B) branches are developed, which are also branched cymosely, and are only slightly poorer than the axillary axis. So the axillary axis (fig. 2b A) is supposed to represent the first and single racemous ramification of a thyrsus. KEY TO THE SPECIES 1. Flowering axes sharply triangular in cross-section. Branches with 2 lateral, basal buds (scars!). Stamens 20-40 1. T. triangulare 1. Flowering axes terete. Branches without basal buds. Stamens up to 15 2. T. paniculatum 1. Talinuin triangulare (Jacq.) Willd. Sp. PI. 2 (1799) 862; Heyne, Nutt. PI. (1927) 612; Poelln. in Fedde, Rep. 35 (1935) 15; W. H. Brown, Useful Pl. Philip. (1950) 520; Backer & Bakh./. Fl. Java 1 (1963) 217. — Portulaca triangularis Jacq. Enum. PI. Carib. (1760) 22. — Portulaca racemosa L. Mant. (1771) 242. — T. racemosum (L.) RoHRB. in Mart. Fl. Bras. 14, 2 (1872) 297; Den Berger, Trop. Natuur 7 (1918) 28, t. 1-3. — Fig. lb, 2. Erect semi-shrub up to c. 1 m. Leaves elliptic to obovate, up to 15 by 5 cm, acute to acuminate; nervation pinnate. Axillary buds with 2 subulate, small cataphylls; the latter always with a dormant axillary bud up to 1 '/2 cm in the side-axes (caducous in dried specimens leaving a scar). Thyrsi terminal, corymboid, up to r. 12 cm 0, the axes sharply triangular with 8-c. 28 flowers. Bracts and brac- teoles subulate. Sepals deltoid, c. 4.6 by 31/2 mm. acuminate. Petals 5, obovate, up to 10 by 4 mm, emarginate, pink. Stamens 20-40; filaments up to 5 mm; anthers c. 0.7 by V2 mm. Style c. 2.7 mm. 2-3-fid. Fruit up to 5 mm 0, 2-3-valved, yellow. Seeds'r-o, c. 1.2 mmo; testa cells radially elongated, smooth, tubercled at the edge. Distr. Pantropic weed, still extending its range, native of tropical America. According to Dr. Leeuwenberg it extends enormously along new- made ways in Africa. Ecol. Waysides, waste places, edges of forests. Uses. A commonly used vegetable, easily propagated by cuttings, profusely used in war prison camp gardens, in Java imported from Surinam {cf. Heyne, Nutt. PI. 1927, 612). Vern. Surinam purslane. Malesia: krokot blanda, poslen, S. 2. Talinum paniculatum (Jacq.) Gaert. Fruct. 2 (1791) 219, t. 128; Heyne, Nutt. PI. (1927) 612; Poelln. in Fedde, Rep. 35 (1935) 10; A. C. Smith, Bull. Torr. Bot. Club 70 (1943) 537; Greenwood, J. Arn. Arb. 30 (1949) 75; Backer &. Bakh./. Fl. Java 1 (1963) 217. — Portulaca paniculatum Jacq. Enum. PI. Carib. (1760) 22. — Portulaca 1971 PoRTULACACEAE (Gecsink) 125 patens L. Mant. (1771) 242. — T. patens (L.) WiLLD. Sp. PI. 2 (1799) 863; Hassk. Hort. Bog. (1858) 74; Drake del Cast. 111. Fl. Ins. Mar. Pac. (1890) 111; F. M. Bailey, Queensl. Fl. 1 (1899) 95; Backer, Ann. Jard. Bot. Btzg Suppl. 3 (1910) 416; Schooifl. (1911) 84; Craib, Fl. Siam. Enum. 1 (1925) 110; Merr. & Chun, Sunyatsenia 1 (1934) 56; F. B. H. Brown, Bern. P. Bish. Mus. Bull. 130 (1935) 77. —Fig. la. Erect semi-shrub up to c. 70 cm. Leaves elliptic to obovate, up to 1 1 by 5 cm, acute to acuminate; nervation pinnate. A.xillary buds with 2 subulate cataphylls. Terminal thyrsi up to c. 18 by 15 cm, with up to r. 10 dichasia, each with up to 30 flowers. Bracts and bracteoles subulate. Sepals suborbicular, c. 1.1 mm, acute. Petals (4-)5(-6), obovate, c. 4 by 2 mm, pink, apex emarginate. Stamens (4-)15; filament c. 2.7 mm; anthers c. 0.4 mm. Style c. 1.6 mm, 3-fid. Fruit c. 3 mm 0, yellow or pink, 3-valved. Seeds c. 1.2 mm o; testa cells radially elongated, shortly tubercled or not, with small pits between the cells. Distr. Pantropic weed, native of tropical America, still extending its range. Ecol. Waysides, waste places, edges of forests. Uses. Cultivated as an ornamental, and with edible leaves. According to Heyne (Nutt. PI. 1927, 611) a decoction of the roots is used by Chinese as a surrogate aphrodisiac. Notes. 1. My cultivated specimens had pink sepals, petals, filaments, and style. According to field-labels, there are in America also forms with white and yellow petals. 2. I received at the Leyden Hortus seed of ob- viously two strains (pure lines?) recognizable by the seed-coat; the one has small tubercles on the cells of the testa, the other has no such tubercles. Specimens raised from these two seed types in the glass-house of the Leyden Hortus on vegetable mould showed difterent characters, as tabulated below: Seed with tubercles : Leaves up to 7 by 4 cm. Panicle c. 8 cm wide. Dichasia with up to 15 flowers. Fruit reddish. Seed without tubercles: Leaves up to 1 1 by 5 cm. Panicle c. 15 cm wide. Dichasia with up to 30 flowers. Fruit yellowish to olivegreen. From the herbarium it appears that these 2 seed types occur almost throughout the range of the species. The correlating characters are difficult to observe in the herbarium, and besides are liable to be influenced by the ecology of the locality (leaves and panicle). There are no transitions and obviously these two strains keep constant as a sort of pure lines. 3. MONTIA LiNNF, Sp. PI. (1753) 87; Swanson, Brittonia 18 (1966) 229. — Fig. Ic, 3. Copiously branched, succulent herbs or waterplants. Basal rosette mostly absent. Leaves ovate to linear, occasionally parallel-veined, without axillary hairs. Fig. 3. Muntia funtana L. a. Flowering branch, xl'/2, h. ditto, y 5, c. flower, X 10, d. sepals, xlO, e. corolla, x 10,/. stamens, x 10, k- young fruit with style, ■ 10, h. burst fruit with 1 seed, x20 {a-h Bra.ss & Meijer Dreps 9972, L). 126 Flora Malesiana [ser. I, vol. 7^ Flowers in axillary and/or terminal cymes or solitary. Calyx persistent. Petals 5. Stamens 3 or 5; anthers 4-celled. Ovary superior; style arms 3. Fruit pyriform to globular, 3-valved, after dehiscence occasionally twisted. Seeds 1-5. Distr. About 50 spp. Temperate and warm-temperate in Europe, Northern Asia, N. & S. America, Central Africa, and in Tasmania and New Zealand (the present species only!); the sole tropical localities in the Old World are on Mt Kilimanjaro in Central Africa and the high mountains of New Guinea. Generic delimitation, e.g. against Claytonia, is not very satisfactory. 1. Montia fontana Linne, Sp. PI. (1753) 87. — 0; the 3 valves twisted after dehiscence. Seeds M. lamprosperma Cham. Linnaea 6 (1831) 565; 2-3, c. 1.2 mm 0; testa cells radially elongated. Merr. & Perry, J. Arn. Arb. 23 (1942) 386; Distr. Northern hemisphere in temperate lo- Geesink, Pac. PI. Areas map IS4 (inedit.). — Fig. calities, Americas (Rocky Mts, Andes), Central Ic, 3. Africa (Mt Kilimanjaro), SE. Australia (Alps and Aquatic herb, in patches up to c. 6 cm high. Mt Lofty Ranges, south of 36"" SL), Tasmania. Leaves opposite, elliptic, up to 4 by 2 mm, not and New Zealand; in Malesia: West New Guinea caducous but decaying. Flowers solitary or 2-3 (Mt Wilhelmina) and East New Guinea (between together with a membranaceous bract, inserted Mt Dickson and Kuputivava). This peculiar dis- in the axil of a leaf. Sepals suborbicular, c. 1.2 tribution in Australasia reminds of that of mm 0, acute to mucronate. Petals 5, obovate to Hydrocotylevulgarish.{cf.Y\.Ma\.\,A, 1949, 116). spathulate, 2 larger, up to 1.6 by 1 mm, 3 smaller, Ecol. Stream banks, 3500-3650 m. up to 1.6 by 0.6 mm, the latter each with an Note. A variable species of which several in- epipetalous stamen. Filaments up to c. 1 mm; fraspecific taxa are distinguished, which to some anthers c. 0.34 by 0.17 mm. Style arms 3, sub- have the status of species, sessile, c. 0.1 mm long. Fruit globular, c. 1.2 mm 4. PORTULACA LiNNE, Sp. PI. (1753) 445; Pax & Hoffm. in E. & P. Nat. Pfl. Fam. ed. 2, 16a (1934) 246; Poelln. in Fedde, Rep. 37 (1934) 240; Legrand, Com. Bot. Mus. Hist. Nat. Montevideo 31, 1 (1953) 1; ibid. 34, 1 (1958) 1; Geesink, Blumea 17 (1969) 275. — Fig. 4-6. Mostly succulent, copiously branched herbs. Leaves linear to orbicular, in most species with axillary hairs. Flowers in (l-)2-30-flowered, terminal capituli; re- ceptacle infundibular, mostly with hairs or scales in the axils of the bracts (and bracteoles), surrounded by a whorl of c. 3-30 involucral leaves (or in some Austra- lian 5/7/?. in axillary and/or terminal dichasia). Sepals occasionally keeled or hooded, persistent or caducous with the petals, stamens and style. Petals 4-6(-8), mostly obovate, occasionally emarginate or mucronate. Stamens 4-c/5, in 1 whorl. Ovary half-inferior, occasionally apparently inferior; style with 2-18 arms. Capsule with a caducous circumscissile operculum. Seeds c/5 (in some Austrahan spp. \-^). Distr. Mainly tropical and subtropical all over the world, not more than c. 40 spp., possibly several very polymorphous, a few worldwide anthropochorous weeds. Ecol. Savannahs, shores, in pastures and ruderal places, along waysides, etc. Uses. P. oleracea is a well known vegetable with cultivars, P. quadrifida and other species are used as medicine (see Caius, J. Bomb. Nat. Hist. Soc. 41, 1939, 369), P. pilosa sens. lat. cultivated for ornamental purpose. Morph. In my opinion (Blumea 17, 1969, 277) the capitulum is a condensed cyme. It is surrounded by 3-18 involucral leaves, not to be confused with the "Involukralblatter" in the sense of Von Poellnitz. The nerved scales between the flowers are considered bracts and bracteoles and have, like the vegetative leaves, axillary hairs. In P. quadrifida the small white tubercles at the base of a tuft of hairs are supposed to represent the bracts and bracteoles. The axillary hairs are mostly homologized with stipulae, but in my opinion (I.e.) they are probably not of stipular nature. They are mostly found in 2 tufts in the leaf-axils. In Malesian spp. they are con- fluent at the base. In P. quadrifida they are present in a whorl around each node. Taxon. In my precursor (I.e.) I have modified and simplified the infrageneric subdivision of the genus. Of the two subgenera, one is confined to Australia (subg. Portulacella). 1971] PoRTULACACEAE (Gcesink) 127 As to the species concept, the one adopted here is distinctly larger than currently accepted, including that used by Von Poellnitz and Legrand. As I have shown, and experimentally checked in all four genera, selfing in bud seems the rule in the family, which explains that in nature pure lines are formed, and that populations may consist of several pure lines keeping distinct for the small characters in which they differ. For those who want to give a name and rank to any constant difference there is hence ample opportunity for distinguishing varieties, subspecies, and even species, especially in variable complexes such as those of P. oleracea and P. pilosa. In view of this genetical situation it seems that according to a normal specific concept there are far less good species than distinguished by Von Poellnitz and Le- GRANT* who recorded some 130 for the world and c. 62 (+21 varieties) for the Americas only respectively. KEY TO THE SPECIES 1. All leaves opposite. Hairs intra- and interpetiolar. 1. Sect. Neossia 1. P. quadrifida 1. At least the middle-cauline leaves spirally arranged. Hairs if present only axillary. 2. Sect. Portulaca. 2. The largest leaves obovate to spathulate. Axillary hairs inconspicuous. Sepals distinctly carinate (20 X magn.). 1. Subsect. Portulaca. 3. Mostly 1-3 involucral leaves with an axillary axis. Capituli (2-)3-30-flowered. Stamens 7-15. Fruit c. 4 mm long 2. P. oleracea 3. Involucral leaves mostly without axillary axes. Capituli (l-)2-3(-6)-flowered. Stamens 18-50. Fruit c. 7 mm long 3. P. lutca 2. The largest leaves linear to elliptic (obovate to spathulate in P. macrorhiza from Timor). Hairs mostly conspicuous. Sepals not carinate (20 x magn.), occasionally with an apical, dorsal, about dome-shaped hood. 2. Subsect. Stellulato-tuberculatae. 4. Largest leaves obovate to spathulate, c. 8 mm wide. Timor 4. P. macrorhiza 4. Largest leaves linear to elliptic, less than 4 mm wide. Pantropic 5. P. pilosa 1. Subgenus Portulaca Geesink, Blumea 17 (1969) 288. — Subg. Euportulaca Spegazzini, Ann. Soc. Ci. Argent. 82 (1917) 17; Poelln. in Fedde, Rep. 37 (1934) 242. Leaves opposite or spirally arranged. Flowers sessile, solitary or in glomeruli. Note. The single other subgenus Portulacella is exclusively Australian, and has peduncled flowers in cymes. 1. Section Neossia Legrand, Com. Bot. Mus. Hist. Nat. Montevideo 34, 1 (1958) 3; Geesink, Blumea 17 (1969) 289. — Subsect. Tuberculatae Poelln. in Fedde, Rep. 37 (1934) 243. — Subg. Enantiophylla Legrand, Com. Bot. Mus. Hist. Nat. Montevideo 31, 1 (1953) 5. Leaves opposite. Hairs or scales intra- and interpetiolar. Bracts and bracteoles reduced to small, white tuberculi at the base of a tuft of hairs or scales (in the capitulum). Distr. About 7 spp., pantropic, 5 in Africa, I in South India and Ceylon. 1. Portulaca quadrifida L. Mant. PI. I (1767)73; PI. (1927) 613; Backer, Onkr. Suikcr. (1930) DC. Prod. 3 (1828) 354; Decni:. Herb. Timor. 242, Atlas t. 255; Poelln. in f-eddc. Rep. 37 Dcscr. (1835) 120; Hassk. PI. Jav. Rar. (1848) (I9.M)275; Backer &Bakii./. II. Java I (1963) 437; Mio. PI. Jungh. (1855) 396; Fl. Ind. Bat. I. I 216; Geesink, Blumea 17 (1969) 290. — P. (1858) 1061; Dyer, FI. Br. Ind. I (1874) 246; mcriJiumi L. /: Suppl. (1781) 248; Bl. Bijdr. King, J. As. .Soc. Bcng. 59. ii (1890) 145; Ba(khr, (1826) 1136; Blanco, FI. Filip. cd. 3. 2 (1878) FI. Bat. I (1907) 79; Voorl. (1908) 18; Sthoolll. 162. /'. quadrilida var. rncrii/iiinii DC. Prod. 3 (191!) 84; Merr. FI. Manila (1912) 200; Int. (1828) 354; Decne, Herb. Timor. Dcscr. (1835) Rumph. (1917) 217; Gihbs, Arfak (1917) 211; 120; Boeri. in Vcth. Midd. Sumatra 4 (1884) 17. Merr. Sp. Blanc. (1918) 142; Ridi . FI. Mai. Pen. -- Fig. 4. 1 (1922) 151; Merr. Fn. Philip. 2 (1923) 136; Creeping herb, rooting at the nodes; nodes RiDL. FI. Mai. Pen. 5 (1925) 289; Heyne. Null. with a whorl of hairs. Leaves elliptic to cordate, 128 Flora Malesiana [ser. I, vol. V ^ VUSMA '6'J Fig. 4. Portulaca quadrifida L. a. Habit of part of a plant, x 1 '/^, b. flowering end of a branch, x 5, c. flower, x5, cf. corolla, x7, e. sepals, x7, /. stamens, x7, g. young fruit, operculum partly removed, x7, h. node with a whorl of hairs, x5 (a-h Weber sm.). 2-20 by 0.8-7 mm with c. 5 mm long axillary hairs. Flowers terminal, l(-3) on an infundibuliform, profusely hairy stem-apex, at edge with 4, seldom more leaves. Sepals c. 3 mm long. Petals 4, obo- vate, up to 5 by 4 mm, yellow. Stamens 8 or 12; filaments up to 3'/2 nim; anthers c. 0.30 by 0.30 mm. Style up to c. 4 mm with (3-)4(-5) arms. Fruit ± obovate, up to c. 2-3 '/^ by 3 mm; oper- culum c. 2/3 the height, shining, straw-yellow. Seeds CO, 0.8-1 mm 0, dull; testa cells elliptic, radially arranged, the surface either convex or with a pyramidal tubercle. Distr. Pantropic, except Australia and the Pacific east of Samoa, throughout Malesia. E c o 1 . Waysides, deforested land, ruderal places, often between gravel or in sand, not on the sandy beach, below c. 300 m. Fl. fr. Jan.-Dec. Uses. A native vegetable and used as a medi- cine. See Heyne, Nutt. PI. (1927) 613; Caius, J. Bomb. Nat. Hist. Soc. 41 (1939) 369; Quisum- BiNG, Med. PI. Philip. (1951) 284; Burkill, Diet, ed. 2, 2 (1966) 1833. Vern. Rumput segan, Mai. Pen.; gelang pasir, kremi, J; Philip.: maraiigalok. Ilk., lb., sayikan. Tag.; djalu djalu bubudo, Ternate. Note. Several authors have determined speci- mens of P. pilosa as P. quadrifida, especially in New Caledonia and in the rest of the Pacific. 2. Section Portulaca 1 . Subsection Portulaca Geesink, Blumea 17 (1969) 291. — Sect. Carinatae Poelln. in Fedde, Rep. 37 1971 PoRTULACACEAE (Gecsink) 129 (1934) 242. — Sect. Portulaphiton Legrand, Com. Bot. Mus. Hist. Nat. Monte- video 31, 1 (1953) 6. At least the middle-cauline leaves spirally arranged. Hairs, if present, only axillary. Bracts and bracteoles membranous. 2. Portulaca oleracea Linne, Sp. PI. (1753) 445; Bl. Bijdr. (1826) 1136; DC. Prod. 3 (1828) 353; MiQ. PI. Jungh. (1855) 396; Fi. Ind. Bat. 1, 1 (1858) 1061; Sum. (I860) 150; Benth. & F. v. M. Fl. Austr. 1 (1863) 169; Dyer, FI. Br. Ind. 1 (1874) 247; Blanco, FI. Filip. ed. 3, 2 (1878) 162 t. 164; King, J. As. Soc. Beng. 59, ii (1890) 144 Backer, FI. Bat. 1 (1907) 80: Voorl. (1908) 18 Schoolfl. (1911) 84; Merr. FI. Manila (1912) 200 Int. Rumph. (1917) 217; Sp. Blanc. (1918) 142 RiDL. FI. Mai. Pen. 1 (1922) 151; Merr. En Philip. 2 (1923) 136; Backer & Sloot. Theeonkr (1924) 110, t. 110; Heyne, Nutt. PI. (1927) 612 Backer, Onkr. Suiker. (1930) 243, Atlas t. 254 PoELLN. in Fedde, Rep. 37 (1934) 258; Backer & Bakh. /. Fl. Java 1 (1963) 216; Geesink, Blumea 17 (1969) 292. — P. diptera Zipp. ex Span. Linnaea 15 (1841) 207, num. nud. Herb, up to c. 40 cm. Leaves spirally arranged to subopposite, obovate to spathulate, up to 40 by 20 mm, with inconspicuous, up to c. 1 mm long a.\illary hairs. Capituli 2-30-flowered. Mostly l-2(-3) of the 2-8 involucral leaves with an axillary axis. Floners surrounded by up to c. 5 by 6 mm long bracteoles and inconspicuous hairs. Sepals up to c. 6 by 6 mm, carina up to c. 3 by 2 mm. Petals (4-)5, broadly obovate, up to 7 by 6 mm, yellow. Stamens 7-10(-15); filaments up to 4 mm; anthers 0.2-0.5 by 0.2-0.4 mm. Style up to c. 5 mm with (4-)5 arms. Fruit ovate, c. 4 by 3 mm; operculum Vi-Vi, as high as the fruit, shining, straw-yellow. Seeds 'n:, '/^-1.2mm o, granulate; testa cells stellulate, with many fine tubercles. Distr. Pantropic, throughout Malesia. Ecol. Waysides, deforested and ruderal places, sandy shores, even on exposed rocks, up to 1800 m. Fl. fr. Jan. -Dec. Uses. Eaten as a vegetable and used as a med- icine. See Heyne, Nutt. PI. (1927) 612; Ochse & Baku. Ind. Groent. (1930) 615; Burkill, Diet. (1935) 1833; Caius, J. Bomb. Nat. Hist. Soc. 41 (1939) 369; W. H. Brown, Useful PI. Philip. I (1950)520. Vern. Purslane, E, postelein, parse lein, D; gelang, S, M, J, gelang pasir, sesegan, M, krokot, J, re-sirijan. Mad., silobar pinggan, M (Sum.), segan jantan, rumput beremi, M (Mai. Pen.); Philip.: aluslman, ausiman, galusiinan, Bik., olasiman, sahikan, u list man, Bik., Tag., golasiman, kolasiman, makabling. Tag., bakbakad, lungum. If., dubdupil, Bon., iigalug. Ilk., kantatdba, Pan- gasinan; djalu djalu kiki, Ternate. Notes. Several authors distinguished the sub- species (or varieties) oleracea (syn. sylvatica) and saliva, the latter is the cultivated form. The dif- ference is probably caused by a diflerent chromo- some number (according to a letter of G. Tisch- LER to O. Degener) added to a Hawaiian specimen in the Bishop Museum: ''sativa'' would be a hexaploid with 2n = 54, "sylvatica^' a diploid with 2n = 18. In the herbarium these two taxa cannot sharply be separated: ""sativd" is generally tall, with large seeds, "sylvatica" is smaller, with smaller seeds, but they are connected by transitional specimens also. For an anatomical comparison of the seeds of these two forms, see Kowal, Monogr. Bot. 12 (1961) I, who also found the characters overlapping . 3. Portulaca lutea Forster [PI. Esc. (1786) 72, nomen] ex Seemann, FI. Vit. (1865) 9; Geesink, Blumea 17 (1969) 291. Herb, up to 70(?) cm. Leaves spirally arranged to subopposite, obovate to orbicular, up to c. 30 mm long, not translucent in the dried state, with up to 6 mm long axillary hairs. Capituli 1-6- flowered. Flowers surrounded by up to 4 by 2 mm long bracteoles and up to c. 2 mm long hairs. Sepals suborbicular up to c. 9 mm long, fleshy in centre, with an up to 4 by 1 '/2 mm long, dorsal, apical keel. Petals 5, broadly obovate to obovate, up to c. 10 mm, emarginate to mucronate, yellow. Stamens 18-f. 50; filaments up to c. 4 mm; anthers up to c. 0.7 by V2 mm, at least occasionally red. Style up to c. 5 mm with (4-)5 arms. Fruit ± ovate, f. 7 by 5 mm; operculum 73 as high as the fruit, shining, straw-yellow. Seeds '^o, up to c. 1 mm 0; testa cells stellulate, flat, convex, whether or not with tubercles or spines. Distr. Pacific Islands, from Samoa and New Caledonia to the Marquesas, not yet recorded from Malesia. 2. Suhsection Stellulato-tuherculatae PoF.i.f.N. in Fedde, Rep. 37 (1934) 242; Geesink, Blumea 17 (1969) 293. — Sect. Rotioulatae Poelln. in Fedde, Rep. 37 (1934) 242. — Sect. Pseudohip.soclasis Lecrand, Com. Bot. Mus. Hist. Nat. Montevideo 31, 1 (1953) 7. — Sect. Catocla- sis Lfgrand, Com. Bot. Mus. Hist. Nat. Montevideo 31, 1 (1953) 8; ihid. 34, 1 (1958) 10, e.xcl. subsect. Squamosae. 130 Flora Malesiana [ser. I, vol. 7^ £ VIJSMA '69 Fig. 5. Portulaca macrorhiza Geesink. a. Branch, nat. size, b. flowering end of a branch, x2, c. flower, X 5, d. sepals, x 5, e. corolla, the frontal petal halved, x 5,/. stamens, x 7, g. young fruit, the operculum partly removed, x7, h. node with axillary hairs, x5 {a-h Zippelius s.n., type, HLB 200, 850, L). 4. Portulaca macrorhiza [Zipp. ex Span. Linnaea 15 (1841) 207; MiQ. Fl. Ind. Bat. 1, 1 (1858) 1060; Pax & HoFFM. in E. & P. Nat. Pfl. Fam. ed. 2, 16c (1934) 247, nom. nud.\ Geesink, Blumea 17 (1969) 293. — Fig. 5. Herb, up to 10 cm. Leaves spirally arranged, obovate to spathulate, up to c. 30 by 8 mm, obtuse to truncate; axillary hairs up to c. 4 mm. Capituli 2-3-flowered. Flowers surrounded by up to c. 3 mm long hairs and c. 1 Vi by 1.7 mm long bracteoles. Sepah ^- 5.3 mm. Petals broadly obovate, c. 7 (by 5?) mm, yellow. Stamens c. 60; filaments up to c. 4 mm; anthers c. 0.6 by 0.4 mm. Style c. 4 rrun with 5 arms. Fruit globose, c. 3 mm 0; operculum about half as high as the fruit, shining, straw-yellow. Seeds oo, elliptic, c. 0.7 by V2 mm; testa cells elliptic, ± stellulate, shining. Distr. Malesia: Lesser Sunda Islands (W. and E. Timor). Ecol. On limestone karst, c. 350 m, in E. Timor (Baucau), only once noted. Fl. fr. Dec. (one record). 1971] PoRTULACACEAE (Gecsink) 131 Uses. As a medicine against gonorrhoea. Vern. Nati biti, Timor. 5. Portulaca pilosa Linne, Sp. PI. (1753) 445; Backer, Schoolfl. (1911) 84; Merr. FI. Manila (1912) 235; En. Philip. 2 (1923) 136; Ridl. F1. Mai. Pen. 5 (1925) 289; Heyne, Nutt. PI. (1927) 612; Poelln. in Fedde, Rep. 37 (1934) 261; Henderson, Mai. Wild FI. 4 (1949) 33; Backer & Bakh. /. FI. Java 1 (1963) 216; Geesink, Blumea 17 (1969) 294. Variable herb. Leaves spirally arranged, obovate to linear, 2-30 nun long, with axillary hairs. Capituli l-12-flo\vered. Flowers surrounded by bracteoles and hairs. Petals 4-6. Stamens (6-)10- 75. Style arms 4-8. Fruit ovate, globular or obo- vate; operculum V2 to I'a as high as the fruit. Seeds cv; testa cells elliptic to stellulate. KEY TO THE SUBSPECIES 1. Ripe seeds bluish, grey, or dull black. 2. Cells of the testa convex to pyramidal, without a nipple. Petals 21/2-I2 by 1.8-11 mm. 1. ssp. pilosa 2. Cells of the testa flat, but at seed-edge nippled. Petals larger, up to c. 25 mm 0. 2. ssp. grandiflora 1. Ripe seeds dark brown or shining black. 3. Cells of the testa stellate, in shape orbicular, sometimes nippled. Flowers orange to red. 3. ssp. sundaensis 1. ssp. pilosa. — Geesink, Blumea 17 (1969) 295. — P. tuberosa RoxB. FI. Ind. ed. Carey 2 (1832) 464; Dyer, FI. Br. Ind. 1 (1874) 246; Backer, Voorl. (1908) 19; Schoolfl. (1911) 85; Poelln. in Fedde, Rep. 37 (1934) 312; Backer & Bakh./. Fl.Java 1 {\9(>'i)l\ 6 {race tuberosa). — P.australis Endl. Atakta Bot. (1833) 7, t. 6; ? J. J. Smith, Teysmannia 10 (1899) 92 {race australls). — P. helianthemoides Zipp. ex Span. Linnaea 15 (1841) 207, nomen {race tuberosa). — P. cincta Fenzl, Nat. Tijd. N. I. 14 (1857) 162; Miq. FI. Ind. Bat. 1, 1 (1858) 1061; Boerl. Handl. 1 (1890) 85; Poelln. in Fedde, Rep. 37 (1934) 312 {race tuberosa). — P. filifolia F. v. M. Fragm. 1 (1859) 169 (race filifolia). — P. sclerocarpa [non Gray, Bot. U.S. Expl. Exp. 1 (1854) 141] Koord. Minah. (1898) 345; Koord.-Schum. Syst. Verz. 3 (1914) 40, record corrected to P. pilosa {race pilosa). — P. pachyrhiza [non Gagn. Bull. Soc. Bot. Fr. 56 (1909) 41] Merr. En. Philip. 2 (1923) 136 (race tuberosa). — P. samoensis Poelln. in Fedde, Rep. 33 (1933) 163; ibid. 37 (1934) 300 (race tuberosa). — P. javanensis Poelln. Rev. Sudamer. Bot. 7 (1943) 273, ind. var. grisea Poelln. {race tuberosa). Herb, 10-c. 30 cm. Leaves elliptic to linear, 4-28 by Vi-A mm; axillary hairs 1-18 mm long. Capituli (l-)2-10-flowered; more or less congested with adjacent quasi-axillary capituli. Flowers sur- rounded by 0.7-2 '/2 by 0.7-2.2 mm long bracteoles and 3-18 mm long hairs. Sepals 2-6 by 1-4 mm, occasionally inconspicuously hooded at apex. Petals 4-6, obovate, 21/2-12 by 1.8-11 mm, yellow or pink. Stamens (7-)20-30(-35?); filament 1-5 mm; anthers globose to elliptic, 0.35-0.7 mm. Style 2-8 mm, 3-7-fid. Fruit ± globose, c. 2-3 (-4) mm 0; operculum 1/2 to 2/3 as high as the fruit, shiny, straw-yellow to olive green. Seeds 0.4-0.7 mm 0, dull light to dark grey, or bluish; testa cells elliptic, elliptic with lobes or stellate; pyra- midal all over the seed, or not. Distr. Pantropical, throughout Malesia. Taxon. A subspecies consisting of several 'races', which are easy to distinguish as shown in the following table. Race Range Petals Stamens Length anthers Seed Colour seed Shape testa cells Surface testa cells pilosa pantrop., excl. Austr. pink (7-)20-30 0.5 0.4-0.6 blue stellate pyramidal tuberosa E. Asia, Malesia, W. Pacific yellow (17-)25-30 0.7 0.6-0.7 dark grey elliptic, lobed, stellate at seed edge convex filifolia Australia, Java: in Hort. Bogor yellow 12-30 0.4 0.6-0.7 bright grey stellate pyramidal P. ausiralis Endl. is entered in the synonymy, but it remains uncertain whether this race was indeed represented by the material J. J. Smith identified from the island Cjroot Kombuis (Bay of Djakarta), as no specimen is available. The Australian race fili)olia is only represented by specimens collected in the Botanic Gardens at Bogor. Uses. A native vegetable and used as a medi- cine. Sec Of USE & Baku. Ind. Groent. (1931 ) 615, andCAius. J. Bomb. Nat. Hist. Soc. 41 (1939)369. Vern. Rose- flowered purslane, E, pt'nawar, M (Mai. Pen.), rvhlia ledkah. Mad.; Philip.: romrwA.M, Ilk.; njali' njali' wolanda, lernatc. 2. ssp. Krandiflora (HuoK.) Geesink, Blumea 17 (1969) 297. - - P. Krandiflora Hook. Bot. Mag. n.s. 3 (1829) t. 2885; Backlk, Voorl. (1908) 19; 132 Flora Malesiana [ser. I, vol. 7^ Fig. 6. Portulaca pilosa L. ssp. sundaensis (Poelln.) Geesink. a. Habit, X 1 V2. ^- flowering end of a branch X 5, c. flower, x 5, d. sepals, x 7, e. corolla, x 7,/. stamens, x 7, g. young fruit, x 7, h. node with axillary hairs, x 5 \a-h Jaag 799, L). Schoolfl. (1911) 85; Bruggeman, Ind. Tuinb. (1938) 84, t. 55; Steen. F1. Sch. Indon. (1949) 176; Backer & Bakh. /. Fl. Java 1 (1963) 216. Herb, up to c. 30 cm? Leaves linear, up to 25 by 3 mm, with c. 5 mm long axillary hairs. Flowers in each capitulum up to c. 5, flowering succedane- ously, each surrounded by up to 10 mm long hairs and deltoid, up to 4.7 by 2 mm long bracteoles; transitions between leaves and bracteoles present. Sepals c. 8 mm long, with a very small apical keel. Petals d= obovate, c. 25 mm 0, pink, red, orange or yellow (cultivated in many colours). Stamens c, 40-75; filaments up to c. 6 mm; anthers elliptic, c. 1.4 by 0.3 mm, 2- and 4-celled. Style up to c. 13 mm, with 5-18 arms. Fruit ± globose, c. 5 mm 0; operculum % to Vi as high as the fruit, shining, straw-yellow. Seeds c. 0.7 mm 0, shining; testa cells ± stellulate, those at the edge of the seed with a central tubercle. Distr. Native of tropical America, cultivated as an ornamental and occasionally escaped elsewhere. 3. ssp. sundaensis (Poelln.) Geesink, Blumea 17 (1969) 298. — P. sundaensis Poelln. in Fedde, Rep. 50 (1941) 105. —Fig. 6. Herb, up to c. 10 cm. Leaves oblong to obovate, up to 7 by 3 mm, with up to 6 mm long hairs. Capituli 2-3-flowered. Flowers surrounded by up to 0.7 by 1 mm long bracteoles and c. 6 mm long hairs. Sepals 5, c. 2.6 by 2.2 mm. Petals obovate, subacute, c. 2.3 by 1.7 mm, orange. Stamens (6-)10-15; filaments c. 1 1/2 mm; anthers c. 0.4 by 0.35 mm. Style c. 1.6 mm, with 5 arms. Fruit ± globose, c. lYzmm 0; operculum ^/^ as high as 1971] PoRTULACACEAE (Gecsink) 133 the fruit, dull, grey-brown. Seeds c. 0.55 mm 0, ± shining; testa cells stellulate. Distr. Malesia: Lesser Sunda Islands (Sum- bawa, Flores, Wetar, Alor), from each island one specimen. Ecol. Close to the coasts, along roads, between rocks. Vern. Tamdsiang dta, Alor. Excluded Portulaca axilliftora (non Pers.) Blanco, F1. Filip. ed. 2, 2 (1845) 285 and P. toston Blanco, Fl. Filip. (1837) 408 are both, according to Mer- rill, En. Philip. 2 (1923) 136, Trianthema portu- lacastrum L. (Aizoac). Portulaca teretifolia L.; F.-Vill. Nov. App. (1880) 15, an American species, probably an in- correct identification. BYBLIDACEAE (C. G. G. J. van Steenis, Leyden) In the former century Byblis was mostly included in the Droseraceae, for example by Bentham & Hooker./. (Gen. PI. 1, 1859, 220); even Engler had it in that position in 1912 (Syllabus ed. 7, 329). Planchon had in 1848 (Ann. Sc. Nat. Ill, 9, 1848, 80, 90) already pointed to affinity with Cheiranthera of the Pitto- sporaceae; Hallier/. merged Byblis and Roridula with Tremandraceae, curiously referring this to an Ochnaceous assemblage (Abh. Gebiete Naturw. Hamburg 18, 1903, 53). About the same time Lang argued (Flora 88, 1901, 179) that on mor- phological and anatomical grounds Byblis cannot belong to Droseraceae, but should be referred to Lentibulariaceae. DiELS (Pfl. R. Heft 26, 1907, 51) and Domin (Act. Bot. Bohem. 1, 1922, 1) definitely concluded to the alliance with Pittosporaceae, and so did Hutchinson (1926, 1959) and Schultze-Menz (Syllabus 1964): resemblance with Drosera is superficial, sympetaly unimportant. Hallier /. and Hutchinson include the S. African genus Roridula also in the family Byblidaceae, but others regard this as an allied family. BYBLIS Salisb. Parad. Lond. (1808) t. 95; Bth. F1. Austr. 2 (1864) 469; Lang, Flora 88 (1901) 179; Domin, Act. Bot. Bohem. 1 (1922) 1; Bibl. Bot. 22 (1929) 702; Diels in E. & P. Nat. Pfl. Fam. ed. 2, 18a (1930) 288; Vester, Bot. Arch. 41 (1940) 563, map 192. — Fig. 1. Erect herbs, viscid, by longitudinal rows of minute, sessile glands and capitate- glandular hairs. Leaves linear to filiform, involute-coiled in vernation, exstipulate, spirally arranged. Flowers axillary, solitary, without bracteoles, 5-merous. Sepals imbricate, persistent, short-connate at the base. Petals larger than sepals, contorted, with a broad, dentate apex and cuneate base, oblique, ± connate at the very base. Stamens 5, sometimes unequal; anthers basifixed, alternipetalous, cells opening with a very short pore-like slit. Disk none. Ovary superior, 2-celled, with CO ovules attached to the axis of the dissepiment about the middle; integument 1; style 1, simple, with a faintly 2-lobed stigma. Capsule ± compressed, 2-celled, loculicid with 2 valves, sometimes the valves later splitting, the dissepiment splitting i halfway. Seeds dark, rugose; embryo elongate, cylindric; cotyledons short, fleshy; albumen present. Distr. Two spp., one in SW. Australia, the other from NW. to NE. Australia, in Malesia: the N. Australian species in South New Guinea. EcoI. This is the fifth genus of insectivorous plants in Malesia, the others being Nepenthes, Utriculuria, Drosera and Aldrovandu. Both species grow in depressions which are swampy on poor soils or which become swampy or water-logged in the wet season. Often gregarious. The way of catching insects (small flies, mosquitoes, moths and ants) superficially resembles that in Drosera, but differs in that the capitate-glandular hairs make no movement towards the prey. Rica LRrfKSON (Austr. PI. 3, 1966, 319, 321) calls it a 'flypaper trap of the passive type'. According to Grieve iihid. I, n. 9, 1961, 23) "insects are usually first caught by the sticky mucilage exuded from the gland- tipped hairs and these tend to collapse or bend as they pour out secretion. The insect is thus also brought into contact with the numerous, minute sessile glands and becomes enveloped in additional secreted fluid. The process of secretion and absorption continues until all of the soft parts of the insect are dis- solved and absorbed, and only the hard, indigestible parts remain. The glands then stop secretion and the stalked ones commence to recover to their normal position. In due course the hard parts of the insect which arc left dry out and fall off." (135) 136 Flora Malesiana [ser. I, vol. 7^ Fig. l.Byblis liniflora Salisb. a. Habit, x 2/3, b. stem tip, x4, c. leaf tip, x\2,d. flower, corolla removed x4, e. anther, x 12,/-g. fruit, x4, h. seed, x8, /. flower, x2>,j. flower, corolla removed, x4, k. anther X 12, /. lengthwise section of ovary {a-h Hoogerwerf 273, /-/ Byrnes 230). It has been suggested that the capitate-glandular hairs secrete a sticky mucilage, but that the secretion of the sessile ones is less sticky and would serve mainly for digesting proteins, but I have no pertinent data to sustain this opinion. The large West Australian species, B. gigantea, is well-known as the 'rainbow plant', a name "believed to be derived from the fact that on looking through the plant towards the setting sun, one can see a spectrum of colours where the edges of the leaves are bordered by the shining drops of liquid on the glands." Anat. Fenner (Flora 93, 1904, 382-388) gave a detailed account of the anatomy of the glands of B. gigantea Lindl. 1971] Byblidaceae (van Steenis) 137 1. Byblis liniflora Salisb. Parad. Lond. (1808) t 95; DC. Prod. 1 (1824) 319; Endl. Iconogr (1841) t. 113; Bth. FI. Austr. 2 (1864) 470 F. V. M. Fragm. 10 (1876) 81; Bailey, Queensl Fl. (1900) 531; Compr. Cat. (1913) 174, f. 145 Britten, 111. PI. Banks 1 (1900) 30, t. 96; Ewart & Davies, Fl. North. Terr. (1917) 117; Domin, Act. Bot. Bohem. 1 (1922) 4; Bibl. Bot. 22 (1929) 703; Specht, Rec. Am. -Austr. E.xp. Amhem Land 3 (1958) 231; Steen. Blumea 16 (1968) 355. — Fig. 1. Unbranched, weak herb, c. 15-40 cm. Rhizome thin. Leaves filiform, very thin, blunt, c. 4-6 cm, spreading. Pedicels already from the base of the plant, usually exceeding the leaves, in fruit up to f. 10 cm, patent, the lower ones reflexed. Pistil and stamens somewhat zygomorphic. Sepals ovate-lanceolate, acute, erect, 3-4 mm, with scarious margin, glabrous, c. 3-4 by 1 Vi mm. Petals oblanceolate, acute, with dentate upper margin, 4-8(-ll)mm. Stamens glabrous, anthers varying from 1 ■/2-3 mm, filaments longest where anthers are shortest. Ovary glandular; style c. 3 mm. Capsule transversely elliptic, with 2 shallow grooves, c. 2 by 4 mm. Seeds ellipsoid, at one end ± pointed, black, 1 mm, lengthwise ribbed, ladder-like tessellate between the ribs. Distr. Queensland to NW. Australia, in Ma- lesia: South New Guinea (near Merauke: Hoo- gerwerf 273), one collection. Ecol. Marsh herb of shallow, seasonal swamps in Melaleuca- Acacia-Tristania- Eucalyptus savannah, at very low altitude. Fr. March. CLETHRACEAE (H. Sleumer, Leyden) 1. CLETHRA LiNNE, Gen. PI. ed. 5 (1754) 188; Sp. PI. (1753) 396; Sleum. Bot. Jahrb. 87 (1967) 55. — Fig. 1-5. Trees or shrubs, evergreen (Mai. spp.)\ leaf-scars large. Leaves crowded towards the end of the shoots, spiral, simple, exstipulate, serrate with glandular teeth, often with an apical gland, more rarely entire; nerves a little decurrent along the midrib, both midrib and nerves + impressed above, dz prominent beneath. In- dumentum of branchlets, leaves and inflorescences consisting of simple, and/or long, fascicled and ± patent, and/or minor, ± depressed stellate hairs. Flowers bisexual, regular, 5(-6)-merous. Inflorescences sometimes simple solitary terminal racemes, but mostly consisting of a terminal raceme and several lower approx- imate racemes, each of the latter from the axil of a ± reduced or caducous leaf, thus forming together a panicle-, fascicle- or umbel-like inflorescence; bracts mostly caducous during anthesis. rarely subpersistent. Calyx lobes 5(-6), persistent, quincuncially imbricate, united at the base only. Petals 5(-6), generally free, some- times cohering to some degree, alternate with the calyx lobes, rather early caducous, generally sweet-scented. Stamens 10(-12) in 2 whorls of 5(-6), the outer whorl opposite the petals, the inner one opposite the calyx lobes; filaments adnate to the corolla at the extreme base; anthers dorsifixed, overturned outwards in bud, erect in anthesis, introrse, upper part of cells ± divergent, opening with apical, slit- like pores; pollen grains single, tricolporate, psilate. Ovary superior, 3-celled, with axile placentation; ovules v:, small, anatropous; style simple, mostly shortly, very rarely hardly divided into three apical lobes, sometimes more deeply so and trifid, each lobe stigmatic at the top. Fruit a 3-valved, loculicidal capsule, the septae of which become loose from the persistent central axis, subtended or i en- closed at maturity by the persistent calyx. Seeds oo, small, subovoid to irregularly angular or subtrigonous, with a foveolate-reticulate testa (all Mai. spp.). Endo- sperm fleshy. Embryo cylindrical. Distribution. A small, monogencric family in the Ericales, of (sub)tropical Asiatic-Malesian, and temperate and tropical American distribution, and with ! sp. in Macaronesia (Madeira, and formerly in Teneriffe). Of the total of 64 spp., 2 temperate spp. are found in North America (C. alnifolia L. and its var. pu- bescens Ait., and C acuminata Michx), iS spp. inhabit (sub)tropical America (Cuba-Jamaica-Central America, in S. America in the Mt Roraima area and in Trinidad, in the Andes from Venezuela to SE. Bolivia and NW. Argentina, and a small separate area in SE. Brazil), \0 spp. are found in SE. Asia (Upper Burma, SW., Central and SE. China, Japan, Indo-China). In Malesia 13 spp., of which 2 in the Malay Peninsula, I in Sumatra, I in Java, 3 in the Lesser Sunda Islands, 4 in the Philippines, 3 in Borneo, 2 in Celebes, I in the Moluccas, and 4 in New Guinea. Fig. 2. Ecology. Clefhra occurs mainly in the lower and upper montane primary (and also secondary) forest. As to climate, the genus prefers the wet montane tropical and subtropical zone, though it extends far into the northern temperate zone, in North America to about 45 ', in Japan to about 42 ' N, whilst its southern limit is about 29 S in SE. Brazil. As the majority of the Ericales, Clethras have preference for acid soils. They occur in Malesia from sea-level to about 3000 m; and up to c. 3800 m in Szechuan and Yunnan, and in the Andes. They are light-demanding, remain sterile in dense forest, and flower in forest borders with open vegetation or exclusively in the latter, also in light secondary plant communities. Dispersal. Nothing is known about the dispersal of Cielhras. The rather small, deeply imprcssed- rcticulate and light seeds of the Malcsian spp. point to dispersal by wind. Anatomy. Dpn Blkoer, Determinatictabel van Malesiii, Veenman, Wagcningcn (1941) 31 (hand (139) 140 Flora Malesiana [ser. I, vol. 7^ Fig. 1. Clethra pachyphylla Merr. a. Habit, X2/3, b. flower, x4, c. petal, x4, d. stamens in bud stage, x4, e. stamens at full anthesis, x4, /. longitudinal section of flower, x4, g. fruit, x4, /;. seeds, x8 (a-/ Chew, Corner & Stainton 1845, g~h Sinclair 9082). 1971] Clethraceae (Sleumer) 141 lens, wood); Den Berger & Bianchi, Tectona 24 (1931) 894-903 (wood); Hara, Bot. Mag. Tokyo 70 (1957) 108-114 and J. Fac. Sc. Un. Tokyo, Sect. 3, Bot. 7 (1958) 367-450 (shoot apex and leaf histo- genesis); Lemesle, Bull. Soc. Pharm. Bordeau.x 104 (1965) 37-48 (localization of tanniferous substances); Lemesle & Duply, C. R. Ac. Sc. Paris 263 (1966) 250-253 (tannins and muciiagenous substances); SuDO, Identification of Japanese Hardwoods, Bull. Govt. Exp. Stat. Meguro 118 (1959); Uphof, Hum- mel & Staesche, Plant Hairs, in Handb. Pflanzenanat. IV, 5 (1962). For genera! surveys of older lit- erature see SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899) 541-551 and ibid. (1908) 195-197 (under Ericaceae), and Metcalfe & Chalk, Anat. Dicot. 2, Oxford (1950) 836-839. The wood shows primitive characters such as narrow solitary vessels with scalariform vessel perfo- rations, fibre-tracheids, apotracheal parenchyma which is diffuse or arranged in short uniseriate bands and heterogeneous rays of two sizes. The petiole is supplied with a strongly incurved to closed vascular strand. Hairs occur as multicellular stellate and uniseriate types. Stomata mainly paracytic. Crystals soli- tary and clustered. Secretory tanniferous cells are present in the parenchymatous tissues of petiole and stem. Metcalfe & Chalk {I.e.) state that Clethra has many anatomical features in common with Ericaceae. — P. Baas. Pollination. The flowers are protandrous. The pollen is shed at least partly before the flowers are open and the stigmas are receptive. Galls. Docters van Leeuwen (Zoocecid. Neth. Ind. 1926, 439, fig. 825 & 826) mentions leaf-galls on C. stimatrana from the Petani ravine near Medan, caused by a Psyllid and a gall mite. On the leaves of C canescens in N. Celebes (Minahasa) globular galls are found. Uses. Temperate species of Clethra, mainly from North America, Madeira and Japan, much less from China, are cultivated as ornamentals in many parts of the world. Not a single species from Malesia so far has been introduced in Botanic Gardens. Phytochemistry. Leucoanthocyanins, caffeic acid, flavonols, taraxeron and ursolic acid are known to occur in species of Clethra (Hegnauer, Chemotax. d. Pfl. 3, 1964, 432-433; Tanabe c.s. J. Pharm. Soc. Japan 86. 1966, 441 ). This spectrum of phytoconstituents is compatible with the generally accepted relationship between Clethraceae and Ericaceae. — R. Hegnauer. Taxonomy. A monograph of the genus has been published by the author (Bot. Jahrb. 87, 1967, 36-175, pi. 1). In this work it was shown, that the North American and the Asiatic-Malesian species of Clethra belong to sect. Clethra by their subovoid to subtrigonous and impressedly reticulate seeds, whilst the Central and South American species and C. arborea in Madeira form a second section, Cuel- laria, distinguished by flat and variously winged seeds. For the determination of Clethras close observation of the indumentum with a lens is necessary. There are (i) simple, mostly rather long, appressed or patent hairs, (ii) fascicled, t obliquely erect, generally elongated hairs, (iii) stellate hairs of generally small size and ± flattish. On most Clethras a combination of 2 or all 3 types of hairs is found, but their density may difi"er greatly on the various organs, resulting in a fine tomentellous, a thicker tomentulose or tomentose to villous tomcntum of white to greyish, pale ferrugineous to dark rusty and even rufous colour. Besides the various kinds of hairs, the discrimination of species is based on floral characters. Collectors should try to collect flowering material and (sterile) branchlets with young, not yet glabrate leaves /ro/// the same specimen, and abstain from collecting sterile or fruiting material. KEY TO THE SPECIES 1. Filaments hairy. 2. Filaments hirsutulous. Style elongate in fruit (up to 12 mm) 1. C. sumatrana 2. Filaments laxly set with longish hairs. Style a little or hardly elongate in fruit (up to 7 mm). 3. Leaves subcoriaceous, practically glabrous in mature state, inflorescence dilutely rusty-tomcntcllous in all parts 2. C. hcndcrsonii 3. Leaves coriaceous, still densely stellate-hairy along midrib and primary nerves in mature state beneath. Inflorescence rusty-villous in all parts 3. C. syniingtonii 1. Filaments glabrous. 4. Younger, not yet glabrate leaves (which still show the full quality and quantity of the tomentum) laxly to more densely hairy only on midrib, nerves and veins, and glabrous on the intcrvcnium be- neath, mature ones glabrcscent, or leaves glabrous from the beginning. 5. Lateral nerves (l5-)l7-20(-23) pairs, * approximate to each other 4. C. javanica 5. Lateral nerves I0-I2(-I4, rarely up to 15) pairs, more distant from each other. 5. C canescens 4. Younger, not yet glabrate leaves tomentellous to tomentose all over the undersurface, mature ones but slightly or tardily glabrescent. 6. Younger leaves : apprcsscdiy tomentellous on the intervenium beneath, i.e. covered there ex- clusively with minute, : flattish stellate hairs. 7. Younger leaves densely covered with minute stellate hairs on the whole undersurface in a coherent layer, and besides more laxly set with stellate-fascicled and darker (generally rusty) hairs on nerves, veins and vein lets 6. C. kcbareasis 7. Younger leaves on the undersurface set with dense minute stellate hairs, which form a coherent pale layer, other larger and/or darker stellate-fascicled hairs very few, or mostly absent. 142 Flora Malesiana [ser. I, vol. 7^ 8. Leaves regularly serrate in the upper 2/3, with C15-)l7-20 pairs of lateral nerves. 7. C. sumbawaensis 8. Leaves subserrate to entire, with 10-14 (rarely up to 15) pairs of lateral nerves. 9. Petals 2-2. 2(-2. 5) mm 8. C. tomentella 9. Petals (3.5-)4(-5) mm 9. C. arfakana >. Younger leaves u: erectly tomentulose or tomentose on the intervenium beneath, i.e. covered there with minute and more coarse stellate (suberect), or with minute stellate and coarse (± erect) fascicled hairs. 10. Racemes generally more slender and elongate (flowers ± laxly arranged), and tomentellous by noinor stellate, rather flattish hairs 10. C. longispicata 10. Racemes generally more robust and shorter (flowers ~ densely arranged) and tomentulose to tomentose by a combination of minor stellate and robust, ± erect fascicled hairs. 1 1. Pedicels 1-2 mm at anthesis. Style 1-1.5 mm at anthesis, hardly 2 mm in fruit. Lateral nerves of leaves ± inarching before the margin H. C. papuana 1 1. Pedicels 2-3(-5) mm at anthesis. Style 2.5-3 mm at anthesis, 3-4 mm in fruit. Lateral nerves of leaves — excurrent along the margin. 12. Ovary tomentose by long erect hairs 12. C. pulgarensis 12. Ovary tomentulose by short hairs 13. C. pachyphylla 1. Clethra sumatrana J. J. S. Ic. Bog. 4, 1 (1910) t. 319; Merr. Philip. J. Sc. 14 (1919) 248; Doct. V. Leeuwen, Zoocecid. Neth. Ind. (1926) 439, f. 825 & 826; Metcalfe & Chalk, Anat. Dicot. 2 (1950) 838, f. 192, F-G (anat.); Sleum. Bot. Jahrb. 87 (1967) 77. — C. pulcherrima Ridl. J. Mai. Br. R. As. Soc. n. 87 (1923) 75; Merr. Contr. Arn. Arb. 8 (1934) 121. — C. longipedi- cellata Merr. Pap. Mich. Ac. Sc. 19 (1934) 179. — C. pubifolia Merr. I.e. 180; Burret, Notizbl. Berl.-Dahl. 15 (1940) 188, in text. Much-branched shrub, or tree with erect or crooked trunk and a dense crown, (3-)5-15 (rarely up to 25) m; bark whitish. Branchlets stoutish, younger parts covered with a rusty stel- late tomentum maybe mixed with simple hairs, early glabrescent. Leaves lanceolate to oblong- or obovate-elliptic, apex shortly acutely acuminate, base cuneate, thin-coriaceous, young ones covered with a thick tomentum of rusty to rufous shorter stellate and longer fascicled hairs at least on mid- rib and nerves, older ones mostly rather persistent- ly so by both or but one type of these hairs on the entire undersurface or only on midrib, nerves and maybe veins, generally distantly serrate with callose teeth especially in the upper part of the lamina, rarely subentire or entire, reddish when young, dark green at maturity, 4-11 (-15) by (1.5-)2-4.5(-8) cm, nerves in 8-13 pairs, curved- spreading, slightly raised beneath as are the trans- verse veins, reticulation not conspicuous in gen- eral; petiole 6-12(-15, rarely up to 30) mm. Racemes elongate, erect-ascendent, 8-30 cm, usu- ally solitary and terminal, more rarely also from the axil of 2 or 3 upper leaves, the latter mostly normal in size and rz persistent, rarely reduced and caducous during anthesis, covered by the same, but generally more dense and more per- sistent indumentum found in the branchlets and leaves. Pedicels rather slender, 3-10 mm (in fruit 10-20 mm), bracts lanceolate-linear, caducous, 8-15 by 1-2.5 mm. Calyx lobes subovate or ovate- oblong, subacute, rusty-tomentellous or -tomen- tose in the upper part inside and all over outside, the fascicled hairs few to numerous, (4-)5-6(-8) by 2-3 mm. Petals obovate, erose-crenulate at apex, glabrous, white-creamy to pinkish, (5-)6-7 (-9) by 4-5(-6) mm. Filaments linear, glabrous at both ends, longish and subappressedly hirsutu- lous between, 4-5 mm; anthers obsagittate, 1.5-2(-2.5) mm, base extended into an acute triangular appendix. Ovary subappressed-hirsute; style slender, glabrous or with a few hairs at the very base, 6-7 mm at anthesis, 8-12 mm in later stages, apex thickened and very shortly trilobed at anthesis, rarely more deeply so in fruit. Capsule subglobose, hirsutulous, 4—5 mm 0. Seeds sub- obovoid-oblong or subtrigonous, variously com- pressed, c. 1.5 mm. Distr. Malesia: Sumatra (Atjeh, Eastcoast, Tapanuli, in the Westcoast Res. only at Brani near Bukit Tinggi). Ecol. In primary and secondary forest, also in Pinus merkusii forest, (390-)900-3000(-3300?) m. Fl. fr. Jan. -Dec. Fig. 2. Distribution of Clethra in Malesia and adjacent SE. Asia. The number above the hyphen indicates the number of endemic species, that below the hyphen the number of non-endemic species in each island or district. 1971] Clethraceae (Sleumer) 143 Fig. 3. Cleihra hendersonii Sleum., Mt Beremb^n, 1720 m, Malay Peninsula, Cameron Highlands (Sleumer 4675) (Sleumer, 1963). Vern. Kumhawan^, Karo, hudi-hudi, holi-boli, darik-darik, hau si martadjaii, simar darih-darih, Toba. 2. C'lethra hendersonii Sleum. Bot. Jahrb. 87 (1967) 79. — C. canescens (nun Reinw. ex Bl.) Riul. J. Fed. Mai. St. Mus. 6 (1915) 158. — FlR. 3. Slender, small or medium-sized, few-branched tree; barit rather smooth, greyish, minutely longitudmally fissured. Branthlets rather slender, dilutcly rusty-slellate-lomfntellous distally. Leaves elliptic-oblong or elliptic, sometimes subovate- clliptic, apex shortly acuminate, base cuncate and slightly inequilateral, subcoriaceous, dark green and dull above, paler and shining below, (lush reddish, denticulate-serrulate, «-!()( -12) by 4-5.5(-6) cm, young ones stcllate-puberulous, early glabrate above, set with sparse simple hairs at midrib and moreover i laxly with small stellate hairs at midrib, nerves and veins below, mature ones finally glabrous, nerves (10 )ll 13 pairs, curved- ascendent, raised beneath, reticulation faint; petiole rather slender, 1.5-2.2 cm. Racemes 2 144 Flora Malesiana [ser. I, vol. 7^ or 3 (-5), subdensely set with flowers, up to 22 cm, tomentellous by rusty-brownish simple and paler stellate hairs in all parts. Pedicels rather robust and 3-4 mm (in fruit 6-7 mm), bracts subulate, 3-4 mm, caducous. Calyx lobes oblong, 4-5 by 2 mm. Petals obovate, apex erose, spreading, white, slightly fragrant, glabrous on both faces, sparsely fimbriate, (5-)6-7 by (3-)4-5 mm. Filaments laxly set with longish hairs, 5(-6) mm; anthers narrowly obcordate, base acute, 1.8 mm. Ovary subsericeous-tomentose; style glabrous, shortly trilobed, 5-6 mm in anthesis, 7 mm in fruit. Capsule depressed-globose, 4 by 5 mm, sub- tended by the spreading sepals, Seeds oblong- ovoid, c. 1.5 mm. Distr. Malesia: Malay Peninsula (Pahang: Cam- eron Highlands and MtTahan; Selangor: Mt Nuang). Ecol. Scattered in forest, often on ridges or summits, 1340-1740 m. Fl. April-July. 3. Clethra symingtonii Sleum. Bot. Jahrb. 87 (1967) 80. Shrub. Tips of branchlets and innovations short- ly rufous-villous. Leaves elliptic-oblong, apex shortly subacutely attenuate, base cuneate, very base sometimes obtuse, slightly inequilateral, co- riaceous, young ones early glabrate above, ± densely set with rather large stellate hairs on mid- rib, nerves and veins beneath as are the petioles, mature ones persistently so, 6-9 by (2.5-)3-4 cm, glandular-subserrate-denticulate in the upper, entire in the lower part, nerves in 9-10 pairs excurrent along the edge, well-raised beneath, veins transverse and prominent beneath, retic- ulation visible; petiole 2-3 cm. Racemes terminal and from the upper 3 or 4 axils (which are mostly defoliate in flowering time) close together forming a panicle, all over with a rusty, almost villous tomentum of numerous longer crisp fascicled and shorter stellate hairs; rachis robust. Pedicels thickish, 2-3 mm (in fruit 5-6 mm), bracts subu- late, caducous, 3 mm. Calyx lobes oblong, c. 5 by 2.5 mm. Petals obovate, apparently cup-like converging, quite glabrous, white, scented, 6 by 3 mm. Filaments laxly long-hairy; anthers narrow- obcordate, base acute, 2 mm. Ovary densely set with erect hairs; style subappressedly hairy in its lower third, glabrous for the rest, 5 mm (in fruit 6 mm), stigma shortly 3-lobed. Capsule sub- globose, c. 4 mm 0. Distr. Malesia: Malay Peninsula (Perak: G. Kerbau For. Res.), once found. Ecol. On open ridge top, 1830 m, not common. Fl.fr. July. 4. Clethra javanica Turcz. Bull. Soc. Nat. Moscou 36, ii (1863) 232; Koord. Album Na- tuurmonum. Ned. Ind. ser. 1 (1918) 3, pi. 9 (phot.); Bull. Jard. Bot. Btzg III, 1 (1919) 148, t. 10; Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 161, p. 1;Back. &BAKH./. Fl. Java 2 (1965) 178; Sleum. Bot. Jahrb. 87 (1967) 83. — C. canescens (non Reinw. ex Bl.) Miq. Fl. Ind. Bat. 2 (1859) 1056, p.p.; Koord. Jungh. Gedenkb. (1910) 184; Exk. Fl. Java 3 (1912) \, p.p.; Back. Bull. Jard. Bot. Btzg II, 12(1913) 11; J.J. S. in K. & V. Bijdr. 13 (1914) 85, as to descr. — C. sp. Zoll. Syst. Verz. 2 (1854) \3S,p.p. var. javanica. Small tree, 3-6(-10)m, trunk up to 15 cm 0, often crooked, or much branched shrub, 2-5 m; branches up to 5 cm a. Branchlets rather robust, covered distally with a rusty hirsute or almost lanate tomentum as are the petioles. Leaves lanceolate to oblong-elliptic, sometimes partly obovate in the same specimen, apex shortly acuminate, base cuneate, subcoriaceous, dark green, shining and glabrous above, much paler beneath, and set there still towards the maturity with both longish simple or fascicled rufous and fewer minor stellate hairs on midrib and nerves, whereas the veins bear only scattered stellate hairs and the intervenium is glabrous, finally glabres- cent, (5-)6-10(-13) by 2. 5-4(-6) cm, rather reg- ularly and deeply (1.5-2 mm) serrate, nerves (15-) 17-20 (-2 3) pairs, spreading, ± approx- imate and subparallel, rather straight from the midrib and partly forked before the edge, ± sharply prominent beneath, veins transverse, ± raised beneath too; petiole robust, 1(-1.5) cm. Racemes panicled or subumbellate, rachis and pedicels covered with a floccose and rusty, almost hirsutulous tomentum, 5-10(-14)cm, flowers rather densely arranged, subpatent. Pedicels rather thickish, 3-5 (-6) mm, bracts subulate, 2-4(-6) mm, caducous. Calyx lobes ovate-oblong, sub- acute, dorsally a little keeled and stellate-tomen- tellous, fimbriate, (4-)4.5-6 by 2.5 mm. Petals obovate, apex slightly erose, thin, glabrous, white or pinkish, (5-)6-7 by c. 3 mm. Filaments glabrous, 2.5 mm; anthers obcordate, base sub- acute, 1.3 mm. Ovary shortly rufous-hirsutulous; style glabrous, 3-3.5 mm, apex shortly trilobed. Capsule subglobose, 2.5-3 mm 0, pedicel elongate to 10 mm, style slender, 3.5-4 mm. Seeds ovoid, subtrigonous or variously compressed, 0.5-0.8 mm. Distr. Malesia: East Java (Mt Jang). Ecol. In mixed primary forest edges, 1830-2300 m. Fl. fr. April-Oct. var. lombokensis Sleum. Bot. Jahrb. 87 (1967) 84. — C. canescens (non Reinw. ex Bl.) Miq. Fl. Ind. Bat. (1859) 1056, p.p. — C. sp. Zoll. Syst. Verz. 2 (1854) 138, p.p. Leaves very similar to those of var. javanica, but thinner, young ones on the undersurface sparsely clothed with fascicled hairs at midrib and nerves, and with stellate ones at the veins, becoming almost completely glabrous at maturity, up to 1 5 by 5 cm, more deeply (up to 4 mm) and more irregularly serrate. Distr. Malesia: Lesser Sunda Islands (Lombok), in montane forest and Casuarina bush of Mt Rind- jani, 1 700-2400 m. Ecol. On dry soil of volcanic debris. Fl. June-Aug. 1971] Clethraceae (Sleumer) 145 5. Clethra canescens Reinw. ex Bl. Bijdr. (1826) 863; G. Don, Gen. Syst. 3 (1834) 842; Miq. F1. Ind. Bat. 2 (1859) 1056, p.p.; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 41; Koord. Minah. (1898) 515; Sarasin, Reisen Celebes 1 (1905) 29, in text.: J. J. S. Ic. Bog. 4 (1910) 61, t. 318; Koord. - ScHUM. Syst. Verz. 3 (1914) 100; Koord. Bull. Jard. Bot. Btzg III, 1 (1919) 148, in te.\t.\ ibid. Ill, 2(1920)255; von MALMin Fedde, Rep. 34 (1934) 284; Sleum. Bot. Jahrb. 87 (1967) 85. KEY TO THE VARIETIES 1. Racemes and pedicels i robust. Petals 3^ mm. 2. Leaves early glabrescent; nerves starting from the midrib at an ~ acute angle. 1. var. canescens 2. Leaves with more persistent minute rusty stellate hairs underneath; nerves starting from the midrib at — right angles. 2. var. luzonica 1. Racemes and pedicels slender or almost so. 3. Leaves underneath on midrib, nerves and veins more laxly to rather densely clothed with subpersistent rather large stellate hairs. Petals 3-4 mm 3. var. dementis 3. Leaves underneath on midrib and nerves, rarely on veins, laxly set with fascicled and/or minute stellate hairs, early glabrescent, or young ones already subglabrous. 4. Racemes generally elongate, rather dense- flowered. Petals 2-3 mm. 4. v'«r. novoguineensis 4. Racemes generally short, rather lax-flowered. Petals (3-)4 mm 5. var. ledermannii 1. var. canescens. Tree up to 20 m, bole up to 40 cm 0, covered with greyish bark; at higher altitudes a shrub, 2-5 m. Branchlets covered with a woolly-floccose rufescent, later greyish tomentum at the tips. Leaves elliptic or obovate-clliptic, apex shortly subacutcly acuminate, base cuneate, thin-coria- ceous, entire below, regularly serrate with callose teeth c. I mm long, younger ones glabrous above, i densely set with minute stellate i rusty hairs on midrib, nerves, veins and veinlets, and with sparse, longer simple or fascicled hairs on the midrib beneath, glabrous on the intcrvcnium beneath, mature ones glabrescent, generally (4-)5-8(-ll) by (2-)2.5-4cm, at higher altitudes constantly smaller, (2.5-)3-5(-6) by (l-)l.5-2 cm, nerves 10-12 (rarely up to 15) curved- ascendent pairs, which rather distinctly inarch before the edge, veins slightly prominent, veinlets rather inconspicuous beneath; petiole :; robust, 0.6-1. 2(1. 5) cm. Racemes few to several, forming an erect panicle, all over covered with a floccose- lanatc, • rusty or finally greyish tomentum, 6-l2(-i6) cm, the numerous flowers rather laxly set along the thickish rachis. Pedicels robust, n-)4-5 mm at anthesis, up to 6 mm in fruit, bract subulate, up to 6 mm, caducous. Calyx lohes ovate-oblong, rather abruptly acuminate, long-ciliatc, pale rusty-floccose dorsally, 3 4 mm. Petals obovatc, crcnulatc all along the margin. glabrous, white (or suffused with red initially), with a rather unpleasant smell, 3-4 by c. 2 mm. Filaments glabrous, c. 2.5 mm; anthers broad- obovate, base almost mucronate, 0.8 mm. Ovary villous; style glabrous, 3-3.5 mm (in fruit 3.5-4 mm), very shortly trilobed. Capsule subglobose, tomentulose, 2.5 mm 0. Seeds irregularly ovoid- angular, 0.8-1 mm. Distr. Malesia: Celebes, Lesser Sunda Islands (Flores); slightly diff"erent forms in the Moluccas (Buru, Ceram and Ambon). Ecol. A tree in mountain forest or forest edge, ascending into more open summit vegetation and shrub-like there, sometimes on volcanic tuff", rather common locally, (1160-)1400-2800(-3000) m. Fl. fr. Jan.-Dec. Vern. Kaju parang, M, madaiisip, Ts. (Mina- hasa), pitjisan djabong, Makassar. 2. var. luzonica (Merr.) Sleum. Bot. Jahrb. 87 (1967) 88. — C. luzonica Merr. Publ. Gov. Lab. Philip, n. 29 (1905) 38; Philip. J. Sc. 5 (1910) Bot. 371; En. Philip. 3 (1923)242. Leaves elliptic, more rarely oblong-elliptic, apex shortly subacutcly acuminate, sometimes more obtuse, base broadly cuneate to obtuse, sub- coriaceous, younger ones laxly set with minute stellate hairs above, densely clothed with rather small floccose stellate hairs on the prominent midrib and nerves, and more laxly so on the veins beneath, glabrous on the intervenium, moreover sparsely set with longer simple or fascicled hairs on midrib and nerves beneath, mature ones glabrescent first above, tardily beneath, subentire or minutely (rarely more deeply) serrate, (3.5-)4-7(-10) by (1.5-)2-4(-5) cm, nerves 10-12(-14) pairs, curved, rather ap- proximate, i parallel and anastomosing; petiole rather robust, 0.3-1 (-1.3) cm. Racemes panicled or almost umbellate, 3-6(-l 1 ) cm, covered with a tomentum of small dark-rusty stellate floccose hairs; flowers ± dense along the thickish rachis. Pedicels robust, l(-2) mm, slightly accrescent in fruit, bract subulate, caducous, 3-4 mm. Petals white orcream, irregularly erose at apex, 3-3.5 mm. Distr. Malesia: Philippines (Luzon: Mountain Prov.; Isabela Prov.). Ecol. In montane Pinus insularis, Quercus or Podocarpus forest, also in mossy forest or in shrubby summit vegetation, l220-2250(-2700?) m, rather common locally. Fl. fr. Jan. -Dec. Vern. Amog, kamueg, Ig., apiit, Bon. 3. var. dementis (Merr.) Sleum. Bot. Jahrb. 87 (1967) 86. — C. dementis Merr. Philip. J. Sc. 13 (1918) Bot. 104; En. Born. (1921) 460. - C. ca- nescens {nan Reinw. ex Bl. s. sir.) Stapf, Trans. Linn. Soc. Bot. 4 (1894) 198; Gibbs, J. Linn. Soc. Bot. 42 (1914) 105. Leaves oblong-lanceolate to oblong, apex acutely acuminate, base cuneate-attenuale, thinly subcoriaceous, younger ones first stellate-hairy, early glabrescent above, L densely set with fas- cicled and stellate, rather coarse hairs at midrib and nerves beneath, laxly or hardly so at the veins, 146 Flora Malesiana [ser. I, vol. 7* intervenium glabrous, mature ones quite gla- brous and a little shining above, glabrescent first at the veins, more tardily so at midrib and nerves beneath, subentire or shallowly glandular-serrate, (6-)7-]4 by (1.5-)2-3(-3.5-4) cm, nerves (10-) 12-14 pairs, curved-ascending towards the edge, rather indistinctly inarching there, prominent beneath, veins but slightly raised beneath; petiole slender, l-1.5(-2)cm. Racemes panicled, slender, 10-20 cm, covered with a dilutely rusty tomentum of both smaller and larger stellate and/or fascicled hairs; flowers rather densely arranged along the robust rachis. Pedicels slender, 1-2 mm (in fruit c. 3 imn). Petals erose-fimbriate at apex, 3-4 mm. Distr. Malesia: Borneo. Ecol. Generally in mountain (also mossy) forest or on forest edges, rarely in lowland (ap- parently secondary) forest, locally common, especially on Mt Kinabalu, (30-)700-1200(-1830) m. Fl. fr. Jan. -Dec. Vern. Kalintiihan, mitindike, Dusun. 4. var. novoguineensis (Kaneh. & Hatus.) Sleum. Bot. Jahrb. 87 (1967) 90. — C. longispicata J. J. S. var. novoguineensis Kaneh. & Hatus. Bot. Mag. Tokyo 56 (1942) 474, f. 3. — C. lancifolia TuRC7. Bull. Soc. Nat. Moscou 36, ii (1863) 231; Merr. Philip. J. Sc. 1 (1906) Suppl. Ill; ibid. 2 (1907) Bot. 292; En. Philip. 3 (1923) 242. — C. alnifolia (non L.) Blanco, Fl. Filip. ed. 2 (1845) 259; ed. 3, 2 (1878) 117; Merr. Sp. Blanc. (1918) 297. — C. canescens {non Reinw. ex Bl. s. str.) F.-Vill. Novis. App. (1880) 121; Vidal, Sinopsis Atlas (1883) 30, t. 60, f. B; Phan. Cuming. Philip. (1885) 123; Rev. PI. Vase. Filip. (1886) 172; Merr. En Philip. 3 (1923) 242. — C. williamsii C. B. Rob. Bull. Torn Bot. CI. 35 (1908) 73, 75. — C. castanea Elm. Leafl. Philip. Bot. 9 (1934) 3182. Leaves lanceolate to oblong, apex shortly sub- acutely acuminate, base ± cuneate, younger ones glabrous above, more densely or laxly set with longish simple or fascicled, ± appressed pale rusty hairs on midrib and nerves, ± laxly and minutely stellate-hairy on veins beneath, mature ones gradu- ally glabrescent, finally sometimes quite glabrous, shortly or rarely more deeply and distantly serrate in the upper part, occasionally subentire, (4-)5-10 (-14, rarely -17) by (1.5-)2-3.5(-4.5, very rarely -8) cm, nerves (8-)10-14 (rarely -16) pairs, curved-ascending, ± joined before the edge, raised beneath, veins often obscure though a fine reticulation is visible; petiole rather slender, 0.5-1 (-1.8) cm. Racemes panicled, (4-)6-15(-24) cm, flowers rather dense along the ± slender rachis, covered with a tomentum of longer, fas- cicled rusty and minor pale stellate hairs (rarely only with the latter ones). Pedicels generally rather slender, rarely more robust, 1-2 imn (in fruit 2-3 mm). Petals irregularly erose-crenulate mainly at apex, (2-)2.5(-3)mm. Distr. Malesia: SE. Celebes, Philippines (Batan Is, Luzon, Mindoro, Samar, Leyte, Biliran, Negros, Mindanao, Catanduanes), NW. New Guinea. Ecol. In the Philippines and in Celebes in mountain (also mossy) forest, mainly on ridges and peaks, (400-)700-2100 m, in New Guinea in open beach forest and in savannah-like thickets on dry hill (300 m) near the coast, presumably mainly in secondary vegetation. Fl. fr. Jan. -Dec. Vern. Alibungog, C. Bis., kalyapi, kayat- buntot. Bag., kamog. Ilk., kamueg, Ig., malaklak. Tag., maratungau, Ibn., tagobahi, P. Bis., mayaorin, Batan. 5. var. ledermannii (Schltr) Sleum. Bot. Jahrb. 87 (1967) 87. — C. papuana Schltr, Bot. Jahrb. 52 (1915) 219, non J. J. S. 1914. — C. ledermannii Schltr, ibid. 55 (1918) 194. Leaves lanceolate-elliptic to lanceolate, apex acutely acuminate, base cuneate, thin-subcoria- ceous, younger and submature ones glabrous above, set with sparse rather coarse rufescent simple or fascicled and/or finer stellate hairs on midrib and nerves, hardly at veins beneath, in- tervenium glabrous, mature ones almost com- pletely glabrous, rather sharply serrate in the upper part, 5-8(-ll) by 2-3 (-4) cm, nerves 12-14(-15) pairs, ± curved, subparallel, rather indistinctly inarching before the edge, prominent beneath, veins ± obscure; petiole rather slender, (0.7-)l-1.5 cm. Racemes panicled, all over cov- ered with a very short tomentum of brownish to greyish stellate hairs, in which a few larger fas- cicled hairs occur, rather lax-flowered, 5-12 cm. Pedicels rather slender, 2-3 mm (in fruit 3-3.5 mm). Petals erose-crenulate, (3-)4 mm. Distr. Malesia: E. New Guinea (Sepik Distr.; Western Highlands). Ecol. In mountain (also mossy) forest, and in regrowths, (200-)400-1830 m. Fl. June-Sept. Vern. Talellpi, Wapi (Torricelli Mts). 6. Clethra kebarensis Sleum. Bot. Jahrb. 87 (1967) 92. — Fig. 4. Laxly branched shrub or treelet, 2-3 (-6) m, trunk often blackish, 4-10 cm 0. Branchlets distally ferrugineous-tomentellous. Leaves obo- vate-oblong or -elliptic, more rarely oblong, apex ± shortly subacutely acuminate, base cuneate, rarely obtuse or even rounded in the same spec- imen, subcoriaceous to firmly chartaceous, young ones very laxly stellate-hairy above, sub- densely covered beneath with short, fascicled rusty hairs at midrib, and all over the under- surface with very dense minute stellate hairs, which form a pale coherent layer, moreover nerves and especially veins and veinlets set with darker rusty stellate-fascicled, tardily evanescent hairs so to say on top of the pale layer mentioned above, mature leaves dark green above, yellowish rusty beneath, early glabrescent above, tardily so beneath, generally shortly to very shortly sub- serrate- or crenate-dentate, (4-)6-9 by (2-)2.5-4(- 5.5) cm, nerves in 14-16(-17) pairs, slightly curved or rather straight from the midrib, rather obscurely inarching, prominent below, veins but slightly raised; petiole 9-14 by c. 1 mm. Racemes 3-6, ± fascicled, partly branched below, erect. 1971] Clethraceae (Sleumer) 147 . .;., Kcbar Valley, Ncrtoi, 700 m, NW. New Ciuinca (\,.,. i;,. Sleumer 6841) (Sleumer, 1961). dense-flowered, 6-12 cm, covered with a short, almost villous, rusty (finally greyish) pubescence of fascicled and stellate hairs; rachis rather robust. Pedicels thickish, 1-1.5 mm at anthcsis, subtending bracts subovatc-subulate, 2 mm, persistent for a while. Calyx lobes ovate, subacuminate, 2-2.5 mm. Petals obovate-spathulate, often connate or coherent in the lower part, crenulate, glabrous, white or cream, (2.5-).l-3.5 by c. 1.5 mm. Fila- ments dilated downwards, glabrous, 2(-2.5) mm; inthcrs obcordatc, c. 0.6 mm. Ovary rusty- •omentulosc; style glabrous, 1.2 1.5 mm (in fruit to 2.5 mm). Capsule subglobosc, c. .1 mm e. Seeds subtrigonou6-ovoid, 0.8 mm. Distr. Malesia: New Guinea (Vogelkop Peninsula). Ecol. In patches of shrubberies within grass- land or fern thickets of Gtcichen'ui, or in forest edges, not rarely in periodically burned vegetation, fairly common locally, sometimes forming pure loose stands on sandy or clayey soil, 700-1200 ( I950)m. H. jr. Jan. Dec. 7. Clcthra sumbawaensis SituM. Bot. Jahrb. 87 (1967)94. Treelet, 4-8 m. Branchlets distally clad with a rusty rather scabrid pubescence, l.cuvcs elliptic- oblong or elliptic, apex shortly and acutely acuminate, base cuneate, thin-coriaceous, younger and submaturc ones glabrous above, subdensely set with longjsh simple and fascicled (or stellate) rusty hairs along midrib and nerves, sparsely so 148 Flora Malesiana [ser. I, vol. 71 on the veins beneath, moreover beneath all over the iindersurface with a dense thin layer of minute stellate and pale (or here and there rusty), finally greyish hairs, apparently very tardily glabrescent, serrate in the upper y\, (5-)6-9 by 2-4 cm, nerves (1 5-) 17-20 pairs, rather straight and sub- parallel, in part forked distally, reticulately dis- solving before the edge, raised beneath, veins transverse, slightly prominent beneath, veinlets obscure; petiole l-1.5(-2)cm. Racemes rather numerous, condensed to a panicle, 5-8(-10) cm; rachis stoutish, covered with a floccose rusty kind of wool. Pedicels rather slender, with a short stellate pubescence as are the sepals, 3-4 mm, bracts subulate, tomentulose, 8-10 mm, caducous after anthesis. Calyx lobes ovate-acuminate, less hairy at the fimbriate margin, glabrous inside, 3 mm. Petals obovate, thin, white, fragrant, gla- brous, minutely fimbriate-crenulate i all along the margin, 5 by c. 2.5 mm. Filaments glabrous, 2 mm; anthers broadly obovate, base apiculate, 1 mm. Ovary tomentose; style slenderly columnar, glabrous, 4-4.5 mm, shortly 3-iobed. Distr. Malesia: Lesser Sunda Islands (Sum- bawa: Mt Batulanteh; Flores: Mt Kanaka). Ecol. In forest, (900-) 1 600-2400 m, on an- desite soil. Fl. April, Oct. 8. Clethra tomentella Rolfe ex Dunn, Kew Bull. (1922) 185; Merr. En Philip. 3 (1923) 243; Sleum. Bot. Jahrb. 87 (1967) 94. Small tree or shrub. Branchlets rather slender, tips grey-tomentellous. Leaves lanceolate to elliptic-oblong, or oblanceolate, apex ± shortly subacutely acuminate, base attenuate, firmly chartaceous, subentire or sometimes sparsely very shortly serrate-dentate in the upper part, younger and submature ones glabrous above, appressedly greyish-stellate-tomentellous all over beneath, mature ones glabrescent tardily, first on the inter- venium, finally also almost completely along midrib and nerves, 6-9(-l 1 ) by (2-)2.5-3(-3.5) cm, nerves (10-)12-14(-15) pairs, prominent beneath, reticulation visible only in the already glabrate parts; petiole rather slender, (5-)6-10(-12) mm. Racemes several, panicled, laxly or more densely set with flowers, 7-14(-20) cm, covered with a short, greyish or rusty-stellate tomentum; rachis slender. Pedicels slender, c. 2 mm (c. 3 mm in fruit), bracts subulate-linear, 3-5 mm, caducous in the beginning of anthesis. Calyx lobes ovate- acuminate, glabrous inside in part, 1.5-2 mm. Petals obovate-spathulate, minutely erose apically, glabrous, white, 2-2.2(-2.5) mm. Filaments dilated towards the base, glabrous, 1 mm; anthers obovate, hardly I/2 mm. Ovary tomentellous; style thickish, glabrous, 1 mm, very shortly 3- lobed. Capsule subglobose, 2.5 mm 0. Seeds sub- trigonous-ovoid, 0.7-1 mm. Distr. Malesia: Philippines (Luzon: Mountain Prov. to Batangas). Ecol. Scattered in thickets or forests on slopes, chiefly on ridges in mossy forest, 600-1800 m. Fl. fr. May-Sept. Vern. Amog, kamug, kamung, Ig., ayusan. Tag. 9. Clethra arfakana Sleum. Bot. Jahrb. 87 (1967) 95. Shrub or erect treelet, 2-3 m, much branched. Branchlets slender, tips rusty-tomentellous. Leaves obovate-oblong or -elliptic, apex shortly acutely acuminate, base cuneate, slightly inequilateral, subcoriaceous to firmly chartaceous, flush red- dish, younger and mature ones dark green and glabrous above, whitish greyish below, i.e. except the sparsely hairy or subglabrous midrib and nerves covered by a short tomentum of minute stellate hairs, very tardily glabrescent, (sub)entire or shallowly dentate, (2-)3-4.5 by (l-)1.3-2 cm, more deeply dentate and up to 7.5 by 3 cm in not flowering new shoots, nerves in 10-12 pairs, i straight from the midrib, curved upwards, slightly prominent beneath; petiole 5-8(-IO) mm. Racemes several, panicled, 5-7(-8) cm, rather lax-flowered, pale-rusty stellate-tomentellous in all outer parts; rachis slender. Pedicels slender, 2-3 mm, basal bracts early fugacious. Calyx lobes ovate-ob- long, long-ciliate, 3 by 1.5 mm. Petals obovate- spathulate, sometimes coherent at the base, glabrous, white, shortly erose-fimbriate all along the edge, (3.5-)4(-5) by 2-2.5 mm. Filaments di- lated towards the base, glabrous, 2.5 mm; anthers obcordate, about 0.5 rrun. Ovary almost hirsute; style glabrous, 1.8 mm at anthesis, hardly ac- crescent in fruit. Capsule subglobose, 2.5-3 mm 0. Seeds oval to subtrigonous, c. 1 mm. Distr. Malesia: New Guinea (Vogelkop Peninsula: Arfak Mts around the Anggi Lakes). Ecol. In edge of Notlwfagus-Myrtaceae- forest or in more open heath formation within devastated montane forest, 2000-2600 m, common locally. Fl. fr. Jan. 10. Clethra longisplcata J. J. S. Bull. Jard. Bot. Btzg III, 4 (1922) 240; Sleum. Bot. Jahrb. 87 (1967) 96. — C. elongata J. J. S. Bull. Jard. Bot. Btzg III, 1 (1920) 398, t. 47, non Rusby, 1907. Shrub or treelet, up to 5(-12)m, trunk up to 10 cm 0, bark green, with shallow pale brown flakes. Branchlets and petioles rusty to greyish tomentellous. Leaves oblong-lanceolate to elliptic- oblong, more rarely elliptic, apex rather shortly acuminate and subacute, base cuneate, rarely almost obtuse, inequilateral, subcoriaceous, young- er ones glabrous above, covered all over beneath by a coherent tomentum of minor stellate cin- namomeous to greyish hairs, and of larger stel- late hairs so to speak on top of that layer, mature ones tardily glabrescent beneath, furtl^ermore sometimes with sparse fascicled hairs on the mid- rib beneath, (4.5-)6-ll by (1.7-)2-3.5(-4) cm, callose-denticulate or subentire, nerves 10-12(-14) pairs, generally starting at an acute angle, curved- ascending towards the edge, a little impressed above, prominent beneath, veins distinctly, veinlets rather inconspicuously raised beneath; petiole 0.6-1. 3(-l. 6) cm. Racemes panicled, rather slender, (15-)20-30(-35) cm, laxly many- flowered, with a short stellate tomentum, in which also some fascicled hairs may be found. Pedicels slender or almost so, 2-3 mm (to 4 mm in fruit), 1971] Clethraceae (Sleumer) 149 Fig. 5. Clethra papuana J. J. S., Kebar Valley, Mt Nettoti, 1900 m, NW. New Guinea (Van Royen & Sleumer 7449) (Sleumer, 1961). bracts subulate, caducous. Calyx lobes narrowly ovate-subdeltoid, l.5-2(-2.5) mm. Petals partly connate at base, broadly spathulate, crenulate. glabrous, white, scented, c. 3 by 1 .5 mm. Filaments glabrous, 1.5 mm; anthers obcordatc, 0.5 mm. Ovary apprcssedly hairy; style glabrous, c. 1 .7 mm (to 2 mm in fruit), very shortly .1-lobed. Capsule subglobosc, c. 2.5 mm 0. Seeds convex- ovoid, 0.7 mm. Distr. Malesia: Borneo, Philippines (Palawan) and Central Celebes. Ecol. Generally in mountain forest, also in secondary vegetation, on clayey ground, (60-) 500 1525 m. Fl. Jan.-Dec. Vcrn. Ponyo, Toradja, lanf;, Iban, yarupoi, Dusun. 11. Clethra papuana J. J. S. Nova Guinea 12 (1914) 169, t. 53; Kaneh. & Hatus. Bot. Mag. Tokyo 56 (1942) 476; Sleum. Bot. Jahrb. 87 (1967)97. — Fig. 5, vur. papuana. Shrub or treelet, (l-)3-6(-8) m, more rarely a tree up to 15 m, bole up to lOm by 30 cm. Branchlcts at tips rusty subvillous-tomentose. Leaves elliptic or obovatc-elliptic, sometimes ob- long-elliptic, rarely elliptic-lanceolate, apex short- ly subacutcly acuminate, rarely rather blunt (more acutely acuminate in sterile shoots), base inequilateral, broadly cuncate to subtruiicate- obtuse, coriaceous, (irm, younger ones with stellate and fascicled hairs mainly along midrib 150 Flora Malesiana [ser. I, vol. 7^] and nerves above, clothed all over beneath by a dark rusty subvillous tomcntum formed by stellate and fascicled hairs, mature ones glabrate above, dark green in the fresh state there, under- surface rusty- to greyish- or (almost silvery-) tomentulose (the large fascicled hairs having mostly disappeared by then), entire or shortly ir- regularly serrate-dentate (more deeply and sharply serrate in sterile shoots), (4-)5-8(-12) by 2-4 (-5.5) cm, variable in size even in the same specimen, sometimes microphyllous, nerves 12-J4 pairs, much spreading, ± inarching at or a little before the edge, well raised beneath, veins not much conspicuous beneath, reticulation faint; petiole rusty-tomentulose, (l-)1.2-1.6(-l.8) cm. Racemes panicled, erect, all over rusty-tomentose, flowers generally densely arranged, rarely up to 15 cm; rachis robust. Pedicels thickish, 1-2 mm (up to 4 mm in fruit), bracts thickish, subulate, caducous. Calyx lobes oh\onz-ovdi\.e, 3-3. 5 (-4) mm. Petals not rarely connate or coherent at base, obovate-spathulate, edge slightly erose, white, sometimes pale cream or suffused with red, gla- brous, slightly scented, c. 3.5 by 2 mm. Filaments glabrous, hardly 2 mm; anthers it obcordate, 0.8 mm. Ovary almost villous; style glabrous, shortly trilobed, (1-)1.2-1.5 mm (hardly 2 mm in fruit). Capsule subglobose, c. 3 mm 0. Seeds ovoid-oblong, irregularly angular, 0.8-1 mm. Distr. Malesia: NW. New Guinea (Vogelkop Peninsula: Kebar Valley and Arfak Mts). Ecol. In (mossy) Notlwfagus-Coniiev-ioresi edge or in open heath vegetation on crest, 1800- 2600 m. Fl. Oct.-Jan. var. trichostyla Sleum. Bot. Jahrb. 87 (1967) 98. Petals (3 mm) laxly hairy along the edge. Style set with a few appressed hairs at base. Otherwise as var. papuana. Distr. Malesia: W. New Guinea (Wissel Lakes area), once found. 12. Clethra pulgarensis Elm. Leafl. Philip. Bot. 5 (1913) 1757; Merr. En. Philip. 3 (1923) 243; Sleum. Bot. Jahrb. 87 (1967) 99. Low, stunted tree, or shrub-like; branches as- cending, rigid, covered with brown bark. Branch- lets short, rather robust, short-hairy. Leaves ob- long to subobovate-elliptic, apex shortly at- tenuate or subacuminate, base broadly cuneate, the very base often obtuse, subcoriaceous, firm, younger ones very densely shortly stellate-hairy above, covered all over beneath with a tomentum of minor and larger stellate, brownish greyish hairs, large fascicled hairs on petiole and midrib below few or absent, mature ones glabrous and a little shining above, rather persistently greyish tomentulose beneath, finely serrate-dentate in the upper part, tips of the serratures usually callose, 5-8 by (2-)2.5^ cm, midrib bold beneath, nerves in 9-11 (-12) pairs, curved, excurrent or divided along or before the edge, raised beneath, reticula- tion rather faint; petiole 0.5-1 cm by c. 2 mm. Racemes panicled, (8-)10-15cm, covered with a fulvous-greyish tomentum of stellate and fascicled hairs; rachis stoutish. Pedicels robust, 2-3(-5) mm, bracts subulate, caducous. Calyx lobes ovate-oblong, 4-5 by 2(-2.5) mm. Petals obovate- spathulate, minutely crenulate at apex, white, glabrous, (4-)5(-6) by c. 2.5 mm. Filaments gla- brous, 3 mm; anthers obcordate, 1 mm. Ovary longish erect-tomentose; style thickish, glabrous, (2.5-)3 mm (3-4 mm in fruit), shortly 3-lobed. Capsule depressed-globose, c. 3 mm 0. Seeds oval, 1 mm. Distr. Malesia: Philippines (Palawan), in low woody vegetation on summit of Mt Pulgar near Puerto Princesa, 1200 m. Fl.fr. May. 13. Clethra pachyphylla Merr. Philip. J. Sc. 13 (1918) Bot. 103; En. Born. (1921) 460; Sleum. Bot. Jahrb. 87 (1967) 99. — Fig. 1. Shrub or mostly small tree, 2-4(-10) m high, trunk up to 20 cm 0, laxly branched, branches obliquely ascending to almost horizontal. Branch- lets robust, rufous-ferrugineous-tomentose at apex. Leaves oblong or obovate-oblong, more rarely oblong-elliptic or elliptic, apex shortly subacutely acuminate, base cuneate, coriaceous, firm, younger ones subdensely floccose- or subvillous-stellate- hairy above, all over beneath with a tomentum of minor pale stellate hairs and a more coarse one of major stellate and fascicled, initially rusty, finally greyish hairs, midrib and nerves mainly with major fascicled hairs, entire or more rarely and but in part irregularly serrulate, 4-9(-14) by (2-)2.5-3.5(^, -5) cm, midrib bold beneath, nerves (9-)10-12(-13) pairs, prominent beneath, curved and i excurrent at the edge, veins slightly raised beneath, reticulation more distinct but in ± mature leaves; petiole 1.2-1.5(-1.8) by 1.5-2 mm. Racemes panicled, rather dense-flowered, robust, all over villous or almost so by minor and major stellate and coarse fascicled rusty hairs, 5-10(-12) cm; rachis stout. Pedicels thickish, 2-3 (-4) mm, bracts subulate, 4-5(-8) mm, caducous. Calyx lobes oblong-ovate, subacuminate, (3.5-)4-5 mm. Petals spathulate to spathulate-oblong, irregularly erose or crenulate at apex, white to cream, scented, glabrous, (3-)4(-5) by 2.5-3 mm. Filaments gla- brous, 2-2.5 mm; anthers subobovate, 0.8-1 mm. Ovary short-tomentulose; style thickish, glabrous, 2.5-3 mm (3.5-4 mm in fruit), shortly 3-lobed. Capsule depressed-globose, 3 mm 0. Seeds sub- trigonous, 1-1.2 mm. Distr. Malesia: NE. Sarawak (Mt Murud area) and North Borneo (Mt Kinabalu). Ecol. Scattered in Quercus havilandii and Tris- tania forest, still not too rare in mossy Lepto- spermum-Dacrydium forest, also in secondary vegetation, (870-)1200-2500(-3050) m, on poor soil. Fl.fr. March-Oct. OXALIDACEAE (J. F. Veldkamp, Leyden) Herbs, sometimes with scaly rhizomes, bulbs, bulbils or stolons, or woody perennials, shrubs, lianas or trees. Leaves penninerved, digitately or pinnately trifoliolate, imparipinnate or paripinnate, basal, alternate, subopposite or apically tufted. Stipules sometimes present. Petioles with basal joint, petiolules articulated. Inflorescences basal, axillary or pseudoterminal, cymose to pseudumbellate, rarely racemose, 1-many-flowered, bracteate and bracteolate. Flowers o, very rarely also 3 specimens (Dapania), actinomorphic, 5-merous, hetero-tri-, -di-, or homostylous, sometimes cleistogamous. Pedicels articulate. Sepals imbricate, free or connate at base, sometimes with apical calli (Oxalis), persistent. Petals contort, quincuncial or cochlear, free but usually cohesive above the base ('pseudosympetal'), clawed (sometimes minutely so), glabrous or inside sometimes with minute papillae or pilose. Filaments 10, obdiplostemonous, connate at base into an annulus, persistent, the epipetalous (shorter) sometimes with a basal gland near the insertion of the petals, or sometimes with 2 scales or dark lines on the annulus (Dapania), rarely without anthers; the episepalous (longer) with a dorsal tooth (Oxalis) or hunchbacked; anthers dorsifixed, versatile, 2-celled, dehiscing extrorsely by longitudinal slits. No disk. Ovary 5-celled, superior; styles 5, terminal, persistent, free, in LF^ and MF erect, in SF patent to recurved, rarely reduced (^ flowers); ovules 1-2-several per cell in 1-2 rows, epi- and anatropous, pendulous, super- posed, bitegmic. Fruit capsular, loculicid, 5-celled, dry, rarely fleshy and indehis- cent. Seeds usually with an aril; endosperm copious, fleshy, rarely absent; embryo straight. Distribution. 6(7?) genera with c. 850 spp. Of the Malesian representatives Oxalis, the largest genus, is most numerous in S. America and S. Africa and Biophytum in S. America and Madagascar; Dapania has 2 spp. in Maiesia and I in Madagascar; Sarcotheca (11 spp.) is endemic in Malesia, while Averrhoa (2 spp. ) assumediy also originated here; it is now cultivated pantropically. In Malesia there are 5 genera with 29 species, of which 14 endemic. The generic distribution of the family offers in the Old World two remarkable patterns. First, that of Dapania which shows the characteristic disjunction between West Malesia and Madagascar. Second, that of O.xalis sect. Acetosellae which almost resembles that of Euphrasia, that is, a temperate genus with two stepping stones across the tropics (Luzon, New Guinea), otherwise bipolar. Fig. 2. Ecology. Many members of the family have sensitive leaves and show sleeping movements, some being also seismonastic which is conspicuous in Biophytum. Dispersal. The fleshy fruits oi Averrhoa, Dapania, and Sarcotheca are no doubt eaten by various animals as bats, birds, and monkeys, and dispersed by them. The seeds of O.xalis and Biophytum have a peculiar ejaculative aril originally enveloping the entire seed which at maturity shoots them away for some distance (ZiEGLER, Ber. Bayer. Bot. Ges. 36, 1963, 61). The cultivated species of O.xalis which set no fruit in Malesia, are locally tenacious weeds through their bulbils. The native O.xalis species propagate also vegctatively, by stolons and root-stocks. All O.xalidaceae have an arillate seed except the species of Sarcotheca and Averrhoa bilimbi. On germination some remarks are made under the genus Biophytum. Morphology. Episeptal rimae are found in some species of O.xalis, in most of Dapania and all Sarcothecas; in A verrhoa they are only inconspicuous, apical furrows. The most primitive state is probably represented in O.xalis corniculala. The septs fail to enlarge towards fructification, whereby the cells of the fruit arc only united by their attachment to the central axis. The walls of the cells are pressed together and slits arc formed, especially conspicuous by transparent ridges. These ridges are wide apart in the Malesian Dapanias and are slightly bent inwards. In Sarcotheca the septs arc developed, at least in the lower half; in the upper half the rimac may be open and papillose inside, or closed and glabrous. Episeptal rimac arc absent in all Biophylums and in Dapania pcntandra from Madagascar. (1) LF = long-styled form, MF - mid-styled form, SF = short-styled form. (151) 152 Flora Malesiana [ser. I, vol. 7^ Dehiscence of the fruits in Biophytum and Dapania is into a 5-rayed star. In Oxalis the valves remain united and only longitudinal loculicid slits are iormcd. Sarcotheca and A verrhoa have indehiscent fleshy fruits. Anatomy. Chauvel, Rech. sur la famille Oxal. (1903) thesis; Moll & Janss. Mikr. 2 (1911) 9 (Averrhoa); Heimsch, Lilloa 8 (1942) 97, 191; Metcalfe & Chalk, Anat. Dicot. 1 (1950) 299. Phytochemistry. Few phytochemical investigations were performed with members of the family. In fact, distinct constituents have been isolated only from a few species of Oxalis. The tendency shown by many species of Oxalis to accumulate large amounts of oxalic acid in water-soluble form is known since a long time. Leucocyanidins and ieucodelphinidins have been demonstrated to be present in the leaves of some species of Oxalis and of Averrhoa carambola L. This agrees with the idea that Oxalidaceae represent the most primitive family of Geraniales. Probably the species described by plant anatomists as possessing 'tannin' cells are the ones which contain leucoanthocyanins in leaves. The bright yellow flower pigments of Oxalis cernua Thunb. are the aurone glycosides aureusin and cernuoside. A yellow quinonoid pigment was isolated from the bulbs of one species of Oxalis (O. purpurata Jacq. ?) and later identified with rapanone, a benzoquinone occurring frequently in myrsinaceous plants. Too little phy- tochemical information is available at present for a chemotaxonomical appreciation of systematic relationships of Oxalidaceae. It might be significant, however, that rapanone does also occur in Con- naraceae. General reference: Hegnauer, Chemotaxonomie der Pflanzen 5 (1969) 255. — R. Hegnauer. Pollination. Heterostyly is a common feature in the family, heterotristyly is observed in Oxalis, Biophytum, and Averrhoa bilimbi. This must be assumed to be the primitive condition, as is found in the allied Connaraceae. From it is derived a heterodistylous condition in Sarcotheca and Dapania, in certain species of Oxalis and Biophytum, and m Averrhoa carambola. A further derived, homostylous condition is found in certain species of Oxalis and Biophytum. Darwin concluded that the heterotristyly in a Ceylonese Biophytum which he examined, was also functional in analogy with Oxalis while in cleistogamous flowers ^ he observed that in some way the incompatibility factor seemed to be removed (The Different Forms of Flowers, ed. 1877, 181, 323). Salter (J. S. Afr. Bot. Suppl. 1, 1944) confirmed by experiments that functional heterotristyly occurs in Oxalis. This was also found in S. American species of Oxalis. Miss P. Mayura Devi (J. Genetics 59, 1964, 41) found in Indian Biophytum in her experiments a significant incompatibility in illegitimate crossings, except for MF selfed with pollen of the long stamens. This means a loss of compulsory he- terotristyly. In a later article {ibid. 1966, 245) she described another mid-homostylous form which proved excellently self-compatible. It is regrettable that she did not conserve voucher specimens bound to her experiments, as the identification of Indian Biophytum is in distinct confusion and more than one species is cited as B. sensitivum. The plants used by Devi certainly do not belong to B. sensitivum, as in her pictures the corolla is much longer than the calyx. Functional heterotristyly is apparently present in the introduced species of Oxalis, O. deppei and O. latifolia, which in Malesia occur as SF only and have never been observed with fruit. O. corymbosa, although present with SF and MF, does not fruit either, but the flowers are often monstruous in this species. All three reproduce very successfully by bulbils and can become obnoxious weeds. Biophytum fruticosum, B. adiantoides and B. microphyllum are also heterotristylous. No experiments have been done. Biophytum sensitivum and B. reinwardtii sens. str. are both in India and Malesia mid-homostylous. Reduction to a single stylar form (LF) is found in Oxalis magellanica and O. acetosella ssp. griffithii; reduction to homostyly (MF) has very far progressed in both O. corniculata where LF occurs rarely (6 out of 105 plants examined; cf. also Eiten, Am. Midi. Natur. 69, 1963, 280) and Averrhoa bilimbi, where LF and SF were only observed once. The introduced Biophytum dendroides is apparently self- compatible, as only one form (LF) has been found at Bogor and in the glasshouses of the Botanical Gardens of Leyden and Groningen, where in the apparent absence of pollinators it is fully fertile and weedy. Heterodistyly (LF and SF) occurs in Biophytum petersianum, Sarcotheca, and Dapania pentandra from Madagascar. It is plausible that the S flowers of the Malesian Dapania originated by an extreme reduction of the pistil in SF-flowers. Averrhoa carambola is heterodistylous for LF and MF. The latter type might well correspond with SF, as the shorter stamens are much reduced and without anthers. More information about heterostyly and literature concerning this matter is given by Ornduff and by MULCAHY (Am. J. Bot. 51, 1964, 307 and 1045). Palynology. See Erdtman, Pollen Morphol. & Plant Tax. 1 (1952) 302; Huang, Taiwania 13 (1967) 70 and Huyhn, Bot. Jahrb. 89 (1969) 272. Uses. Several species of Oxalis are cultivated as ornamentals, Averrhoa for its edible fruit. The wood of the ligneous Oxalidaceae is useless as timber. In Malaya the fruits of Sarcotheca are sometimes eaten. See also Heyne, Nutt. PI. (1927) 850. Taxonomy. In Engler's Syllabus (2, 1964, 248) Scholz divided the genera in two groups, A and B, on the aestivation and the number of ovules per carpel. Through this Averrhoa was joined to the affinity of Oxalis and Biophytum. However, the aestivation is inconstant. Moreover, the affinity of Averrhoa is (1) In Biophytum I found also in not cleistogamous flowers opened anthers with good pollen already in bud (protrandry). 1971] OxALiDACEAE (Vcldkamp) 153 doubtless with Sarcotheca and Dapania in all other characters. These three genera form a very clear reticulate affinity. The family is considered to be the most primitive of the Geraniales by Hallier/. (Arch. Neerl. ser. 3B, 1, 1912, 109) andENGLER (Nat. Pfl. Fam. ed. 2, 19a, 1931, 10). This primitive position is possibly the reason for the affinity with the Connaraceae, at which some authors have pointed (Hallier /. New Phyt. 4, 1905, 158; Arch. N^erl. ser. 3B, 1, 1912, 109; Schellenberg, Pfl. R. Heft 103, 1938, 127; Takhtajan, Evol. Angiosp. 1959, 236). The latter family is usually placed near the Leguminosae. As a matter of fact specimens of Rourea and Sarcotheca were often confused and are very similar, the pistil and fruit excepted. The closest allied family is Geraniaceae in which they were merged by Bentham &. Hooker /! In fact the American Hypseocharis seems to link both families. The only constant character was said by Hallier /. (Beih. Bot. Centralbl. 39, ii, 1921, 172) to be the tenuinuceilate ovules in Oxalidaceae and crassinucel- late ovules in Geraniaceae, but Hypseocharis and many species of other genera have not been checked on the general validity of this character, as Hallier /. himself admitted. And one must be very careful in this respect; for example Hutchinson (Fam. FI. PI. ed. 2, 1, 1959, 494) stated that Oxalidaceae have albuminous seeds, but some species of Oxalis are exalbuminous (cf. Salter, J. S. Afr. Bot. Suppl. 1, 1944, 26); he stated also that Oxalidaceae are exstipulate, but stipules are found in all Biophytums and also in some species of Oxalis (sect. Acetosella, O. corniciilata, etc.). Hutchinson's division of Oxalidaceae into three different families belonging to three different orders seems not to have a reliable basis as Averrhoa cannot be divorced from Sarcotheca and Dapania and there is no reason for admitting a preponderant importance to their ligneous habit; besides many species of Biophytum and Oxalis are dwarf shrubs. KEY to the genera 1. Herbs or dwarf shrubs, rarely up to 1 '/t m high. Leaves 3(-4)-foliolate, or paripinnate, herbaceous. Fruit a dry capsule. 2. Leaflets 3-4. Capsule with the valves remaining attached to the central axis 1. Oxalis 2. Leaflets 6 or more, pinnate. Capsule dehiscing into a 5-rayed star, without leaving a columella. 2. Biophytum 1. Shrubs, trees or lianas, much higher. Leaflets 1 or 3, or leaves imparipinnate, 2-many-jugate. Fruit fleshy, dehiscent into a 5-rayed star or indehiscent. 3. Leaflets 1 or 3, chartaceous or subcoriaceous. Lateral petiolules articulate, after dropping leaving a short stalk on the rachis. Ovules 1-2 per cell. 4. Lianas. Inflorescences racemose. Petals inside glabrous. Fruit dehiscent into a 5-rayed star. Seeds arillate 3. Dapania 4. Shrubs or trees. Inflorescences paniculate. Petals minutely papillose inside. Fruit indehiscent (but episeptal rimae sometimes open!). Seeds exarillate 4. Sarcotheca 3. Leaves imparipinnate, 2-many-jugate, herbaceous to papyraceous. Lateral petiolules after dropping not leaving a stalk on the rachis. Ovules 3-7 per cell 5. Averrhoa 1. OXALIS LiNNE, Gen. PI. ed. 5 (1754) 198; Sp. PI. (1753)433; Knuth, Pfl. R. Hefl 95 (1930). - Fig. 1. Annual or perennial herbs (or dwarf shrubs, cxlra-Mal.), some stemless with rhizomes or bulbs. Leaves digitately or pinnalely 3(-4)-foliolate; leaflets in Mai. spp. (except O. harrelieri) i sessile. Stipules when present adnate with the base of the petiole. Peciuncles basal from bulbs or solitary and axillary. Inflorescences cymose to umbellate, 1 -many-flowered. Bracteoles 2-several, sometimes with apical calli. Pedicels articulate at base and sometimes beneath the calyx. Sepals shortly connate at base, with or without apical calli. Petals coherent above the claw, contort, glabrous. Filaments: longer ones sometimes with a dorsal tooth. Stif^ma cylindric and minutely bilobed to peltate, sometimes papillose. Ovules 1 to c. 10, in 1-2 rows per cell. Capsules loculicid by longitudinal slits, sometimes with episeptal rimae. Seeds 1-r. 10 per cell, usually few; aril bivalved, ejaculatory; testa smooth, or with transverse ridges or longitudinal furrows. 154 Flora Malesiana [ser. I, vol. 7^ Fig. 1. O.xalis magellanica Forst./. a. Habit, x\,b. 'callus' of leaf, X 12, c. apex sepal, X 12, J. bracteoles, from both sides, X 4. — O. corymbosa DC. e. Leaf, X 1 . — O. barrelieri L. /. Leaf, nat. size, g. dentate stamen, x 12. — O. acetosella L. ssp. griffithii Hara. h. Leaf, nat. size, /. bracteoles, from both sides, x4 (fl-i/BALGOOY 252, e Backer 37150,/-^ Boerlage s.n. a. 1880, h-i Steiner 1990). Distr. Cosmopolitan, at least 700 spp., mainly from S. America and the Cape, in Malesla 3 native spp.; others introduced and escaping, sometimes becoming weedy. Fig. 2 (§ Acetosellae). Ecol. O. acetosella ssp. griffithii and O. magellanica are (in Malesia) both characteristic mountain plants, above 2200 m. O. corniciilata is rather indifferent to altitude. Morph. The longer filaments are frequently thickened in the basal part, as in Biophytiim. The apex of the thickening may emerge as a small tooth, e.g. in O. latifolia, O. deppei, and O. barrelieri. This tooth does not carry a vascular trace (Narayana, J. Jap. Bot. 41, 1966, 321). For heterostyly see under the family. An important study of the morphological diversity and taxonomy of the S. African species is provided by Salter (J. S. Afr. Bot. Suppl. 1, 1944). Notes. Pending a revision of the S. American species, Miss Lourteig (Paris) advised me to accept the names of the introduced species in the current sense. Sizes of petiole and pedicels are given only for the portion above their articulation, as the part below it is too variable. The length of a 2-lobed leaflet is measured from the base to the apex of the lobe. The length of the filaments includes that of the basal, fused, annular part. 1971] OxALiDACEAE (Veldkamp) 155 KEY TO THE SPECIES !. Leaves along a distinct, supraterranean, creeping to erect stem. No bulbs. 2. Shrublet. Petioles phyllodial, leaflets 0-3, often minute. Peduncles abortive. Flowers 1-9 in a fascicle, yellow. Cultivated in Java (§ Heterophylhim) (O. rusciformis Mikan) . . . . O. fruticosa Raddi 2. Herbaceous. Petioles not phyllodial, leaflets not reduced. Peduncle distinct. 3. Leaf pinnately 3-foliolate (rachis developed under terminal leaflet). Leaflets elliptic to oblong, apex not notched. Petals pink with yellow base (§ Thamno.xys) 1, O. barrelieri 3. Leaf digitately 3-foliolate. Leaflets obcordate. Petals yellow (§ Corniculatae). 2. O. corniculata I. Stemless herbs with bulbs or subterranean rhizomes; leaves all basal. 4. Rhizome, no bulbs. Inflorescence 1-flowered. Sepals with 0-1 greenish, round apical callus. Petals white (§ Acetosellae). 5. Stipules conspicuous, much wider than the petiole. Leaflets up to \Va by 1 Vi cm, obcordate; beneath whitish and glaucous and with a pale callus in the notch. Bracteoles 2, free, not vaginate, subglabrous. Sepals pubescent, with 1 pale apical callus 3. O. magellanica 5. Stipules narrow, slightly broader than petiole, not very conspicuous. Leaflets 1-3 by 1 '/4-4 cm, fishtail-shaped; beneath not whitish and glaucous, ecallosc. Bracteole 1, emarginate to bifid, va- ginate, with an apical brown hair-tuft. Sepals subglabrous, ecallose. 4. O. acetosella ssp. grifBthii 4. Bulbs, no rhizome. Inflorescence 2-many-flowered. Sepals with 2-4 orange apical calli. Petals red or purple with greenish to yellowish base. 6. Outer tunics many-nerved, fibrous-withering. Leaflets 4(-6), obdeltoid, not notched, not punctate, at apex sometimes with 2 minute calli. Umbel with many bracts. Filaments crispy-ciliate, the longer with a dorsa! tooth. Sepals with 4 linear calli. Petals red (bluish when dry) (§ Polyoxalis). 5. O. deppei 6. Outer tunics 3-several-nerved, not fibrous. Leaflets 3, obcordate or fishtail-shaped. Sepals with 2 oblong apical calli. Petals purple (bluish when dry) (§ JonoxaHs). 7. Stolons apically bulbiferous. Outer tunics membranaceous, transparent, several-nerved. Leaflets fishtail-shaped, not punctate, notch often with 2 minute calli. Bracts of umbellate inflorescence 2, opposite. All filaments moderately ciliate, the longer with a minute tooth. ... 6. O. latifolia 7. Stolons absent, bulb consisting of many small bulbils; outer tunics papyraceous, brownish, 3-nervcd. Leaflets broadly obcordate, minutely punctate all over the surface, especially along the margins, ecallose. Bracts of the cymose, contracted inflorescence many. Shorter filaments glabrous, the longer ciliate, without a tooth 7. O. corymbosa 1. O.xalis barrelieri Linne, Sp. PI. ed. 2 (1763) 624; RiDL. Fl. Mai. Pen. 1 (1922) 330; Knuth, Pfl. R. Heft 95 (1930) 65; Henderson, Mai. Wild Fl. (1959) 47, fig.; Backer & Bakh. /. Fl. Java 1 (1963) 245. — O. sepium St. Hil. Fl. Bras. Merid. 1 (1825) 89; Progel, Fl. Bras. 12, 2 (1877) 505, incl. var. picia Progel; Knuth, Pfl. R. Heft 95 (1930) 64; Heyne, Nutt. PI. (1927) 151. — Fig. If-g. Erect herb, stem up to I Vi m. without bulbs or stolons. Stem branched, sparsely patent to ap- pressed-rcflexed pubescent, glabrescent; hairs simple, straight or bent below the middle, cglan- dular. Leaves usually more or less opposite, pinnately 3-foliolatc, cxstipulate; petiole 2-9 cm, apprcssed-puberulous; leaflets elliptic to oblong, base cuncate to emarginate, apex obtuse to rounded, not notched, ecallosc, glabrous above, margins strigosc, especially at base, beneath pale and glaucous, sparsely to moderately appressed pubescent, terminal leaflets largest, l-3'/2 by •/2-2'/2 cm. Peduncles 3-5 '/2 cm, pubescent, once or twice dichasially forked; branches up to 4'/2 cm, 4-16-flowcrcd, the bracts opposite the flowers. Bracteoles minute, ciliate, ecallosc. Pedicels I '/2-3 mm, glabrous. Sepals 2-4 by Vj-^Va fnm, ovatc- lanccolatc, acute, glabrous or with a few hairs, ecallosc. 3-ncrvcd. Petals 6-9 by 2-2 'Z: mm. obovutc-lanccolatc, apex rounded, after anthcsis rolling inwards, pink, lower half greenish with yellow spots, glabrous. Filaments (MF) '/^-l and 1 ^4-214 nun, the shorter glabrous, the longer with a dorsal tooth, patent-hairy. Ovary 1 by '/2 mm, glabrous; styles (MF) 1-1 '/2 mm, ascendingly stri- gosc; stigmas small, capitate; ovules 4 per cell, in 1 row. Fruit 5-10 by 3-5 mm, slightly ovoid, 5-angular, apex and base 5-lobed, glabrous; episeptal rimae present. Seeds 3-4 per cell, 1 'Z: by 1 mm, ± flattened-ovoid; testa transversally ridged. Distr. Native of tropical S. America, cul- tivated and established in many places, the oldest collected specimen from Bogor dates from BoERLAGE, a. 1888; in Malesia: Sumatra, Banka, Malaya, Java, Papua (Central Distr.). Ecol. Around gardens, along roads, in hedges, fields, and village groves, along rivers, grassy places with shade, up to 1500 m. Vern. Tjalingtiing, J, bilimbing tanah, Banka, kopomani, Tamil. Uses. The leaves are eaten for their sour taste. Note. Although only the MF form is found in Malesia. fertile seed is formed; there is apparently no functional hctcrostyly. 2. Oxalis corniculata Linn^;, Sp. PI. (1753) 435; Zurc. Abh. Ak. Wiss. Munch. I (1830) 230, incl. var. repens (Thunh.) Zucc; Planch, in Houttc. Fl. Serres 12 (1857) 205. incl. var. alro- purpurea Planch.; Miq. Fl. Ind. Bat. I, 2 (1859) 135; EiMiEW. & Hook./. Fl. Br. Ind. 1 (1874) 436; F.-ViLL. Novis. App. (1880) 32; Guillaumin, 156 Flora Malesiana [ser. I, vol. 7^ Fl. G^n. I.-C. 1 (1911) 610; Back. Schoolfl. Java (191 1) 170, incl. var. javanica (Bl.) Back.; Knuth, Notizbl. Beri.-Dahl. 7 (1911) 300, incl. var. sericea Knuth; Ridl. Fl. Mai. Pen. 1 (1922) 330, incl. var. villosa Hook. /. ex Ridl. [spluilm. ? var. villosa HoHENACKER, cf. YouNG, Watsonia 4 (1958) 57]; Wiegand, Rhodora 27 (1925) 113; Knuth, Pfl. R. Heft 95 (1930) 146; Masamune, Fl. Geogr. Stud. Yakushima (1934) 257, incl. ssp. repens (Thunb.) Masam.; Knuth in Fedde, Rep. 48 (1940) 3, incl. var. papuana Knuth; Hara, Enum. Sperm. Jap. 3 (1954) 8; Eiten, Taxon 4 (1955) 99; Am. Midi. Natur 69 (1963) 257; Back. & Bakh. /. Fl. Java 1 (1963) 246; Veldk., Fl. Thail. 2 (1970) 17. — O. repens Thunb. Diss. Oxai. (1781) 16, fig.; Bl. Bijdr. (1825) 243; B. L. Robinson, J. Bot. 44 (1906) 311; Merr. En. Born. (1921) 311; En. Philip. 2 (1923) 323. — O. javanica Bl. Bijdr. (1825) 243; MiQ. Fl. Ind. Bat. 1, 2 (1859) 135. — O. acetosella (non L.) Blanco, Fl. Filip. (1837) 388. — O. boridiensis Knuth in Fedde, Rep. 48 (1940) 3. Perennial herb, ascending to erect, rooting at the nodes; main root sometimes much thickened and woody; stems several from the main root, branch- ing above the ground, puberulous to pubescent (hairs mainly 1 -celled). Leaves scattered, distant, or in small tufts. Stipules indistinct to small, some- times conspicuous, up to 3 by 1 mm, rectangular. Petiole 1-5V2(-10) cm, appressed to patently pu- berulous. Leaflets broadly to elliptic-obcordate, 4-20(-25) by 5-18(-25)mm, incised up to half- way, ecallose, lobes rounded, rarely obtuse; upper surface glabrous to sericeous; beneath paler, sometimes glaucous, sparsely pubescent to seri- ceous. Peduncles up to 20 cm, usually much shorter, sparsely puberulous to sericeous. Inflores- cence cymose to pseudo-umbellate, l-5(-8)- flowered. Flowers usually MF, rarely LF. Bracts 2-several, subopposite to whorled, ovate-lanceo- late, acute, puberulous, sometimes with septate hairs. Pedicels up to 20 mm, articulate at base and beneath calyx, in fruit straight to sharply bent at the articulations, but the fruit always erect. Sepals lanceolate, obtuse to rarely obliquely retuse with narrow pale margins, 2-6 by Vi-I mm, sparsely puberulous to sericeous, sometimes with septate hairs. Petals spathulate-oblong to -lanceo- late, 3 '/2 -1 by 1 -7 mm, apex rounded to emarginate, after anthesis apically crumpled, yellow, with darker or lighter base. Filaments glabrous, the longer edentate, in MF 1-4 and 3-6 mm, the shorter rarely with abortive anthers, in LF 23/4-31/2 and 31/2-41/2 mm. Ovary 1 1/2-2 by 34-I mm, ellipsoid to cylindric, puberulous; styles in MF 1-4 mm, in LF 3-4 mm, minutely ciliate, sometimes mixed with minute septate hairs; stigmas small, cylindric, sometimes flattened and minutely bifid, papillose; ovules (l-)5-lI per cell, in 1 row. Fruit 9-20(-24) by 2-4 mm, usually linear-cylindric, sometimes ellipsoid, pentagonal, acuminate, minutely puberulous, hairs reflexed or patent to ascending in upper half, mixed with patent, septate hairs; episeptal rimae closed, in- conspicuous; cells inside sparsely to moderately strigose. Seeds (0-)5-l 1 per cell, 1 by % mm, flattened-ovoid; testa with c. 3 regular rows of 7-10 transversally connected rows of ridges. Distr. Cosmopolitan, origin unknown, in Ma- lesia several forms occur; throughout Malesia, three times collected in Celebes and Borneo and scarce in Malaya (Malacca, Perak, Penang; a common weed in Singapore Bot. Gardens). Ecol. In many islands a common weed on all sorts of disturbed soil, in grassfields, gardens estates, along roads and river-banks, on walls, etc., in Java up to 2200, in New Guinea to 3000 m. Vern. Sikap dada, M (Sing.), asim-asim, atim-atiin, lela, Atjeh, daun ascm (ketjil), (djukut) tjalintjing, semanggi, semanggen, J, tjembitjena. Mad., mala mala, Ternate, kuja kawiana, Alor; Philip.: daraisig, Bik., iayo, kungi, malabalugbug- dagis, Pamp., kanapa, Ig., marasiksik. Ilk., pikhik, Iv., salamdgi, Bon., susokoyili, taingangdaga, Tag.; New Guinea: keketi, Tumba, puggepagl, Yoowi, songongom, Wapi, girobi, Musa, Safia, jampijamp, Enga, Yogos, gagari, Eng, Kapilam, kwibant, Maring, kale, Rabaul, tenaquipo, Jimi. Uses. Cf. Heyne, Nutt. PI. (1927) 850. Ac- cording to ViNK (n. 16308) from Uinba, Nona- Minj Divide, Kubor Range, W. Highlands, New Guinea (20-8-1963) used 1) "In marrying cere- mony the young woman takes fresh leaves and makes a gag of it with salt and cold water; the gag is chewed and the juice is spit on pigmeat, which is given to the new husband. 2) When the garden gives a bad production of sweet potato, the woman looking after the garden puts a bundle of the leaves in her girdle to get a higher production." Note. A most complex and variable species. Many infraspecific taxa have been described, but all appear to be linked by intermediate forms. The following extremes can be recognized; 'var. repens Thunb.' Small, decumbent plants. Leaflets rather dark, small, glabrous to moderately strigose. Inflorescences few-flowered, flowers small. Throughout Malesia, a form of exposed, sunny places. 'var. atropurpurea Planch.' Plant brownish to purplish in vivo, dark green when dry; petals more or less flushed or blotched with reddish brown in vivo, fading to whitish or yellowish in drying. Cultivated and escaping. 'var. sericea Knuth' (= 'var. trichocaulon Levl.' according to Hara, 1954; no material of this seen). Large pubescent plant, foliose apices of stems erect or ascending; stipules minute, leaflets strongly pubescent, terminals large, longer than broad, incised for V9-V5 of leaflength, floral parts relatively large; ovules and seeds 1-5 per cell; fruit oblong, stout, pubescence ascending at least in upper half. Eastern New Guinea and New Britain (Formosa? Japan? Korea?). Shaded river- banks, open places in forest, edges of paths and trails, grassfields. 3. Oxalis magellanica Forst. f. Comm. Gott. 9 (1789) 33; Hook./. Fl. Nov. Zel. 1 (1853) 42, fig.; Fl. Tasm. 1 (1860) 59; Benth. Fl. Austr. 1 (1863) 300; Hook./. Handb. N. Zeal. Fl. (1864) 1971 OxALiDACEAE (Vcldkamp) 157 38; Reiche, F1. Chile 1 (1896) 339; Knuth, Pfl. R. Heft 95 (1930) 230. — O. lactea Hook. Comp. Bot. Mag. 1 (1836) 276; Skottsb. The Plant World 18 (1915) 129: Cheeseman, Man. N. Zeal. Fl. ed. 2 (1925) 536; Curtis, Stud. Fl. Tasm. 1 (1956) 96; All.an. Fl. N. Zeal. 1 (1961) 238. — Fig. la-d. Stemless herb with stolons, without bulbs. Rhizome pink to brownish, glabrous with distinct, amplexicaul scale-like remains of leafbases. Stipules much broader than the petiole, con- spicuous, membranaceous, brown, glabrous. Pet- ioles 1-8 '/2 cni. reddish. Leaflets obcordate, 4-13 by 4-12 mm, incised up to \i of the length, lobes rounded; upper surface usually glabrous; beneath pale, glaucous, appressed-strigose gla- brescent, with a — prominent, greenish to brown- ish callus at the notch. Peduncles few, below basal articulation I;-- ^m, above 1-6 '/2 cm, sparsely ap- pressed- to patently pubescent; bracteoles 2, sub- opposite, 2'/2-7 by l-lV2nim, lanceolate, acute, free, very sparsely strigose to glabrous, ecallose, placed in upper */5-*/5th. Buds erect. Flowers solitary, only LF. Sepals elliptic to obovate- oblong, 3-5 by 1 Vi-^l^ mm, acute to truncate and minutely 3-lobed (apex or middle lobe cucullate, sometimes inconspicuous by the hairs, seemingly callose), sparsely to moderately pubescent and somewhat glaucous, margined, slightly shorter to equal to the fruit. Petals white, 8-1 1 '/t by 4-6 mm, spathulate, somewhat oblique, apex rounded to emarginate, glabrous to — ciliate. Filaments (LF) 3-4'/2 and ^Vi-^Vi mm long, glabrous, edentate. Ovary 1-2 by 1-2 mm, glabrous; styles 3-5 mm, glabrous; stigma disk-shaped; ovules 2-5 per cell, in 1 row. Fruit 3-6 by 3-5 mm, glabrous, finally nodding. Seeds 1-3 per cell, 1 Vi by 1 mm, ^ flattened-ovoid, shiny, brownish, smooth to slightly lengthwise furrowed. Distr. Southern Andine S. America, New- Zealand, Tasmania, and the Victorian Alps; in Malesia: New Guinea, on the high mountains (Wilhelm, Otto, Kubor Range, Finisterre Range, Sarawaket Range, Wilhelmina), 2200-3700 m. Ecol. Subalpine to alpine, on shaded, moist to wet, humous soil, among grasses and mosses, sometimes on tree-trunks or rocks. Fl. May-Nov. Vern. Chimbaemagl, Chimbu. Notes. According to Skottsberg I.e. the S. American specimens would differ in having less distinctly obcordate leaflets, peduncles not ex- ceeding 1 cm, and flowers only c. 5 mm across. The New Zealand botanists accordingly call the species in New Zealand O. lactea, but Allan added that some specimens from Chile (in K) approach the New Zealand form. A comparative study of Chilean, .Australasian, and Papuan specimens showed that Skottsberg's differentiation does not hold and that only one species is concerned. 4. Oxalis acetosella Linne, Sp. PI. (1753) 433. ssp. griffithii (Edgew. & Hook. /.) Hara, J. Jap. Bot. 30 (1955) 22; Fl. E. Him. (1966) 168, 638, 661. — O. griffithii Edgew. & Hook. f. Fl. Br. Ind. 1 (1874) 436; Knuth, Pfl. R. Heft 95 Fig. 2. Distribution of Oxalis § Acelosellae. la. O. acetosella L. ssp. acetosella, lb. ditto, ssp. griffithii Hara, Ic. ditto, ssp. monlana HufrtN, 2. O. obtriangulata Maxim., 3. O. magellanica Forsi./., 4. O. oregana Nun. 158 Flora Malesiana [ser. I, vol. 71 (1930) 234; Ohwi, F1. Japan (1965) 580. — Fig. Ih-i. Stemiess herb with stolons, without bulbs. Rhizome dark brown, usually scaley, sparsely pubescent to glabrous. Stipules shorter and ± broader than the basal part of the petiole. Petioles 41/2-22 cm, greenish, red-tinged, pubescent; below articulation flattened, thickened, persistent, final- ly ± woody, brownish when dry, moderately to densely pubescent. Leaflets 1-3 by 1 '/4^ cm, broadly obtriangular, fishtail-shaped, the straight upper edges forming a very wide angle, incised up to halfway, lobes rounded to obtuse; upper sur- face subglabrous to sparsely appressed-strigose; beneath paler, not glaucous and with denser pu- bescence, midrib i thickened in notch. Peduncles 1-2, below basal articulation up to '/t cm, above 5-1 1 cm (shorter in cleistogamic flowers), sub- glabrous to densely pubescent, especially distally; bracteoles zb halfway or higher, partly connate- vaginate, midrib dorsally pubescent, apex with hairy brown tuft, ecallose. Sepals 3-6^4 by 1 '/2-2I/2 rrun, oblong, rounded to emarginate, not cucullate, ecallose, subglabrous, edge closely ciliate. Petals 8-19 by 4—8 mm, spathulate-oblong, truncate to emarginate, often oblique, white, often with lavender veins and yellowish base in vivo. Filaments (LF) V/n-'iVi and 3-6 mm, glabrous, edentate. Ovary 2-3 '/2 by 1 '/^ mm, glabrous; styles (LF) 3-6 mm, glabrous; stigma minute, more or less hook-like, entire; ovules 1-5 per cell. Fruit 6 by 5 mm, ovoid, acute, glabrous, nodding. Seeds 1-2 per cell, 2 V^ by 1 V2 mm, flattened-ovoid, ± smooth to lengthwise ridged, light brown. Distr. From India (Sikkim, Bhutan, Khasia) through China to Japan and Formosa; in Ma- lesia: Philippines (N. Luzon: Mt Pulog), two col- lections (Steiner 1990, March 1961; Jacobs 7361, Jan. 1968). Ecol. Mossy forest ravine, probably 2500- 2900 m. Fl. Jan-March. Fig. 2. Hara (1966) stated that it is found more to the south and at lower altitudes than ssp. acetosella, both in E. Himalaya and in Japan. Notes. Fernandez-Villar (Novis. App. 1880, 32) recorded true O. acetosella L. from a medical garden at Manila. 5. Oxalis deppei Lodd. Bot. Cab. 15 (1828) 1500; Knuth, Pfl. R. Heft 95 (1930) 288; Back. & Bakh. /. Fl. Java 1 (1963) 246; Matthew, Rec. Bot. Surv. India 20 (1969) 56. — O. tetra- phylla (non Cav.) Back. & Sloot. Handb. Thee (1924) 157, t.; Heyne, Nutt. PI. (1927) 851. Stemiess herb from bulbous base, no rhizome, no stolons. Bulb up to 2'/2 by 2 cm, ovoid, acute, with lateral, sessile bulbils; tunics up to 2 14 by 1 '/2 cm, ovate, acute, acuminate or bilobed with terminal leaf, brown, many-nerved, outer fibrous, margins bearded, rest glabrous, inner becoming fleshy. Innovations brownish pilose. Petioles up to 40 cm, glabrous or with a few hairs. Leaflets 4(-6), unequal, 2 V2-6V2 by 1 3/4-61/2 cm, obdeltoid, entire to slightly retuse and ± apiculate (tip often ± folded and occasionally with 1-2 minute calli underneath), glabrous to sparsely pubescent, usually with dark V-shaped marking in lower third, epunctate. Peduncles up to 50 cm, ± glabrous. Inflorescence pseudo-umbellate, 6-25-flowered. Bracts many, up to 21/2 rrmi, ovate, acute to acu- minate, glabrous or with strigose margin, ecallose or with minute apical calli. Pedicels up to 3 cm, glabrous. Sepals 5-7(-IO) by 11/2-3 mm, elliptic to lanceolate, obtuse to emarginate, glabrous, ciliate on edge, c. 7-nerved, apical calli 2-6, usually 4, linear, inconspicuous, orange. Petals 11/2-2 1/2 by V^-l Va cm, spathulate-oblong, rounded to retuse, often oblique, glabrous, dark red, when dry often bluish with greenish base. Filaments (SF) 31/2^1/2 and6-6i/2mm, (LF) 6 and 10 mm, the shorter with a few, the longer with many patent, crisped cilia, dentate. Ovary and stamina! tube c. Y2 mm stipitate. Ovary 1 1/2-5 by 1-3 mm, ellipsoid, glabrous; styles (SF) 1 mm long, (LF) c. 15 mm, glabrous, dark when dry; stigmas cy- lindric, ± bilobed, later ± peltate, not papillose; ovules 3-5 per cell, in 1-2 rows. Distr. Native of Mexico, cultivated and escaping; in Malesia hitherto only found in Java (Tjinjiruan); introduced before 1911, now quasi- spontaneous. Ecol. Locally naturalized in mountain Cin- chona estates, c. 1 600 m, sometimes gregarious, propagating by bulbils; fruit unknown from Java. Apparently functionally heterostylous. In Malesia only the SF has been found so far. Dimensions of the LF have been taken from Matthew, I.e. Vern. Tjalingtjing badak, tj. gede, S. Uses. Ornamental plant and for ground-cover, but difficult to eradicate; leaves eaten as vegetable (Heyne, 1927). Note. O. tetraphylla Cav., with which this species has often been confused, has bulbiferous stolons, emarginate, not apiculate leaflets, and smaller flowers. 6. Oxalis latifolia H. B. K. Nov. Gen. Sp. 5 (1821) 184, t. 467; Knuth, Pfl. R. Heft 95 (1930) 273; Heyne, Nutt. PI. (1927) 851; Symon, Trans. R. Soc. S. Austr. 84 (1961) 75; Young, Watsonia 4 (1958) 63. — O. intermedia {non A. Rich. ?) Back. & Bakh./. Fl. Java 1 (1963) 246. Stemiess herb from bulbous base, no rootstock; bulb ovoid, up to 5 by 2 cm, acute, with numerous, basal, n= erect stolons, with a few small scales, ending in ovoid, acute bulbils pale brown when dry; tunics many, outer up to 5 by 2 cm, ovate, acuminate or with terminal leaf, membranous, transparent, white, nerves 3 to several, orange; inner becoming fleshy. Petiole up to 20 cm, ± glabrous. Leaflets ± equal, 1 1/2-71/2 by 2-8/2 cm, broadly obdeltoid, fishtail-shaped, incised up to halfway, glabrous, beneath subglaucous, often with 2 orange calli in notch, epunctate. Peduncles up to 25 cm, slightly hairy. Inflorescence umbellate, 5-13-flowered. Bracts 2, c. 1/2 mm, ovate, margin glabrous to strigose, minutely callose or ecallose. Pedicels up to 2 cm, glabrous, filiform. Sepals 4-4i4(-6) by li/2-2(-3) mm, oblong, obtuse, glabrous, indistinctly 3-5-nerved; apical calli 1971] OxALiDACEAE (Vcldkamp) 159 2, orange, not confluent, c. 1 mm long, minutely hastate. Petals 10-20 by 3-6(-8) mm, narrow- ly obtriangular, truncate, — oblique, red-purplish with greenish base, crumpled after anthesis. Filaments (SF) 2 1/2-3 M) and 4-5 (-6) mm, puberulous, the longer with a minute tooth. Ovary and staminal tube c. '/: rnni stipitate. Ovary 1 >/^ by -/4 mm, apically sparsely ciliate on the ribs; styles (SF) c. 1 mm, sparsely ciliate; stigma peltate, c. V4 mm 0, not papillose; ovules 3-6 per cell, in 1-2 rows. Distr. Native of Central and tropical S. Ameri- ca; cultivated and escaping, e.g. in Malesia: Java (Preanger Mts; W. Java: Gedeh, Lembang, Tjinjiruan). Ecol. In gardens, fields, and estates, 1250-1550 m. No fruits are recorded from Java, the plant is apparently functionally heterostylous. It is difficult to eradicate because of the many bulbils. Vern. Tjalingtjing, S. Note. O. intermedia A. Rich. (Ess. F1. Cuba, 1842, 315) is said to be more pubescent and to have edentate, longer filaments. As the teeth are minute, however, they might easily be overlooked. 7. Oxalis corymbosa DC. Prod. 1 (1824) 696; RiDL. Fl. Mai. Pen. 1 (1922) 330; Back. & Sloot. Handb. Thee (1924) 155, t.; Heyne, Nutt. PI. (1927) 851; Symon, Trans. R. Soc. S. Austr. 84 (1961) 74; Back. & Bakh./. Fl. Java 1 (1963) 246. — O. martiana Zucc. Denkschr. K. Ak. Wiss. Munch. 9 (1824) 144; Back. Schoolfl. Java (1911) 170; Merr. En. Philip. 2 (1923) 323; Knuth, Pfl. R. Heft 95 (1930) 250. — O. violacea (non L.) Hall. /. Med. Rijksherb. Leiden 12 (1912) 19. — Fig. le. Stemless herb from bulbous base, no rhizome, no stolons; bulb globose, c. 1 cm o; bulbils many, clustered, globular to ovoid, acute; outer tunics brown, papyraceous, up to 20 by 6 mm, ovate to oblong, acuminate or with terminal leaf, margins glabrous to long-bearded, distinctly 3-nerved; inner pale and fleshy. Petiole up to 30 cm, patently villose. Leaflets — equal, 1 Vi-^Vi by P/4-5'/2 cm. broadly obcordate, incised for ^/j, lobes rounded, sometimes i overlapping; upper surface sub- glabrous, beneath appresscd-puberulous; minutely orange-punctate all over the surface, especially along the margins. Peduncle up to 35 cm, hairy in various degree. Inflorescence often up to twice forked; pseudo-umbels 2-12-flowered. Bracts many, c. 1 mm long, elliptic, rounded, pale, with strigose margin and 0-3 orange, linear calli in the middle. Pedicels up to 2'/2 cm, ascendingly ap- prcsscd-strigosc. Sepals 3'/2-5(-6) by 1-2 mm, oblong, acute, apex minutely bifid to the 1-3 or- bicular to oblong, with 2 orange, apical calli, ;- puberulous, 3-5-ncrvcd. Petals 1 1-20 by 4-7 mm, spathulatc-oblong to -lanceolate, obtuse to trun- cate, often oblique, light reddish purple with darker veins, yellowish at base Filaments (MF) 2-3 and 5-6 mm, (SF) 3' 2^ and 5-6 mm; the longer dorsally ciliate, edentate. Ovary and staminal tube c. -^4 mm stipitate. Ovary 2 by 1 mm, abundantly ascendingly ciliate to glabrous; styles in MF 1 5/4-2 mm., in SF 1-1 W mm long, abundant- ly ciliate; stigma bilobed, papillose, c. 1 mm 0; ovules 3-8 per cell, in 1-2 rows. Fruit not seen. Distr. Native in tropical S. America, natural- ized in many parts of the World; in Malesia cul- tivated and escaping in Java (introduced from Sydney before 1848), W. Sumatra, Malaya, and Philippines (Luzon). Ecol. In fields, road-sides, and estates, often as a gregarious weed, 400-1450 m. No seed is set; propagation is by the many bulbils, which make it a difficult plant to eradicate. Vern. Tjalingtjing beureum, tj. gede, tj. tSgal, S, kembang gelas, J, asam-puja, Padang. Uses. Occasionally cultivated; the leaves are sometimes used as a substitute for tamarind (Heyne). Terat. No fruits have been recorded, although two stylar forms occur, possibly in the same populations, but certainly in the same environ- ment. The reason for inability of fruit-setting probably lies in the common occurrence of mon- struous flowers. Sometimes the filaments broaden and form petaloid structures, occasionally still with the anthers present in reduced state. More often there is a transition of the anthers towards carpel-like structures. In the least monstruous forms the con- nective is flabelliform with patent outer walls of the anthers. The next step is an apical elongation and even more developed anther-cells. The elon- gation may bear a stigma-like structure. Especially when there is an extra filamentous whorl between the inner (longer) filaments and the pistil extreme cases occur. These 'filaments' may finally resemble stipitate, free carpels with a full grown style and papillose stigma; they are apparently open on the introrse side. Ovules have not been seen, but may be present as they are very small and transparent even in the normal plant. It is possible that these 'filaments' fuse with the pistil, as occasionally more than 5-merous pistils carrying ovules are observed with the stipitate alongside. Thus there seem to be transitional stages between the anthers and pistil! Once a sepal was observed to arise from the cen- tre of a deformed pistil. Notes. O. articulata Savigny (in Lamarck, Encycl. Bot. 4, 1798, 686) from Argentina, is often confused with this species. O. articulata diff"ers in the presence of a tuberous rhizome, by the deep- er obcordate leaflets that have large spots, and the denser pubescence. O. violacea L. (Sp. PI. 1753, 434) from N. America, has cmarginate, obdeltoid leaflets, epunctate with two brown calli at the notch. 2. BIOPHYTUM DC. Prod. 1 (1824) 689; Edgf.w. & Hook./ Fl. Br. Ind. 1 (1874) 436; Knuth, Pn. R. Hcfl 95 (1930) 391; Steen. Bull. Jard. Bot. Bl/g 111, 18 (1950) 449. — 160 Flora Malesiana [ser. I, vol. 7^ Oxalis sect. Biophvtum Endl. Gen. PI. (1839) 1172; Miq. F1. Ind. Bat. 1, 2 (1859) 134; Progel, FJ. Bras. 12, 2 (1877) 482. — Fig. 3-4. Erect annual herbs or usually sympodially branched dwarf shrubs. Leaves paripinnate in tufts at the end of the stem or branches, with setaceous stipules; leaflets opposite, subsessile, terminal pair mostly different from the others, their acroscopical half of base cuneate, basiscopical half rounded to truncate, rachis prolonged in a mucro. Flowers terminal in a usually peduncled, bracteate pseudo- umbel, heterodi-, tri- or homostylous. Pedicels articulate at base. Sepals glabrous inside, i free. Petals contort, glabrous, coherent above the claw. Filaments: shorter ones with a more or less pronounced callus at base, longer ones always edentate. Styles in LF and MF ciliate; stigmas terminal, subulate and entire or spoon-shaped to flattened, crenate to bifid; ovules 3-6 per ceH, alternating in 2 rows. Capsule finally loculicid to the base, forming a 5-rayed star. Seeds 1-6 per cell, aril white, thin, at maturity bivalved and ejaculatory. Distr. Pantropical, possibly some 70 spp., in Malesia 7 spp. among which one escaped and another introduced. Ecol. B. dendroides is obviously escaped from the Bogor Botanic Garden and became naturalized in the immediate vicinity of Bogor. B. reinwardtii and B. sensitivuin are weeds in anthropogenic places; it is doubtful whether the former is indigenous. The three ligneous species, B. adiantoides, B. fruticosum and B. microphylliim are forest dwellers. B. petersianum is a widely spread annual which prefers places subject to a pronounced dry season, as also shown by its absence from the great ombrogenous forest belt in Malaya, Borneo and Sumatra and scarcity in West Java (some local spots N of Bandung). For heterostyly see under the family. Germination is insufficiently known and seems variable. Shetty (Proc. Symp. Recent Adv. Trop. Ecol. Varanasi, 1968, 213-224) found that in B. sensitiviim (a name not checked by me through voucher specimens) seed collected at Varanasi (Benaras, at c. 25° NL) at the beginning of the (dry) winterseason failed to germinate 'under any condition' for the first 8V2-9 months and showed hereby to possess a dormancy period. Germination capacity decreased again considerably after 11-12 months. During the first month of the germination period it needed thorough washing by which Shetty concluded to the presence of a water-soluble inhibitor; besides optimal temperature needed for germination was found to be 30-40° C. Whether this behaviour holds also for Malesia is unknown; it is common in regions with- out a dry or cooler season. I have made germination tests with B. dendroides in the Leyden Hortus where the seeds germinated in a very short time. Morph. I have raised seedlings of 5. dendroides of which the germination is epigaic. The first leaves, which bear successively 2, 4, 6, and more leaflets, originate from the plumule which remains between the cotyledons. Simultaneously the hypocotyl starts to lengthen, forming the initial stem carrying the leaf tuft at its apex. When this cycadoid first stage is formed, one finds below the tuft usually a collar of reflexed bristles below the leaf-scars of fallen leaves. In the annual species, characterized by a pithy (compressible) hypocotylar stem, there is usually one apical tuft, exceptionally there are two, both ses- sile. In the perennial ligneous species other tufts are formed, either on short-shoots or by genuine lateral distinctly sympodial branching. The sympodial branching of the suff"ruticose species is very characteristic; it is partly by thick short- shoots on which the cortex is covered by leaf-scars, withering with age, and partly by long-shoots which carry always under the pseudo-umbel of leaves a reflexed tuft of hairs, the shoot itself showing no leaf- scars. In the tuft are many, often finally reflexed narrow to needle-like stipules. The peduncle of the in- florescence resembles the 'internode' of the long-shoots, but it carries on top a fascicle of narrow cuspidate bracts, the inner ones of which bear flowers. In B. reinwardtii and B. adiantoides it may happen that in the pseudo-umbel some reduced leaves occur. This can be found also in B. fruticosum (Kaudern 438); of this species a still more remarkable specimen has been collected at Sse Mao (PI. Yunnan & Mekong), Prince H. d'ORLEANS (in P) where the pseudo-umbel has produced a new umbel with reduced leaves and flowers, exactly similar to a normal long-shoot. This also occasionally occurs in B. reinwardtii (Rant s.n., 1922, do, 1924, Beumee 4848, KooRDERS 29864). A remarkable case of proliferation difficult to account for. ^Callus'' of the shorter filaments. At the base of the annulus below the shorter (epipetalous) filaments there is a usually dark coloured tumidity, sometimes cup-shaped, in front of the petals. Similar ones have already been noticed in Hypseocharis by Baillon (Adansonia 10, 1873, 363; liist. PI. 5, 1874, 26, 41, f. 56, 57) and in some species of Oxalis by Baillon (1874) and Progel (F1. Bras. 12, 2, 1877, t. Ill and 113, 6). Hallier /. discussed their taxonomical value as similar structures occur in Geraniaceae, 1971] OxALiDACEAE (Veldkamp) 161 Limnanthaceae and Linaceae (Beih. Bot. Centralbl. 39, ii, 1921, 165). Narayana (in lift.) found they have no vascular trace in B. candoUeanum (called 'fi. intermediurn, in J. Jap. Bot. 41, 1966, 321). The callosities are possibly homologous with the scales found in Dapania on the annulus at the base of the shorter filaments (Veldkamp, Blumea 15, 1967, 523). The indument is made up of setaceous, simple hairs. In the inflorescences small, septate, capitate, glandular hairs and/or minute, red, club-shaped glands may be present. Syst. I agree with Van Steenis that the subdivision of the genus as given by Knuth is unsatisfactory and new criteria must be considered. The first provisionally proposed sect. Sensitiva (with the generic type species B. sensitivum) for annual species and sect. Prolifera (type species B. proliferum) for the suf- fructicose species. But the situation might be more complicated and a subdivision has to wait for a complete revision of the genus. KEY TO THE SPECIES 1. Leaves 3-9-jugate; leaflets orbicular to elliptic; veins ± perpendicular to the midribj few. Flowers sessile in centre of tuft, (in Mai.) rarely on a peduncle up to 1 Vi cm. Unbranched annual. 1. B. petersianum 1. Leaves 7-32-jugate; leaflets elliptic to lanceolate; veins oblique to the midrib, many. Flowers in distinctly peduncled pseudo-umbels. 2. Midrib of leaflets (not terminal pair) at base i median. 3. Bracts of pseudo-umbel bushy, setaceous, 3-6 mm. Calyx with long, simple hairs. Perennial, finally (always?) branched; stem woody, not compressible, old parts reflexed. Petals ±: equalling the sepals. Sepals 1 Vz-l times as long as the fruit 2. B. dendroides 3. Bracts of pseudo-umbel ovate-attenuate, 1-3 mm. Calyx puberulous and with septate-glandular hairs. Annual, never branched; stem soft, pithy, compressible, old parts caducous. 4. Sepals — equalling the petals, in fruit 4—7 mm long, longer than the pedicels, 1 Vi-l times as long as the fruit 3. B. sensitivum 4. Sepals zr half as long as the petals, in fruit 2'/i-4 mm long, shorter than the pedicels, ± as long as the fruit 4. B. reinwardtii 2. Midrib of leaflets (not terminal pair) at base excentric, in basiscopic corner. Perennials, ligneous, at least finally branched. 5. Leaves 14-32-jugate. Terminal leaflets 3-6 '/2 by 1 '/2-3 mm, obovate-oblong, as long as the pre- ceding. Sepals in fruit 3-7-nerved, 1-2 times as long as the fruit. Fruit puberulous in upper half. 5. B. microphyllum 5. Leaves 18-27-jugate. Terminal leaflets 9-22 by 3-8 mm, oblanceolate, as long as the preceding. Sepals in fruit 5-12-nerved, l!4-2 times as long as the fruit. Fruit glabrous. 6. B. adiantoides 5. Leaves 7-17-iugate. Terminal leaflets 5-20 by 3-8 mm, obovate to obovate-oblong, and as long as the preceding (rarely elliptic-oblong and longer than the preceding: var. papuanum). Sepals in fruit 3-6-nerved, slightly exceeding the fruit or i equalling it {yar. papuanum). Fruit apical ly puberulous 7. B. fruticosum 1. Biophytum petersianum Klotzsch in Peters, obovate, i oblique, midrib excentric; other Reisc .Mossamb. Bot. 1 (1862) 81, t. 15; Steen. leaflets triangular to orbicular-elliptic, midrib ± Bull. Jard. Bot. Btzg III, 18 (1950) 452; Back. & median; apex rounded to obtuse, glabrous or with Baku./. FI. Java 1 (1963)246; Exell, FI. Zamb. sparsely ciliate margin; nerves few, ± perpen- 2 (1863) 158; Veldk. FI. Thail. 2 (1970) 18. — dicular to midrib, conspicuous. /'<'cllowish appresscd to patently pubescent; leaflets rarely overlapping, terminals 9-22 by 3-8 mm, oblong to lanceolate, widest at or above the middle, • as long as the preceding, midrib median; others asymmetric, elliptic to oblong, basiscopical half of base rounded, acroscopical half truncate, more or less drawn out, margins otherwise ::: parallel, midrib excentric at base; apex central, rounded, apiculate; above sparsely strigose, hairs often in three rows, to glabrcscent, beneath more strigose; nerves many. Peduncle 5-191/2 cm, up to 9-flowered, puberulous, some- times with a few septate-glandular hairs. Pedicels 5-17 mm, puberulous and with a few gland hairs. Sepals 4'/2-6 by \-\Y2mm, lanceolate, acute, sparsely strigose to glabrous, sometimes with a few septate gland hairs, Y2--A as long as the corolla, in fruit 5-12-nervcd, half to r equally as long as the pedicel, l'/4-2 times as long as the fruit. Petals 9-10 by l-2'/2 mm, lanceolate, apex rounded to truncate, white with yellowish base. filaments in SF l'/2-3'/2 and 4'/2-6'/2 mm long, in MF 1 '/2 anil SVi mm long, in LF 1-1 '/2 and 2-V4-4 mm long, the longer ciliate. Ovary I by Vz-V* nim, glabrous; styles puberulous, in SF '/2-1 1/4 mm, in MF 2'/2 mm. in LF 4-6 mm long; stigma flattened, crenate to bifid; ovules 2-4 per cell. Fruit 3-4 by 2-3 mm, glabrous. Seeds 2-3 per cell, 1-1(4 by '/i-1 mm, with transverse tuberculated ridges. Distr. S. Vietnam, Cambodia. Thailand, Tenasserim, and in Malvsia: northern half of the Malay Peninsula (Pcrlis, Perak, Pahang). Fig. 5. Ecol. In crevices of (limestone) rocks 166 Flora Malesiana [ser. I, vol. 71 along rivers and in open woods, up to 300 m. Vern. Daun pajong, maiong, pajong ali, M. Uses. Given to small children against stomach trouble in Perak. 7. Biophytum fruticosum Bl. Bijdr. (1825) 242; Knuth, Pfl. R. Heft 95 (1930) 412 in annot. sub B. nudum; Steen. Bull. Jard. Bot. Btzg III, 18 (1950) 454; Reinwardtia 1 (1952) 477, p.p. — B. esquirolii Lev. in Fedde, Rep. 12 (1913) 181; Knuth, Pfl. R. Heft 95 (1930) 413; Merr. Lingn.Sc. J. 13 (1934) 31 ; Rehder, J. Arn. Arb. 18 (1937) 209. — B. thorelianum var. sinensis Guil- LAUMiN, Not. Syst. 1 (1909) 25; Bull. Mus. Hist. Nat. Paris 15 (1909) 128; Knuth, Pfl. R. Heft 95 (1930) 413. — Oxalis blumei Zucc. Abh. K. Ak. Wiss. Munch. 1 (1830) 276; MiQ. Fl. Ind. Bat. 1, 2 (1859) 134. var. fruticosum. Perennial. Stem woody, branched, up to 50 cm. Leaves 7-17-jugate; rachis 3-9 cm, appressed to patently strigose, especially on the upper surface; leaflets rarely overlapping; terminals 5-12 by 3-8 mm, i as long as the preceding, obtriangular to obovate-oblong, ± symmetric, midrib median; other leaflets elliptic to lanceolate, basiscopical half of base rounded, acroscopical half truncate, more or less drawn out, midvein excentric; apex rounded, apiculate, central; above densely strigose, glabrescent, then hairs often in three rows, lower surface less strigose; nerves many. Peduncle up to 13 cm, strigose, no septate glandular hairs, up to 10-flowered. Pedicels up to 9 mm. Sepals 31/2-6 by 1-11/2 rnm. ovate-lanceolate, densely to sparsely strigose, no septate glandular hairs, in fruit 3-7-nerved, ^/g as long as the pedicel, slightly exceeding the fruit, c. 2/3 as long as the corolla. Petals 5I/2-IO by 1-21/2 mm, lanceolate, obtuse, white to pink. Filaments in SF 2-3 and 3 1/2-4 mm, in MF 2-21/2 and 3 mm, in LF 1 1/2-4 and 21/2-5 mm long, the longer always ciliate. Ovary i/^-l by 1/2-I mm, apically with a few cilia to glabrous; styles in SF 1/2 mm. stigma bifid, flattened; in MF 21/^-4 mm, in LF 2-5 mm, stigma subulate to slightly flattened, entire; ovules 2-4 per cell. Fruit 21/2-4 by 21/2-3 mm, puberulous but eglan- dular in upper half. Seeds 2-4 per cell, 1 by Y^ mm, transversely tuberculate and ridged. Distr. SW. China (Yunnan, Hupeh, Kouy- Tschiou, Canton, Hainan), Indo-China (Mekong), NE. India (Silhet); in Malesia: Philippines (Bohol; Sulu Is.: Bengao I.), E. Celebes, and Lesser Sunda Islands (S. Central Timor; S. Wetar). Fig. 5. Ecol. In crevices of rocks, on river-banks, in thickets and shaded cultivated areas on sand, up to 800 m. var. papuanum, var. nov. — Oxalis albiflora F. v. M. Vict. Natur. 8 (1892) 164, nom. nud. — B. albi- florum F. v. M. Vict. Natur. 9 (1893) 112; J. Bot. 31 (1893) 325; Knuth, Pfl. R. Heft 95 (1930) 397; Steen. Bull. Jard. Bot. Btzg III, 18 (1950) 455. — Oxalis papuana F. v. M. Vict. Natur. 9 (1893) 1 12, nom. altern. Terminal leaflets 14-20 by 6-8 mm, elliptic to oblong, ± 1.3 times as long as the preceding. Pedicels 9(-25) mm. Sepals in fruit ± ^/j as long as the pedicel. Fruit ± as long as the calyx. Distr. Malesia: SE. New Guinea (Owen Stanley Range), two collections (type not seen). Fig. 5. Ecol. Stony stream-banks in forests. Note. In his preliminary revision Van Steenis accepted a much wider specific concept of B. fruticosum than handled here, in merging B. adiantoides with B. fruticosum and also referring the specimens here distinguished as B. micro- phyllum to it. In fact all three native Malesian fruticose Biophytums and B. thorelianum Guillau- MIN from Indo-China (fig. 5) differ from allied continental species {B. nudum, B. proliferum, B. intermedium, and B. polyphyllum) in having the midrib obliquely inserted at the base and the Asian ones having them median. However, the Malesian material can be sorted into three taxa, although it must be admitted that the great scarcity of collections makes it difficult to find the parameters of their variability. 3. DAPANIA KoRTH. Ned. Kruidk. Arch. 3 (1854) 381; Hall./. Beih. Bot. Centralbl. 34, ii (1917) 25; Knuth, Pfl. R. Heft 95 (1930) 419; Veldkamp, Blumea 15 (1967) 523. - Fig. 7i. Glabrous lianas. Leaves estipulate, unifoliolate; petioles articulate with a constriction (in herb.). Inflorescences racemose, ramiflorous and axillary, solitary to fascicled. Flowers (in Mai.) androdioecious. Sepals connate in lower half, margins ciliate, glabrous inside. Petals apotact or paratact, rarely quincuncial, (in Mai.) free and glabrous inside, minutely clawed, red to white. Filaments (in Mai.) antheriferous, between the filaments with scales on the annulus, these sometimes reduced to dark lines. Ovary glabrous, reduced and sterile in o plants. Ovules in ^ 1-2 per cell, in S 0-1. Capsule fleshy, yellowish green (red when dry), 1971] OxALiDACEAE (VeldkaiTip) 167 loculicid to the base, 1-6-seeded, valves patent; (in Mai.) episeptal rimae present, open to base. Seeds up to 2 per cell; testa smooth, hard, yellowish red, terminally thickened around the micropyle, splitting lengthwise under pressure; aril present, enveloping the seed, attached to the entire length of the adaxial raphe, fleshy, bright to whitish yellow, margin irregular crenate, with oily drops; embryo with the cotyledon 1 Vi-l times as long as the straight, oblique radicle. Distr. Madagascar (1 sp.), in Malesia (Sumatra, Malaya, Borneo) 2 spp. Ecol. Lianas in forests, swamps, near rivers, obviously usually on poor soil, at low altitude. KEY TO THE SPECIES 1. Leaves up to 15 cm, base cuneate to rounded. Sepals glabrous, except the ciliate margin. 1. D. racemosa 1. Leaves longer than 15 cm, base deeply emarginate. Sepals puberulous outside. 2. D. grandirolia 1. Dapania racemosa Korth. Ned. Kruidk. Arch. 3 (1854) 381; Planchon, Ann. Sc. Nat. Bot. IV, 2 (1854) 266; MiQ. Fl. Ind. Bat. 1, 2 (1859) 134; Staff in Hook./. Ic. PI. Ill, 10 (1891) t. 1997; Hall. /. Beih. Bot. Centralbl. 34, ii (1917) 25; Knlth, Pfl. R. Heft 95 (1930) 420; Veldkamp, Blumea 15 (1967) 552, f. Id-g. — D. scandens Staff in Hook./. Ic. PI. Ill, 10 (1891) t. 1997; King, J. As. Soc. Beng. 62, ii (1893) 201; Hall. / Beih. Bot. Centralbl . 34, ii (1917) 25; Ridl. Fl. Mai. Pen. 1 (1922) 334; Knuth, Pfl. R. Heft 95 (1930)420, f.28a-e. Large liana, up to 30 m long, 20 cm 0, rather profusely branched. Leaves 5^/4-1 5(-25) by 2'/2-^^/i cm, oblong to lanceolate, pergamenta- ceous to subcoriaceous, acute to cuspidate, base cuneate to rounded; petiole 3-10 by 1-2 mm, petiolule 2-6 by 1-2 mm. Racemes solitary to fascicled, in ^ 1-3 together, 3-1 3 '/2 cni, in S 1-15 together, 2-5 1/2 (-1 2 Vi ) cm; rachis puberulous, glabrescent. Pedicel (lower joint 0-1 mm, upper '/4-'/2 mm) not covered by the minute, broadly ovate, acute bract V*~V2 by V2 mm. Calyx 1 '/2-2 mm high, glabrous to sparsely puberulous out- side. Sepals Vi-^Vi hy '4-l'/4 mm, broadly ovate to elliptic, rounded to emarginate. Petals 3-4 by l-lUnim, obovate-oblong to -lanceolate, often rolled back at anthesis, obtuse to rounded, darker. Filaments in 6 V^-Vi and ^/i-l mm, in ^S 1-3 and l'/i-4mm long. Pistil in '^ l'/2-2'/2mm, in S Vi-Y* mm; styles in ^ 1-1 V4 mm, in ^ very short, acute. Fruit obovoid with cuneate base, 5-1 1 by 6-10 mm, after dehiscence 9-22 mm 0. Seeds 4-5 by 1 '/: mm; radicle 1 '/4-P/i mm long, cotyledons 2-2 '/2 by Va-1 mm. Distr. Malesia: Sumatra, Malaya, Borneo. Fig. 6. Ecol. Dense jungle, swamp forest, on poor soils, up to 750 m. Fl. fr. Jan. -Dec. Vcrn. Batieg-hatieff, dapan, Sumatra; hiUimhing bail, M, jaloai, Brunei, kara-raka, I ban; pau- kianfi, Malacca, sclamhun akar, Sclangor. Note. A large-leaved form occurs in North Sumatra. 2. Dapania grandifolia Veldkami', Hlumca 15 (1967) 525. f. Ih-p. — Fig. 71. Fig. 6. Distribution of Dapania racemosa Korth. (#) and D. grandifolia Veldk. (A). Liana, or treelet, up to 27 m, 7 cm 0. Leaves coriaceous, (10'/^-)16'/2-31 by 4'/2-Il cm, oblong to lanceolate, widest at or below the middle, base retuse to deeply emarginate, apex acute to at- tenuate; petiole 2-6 by 2-3 '/2 mm, stout, petiolule 1-3 by 2-3 1/2 mm. Racemes -_■- densely and — patently puberulous, not glabrescent, in rj 1-5 fascicled, 3-5 cm long, in j 1-2 together, 4-8 cm long. Bracts boat-shaped, sometimes recurved, V4-I by '/2-I Tim. broadly ovate, acute, covering the minute (c. V2 mm long) pedicel, but not the base of the calyx. Calyx 1^4-2 '/2 mm high, pale to brown puberulous. Sepals 1-1 '/2 by ^/4-l Vi mm, suborbicular to ovate, acute to round- ed. Petals 4-5 by '/2-I t^ mm, obovatc-lanceolate, rounded, base attenuate. Filaments in S 2-3 '/i and 3-4 mm, in -^ V2-V* and J^-I mm. Pistil in cJ reduced, sterile, Vi-V* by V*-V2 mm, ovoid; styles minute, acute; $: ovary P4 by 1mm, styles 2 mm. Fruit before dehiscing 9-1 1 by 6-8 mm, suborbicular to ovoid, after dehiscence 20-22 mm 0. Seeds AVj-^ by 1 '/2-I -^ mm; embryo 4 mm long, radicle I '/i by '/s mm, cotyledons 2'/2 by 1 '/i mm. Distr. Malesia: NE. to SE. Borneo. Fig. 6. Ecol. Primary forest, on flat to undulating land on clay near rivers. Fl.fr. Jan. -Dec. Vcrn. Salung kapll, Dusun. 168 Flora Malesiana [ser. I, vol. 7^ 4. SARCOTHECA Blume, Mus. Bot. Lugd. Bat. 1 (1850) 241; Hall./. Med. Rijksherb. Leiden 1 (1911) 1; Knuth, Pfl. R. Heft 95 (1930) 420; Veldkamp, Blumea 15 (1967) 527 — Roucheria MiQ. Fl. Ind. Bat. 1,2 (1859) 136. — Connaropsis Planch, ex Hook. /. Trans. Linn. Soc. 23 (1860) 166. — Fig. 7a-h. Fig. 7. Sarcotheca glauca Hall./, a. Habit, X2/3, b. flower, x4, c. petal, x6, d. stamens and ovary (SF), xlO, e. ovary (SF), xlO, / stamens and ovary (LF), x 10, g. ovary (LF), XlO, h. fruit, x2. — Dapania grandifoUa Veldk. /. Annulus with scales and filaments, X 10 {a, h Puasa BNB 5484, b-c, f-g Lajangah SAN 33612, d-e Mujin SAN 33562, / Kostermans 10104). Shrubs or trees; innovations pubescent. Leaves estipulate, 1- or 3-foliolate; petioles articulate, with a constriction (in herb). Panicles axillary or pseudoter- minal, one to few together; flowers in more or less stalked cymes, scattered along a simple or sparsely branched rachis; cymes subtended by small caducous bracts, occasionally by a reduced petiole (petioloid), or rarely a small leaf. Flowers heterodistylous. Sepals unequal, shortly connate at base, inside appressed strigose, persistent (except in S. diversifolia). Petals contorted, sometimes paratact, inside with minute papillae in the upper half. Filajnental annulus without scales. Ovules 1971] OxALiDACEAE (Veldkamp) 169 2 per cell. Fruit fleshy, red at least when dry, with i distinct episeptal rimae sometimes lighter and minutely papillose inside (at least when dry). Seeds exaril- late; testa smooth to transversely rugose, hard, reddish, splitting irregularly by pressure; embryo with a straight radicle in line with the cotyledons which are 3-5 times as long as the radicle. Dislr. W. Malesia: Sumatra, Malaya, Borneo, Celebes. Fig. 8. Ecol. Primary and secondary forest on poor soil at low altitude. Fl.fr. Jan.-Dec. Uses. The fruit, although acid, is eaten in curry, sajur, and manisan and is said to be a remedy against coughing. The timber is light and of small dimension, neither very strong nor durable, sometimes used for roofs. Notes. The genus has often been confused with Rourea Aubl. (Connaraceae) which differs from Sarcotheca in having free carpels, 2 collateral ovules, seeds with an aril, and a dry, indehiscent, 1-celled and 1 -seeded fruit. The species of the genus are closely related and most of them occupy small ranges; in several cases close allies show replacing areas. With a broader species concept several species would be reduced to sub- specific rank, notably the triad 5. monophylla from Malaya, S. glauca from Borneo, and S. celebica from Celebes. KEY TO THE SPECIES 1. Leaves trifoliolate, laterals sometimes caducous, leaving a scar. Mature fruit greenish yellow when fresh (red when dry!); more than 1 'A cm long. 2. Calyx 3-5 mm high, outside pale puberulous to glabrous, not persistent in fruit. 1. S. diversifolia 2. Calyx 2 '/2-3 mm high, outside brown puberulous, persistent in fruit 2. S. griffithii 1. Leaves unifoliolate; no lateral scars. Mature fruit red when fresh; less than 1 '/2 citi long. 3. Mature leaf puberulous to pubescent beneath, also between the nerves. 4. Panicles not or barely exceeding the leaves. Petals 4-7 mm long. Rimae obscure, glabrous inside. 5. Leaves 8-23 by 3-8 '/2 cm, margins never paler. Calyx 1 '4-2 mm high 4. S. laxa 5. Leaves 5-11 ',4 by 2-3 '/2cm, margins usually paler. Calyx 2-3% mm high. . . 8. S. ferruginea 4. Panicles exceeding the leaves. Calyx 3-3|/2mm high. Petals 6-8 mm long. Rimae conspicuous, lighter and minutely papillose inside 10. S. ochracea 3. Mature leaf beneath at most sparsely puberulous on the nerves. 6. Calyx (and often panicle also) glabrous to finely pale puberulous. Rimae obscure and glabrous inside. 7. Panicle glabrous, stout. Cymes sessile, flowers in glomerules along the rachis. 3. S. glomerula 7. Panicle puberulous, slender. Cymes stalked 4. S. laxa 6. Calyx at least at the base (and panicle) rusty puberulous. Rimae conspicuous, inside lighter and minutely papillose. 8. Leaves 4-13 '/) cm long. Panicle up to 13 cm, erect, ~_ compact (lax in S. celebica). 9. Cymes not : secund. Basal parts of pedicels all ^ equal. 10. Leaves when dry brown to reddish brown. Basal parts of pedicels 0-2 mm long. Calyx l-%-2|4 mm high, persistently puberulous outside, reddish brown when dry. 11. Leaves 4-10 cm long, acuminate to cuspidate. Panicle ± dense, pedicels with shorter and reduced upper joint (c. \\ mm) 6. S. monophylla 1 1. Leaves 6-1 3 '/2 cm long, cuncatc-acutc to faintly acuminate; venation above more prominent than in the other two species. Panicle slender, lax, joints of pedicel equal (up to 1 V2 mm). 7. S. celebica 10. Leaves when dry palish to olive green, 4'/2-l ' cm long, acuminate. Basal part of pedicels 2-6 mm long. Calyx 2-3 mm high, outside glabrescent in fruit, except for base and margins, crimson when dry 5. S. glauca 9. Cymes :: sccundly branched. Basal part of one pedicel per cyme usually elongated, up to 5 mm. Nerves of leaf beneath often reddish when dry 9. S. rubrincrvis 8. Leaves 7'/2-28'/2 cm long. Panicle usually pendulous, slender, usually much longer than 13 cm. . 12. Leaf widest at the middle, apex gradually acute to caudate, base obtuse to roundcil. Nerves often reddish when dry. F'etiolulcs 2 4 by I -2 mm. Cymes sccundly branched, basal part of one pedicel elongated, up to 5 mm. Claw of petals 2/3-1 mm long 9. S. rubrincrvis 12. Leaf widest at or above the middle, margins ; parallel, apex abruptly acuminate to caudate, base truncate to cmarginatc. Nerves concolourous with intcrvenium when dry. Pctiolules 3-9 by 1-3 mm. Cymes not ; sccundly branched, basal part of pedicels ± equal. Claw of petal W-'/2 mm long 11. S. macrophylla 170 Flora Malesiana [ser. I, vol. 7^ Fig. 8. Distribution of Sarcotheca Bl. 1. Sarcotheca diversifolia (MiQ.) Hall. /. Med. Rijksherb. Leiden 1 (1911) 2; Veldkamp, Blumea 15 (1967) 529, f. 2a-c. — Rourea diversifolia MiQ. Fl. Ind. Bat. Suppl. 1 (1860) 528. — Con- naropsis diversifolia KuRZ, J. As. See. Beng. 39, ii (1870) 69, excl. syn. C. griffithii. — Santalodes diversifolium O. Kuntze, Rev. Gen. PI. 1 (1891) 155. — Connaropsis acuminata Pearson, Kew Bull. (1906) 2. — S. acuminata Hall. /. Beih. Bot. Centralbl. 34, ii (1917) 27. — S. subtripli- nervis Hall. /. I.e. — Connaropsis grandiflora RiDL. Kew Bull. (1930) 75. Shrub or tree, up to 27 m high, to 90 cm 0, often buttressed (90 cm high, 135 cm out, 5 cm 0). Leaves trifoliolate, papyraceous to subcoriaceous, ovate- to elliptic-oblong to -lanceolate, glabrous, acute to caudate, base acute to truncate, venation not very conspicuous; nerves 1-5 pairs, usually ± prominent, veinlets not as finely reticulate as in S. griffithii; lateral leaflets 3-9 1/2 by 1-33/i cm, terminal ones 5V2-I8 by 2-63^ cm; petiole 6-25 by 1-2 '/2 mm, rachis (5-)9-27 by %-2 mm; petiolules 4-7 by y^-l mm. Panicles shorter than subtending leaf, 1-4 together, loosely branched, l-8V2(-13'/4) cm, ferrugineous-puberulous, gla- brescent; branches flattened, 2-30(-40) mm long; bract or petioloid ovate, acute, up to 7 mm. Pedicels: lower joint 2'/i-5mm, upper 1-3 mm; Calyx 3-5 mm high, outside pale puberulous to glabrous, purplish, ± caducous in fruit. Sepals 214-5 by 1 '/4-3 mm, broadly ovate to oblong, acute to emarginate. Petals 7-1 1 by 2-5 mm, obovate-oblong to -lanceolate, rounded to emar- ginate, 1-2 mm clawed, lilac or pink to scarlet or red. Filaments in SF 2%-3i/2 and 3Y4-4y2mm, in LF l'/^-2i4 and 2'/^ -3 mm, the longer oc- casionally without anthers. Pistil pale puberulous to glabrous, in SF 1 14-2 '/4 mm, in LF 31/2-5 mm; styles in SF V^-1 mm, in LF 2'/2-4; ovary 1-2 by %-l 14 mm, ellipsoid. Fruit white to greenish when fresh (red when dry), ellipsoid, glabrous, 16-31 by 9-20 mm; rimae inconspicuous. Seeds IV2 by 5 mm, testa smooth; cotyledons 42/3 by 4 mm, radicle l'/3-l% by ^/j mm. Distr. Malesia: N. Sumatra (off Sibolga: Morsala L), Borneo. Ecol. First and second storey of primary and secondary forest on wet, well-drained, poor soil (e.g. podsolized sand) on undulating to flat land, up to 900 m. Vern. BSlimbing bulat, M, buah piang, Iban, iba jantan, Suluk, kandis(-daham), Tidong, kSrapa-kSrapa, perapan macas, Tutong Dusun, kadazan, tabaus, tSbarus, Brunei. 2. Sarcotheca griffithii (Planch, ex Hook. /.) Hall. /. Med. Rijksherb. Leiden 1 (1911) 2; Knuth, Pfl. R. Heft 95 (1930) 425; Veldkamp, Blumea 15 (1967) 531, f. 2e-f. — Connaropsis griffithii Planch, ex Hook. /. Trans. Linn. Soc. 23 (1860) 166; Edgew. & Hook./. Fl. Br. Ind. 1 (1874) 440; Kurz, J. As. Soc. Beng. 39, ii (1870) 69; King, ibid. 62, ii (1893) 200; Ridl. Fl. Mai. Pen. 1 (1922) 332, fig.; Kochummen, Res. Pamphl. (For. Res. Inst. For. Dep. Mai.) 43 (1963) 6, 38. — Dapania griffithii Knuth, Bot. Jahrb. 50 (1914) 234. Tree, up to 42 m, 1 m (buttresses up to 4 m, 1 m out). Leaves trifoliolate, glabrous, pergamen- taceous, often crisped when dry, venation rather inconspicuous, veinlets above finely reticulate; leaflets elliptic-oblong to -lanceolate, occasionally ± obovate, tapering acuminate to caudate, base obtuse to acute, lateral ones 2-7 by Y^-l'^A cm, nerves 1-5 pairs; terminal leaflet 4-11'/^ by IV2-4 cm, nerves 4-6 pairs; petiole 5-12 by 114-2 mm, rachis 3-16 by 3^-1 mm; petiolules 3-7 by 3/4-1 1/4 mm. Panicles shorter to ± longer than the subtending leaf, stout, densely branched, ferrugineous-puberulous, 2-1 cm; bracts small, ovate or linear, petioloid up to IV2 mm. Flowers pectinately clustered, subtended by minute bracts. Lower joint of pedicel 1-21/2 mm, upper 1/^-1 mm. Calyx IVi-'^ mm high, dark red, outside brown puberulous. Sepalsl-iy^ by 1 y2-2^U mm, broadly ovate to -obovate, acute to emargi- nate. Petals ± glossy, blackish red, whitish at base, 4 1/2-8 14 by 2-2 V^ mm, obovate-lanceolate to -oblong, rounded to obtuse. Filaments in SF 4-41/2 and 5-51/2 mm, in LF 1 1/2 and 2 mm; the longer sometimes without anther and club-shaped, red. Pistil in SF ly^ mm, in LF 3-4 mm; pu- berulous; styles in SF 34 mm, in LF 2-3 mm; ovary 1 by 34-I mm, subglobose. Fruit 18-32 by 10-23 mm, ellipsoid, glabrescent, greenish yellow at maturity, inside yellow, mesocarp glassy; rimae inconspicuous. Seeds 5-7 1/^ by 2-4 mm; testa smooth; radicle 1 V^ by 1/2 mm, cotyledons 5 1/2 by 3 Y2 mm. Distr. Malesia: Sumatra, Malay Peninsula. Ecol. First storey of primary forest on flat to undulating sandy clay on dry to temporarily flooded land. Vern. As(s)am pupy (or pupoii), belimbing (hutan), kaju manau, Lampong, kukui, Oesoe, lain jenis, jintek-jintek, kupoyi, pandija, N. Su- matra, poko pupoe, pupui, pupoi, pupoy. Notes. Leaves of seedlings are 1-foliolate. According to Edgeworth & Hooker /. the leaves are irritable to the touch. 3. Sarcotheca glomerula Veldkamp, Blumea 15 (1967) 532, f. 2i-j. — Connaropsis macrophylla King, J. As. Soc. Beng. 62, ii (1893) 201, non S. macrophylla Bl. 1850; Ridl. Fl. Mai. Pen. 1971] OxALiDACEAE (Vcldkamp) 171 1 (1922) 334. — Dapania macrophylla (King) Knuth, Bot. Jahrb. 50 (1914) 234. — S. macro- phylla [non Bl.) Knuth, Pfl. R. Heft 95 (1930) 424, pro specim. inalay., excl. specim. born, et syn. Miq. Shrub or small tree, up to 9 m, 8 cm o. Leaves unifolioiate, (6-)l 11/2-28 by (2'/4-)4-9 cm, per- gamentaceous to subcoriaceous, glabrous, oblong to lanceolate, margins — parallel, contractedly acute to cuspidate, base broadly truncate to emarginate, not glaucous; nerves 8-13 pairs, branching oft" with wide angle (lower ~ 65-90°), basals usually recurved; petiole 4-11 1/2 by 1 '/2-3 mm, petiolule 3-8 by 2-3 mm, glabrous. Panicles subterminal or axillary, 1-few together, ± erect, glabrous, stout, 4-19 cm; branches reduced, wart- like, up to 2'/2 mm. Flowers in few-flowered fascicles. Lower joint of pedicel — absent, upper 1-3 by l\ mm. Calyx 1 V2-2 mm high, glabrous outside e.xcept the ciliate margin. Sepals 1 '/4-I Vi by 1 '4-I Vi miTi. broadly ovate, rounded to obtuse. Petals 6-8 1/2 by l-%-2'/3mm, lanceolate-oblong to obovate-lanceolate, rounded, red to crimson, tube lighter. Filaments in LF 1 and P/4 mm, in SF 2V'2-3 and l>V2-^mxn. Pistil sparsely puber- ulous, in SF 2 mm, in LF 4 mm; styles in SF ^4-1 mm, in LF 2'/2-3 mm; ovary 1 by Vi-V^i i"ni, subglobose, pubescence mainly apically. Fruit 8-16 by 5-10 mm, ellipsoid to ovoid, acute to acuminate, recurved, glabrous, red; rimae in- conspicuous, in the upper half, neither lighter nor papillose inside. Seeds 10 by 4 mm; testa trans- versely rugose; cotyledons 7|4 by 3 mm; radicle c. 1 mm long, stout. Distr. Malesia: Malay Peninsula. Ecol. Primary and secondary forest on flat to undulating land. Vern. (Asam) (se-)tundok, belimbing bhi, b. hutan, sSndok, tStindok. 4. Sarcotheca laxa (Ridl.) Knuth, Pfl. R. Heft 95 (1930) 422; Veldkcamp, Blumea 15 (1967) 533. — Connaropsis laxa RiDL. J. Str. Br. R. As. Soc. n. 75 (1917) 9; Fl. Mai. Pen. 1 (1922) 333. — Con- naropsis glabra Ridl. J. Str. Br. R. As. Soc. n. 75 (1917) 9; Fl. Mai. Pen. 1 (1922) 332. — S. glabra KNUTfj, Pfl. R. Heft 95 (1930) 422. — Connaropsis sericea Ridl. J. Fed. Mai. St. Mus. 10 (1920) 121; Fl. Mai. Pen. 1 (1922) 334. — S. sericea Knuth, Pfl. R. Heft 95 (1930) 424. — Connaropsis sim- plicifolia Ridl. Fl. Mai. Pen. 1 (1922) 334. — 5. simplicifolia Knuth, Pfl. R. Heft 95 (1930) 424. Shrub or tree, up to 23 m high, 45 cm 0. Branches glabrous or densely fcrrugincous- tomcnlosc, glabrcscent. Leaves unifolioiate, (5-) 8-23 by (2-)3-8'/2 t^m. oblong to lanceolate, elliptic to obovate, acuminate to caudate, base broadly cuncatc to truncate, above sparsely silky pubescent, mainly at base and on the midrib, beneath sometimes subglaucous, glabrous or pale to fcrrugincous pubescent; nerves 5-1 1 pairs, basals sometimes at 90 ; petiole 5-25 by I -3 1/2 mm, peti- olule 3-7 by 1 -3 mm, glabrous or pubescent, /'rt/z/r/w 1-3 together, up to 30 cm, erect, pendent in fruit, glabrous, or fmcly pale pubcrulous, or ferrugineous velvety; branches patent to recurved in fruit, up to 14 mm, sometimes once forked near the top, subtended by a bract, a petioloid, or a small leaf. Lower and upper joints of pedicel subequal, 1/2-1 mm. Calyx 1 '4-2 mm high, outside gla- brous or very shortly pale puberulous or ferru- gineous puberulous. Sepals \V^-1 by 1-2 mm, broadly ovate to spathulate, obtuse to emarginate. Petals 4-7 by 1 V*-~ mm, obovate-lanceolate, rounded to emarginate, whitish to dark red. Filaments in SF l-2'/2 and 2-3 '/2 mm, in LF •/2-1 and 1-1% mm. P/^r// glabrous to appressed- strigose, in SF 1-1 '/i mm, in LF 3^ mm long; styles in SF Vi mm, in LF 2-3 mm long; ovary %-l by 1/2-2/3 mm, ellipsoid. Fruit 6-13 by 4-9 mm, ovoid to ellipsoid, recurved, rounded to acuminate, glabrescent; rimae inconspicuous, closed not lighter nor glandular inside. Seeds 5-9 by 3-4 mm; testa smooth to transversely rugose; cotyledons 4-5 by l-lVi inni, 3-4 times as long as the radicle measuring 1-1 '/4 by V3 mm. Distr. Malesia: Malay Peninsula, ? NE. Sumatra. Ecol. Forest edges and swamps at low altitudes. Note. A collection from Sumatra (Tenajan R., Upper Riouw, Soepadmo 154) of which the leaves and also the fruit somewhat resemble the Bornean S. rubrinervis, but of which the panicle is like the Malayan 5'. laxa var. sericea, might be a new variety of S. laxa, but better material is needed to reach a definite conclusion whether this species occurs also in Sumatra. a. var. laxa. Veldkamp, Blumea 15 (1967) 535, f. 3a-b. — S. laxa (Ridl.) Knuth. — S. glabra (Ridl.) Knuth. Nearly glabrous throughout. Nerves 5-9 pairs. Branches of panicle distinct, slender, 2-14 mm, near top once forked. Petals retuse to trun- cate, 41/2-4^/4 by 114-1 V2 iTim, claw 0-'/2 mm, stout. Fruit 6-7 mm long. Seed 5 by 3 mm. Distr. Malesia: NW. Malay Peninsula. Vern. MisHkam. b. var. sericea (Ridl.) Veldkamp, Blumea 15 (1967) 535, f. 3c-d. — S. sericea (Ridl.) Knuth. — S. simplicifolia (Ridl.) Knuth. Leaves glabrous above, sparsely puberulous to glabrous beneath; nerves 8-1 1 pairs. Panicles densely, shortly puberulous; branches reduced, not forked, 1-6 mm. Petals obtuse to truncate, 5-7 by 1 2/3-2 mm, claw slender, 2/3-! mm. Pistil mainly apically, sparsely strigose. Fruit 8-10 mm long. Seed 7'/2-8 by 3-4 mm. Distr. Malesia: East coast of the Malay Peninsula. Vern. BSlimbing chcrclick, b. liulan, gtriji, m^dang, si^tundok. Uses. Roots for poulticing wounds externally (BURKILL & Haniff SFN 17611). c. var. hirsuta Veldkamp, Blumea 15 (1967) 535, r. 3c-g. Twigs fcrrugineous-tomcntose, patchily gla- brescent. Leaves beneath pale- to rusty-pubescent. 172 Flora Malesiana [ser. I, vol. V above sparsely and silky so, mainly at base and along the midrib; nerves 5-8 pairs. Panicle rusty- pubescent, branches reduced, stout, 1-3 mm. Petals obtuse, sometimes notched; claw I/2-I nim. Pistil densely appressed-strigose. Fruit 9-13 mm long. Seed 9 by 4 mm. Distr. Malesia: Malay Peninsula (Johore). V e r n . Be limbing besi, b. burong, b. hutan, resak rambai daiin. 5. Sarcotheca glauca (Hook. /.) Hall. /. Med. Rijksherb. Leiden 1 (1911) 2; Knuth, Pfl. R. Heft 95 (1930) 421, f. 28f-n.; Anderson, Card. Bull. Sing. 20 (1963) 162 (ecol.); Veldkamp, Blumea 15 (1967) 535, f. 4a-c. — Connaropsis glauca Hook./. Trans. Linn. Soc. 23 (1860) 166. — Fig, 7a-h. Shrub or tree, up to 2 1 m, 30 cm 0; no buttresses. Leaves unifoliolate, AYz-W by ly^-Ay^ cm (on sterile twigs larger, up to 13'/2 by 6V2 cm), elliptic to oblong, acute to acuminate, base obtusely cu- neate to emarginate, glabrous; beneath whitish green, dull, subglaucous; nerves 5-7 pairs; petiole 10-27 by V2-I n^rn, petiolule 3-6 by ^4-1 '/2 mm. Panicles 1-2 together, erect, l'/2-13cm, rusty-puberulous; branches up to 10 mm, usually many-flowered, rather closely placed. Pedicels pectinately clustered, lower joint 2-6 mm, upper !4-'/2 mm. Calyx rusty-puberulous outside, g!a- brescent in fruit except for margins and base, 2-3 mm high, crimson when dry. Sepals P/4-3 by Ya-I Ya mm, ovate to oblong. Petals red, darker at apex, 33^-6 by 1-2 mm, oblong to lanceolate, sometimes obovate. Filaments in SF 2-2^4 and 21/2-32/3 mm, in LF l'/4-l!/2 and lV2-iy4 mm. Pistil in SF 1-1 '/a mm, in LF 3V^-3i/2 mm, ap- pressed rusty-strigose; styles in SF V^-3/4 mm, in LF 2'/2-3 mm; ovary V2-I by '72-% mm, ellipsoid. Fruit bright pink to dark red, 8-12 by 5-11 mm, subglobose to ellipsoid with obtuse to rounded apex, glabrescent; rimae open and conspicuous, inside lighter and papillose. Seeds 6 by 3 1/2 mm; testa shiny, smooth; cotyledons 5 by 2/2 mm, elliptic, radicle 1 mm. Distr. Malesia: NW. Borneo (Sarawak, Brunei, Sabah). Ecol. Rare to very rare in undisturbed forest, heath forest on well-drained humus podsols, ground-water podsols, peaty 'Hochmoors' and sand covered clay on terraces and sand stone. Oc- casionally in secondary forest (Brijnig). Vern. Aremajuh, Dajak, asam daham, barus, belimbing, b. daham, gitan gizu, kandis daham, Brunei, medang, piang, Iban, rangkas-rangkas, Dusun Kinabatangan, segot, Baju, tampusi, temposi(s), Kedayan. The vernacular epithet daham might point at the use of this plant against coughing. Notes. This species is closely allied to S. ru- brinervis Hall. /. and differs in having a less puberulous, when dry dark red to crimson calyx, a short compact panicle with very reduced primary branches, while the way of branching in the cyme is indistinct. The pistil in LF is 3'/^-3i/2 mm long, the filaments in SF 2-2 3^ and 21/2-32/3 mm. The cotyledons are lanceolate. 6. Sarcotheca monophylla (Planch, ex Hook. /.) Hall. /. Med. Rijksherb. Leiden 1 (1911) 2; Knuth, Pfl. R. Heft 95 (1930) 422, f. 27; Veld- kamp, Blumea 1 5 (1967) 536, f. 2g-h. — Connarop- sis monophvlla Planch, ^a- Hook./. Trans. Linn. Soc. 23 (1860) 166; Edgew. & Hook./. F1. Br. Ind. 1 (1874) 440; King, J. As. Soc. Beng. 62, ii (1893) 200; Ridl. F1. Mai. Pen. 1 (1922) 332, f. 34. — Dapania monophylla Knuth, Bot. Jahrb. 50 (1914) 648. Shrub or tree, up to 30 m, 38 cm 0, bole oc- casionally buttressed and crooked. Leaves uni- foliolate, 4-10 by 1/2-31/4 (-7) cm, elliptic to oblong, rarely suborbicular, acuminate to cuspi- date, base rounded to cuneate, glabrous, dark when dry, subglaucous beneath; nerves 5-7 pairs; petiole 6-31 by 1/2-^/4 mm, petiolule 3 by 2/1-I mm. Panicles 1-2 together, erect, 1-9 1/2 cm, rusty-puberulous, i dense; branches up to 4 mm long. Pedicels pectinately clustered, upper joint up to 1/3 mm, lower 1-2 mm. Calyx l-V4-2mm high, outside rusty-puberulous, apical ly less so, red- dish-brown when dry. Sepals 1 1/2-2 by 1-1 1/2 mm, ovate to rectangular, acute to emarginate. Petals deep red, 3-5 by %-l V^ mm, lanceolate, emar- ginate. Filaments in LF 1-1 1/4 and 1 i/^-l I/2 mm, in SF 1 1/2 and 2 mm. Pistil densely appressed rusty-strigose, in LF 3-3 1/4, in SF 1 mm; styles in LF 2-21/2 mm, in SF I/2 mm long; ovary Vi-l by 1/2-2/3 mm, ellipsoid. Fruit up to 13 by 13 mm, pale to bright red, subglobose to ovoid, rounded to acute; rimae conspicuous, open, lighter and papillose inside. Seeds 7I/2 by 3 mm; testa smooth; cotyledons A by IV^ mm, radicle 1 1^ by 1/3 mm. Distr. Malesia: W. and Central Malay Pen- insula. Ecol. Secondary forest, open jungle on flat to undulating land on rich soil at low altitude. Vern. (Asam-)pupoy, belimbing akar, b. besih, b. bulat, b. burong, b. hutan, b. keris, b. k(e)ra, b. penjuru, b. pipit (pepit, pipet), bibit (Sakai), kulat pipit, setimdok. Note. The type (Cuming 2324) came from Malaya, not from the Philippines as Knuth supposed. 7. Sarcotheca celebica Veldkamp, Blumea 15 (1967) 537, f. 4f-m. Shrub or tree, up to 35 m, 17 cm 0. Leaves unifoliolate, (3 i/2-)6-l 3 1/2 by 2-5 cm, elliptic to lanceolate, acute-cuneate, base cuneate to ± truncate, above with prominent, rather coarse venation, glabrous, subglaucous, sparsely pu- berulous; nerves 5-6 pairs; petiole 6-21 by V^-l mm, petiolule 2-5 by 1/2-I !4 mm, sparsely strigose. Panicles 1-2 together, 1-7 cm, erect, slender, rusty-puberulous; branches up to 2(-7) mm, distant. Pedicels few, close together with equal upper and lower joints, up to 1 Vi mm. Calyx 2-2 14 mm high, rusty-puberulous outside, red- dish-brown when dry. Sepals 1 1/2-2 by 1-1 1/3 mm, ovate to rectangular, acute to truncate. Petals red, 4-414 by 1-1 V4 mm, lanceolate, rounded to 1971] OxALiDACEAE (Vcldkamp) 173 retuse. Filaments in SF 1-2 and 2-2 '/4 mm, in LF H and 1 mm. Pistil appressed-strigose, in SF 1 mm, in LFZ': mm; styles in SF '4 mm, in LF 2 mm; ovary '2—^4 by V2— ^4 mm, subglo- bose to ellipsoid. Fruit 8-13 by 6-9 mm, ellipsoid rounded to acutish. glabrescent, red; rimae open from the top to below the middle, ± conspicuous, lighter and papillose inside. Seeds 4-5 by 3 mm; testa slightly transversely rugose to — smooth; cotyledons 3 by 2-^4 mm, radicle 1 mm. Distr. Malesia: Central Celebes (Malili) and Kabaena 1. (S of SE. Celebes). Ecol. Rather open country, primary forest on stony to clayish. flat to undulating land at low altitude. Vern. Ko(e)ngilti {{mo-)piite). To Beia, To Padoe, To Tambee. Note. This new species is best distinguished by its lax, narrow panicles with thin rachis and the coarser and more prominent venation on the upper side of the leaf as compared with its nearest allies S. glauca and S. monophyUa. 8. Sarcotheca ferruginea Merr. Pap. Mich. Ac. Sc. 19 (1933) 160, fig.; Veldkamp, Blumea 15 (1967) 538, f. 3h-i. Tree. Twigs velvety rusty-pubescent, later patchily glabrescent and dark. Leaves shiny golden pubescent when young, upper surface soon gla- brous or with a few appressed, pale hairs at the basis and along the midrib, beneath densely ferrugineous-pubescent, not glaucous, (3'/2-)5- l'/4 by (l-)2-3i;. cm, oblong to oblanceolate, acute to caudate, base broadly cuneate to rounded, margins usually paler; nerves 5-7 pairs; petiole 4-6'/2 by '/2-I mm, petiolule 2-4 by Va-IVz mm, rusty-velvety, glabrescent. Panicles 1-2 together, short, slender, up to 7'/2 cm, velvety; branches distant, '/2-2 mm. Pedicels with unequal lower joint, one per cyme elongated, 3-4 mm, others f. I mm; upper joint Y2-I mm. Calyx 2-3Vi mm high, rusty-puberulous outside. Sepals P/4-3'/i by '/2-I mm, ovate-lanceolate, acute. Petals red, 4-6 by I '/2-2 mm, obovatc-elliptic to -lanceolate, rounded to obtuse. Filamenls in LF 1 and 1 Yi mm. Pistil in LF 3-3 1/2 mm, appressed-strigose; styles in LF l-lVi mm; ovary I by 1 mm, sub- globose. Fruit 7-14 by 6-12 mm, ovoid to sub- globose, rounded, somewhat shiny, reddish; rimae inconspicuous, not glandular or lighcr inside. Seeds 7 by 4'/2 mm; testa transversely rugose; cotyledons 5 by 3 mm, radicle I mm. Distr. Malesia: N. Sumatra (West and West Coast Res.), apparently local and rare. Vcrn. Kaju kundis. Note. I have not seen the short-styled form. 9. Sarcotheca rubrincrvis Hail. /. Bcih. Bot. Ccntralbl. 34, ii (1917) 29; Knufh, F'fl. R. Heft 95 (1930) 425; Veldkami-. Blumea 15 (1967) 539, f 4d-c. — S. ohlonuifolia Mhkr. Univ. Cal. Publ. Bot. 15 (1929) III; Knliii. Pfl. R. Heft 95 (1930)421. Shrub or tree, up to 19 m, bole up to 10 m, 30 cm 0. Leaves unifoliolatc, 5 IH'/j by I '/4-7 cm, oblong to lanceolate, glabrous, acute to caudate, base obtuse to rounded, beneath not or slightly glaucous; nerves 4-11 pairs, often with reddish tinge; petiole 6-20 by 1-2 mm, petiolule 1-A by 1-2 mm. Panicles 1-2 together, elongated, lax, pendulous, rusty-puberulous, 7-381/2 cm; branches elongated, slender, patent, + monochasially branched, up to 8 (-21) mm. Pedicels with un- equal lower joint, usually one per cyme elongated to 5 mm, others up to 3 mm; upper joint '/4-I mm. Calyx 2-3 mm high, rusty-puberulous, at least at base. Sepals 12/3-2^/4 by 1-1'/^ mm, ovate to lanceolate, acute to emarginate. Petals 3%-5V2 by 1-1 1/2 mm, lanceolate, rounded to emarginate, pale red to red, occasionally white, apex usually darker, claw -/3-I mm. Filaments in SF 1 2/3-2 and 2'/2-3 mm, in LF Vi-^/3 and 1-1 V4 mm long. Pistil in SF 1-1 1/4 mm, in LF 21/4-23/5 mm; styles in SF 1,-2 mm, in LF 1 ^/^-P/j mm; appressed- puberulous; ovary Vi-^U by 1/2-2/3 mm, sub- globose, densely appressed-strigose. Fruit 6-10 by 5-10 mm, subglobose, rounded, glabrescent, glau- cous to glossy, pink to bright red; rimae not al- ways conspicuous, lighter and papillose inside. Seeds up to 6-^4 by 5 mm; testa smooth to ± transversely rugose; cotyledons elliptic, 4% by 1 1/2 mm, radicle 1 by ^/j mm. Distr. Malesia: E. Borneo (from Tawao to Balikpapan). Ecol. Primary and secondary forest near rivers on loam, flat to undulating land. Vern. Aslm-asc^m, Dusun Kinabatangan, iba talon, Bajau, ira prumpuan, Suluk, kajo badjuk, Kajam-Dajak, kandis daliam, Brunei, lampyos, Dusun Penompang, pinggoh, pinguh, Klabakan, pingo, Tidon, tcnggoh, Kutai. Note. Sometimes difficult to distinguish from 5. glauca (see there). 10. Sarcotheca ochracea Hall. /. Beih. Bot. Centralbl. 34, ii (1917) 28; Knuth, Pfl. R. Heft 95 (1930) 424; Veldkamp, Blumea 15 (1967) 541, f. 5c-e. Treelet, up to 9 m, 10 cm 0. Twigs velvety, glabrescent. Leaves unifoliolatc, 71/2-23 1/2 by 31/2-1014 cm, elliptic to oblong, abruptly acu- minate to cuspidate, base obtuse to truncate, above glabrous, beneath rusty-pubescent to -velvety, sometimes subglaucous; nerves 6-1 1 pairs; petiole 5-24 by 1 1/2-3 mm, petiolule 3-7 by 11/2-3 mm. Panicles 1-2 together, 7-70 cm, erect to pendulous, rusty-velvety, compact to elongated. Branches very short to elongated, often distant, pectinately branched, many-flowered, patent to recurved, up to 30 mm long, rusty-velvety. Lower joint of pedicel 4 8 mm, upper '/j-l mm. Calyx (2i/4-)3-3i/2 mm high, rusty-puberulous outside. Sepals suborbicular to obovate, rounded to retu.se, 1Va-7) by \y\,-'S mm. Petcls crimson, apically darker, (4i/4-)6 « by (I )|i/2-IV4 mm, obovate- lanceolate to lanceolate, obtuse to rouiulcil. Fila- menls in SF 2 1/2 and 3 3l^ mm, in LF 1 1/2-2 and 2-2 1/2 mm. Pistil densely rusty-strigose, in SF iy4-2, in LF 3-4 mm; styles red, in SF 1/3-^/4 mm, in LF 2-2'/2 mm long; ovary l-l '/z by I mm. 174 Flora Malesiana [ser. I, vol. 7* subglobose. Fruit 8-15 by 6-15 mm, subglobose to oblong, rounded, glabrescent, bright red; rimae conspicuous, lighter and/or papillose inside. Seed 1-2 per fruit, 5|4-7 by 3|4-4'/2 mm; testa trans- versely rugose. Mature embryo not seen. Distr. Malesia: Borneo (Sarawak: Bintulu). Ecol. Along streams in forest on clay. Vern. Ikor mata, Iban, pechi mata. 11. Sarcotheca macrophylla Blume, Mus. Hot. Lugd. Bat. 1 (1850) 242; MiQ. Fl. Arch. Ind. III. (1870) 70, t. 30; Baillon, Adansonia 10 (1873) 364; Hist. PI. 5 (1874) 26, 47; Hall./. Beih. Bot. Centralbl. 34, ii (1917) 29; Knuth, Pfl. R. Heft 95 (1930) 424, /7ro spec. Born.; Veldkamp, Blumea 15 (1967) 541, f. 5a-b. — Roucheria macrophylla MiQ. Fl. Ind. Bat. 1, 2 (1858) 136; Suppl. 1 (1860) 162. Shrub or tree, up to 15 m, lOcmo. Leaves unifoliolate, (5-) 16-28 by 6-10 cm, oblong to oblanceolate, margins ± parallel, abruptly acu- minate to cuspidate, base truncate to emarginate, subcoriaceous, above glabrous, beneath pu- berulous on the veins, sometimes subglaucous; nerves 6-13 pairs; petiole 5-12(-25) by 1 Vi-3 mm, petiolule 3-9 by 1-3 mm. Panicles 1-4 together, slender, lax, 12-85 cm, pendulous, brown-pu- berulous; branches elongated, 6-25(-50)mm, often flattened. Lower joint of pedicel up to 2 mm, upper 1/^-1 mm. Calyx 1 1/^-3 mm high, ferru- gineous-puberulous outside. Sepals l'/^-2i/4 by 1-1% mm, obovate to ovate, rounded to obtuse. Petals dark red, 314-5 by 1-1 ^4 mm, obovate- oblong to -lanceolate, obtuse to emarginate, claw 14-!^ mm. Filaments in SF 1-2 and l%-3 mm, in LF^-lV^mm and l'/^-134mm. Pistil appressed-strigose, in LF 2-3 mm, in SF 1 !4-l% mm; styles in SF 14-% mm, in LF 1-2 mm; ovary 2/3-I by V2-% mm, subglobose to ellipsoid. Fruit 6-1 1 by 5-8 mm, subglobose, rounded, gla- brescent, shiny, dark red; rimae inside lighter, papillose, conspicuous. Seeds up to 9 by 4 mm; testa transversely rugose; cotyledons 4 by 2I/2 mm, radicle 1 mm. Distr. Malesia: Borneo (Indonesian Borneo; Sarawak: Marop). Ecol. Primary and secondary forest on sand. Vern. Belimbing manik, Bakumpai-Dajak, kaju kim, krumbai merah, mim, pengu, Dusun, ram(jb)- ajan. Notes. Blume cited this species also to occur in Sumatra, but this seems to be erroneous. It has been confused with Connaropsis macro- phylla King = S. glomerula. Dubious Sarcotheca philippica (F.-Vill.) Hall. /. Med. Rijksherb. Leiden 1 (1910) 2. — Connaropsis philippica F.-Vill. Novis. App. (1880) 33; Merr. Sp. Blanc. (1918) 19, 195; Knuth, Pfl. R. Heft 95 (1930)417. Subscandent. Innovations dark purple. Leaves imparipinnatc, 1-4-jugate; leaflets 6-10 by 2-3 cm, oblique-ovate, acute, subcoriaceous, pubes- cent when young, later glabrous. Panicle axillary and terminal, shorter than the subtending leaf. Sepals 2-4 mm long. Petals twice as long, connate at base when young, later free. Filaments alter- natingly shorter ("alterna hreviora'), often without anthers; styles 5, erect, adpressed; stigmas oblong, apiculate; ovary ovate, pilose, 5-locular, ovules 2 per cell. Fruit 2-3 by V2-I cm, orange-red, pilose, 5-angular, 5-locular. Seed 1 per cell, ovate, with white aril (translated from type description). Distr. Malesia: Philippines (Bugney near Igbards, Iloilo). Vern. Balabangquilin, malabangquilin. Note. Merrill noted that he has seen no rep- resentative of this genus in the Philippines and none is known to me. Although he is very critical of F.-Villar's work (c/. I.e. 14-18), he thinks it to be '. . . apparently a true . . . Sarcotheca . . .'. Knuth referred it to Averrhoa carambola L., with which the description agrees for the greater part, diff"ering, however, by the subscandent habit, the subcoriaceous leaflets, the 2 ovules per cell, and the rather small, orange-red, pilose fruit. F.-Villar stated he has seen the plant alive, so presumably his remarks pertaining to habit and fruit are correct; the more so as in his work this species follows the description of Averrhoa carambola. As long as no representative is found the status of this species will remain in doubt. Excluded Connaropsis rubescens Ridl. J. Bot. 62 (1924) 295 = Rourea minor (Gaertn.) Leenhouts, Blu- mea 12 (1963) 20 (Connaraceae). Sarcotheca paniculata Ridl. Trans. Linn. Soc. Ill, 2 (1893) 282 = Rourea minor (Gaertn.) Leenhouts, Fl. Mai. I, 5 (1958) 5 1 5a {Connara- ceae). Sarcotheca pinnata Merr. J. Str. Br. R. As. Soc. n. 86 (1922) 314 = Rourea pinnata (Merr.) Veldkamp, Blumea 15 (1967) 543 {Connaraceae). Sarcotheca varians (Craib) Knuth, Pfl. R. Heft 95 (1930) 425. — Connaropsis varians Craib, Kew Bull. (1926) \ 5% = Rourea minor (Gaertn.) Leenhouts, Fl. Mai. I, 5 (1958) 515b {Connaraceae). 5. AVERRHOA LiNNE, Gen. PL ed. 5 (1754) 196; Sp. PI. (1753) 428; Endl. Gen. PI. (1839) 1173, ind. sect. Bilimbi et sect. Carambola Endl.; Knuth, Pfl. R. Heft 95 (1930) 417; BACK.& Bakh./. Fl. Java 1 (1963) 247. — Averrhoaceae Hutchinson, Fam. Fl. PI. ed. 2, 1 (1959) 356. — Fig. 9. 1971] OxALiDACEAE (Veldkamp) 175 Evergreen shrubs or trees. Innovations pubescent, glabrescent. Leaves spirally arranged to terminally clustered, estipulate, imparipinnate, herbaceous; leaflets entire, subopposite, subsessile, terminal largest. Panicles axillary or cauliflorous. Flowers in scattered, stalked, loose cymes, heterodi- or -tristylous. Bracts small, caducous. Sepals quincuncial or paratact, shortly connate at base. Petals contort or paratact, free or coherent above the claw, creamy to dark red with white markings. Ovary appressed-strigose; ovules (2-)3-7 per cell, pendulous. Fruit large, yellowish green and semitransparent in vivo, red when dry; rimae small, reduced to absent (?). Seeds elUptic, flattened; aril present in one species, attached to the entire adaxial raphe, bilabiate, enveloping the seed, fleshy, with oily drops; endosperm present; cotyledons 6-8 times as long as the oblique and stout radicle. Distr. 2 spp., probably native in Malesia, cultivated pantropically at low altitude for their fruit trees, often escaping. It is often assumed that Averrhoa is possibly of American origin. Both species are mentioned in pre- Linnean literature always from Indo-Malesia (Bauhin, 1623, being the oldest). The closest relatives are Malesian-Malagasian. Though I can not forward proof, for or against, an Indo-Malesian origin seems most probable. Ecol. Irritability of the leaves has been noted as early as 1785 by Bruce (Phil. Trans. 356), while Darwin made a detailed study of it {A. bilimbi L.) in his 'The Movements of Plants' (1880) 330, 447. Uses. Mainly for the fruit; see under the species. Syst. The two species are remarkably different in important characters which are almost of the same value as those separating the allied genera Sarcotheca and Dapania as appears from the key to the species. One is tempted to raise them to the status of sections, as Endlicher proposed (Gen. PI. 1839, 1173). At the same time this shows the intimate relationship of the three genera. This defeats Hutchinson's rigorous attempt to assign Averrhoa to a separate family Averrhoaceae, even accommodating it to another order (Fam. Fl. PI. ed. 2, 1, 1959, 356). The reason for this rests on the supposed presence of a disk and horizontal ovules; the first I can not find and the ovules are pendulous. KEY TO THE SPECIES 1. Leaves 3-6-jugate. Panicles axillary, rarely some ramiflorous. Petals up to 8 mm, minutely puberulous inside, coherent above the claw. Shorter stamens without anther. Ovules 3-5 per cell. Fruit stellate in 0. Seed arillate 1. A. carambola 1. Leaves 7-19-jugate. Panicles cauliflorous, rarely some axillary. Petals 10-20 mm, glabrous inside, not coherent above the claw. All stamens antheriferous. Ovules 4-7 per cell. Fruit terete, slightly lobed. Seed exarillate 2. A. bilimbi 1. Averrhoa carambola LiNN^, Sp. PI. (1753) 428; stellatum Rumph. Herb. Amb. 1 (1750) 115, f. Cav. Diss. 7 (1789) 374, f. 220; DC. Prod. 1 35. — Fig. 9. (1824) 689; Bl. Bijdr. (1825) 242; W. & A. Tree, up to 14 m by 30 cm. Branches often Prod. (1834) 141; Blanco, Fl. Filip. (1837) 391; drooping. Innovations yellowish to reddish pu- MlQ. Fl. Ind. Bat. 1, 1 (1859) 133; Edgew. & bcsccnt, glabrescent. Leaves 3-6-jugate, not Hook. /. Fl. Br. Ind. I (1874) 439; Progel, crowded terminally; rachis up to 20 cm; leaflets Fl. Bras. 12, 2 (1877) 520, incl. var. anguslisepala; up to 10 by 4 cm, variable in shape, lowest not Trimen, J. Linn. Soc. Bot. 24 (1887) 129 (typi- rcflcxed, acute to acuminate, glaucous beneath; fication); Ningrat, Teysmannia 3 (1892) 754; nerves 4-10 pairs. Panicles axillary, rarely rami- K. & V. Bijdr. 9 (1903) 106; Back., Fl. Bat. florous, 1-fcw together, ascending, up to 7 cm (1907) 229; Koord. Atlas 4 (1918) 603; Merr. long, flowers hcterodistylous (LF, MF), said Sp. Blanc. (1918) 194; Ridl. Fl. Mai. Pen. I (1922) to be scentless, melliferous. Pedicels 3-6 mm, 332; Heyne, Nutt. PI. (1927) 853; Knuih, Pfl. articulated 1-2 mm below the flower. Sepals up R. Heft 95 (1930) 417; Och.se & Bakh. Vrucht. to 4 by 2 mm, bright red, yellowish brown with (1931) 91, tab. col.; Burk. Diet. (1935) 271; pale margins when dry, obovatc-oblong to trian- Qtis. Med. PI. Philip. (1951) 439; Ba{ K. & gular, acute to obliquely emarginate, subglabrous. Bakm./. Fl.Javal (1963) 247; Vkldk., Fl. Thail. 2 Petals up to 8 by 2 mm, obovate to lanceolate, (1970) 21. — A. acutangula SroKE.s, Bot. Mat. coherent, claw c. I '/2 mm, glabrous, but inside Med. 2 (1812) 543. — A. peniandra Bi.ANf o, with minute septate-glandular hairs. Shorter sta- M. Filip. (1837) 392. — Tamara tonga Rheede, mens ananthifcrous, lilaments subulate with often Hort. Mai. 3 (1682) 51, f. 43, 44. - Prunum swollen base, up to \\'^ mm, longer ones in LF 176 Flora Malesiana [ser. I, vol. 71 Fig. 9. Averrhoa carambola L. (Courtesy T. B. Worthington). up to 3 mm, in MF up to 4'/^ mm. Ovary elliptic, up to 2 1/2 mm long, appressed-puberulous, mainly on the ribs; styles in LF up to 2 mm, in MF V2 rnm; ovules 3-5 per cell. Fruit up to ]2'/2 by 6 cm, ovoid to ellipsoid, stellate in 0, lobed at both ends, apically with minute rimae on the ribs. Seeds up to 10, 12 by 5 mm; aril fleshy, bilabiate, enclosing the seed, lobed at base; cotyledons up to 6'/i by 31/2 mm, elliptic. Distr. Native country unknown, possibly Cen- tral and East Java, but escaping freely, often a relict of former cultivation. Ecol. At low altitude, up to 1000 m, along rivers, in ravines, primary and secondary forest. Dispersal. Fruits eaten by bats, mice and Calotes spp. (Agamidae). Cf. Ningrat (1892). Vern. Variations on belimbing. Sumatra: asom djorbing. Bat., balimbing manis, b. segi, b. sajur, b. batu, kembola, b. besi, b. keris, Mai., kembang bua, kaping, Palemb., b. manih; Java: balingbing amis, tjalingtjing amis, S, b. legi, b. linger, b. lingir, b. manes. Mad., libi melai, Sawu; Borneo: b. pesegi, b. penjura, Kuching, b. amamas, Sabah; Celebes: lumpias mamis, Bentenan, rumpiasa, Bantik, lompiat morominit, Mongodow, dumpias tariis, lopias erne, Tonsa- wang, lumpias tombal, Tontembuan, Tonsawang, lembetue lombiata, Goronta, lombituko gula, Buol, takule, Baree, bainang sulapa, Makass., kulirang taning, pulirang taning, N. Salajar, baknil pasaki; Sepa, S. Ceram, ifel emroro, Masareta, Buru, malibi totofuo, Weda, S. Halm., balibi totofuko, Galelas, Tobelor, totofuko, t. lemo, Ternate, tufuo, Tidore; Philip.: daligdn. Ilk., dalihan, galurdn, Ibn., galangan, P. Bis., gardhan. Bis., malimbin, S. L. Bis., sirinate. Ting.; E. New Guinea: ibeid, Tehid, painayangara, Tumbunke. Uses. The fruit is eaten fresh or pickled. Leaves and fruits are said to be good against fevers, aphtha, angina, chickenpox, ringworm, headaches, 1971 OxALiDACEAE (Vcldkamp) 177 scurvy, dysentery, piles and affections of skin and eyes; they excite the appetite, but may cause vomiting. It is antiphlogistic and a sialogogue. The fruit removes stains f^rom laundry, hands and weapons. See Ningrat (1893), Koorders & Va- LETON (1903). Heyne (1927), Ochse (1931), BuRKiLL (1935), and Qlisumbing (1951). Phytochem. De Peralta (Philip. Agr. 17, 1928, 334) has shown the presence of HCN in leaves, roots and stems; Bate-Smith (J. Linn. Soc. Lond. Bot. 58, 1962, 134) the presence of cyanidin, p-cumaric acid and traces of sinapic acid, and (ibid. 55, 1957, 687) tannin, leuco- anthocyanin and leucocyanidin. Syst. This species is widely cultivated for a long time and many cultivars are e.xtant (c/. Ningrat, 1893), differing in taste and size of the fruit. A /. acida and/, dulcis have been described by Koor- ders & Valeton, and material bearing these names has been e.xamined, but no characters could be found to distinguish them. Progel's var. anguslisepala from S. America seems not to war- rant distinction, as the shape and pubescence of the sepals in Malesia is very variable. This variety has possibly been the reason for an assumed Amer- ican origin of the genus time and again found in literature. 2. Averrhoa bilimbi Linne, Sp. PI. (1753) 428; Cav. Diss. 7 (1789) 373, f. 219; DC. Prod. 1 (1824) 689; Roxb. FI. Ind. ed. Carey 2 (1832) 451; W. & A. Prod. (1834) 142; Blanco, F1. Filip. (1837) 391; Endl. Gen. Pi. (1839) 1173; MiQ. FI. Ind. Bat. 1, 1 (1859) 139; Edgew. & Hook. /. FI. Br. Ind. 1 (1874) 439; Progel, FI. Bras. 12, 2 (1877) 520; Trimen, J. Linn. Soc. Bot. 24 (1887) 129 (typification); K. & V. Bijdr. 9 (1903) 111; Koord. Atlas 4 (1918) 601, 602; Merr. Sp. Blanc. (1918) 195; En. Born. (1921) 311; RiDL. FI. Mai. Pen. 1 (1922) 331; Heyne, Nutt. PI. (1927) 852; Knuth, Pfl. R. Heft 95 (1930) 418, ind. f. papuana Knuth; Ochse & Bakh. Ind. Groenten (1931) 546; Benthall, Trees of Calcutta (1933) 70; Burk. Diet. (1935) 270; Sorgdrager, Pharm. Tijd. 1 (1941) repr. 4 pp. (anat., uses); Quis. Med. PI. Philip. (1951) 438; Back. & Bakh./. FI. Java 1 (1963) 247; Veldk., FI. Thail. 2 (1970) 22. — A. oblusangula Stoke.s, Bot. Mat. Med. 2 (1812) 542. — Bilimbi Rheede, Hort. Mai. 3 (1682) 55, f. 45, 46. — Blinbitifiun teres, Blimbing bula Rumph. Herb. Amb. 1 (1750) 118. Shrub or tree, up to i5m, 30 cm 0. Branches erect. Innovations and green parts long-persisting pale yellowish to rusty velvety. Leaves 7-19- jugatc, usually terminally tufted; rachis 17-57 cm; leaflets up to 12 by 4 cm, variable in pubescence, size and shape, lower reflexed, acute to acumi- nate, not glaucous beneath; nerves 6-14 pairs. Panicles cauliflorous on tubercles, nearly down to ground-level, fasciculate and pendulous up to 20 cm, rarely axillary, solitary and erect, /lowers hctcrotristylous, usually MF. Pedicels 4-17 mm, articulated near or below the middle. Sepals 3-8 by I '/2-3 mm, yellowish red to purplish, sparsely appressed-puberulous outside mainly at base, inside glabrous or subglabrous, elliptic to lan- ceolate or spathulate, acute to rounded. Petals free, 10-20 by 3^ mm, lanceolate-spathulate, inside glabrous, claw 3-6 mm. Stamens all fertile, in SF 31/2^ and 7 rrun, in MF 2-5 and 9-12 mm, in LF 5 and 7'/2 nim, bases not thickened. Ovary densely appressed pale strigose and with short, septate-glandular hairs, in SF2-2V2 by 1 mm, in MF 3-4 by 1-1 '/2 n^m. in LF 4 by 1 mm, elliptic; styles in SF VS-^/4 Trim, in MF 2-4 mm, in LF 6'/2-9mm; ovules 4-7 per cell. Fruit terete- obtusangular, up to 10 by 5 cm, elliptic to obovate, obtuse, rimae present (?), base tapering. Seeds up to 14, 6-7 by 4-6 mm, exarillate; cotyledons 4-6 by 3 Vz-S mm, orbicular, cordate. Distr. Native country unknown, widely cul- tivated, escaping freely, often a relict of former cultivation, for instance in East Malesia. E c o 1 . Along rivers, lowland primary and secon- dary forest. Vern. Bclimbing and many variations; Sumatra: limeng, sHimeng, thlimeng, Atjeh, asom, Batak; Mai. Pen.: b. asam, b. bcsi, b. botol, b. bulolt, b. keris, b. masam; Borneo: beliwit, Dajak Busang, iba, Bajan, blimbing puchung, Kedayan; Java: tjalingtjing, t. wulSt, S; limbi, Bima; libi, Sawu; belerang, Sangi; Celebes: liimpias, Bentenan, Po- nosakan, Tombulu, Tontamhudn, rump^asa diireng, Bantik, lompiat litod, Mongodow, dumpias, d. tuwama, Tonsawang, wuli(n)dan, Tontembuan, lembetue, Gorontalo, lombituko, Buol, sanggulera, Parigi, tangkiirera, Bar^e, balnang, Makassar, kiilirang, pulirang, N. Salajar, tjalene, Bugin.; niniloe dad lok, Roti, kerbol, Timor, baknil, Kai, ahurela, Atamano, W. Ceram, haiirela, Amahai, S. Ceram, taulela, Nuaulu, takurela, Ambon, tahurela, Sapurua, Ulias, taprera, Kajeli, Buru, ifel milo, Masarete, Buru, malibi, Weda, S. Halm., balibi, N. Halm., Ternate; Philip.: iba. Tag., Sul., P. Bis., ibag, Mbo., kalamids, kala- nuas, kamids, kolonanas, kolonauas. Tag., kalin- giua. Bis., kiling-iba, Bik., ibe. Yak., pids. Ilk., puis, Ig.; New Guinea: miri-miri, Kapur, utSkd, Mimika & Atuka R., ollololt, Sentani. Uses. In the Malay Peninsula the leaves are used as a paste against itches, eaten against syphi- lis and a decoction is drunk after childbirth. In Java the leaves arc used against mumps, rheu- matism and pimples, against piles; a decoction of the flowers is used against cough and thrush. The fruit juice is generally used against fevers, scurvy, beri-beri, biliousness, coughs and piles; it also removes stains from laundry, hands and weapons. See Koorders & Valeton (1903), Heyne (1927), Ochse & Bakhuizen (1931), Burkill (1935), and Quisumbing (1951). Fruits and flowers arc edible but more acid than those of A. carambola. Syst. The species has always been regarded as possessing only an MF flower. Short-styled or SF (IJ.MER 15122) and long-styled or LI" (Hallier /. s.n.) forms occur but very rarely so. It is doubtful whether /. papuana Knuth merits distinction; his detinition reads: "In omnibus partibus fere duplo major". Material showing these sizes is 178 Flora Malesiana [ser. I, vol. 71 ] found on most islands throughout its range in Malesia. Excluded Averrhoa acida Linne, Sp. PI. (1753) 428 = Phyllanthus acida (L.) Skeels, Bull. U.S. Dep. Agr. Bur. PI. Ind. 148 (1909) 17 {Euphorbiaceae). Averrhoa frondosa Salisbury, Prod. (1796) 318 = Phyllanthus acida (L.) Skeels, Bull. U.S. Dep. Agr. Bur. PI. Ind. 148 (1909) 17 (Euphorbiaceae). Averrhoa microphylla Tardieu-Blot, Not. Syst. 1 1 (1943) 133; Fl. G^n. l.-C. 1, Suppl. (1945) 547, f. 66, is according to Leenhouts = Rourea harmandiana Pierre (Connaraceae). Averrhoa minima Perrottet, M6m. Soc. Linn. Paris 3 (1824) 101. "Espece nouvelle venue de la Chine, qui s'^leve a 1 metre (3 pieds) au plus (Mascareigne)". Type-material could not be found in the Paris Herbarium. Probably it does not belong to Oxalidaceae, but to Euphorbiaceae? Averrhoa sinica Hance in Walp. Ann. 2 (1852) 241 = Rourea minor (Gaertn.) Leenhouts, Fl. Mai. L 5 (1958) 514 (Connaraceae). VIOLACEAE (M. Jacobs, Leyden; D. M. Moore, Reading) Shrubs, small trees, or lianas, in Malesia evergreen, or herbs. Stipules present. Leaves in Malesia spirally arranged, sometimes distichous, simple, the margin often shallowly incised; generally stalked. Inflorescences axillary variously mo- dified bundles, or racemes, or panicles, sometimes terminal, or flowers solitary in the leaf axils; bracts small; pedicels often articulated, whether in the lower or in the upper part; bracteoles, if present, small and in the lower part of the pedicel. Flowers bisexual or rarely dioecious, actinomorphic or zygomorphic, particularly in the corolla; the parts often persistent in fruit. Sepals 5, the median one adaxial (posterior), free or occasionally for a small portion connate, often ciliate. Petals 5, free, generally sessile, the median one abaxial (anterior), often longer and diffe- rently shaped, the base then mostly with a sac or spur. Androeciwn often cyhndrical, stamens 5, episepalous; filaments often more or less connate into a tube, in the Malesian genera with zygomorphic flowers, those near the odd petal with a re- curved fleshy appendage; anthers introrse. in Malesia nearly always the connective at the top produced into an approximately triangular membranous appendage converging with the others, cells sometimes with a small appendage at the top. Gynoecium superior, sessile, ovary small, subglobose, one-locular, with generally 3 carpels, the median one adaxial, each carpel with a parietal placenta in the middle bearing 1-many anatropous ovules; style straight or, in the zygomorphic flowers S-shaped with the stigma curved towards the odd petal and club-shaped with variations. Fruit in Malesia capsular, the carpels thickened to boat-shaped leathery or woody valves (in the latter eventually the endocarp separated from the pericarp) which spread and often compress upon dehiscence. Seeds 1-many, sessile, one to a few mm in size, often with distinct raphe, sometimes with funicular outgrowths; rich in endosperm; embryo straight. Distribution. A pantropical family; only Viola is cold-loving. Hyba nth us extends, into the subtropics' so does Melicytus (Pacific Plant Areas n. 103, Blumea Suppl. 5. 1966) in Polynesia and New Zealand. Hymenanthera (congeneric with the former? I.e. n. 104) is temperate in SE. Australia and New Zealand. Number of genera 16, 8 of them American; the largest are Viola, currently credited with c. 400 spp., Rinorea with c. 200, Hyhanl/iits with perhaps 70, and there are about 50 more in the other genera alto- gether. Total number of species c. 720, in Malesia 31, two of these introduced. Ecology. Viola is of temperate origin and occurs in Malesia in the mountains. The other Malesian genera occur in the lowlands; Rinorea and Agalea in everwet forests, Hyhanthus in monsoon regions. Ph ytochemistry. Accurate chemical information about Violaceae is scanty notwithstanding the fact that members of this family are used in popular medicine all over the world. The present summary of chemical characters, therefore, must be considered as a very preliminary one. DifTcrent types of crystals of oxalate of lime occur commonly. Members of the genera .4mphirrlw.\ and Allexis accumulate aluminium according to Chenkrey (Kew Bull. 1948, 173). Leaves and flowers contain rather large amounts of acidic mucilage in many instances; usually the mucilage is present in epidermal cells. In some taxa cells with a yellow or reddish 'resin-like' content replace mucilage cells; these latter cells also occur in the mcsophyll and in the cortex, phloem and pith of stems. The chemical nature of the content of these idioblasts is not known. However, the fact that leucoanthocyanins are rather widespread in Violaeeae suggests that the yellow to reddish cell contents may represent so-called 'myriophyllin" or 'incluses'. Personal observation on rhi/omes of Viola mirahilis L. conlirm this supposition. According to Pec kolt (Ber. Dcut. Pharm. Ges. 7, 1897, 97) fresh leaves of l.eonia ulycycarpa R. & P. are used for the pre- paration of a bird-lime in Brazil; this suggests that at least some members of the family may have rubber- containing idioblasts. Roots of several species of Hyhanthus ( lonidiunt) store inulin-like fructans instead of starch; other J//Vy/«fr«f store essentially sugars or starch (Kraus, Sit/. Ber. Naturl". Cies. Halle, 1879, 6). Preliminary observations about polyphcnolic constituents of leaves (Llhketon-Bouchez, Phyto- (179) 180 Flora Malesiana [ser. I, vol. 7^ chemistry 6, 1967, 1601) showed a wide range of compounds; leucoanthocyanins, flavonol and flavones may be present in various combination ; eliagic acid, however, was observed only in trace amount in three species of Viola. Saponins seem to be rather common in violaceous plants; they have been de- monstrated to be present in many species of Viola and in some species of Hymenanthera, Hybanthus, and Melicytus; none of the saponins has been investigated chemically hitherto. There is much confusion about alkaloids in literature; roots and rhizomes of species of Hybanthus and of Viola odorata L., and other species of Viola were used formerly as a substitute for ipecacuanha root. Some authors claimed to have detected emetin or emetin-like compounds (e.g. viola-emetin; violin) in such crude drugs. Other authors, however, could not find emetin though some of them isolated minor amounts of alkaloids {e.g. anchietin, ionidin; compare Peckolt, I.e.; Linde, Apoth. Z. 34, 1919, 37). Most probably many Violaceae contain small amounts of alkaloids; species of Anchieta, Hybanthus, Hynienanthera, Rinorea, and Viola are listed in literature as alkaloid-bearing plants; however, the struc- tures of violaceous alkaloids are totally unknown at present. Most chemical work has been performed with European species of Viola. Glycosides of delphinidin and cyanidin occur constantly in blue and purple flowers; violanin is a delphinidin-derived anthocyanin acylated by p-cumaric acid. Rutin occurs in flowers and in leaves of several species of Viola. From the herb of Viola tricolor L. Horhammer c.s. (Tetrahedron Letters 1965, 1707) isolated apigenin-6,8-di-C- glucoside, which was called violanthin. The flavone glycoside linarin is the main flavonoid constituent of leaves of Viola papilionacea Pursh (V. Plouvier, C. R. Ac. Sc. Paris 264 D, 1967, 145). Violatoside is an arabinoglucoside of methyl salicylate; it occurs in species of the section Melanium but seems to be lacking in other sections of Viola (Picard, Bull. Soc. Chim. Biol. 8, 1926, 568). From roots and rhizomes of Viola odorata L. Pailer and Novotny (Naturwissenschaften 45, 1958, 419) isolated 0.01-0.02 3o of nitropropionic acid. Concluding this survey it must be stated that a thorough chemotaxonomic discussion of Violaceae is not yet possible. At present no chemical characters are known which contradict the generally accepted relationships between Flacourtiaceae and Violaceae. — R. Hegnauer. Notes. Bentham & Hooker's subdivision of the family (Gen. PI. 1, 1862, 115) was extended and modified by Melchior in E. & P. Nat. Pfl. Earn. ed. 2, 21 (1925) 346; for an account in English of the latter's phylogenetic considerations, see Exell, J. Bot. 63 (1925) 330-333. Dr. D. M. Moore's contribution consists of the genus Viola; he also checked the family description. Indexes to the examined specimens were published in the series 'Identification Lists of Malaysian Specimens', n. 27 Rinorea (1966) and «. 28 Viola (1967). KEY TO THE GENERA 1. Flowers actinomorphic, in axillary bundles or racemes. Seeds not winged. Shrubs or trees. 1. Rinorea 1 . Flowers zygomorphic. 2. Woody climbers. Flowers in elongate inflorescences. Seeds winged 2. Agatea 2. Herbs, sometimes with woody base. Flowers solitary in the leaf axils. Seeds not winged. 3. Leaves (sub)sessile. Pedicels jointed. Seeds ribbed lengthwise 3. Hybanthus 3. Leaves stalked. Pedicels not jointed. Seeds smooth 4. Viola 1. RINOREA AUBL. Hist. PI. Guian. Fr. 1 (1775) 235; Taub. in E. & P. Nat. Pfl. Fam. 3, 6 (1895) 329; Melch. ibid. ed. 2, 21 (1925) 349; Jacobs, Blumea 15 (1967) 127; Taton, F1. Congo, Viol. (1969) 2. — Pentaloba Lour. F1. Cochinch. (1790) 154. — Medusa Lour. I.e. 406. — Alsodeia Thouars, Hist. Veg. Isl. Austr. Afr. (1806) 55; Endl. Gen. PI. (1839) p. 911, n. 5047; B. «fe H. Gen. PI. 1 (1862) 118; Boerl. Handl. 1 (1890) 66. — Vareca Roxb. F1. Ind. ed. Wall. 2 (1824) 445. — Jurgensia Spreng. Syst. 3 (1826) 50. —Prosthesia Blume, Bijdr. (1826) 866. — Dioryktandra Hassk. Retzia (1855) 125. — Scyphellandra Thw. En. PI. Zeyl. (1858) 21. — ''Imhofia Zoll.' ex Taub. (1895) I.e., nomen in synon. Small shrubs to small trees; innovations mostly laxly pubescent, the hairs some- times persistent; twigs pithy, generally angular when young, terete when older. Leaves distichous, rarely in a spiral. Petiole comparatively short, or rarely wanting, lacking distinct joints at base or top. Stipules caducous to long-persistent, some- 1971] ViOLACEAE (Jacobs & Moore) 181 Fig. 1. Rinurea horncri (Korth.) O.K. a. Habit, vVi, h. venation undcrncalli, < '/,, c. stipule. :■ 3, d. flower, -4, e. gynoccium surrounded by stamens, > 6, /. stamen from inside, • \2, k- gynoeeium, /6, h. fruit, subtended by calyx and corolla, - 1, /. seed, x I (a, iI-k Cur us 1898, h C'l kmi n.s 9961. r SAN 42087, h Pleyte 232). 182 Flora Malesiana [ser. I, vol. 7^ times quite conspicuous, often striate lengthwise. Leaf blade more or less dis- tinctly acuminate, domatia occasionally present in the primary vein axils, the margin shallowly incised to subentire, surfaces generally dull. Inflorescences lateral, the flov/ers mostly in bundles, sometimes distichous along a short rachis, sometimes in a corymb or panicle on a short peduncle, rarely in a raceme or on brachy blasts; bracts smaller than the stipules; pedicels articulated. Flowers bisexual, rarely di- oecious by reduction, actinomorphic, calyx quincuncial, covering the bud halfway to completely, corolla generally contorted in the same direction as the third sepal. The sepals (almost) equal, fimbriate. Petals in Malesia (sub)equal, free, sessile, thin in texture, to 9 mm long. Androecium nearly as long as the petals, filamental tube more or less fleshy and sometimes 5-lobed, the filaments inserted on its inner margin, or seldom the filaments free with only a pair of scales outside at their base; anthers converging, glabrous or hairy, the connective produced into an entire dorsal membranous appendage smaller to longer than the thecae, sometimes bearing each a ventral appendage at the top smaller than the dorsal one. Gynoecium as long as the androecium, the 3 (—4) placentas with 1-3 ovules each; style straight, with a terminal more or less distinctly 3-lobed stigma. Fruit capsular, approx- imately globose before dehiscence, %-4 cm , generally subtended by the dried up calyx (which rarely expands), corolla, and androecium, the valves leathery, when large two-layered with reinforced apex, mostly smooth and glabrous, rarely en- veloped in a mass of appendages, or hairy. Seeds sessile, 3-6, rarely 1 or 9, ellipsoid, glabrous, with leathery testa and often clearly defined hilum and raphe. Distr. Pantropical, richest in Africa with c. 107 spp. on the mainland and 25 spp. in Madagascar; in the New World with 50-60 spp.; in Indo-Malesia l2-\ 3 spp., altogether extending from S. India/ Ceylon, Assam, Burma, Hainan, to N. Australia, Melanesia, the Carolines; estimated total about 200 spp. Fig. 3. Ecol. Predominantly in the understorey of primary rainforests, at very low altitude, occasionally up to 1000 m, on various soils including limestone. Vern. Common and reliable names seem to be wanting. Burkill, Diet. (1935) 1912, recorded for the Malayan species some vernacular names, and some unimportant medicinal uses. Taxon. A satisfactory subdivision of the entire genus has not yet been made. The one in the Flora of British India is good for the present area, but is to be reconsidered when the neotropical and African species will have been comprehensively studied. Alsodeia is there subdivided into 3 sections, viz. I. Doryctandra (Hassk., genus, actually Dioryklandra) Hook./. & Th. with stamens "exserted" (not thus found by me), filaments slender (not a character), anthers cohering in a cone (notably by peculiar inter- tangling hairs). Contains R. heteroclita. II. Unnamed, with stamens included, anthers free, disk cupular (our filamental tube). Contains most of our species, and the former genera Pentaloba and Prosthesia. III. Scyphellandra (Thw., genus) Hook. /. & Th., with stamens included, anthers free, disk reduced to 5 scales, one at the base of each anther (and, we may add, dioecious). Contains our R. virgata. King, J. As. Soc. Beng. 58, ii (1889) gave another subdivision of Alsodeia; on page 400, Sect. I, Prosthesia (Bl.) King, with 7 spp.; on page 404, Sect. II, Pentaloba (Wall.) King, with 6 spp.; on page 407, Sect. Ill, unnamed, with 1 sp. Various "species" placed by King in diff"erent sections have in tht present revision been merged. Notes. The taxa in Rinorea east of the Indus have all been worked up in the same way, but the date have been sorted out for publication between the Flora Malesiana and its precursor in Blumea 15 (1967) 127-138. The precursor accounts for all names of all taxa, with their first references and their type specimens, but gives descriptions, distribution and ecology only of non-Malesian taxa. The Flora Ma- lesiana accounts for Malesia as far as names and literature are concerned, later references included, but the descriptions cover the taxa found in Malesia over their full range, and the same holds for distribution and ecology. The key in the Flora Malesiana is also complete for all taxa. Six species belonging to other families have been ascribed to Rinorea; see under Excluded. KEY TO THE SPECIES 1. Stipules caducous. Seeds 6 or fewer. 2. Primary veins curving approximately parallel towards the margin at distances of 1-2 cm; secondary venation scalariform. 1971] ViOLACEAE (Jacobs & Moore) 183 3. Inflorescences fasciculate (rarely stalked in R. angiiifera). 4. Dorsal appendage of the anthers small. Style hairy. Fruit enveloped in moss-like appendages. 1. R. anguifera 4. Dorsal appendage of the anthers distinct. Style glabrous. Fruit smooth. 5. Anthers with small ventral appendages. Ovules 3. Fruit subglobose, with 3 seeds. 2. R. bengalensis 5. Anthers with ventral appendages about as long as the dorsal one. Ovules 6. Fruit conical, sub- tended by the expanded calyx, with 1 seed 3. R. horneri 3. Inflorescences on a peduncle (if racemose, see R. longinicemosa). 6. Gynoecium glabrous. Petals to 4 mm long. Anthers with small dorsal appendage. Fruit triangular with rounded corners, 1-3 cm long, valves thick 4. R. sclerocarpa 6. Gynoecium hairy. Petals to 9 mm long. 7. Pedicels jointed near the base. Anthers with small dorsal appendage. Fruit subglobose to ovoid, 8-1 1 mm long, hairy, with a hairy style remnant 5, R. lanceolata 7. Pedicels jointed about the middle. Anthers with a dorsal appendage 1-2 times as long as the cells. (Fruit unknown.) 6. R. macrantha 2. Primary veins few, independently curved towards the margin, secondary venation irregular. 8. Inflorescences more or less elongate. Anthers with distinct ventral appendages. Fruit 1 '4-4 cm o. 9. Stipules l-3'/2 rnm long. Inflorescence axis V2-~Vi cm long. Leaves often obovate. 7. R. javanica 9. Stipules (2-)4-21 mm long. Inflorescence axis to c . 10 cm long. Anthers often bearded at the base. 8. R. longiracemosa 8. Inflorescences fasciculate, or flowers densely set on a short rachis. Anthers with small or no ventral appendages. Fruit Vi-l Vi cm 0. 10. Leaves petiolate, more or less concolorous. Anthers with a dorsal appendage about as long as the cells or longer. 1 1. Leaves l-7(-14) cm long. Petiole 1-5 mm. Plant dioecious; in the o flowers only a style, in the ^ flowers the anther cells vestigial, ovules 6 9. R. virgata 1 1. Leaves 6-36 cm long. Petiole 5-35 mm. Flowers bisexual. Ovules 3. 1 2. Stipules 1 '/2-6 mm, scarcely striate. Leaves dark-coloured in the dried state. Fruit sparsely hairy. 10. R. macrophylia 12. Stipules 4-16 rrun, distinctly striate. Leaves greenish in the dried state. Fruit glabrous. 2. R. bengalensis 10. Leaves subsessile, discolorous, pale underneath, 3i/2-6cm long. Anthers with a small dorsal appendage. Style hairy in the middle only. Ovary glabrous. Andamans, India? R. heteroclita I. Stipules long-persistent. Ovules and seeds 9 II. R. iliaspaiei 1. Rinorea anguifera (Lour.) O.K. Rev. Gen. PI. 1 Merr. Philip. J. Sc. 11 (1916) Bot. 100. — R. (1891) 42; Craib, FI. Siam. En. I (1925) 88; como5« (King) Merr. En. Born. (1921 ) 410. — /?. Merr. Comm. Lour. (1935) 270; Gagn. F1. G6n. echinocarpa Burk. Gard. Bull. S. S. 6 (1930)172. I.-C. Suppl. 1 (1939) 187; Jacobs, Blumea 15 Young twigs terete, generally straight, densely (1967) 127. — Medusa anguifera Lour. FI. covered with a persistent brown tomentum; Cochinch. (1790) 406. — Alsodeia echinocarpa pith narrow. Leaves distichous (spirally arranged Korth. Ned. Kruidk. Arch. 1 (1848) 360; MiQ. on terminal or crooked branches). Stipules (4-) FI. Ind. Bat. I, 2 (1859) 116; Oudem. in Miq. 5-9(-il) mm long, triangular, not appressed, Ann. 3 (1867) 68; Hook, y! & Th. in Hook./. with a distinct keel sometimes protracted into a FI. Br. Ind. 1 (1872) 188; King, J. As. Soc. Beng. slender tip, more or less striate, in Borneo per- 58, ii (1889) 406; BoissiEU, FI. Gdn. I.-C. I sistent, otherwise the scars not conspicuous. (1909) 214. f. 20: 6; Ridl. FI. Mai. Pen. I (1922) Petiole (l-)2-8(-10) mm. Blade thin-coriaceous, 134, f. 12. — Pentaloba corylifolia TuRCZ. Bull. to (8-)l4-28(-37) by (3-)4-8(-IOi/2) cm, widest Soc. Nat. Moscou 27, ii (1854) 341. — Alsodeia at the middle to above, index 2.5-3.7(-4.4); corylifolia (TuRCZ.) TuRfZ. Bull. Soc. Nat. base acute to (mainly in Borneo) narrowly cor- Moscou 36 (1863) i 559. - Alsodeia capillaia date, top acuminate; midrib and veins flat above, King, J. As. Soc. Beng. 58, ii (1889) 407; KiN(i, prominent beneath, sometimes scarcely dilferent Ann. R. Bot. Gard. Calc. 5 (1896) 127, pi. I46A; in colour, sometimes lighter; no domatia; priniary Ridl. FI. Mai. Pen. I (1922) 133. — Alsodeia veins fairly regular and parallel, with distances in ( omasa KiNti, J. As. Soc. Beng. 58, ii (1889) 407; the order of I cm between, no smaller intermediate KiN(i, Ann. R. Bot. Gard. Calc. 5 (1896) 127, veins, secondary venation mostly distinct, scalari- pl. I46B; BoissiFU, FI. G